Whaler Sharks - seafdec.org.my

SHARKS and RAYS of 

BORNEO 

Peter R. Last 4 

William T. White 4 

Janine N. Caira 4 

Dharmadi 4 

Fahmi 4 

Kirsten Jensen 4 

Annie P. K. Lim 4 

B. Mabel Manjaji-Matsumoto 4 

Gavin J. P. Naylor 4 

John J. Pogonoski 4 

John D. Stevens 4 

Gordon K. Yearsley 4

SHARKS and RAYS of 

BORNEO 

Peter R. Last 4 

William T. White 4 

Janine N. Caira 4 

Dharmadi 4 

Fahmi 4 

Kirsten Jensen 4 

Annie P. K. Lim 4 

B. Mabel Manjaji-Matsumoto 4 

Gavin J. P. Naylor 4 

John J. Pogonoski 4 

John D. Stevens 4 

Gordon K. Yearsley 4

© CSIRO 2010 

All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 

and subsequent amendments, no part of this publication may be reproduced, stored in a 

retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, 

recording, duplicating or otherwise, without the prior permission of the copyright 

owner. Contact the author for all permission requests. 

The National Library of Australia Cataloguing-in-Publication entry 

Last, Peter R. (Peter Robert) et al. 

Sharks and rays of Borneo/ Peter R. Last, William T. White, Janine N. Caira, 

Dharmadi, Fahmi, Kirsten Jensen, Annie P.K. Lim, B. Mabel Manjaji-Matsumoto, 

Gavin J.P. Naylor, John J. Pogonoski, John D. Stevens, Gordon K. Yearsley. 

9781921605598 (pbk.) 

9781921605642 (pdf) 

Includes index. 

Bibliography 

Sharks – Borneo. 

Rays (Fishes) – Borneo. 

597.3095983 

Produced and distributed by 

CSIRO PUBLISHING 

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Cover design by Louise Bell 

Illustrations by Georgina Davis, Lindsay Marshall and Stephanie Barnes 

Book design by William White and Peter Last 

Layout by William White 

Printed in Australia by Ligare

iii 

Foreword 

Slow growth, delayed maturity and low fecundity make chondrichthyans 

particularly vulnerable to the many sources of mortality accompanying 

humankind’s spreading influence on the planet. Habitat degradation, overfishing 

and unsustainable fishing practices, including dynamiting, cyanide poisoning and 

finning, all contribute to the pressures imposed on this unproductive group of 

fishes. About a fifth of Indo-Australian chondrichthyans are considered threatened 

and forty percent of conservation concern in the 2009 IUCN Red List, so it 

is critical that regional faunas are well defined and guides are available to help 

make accurate identifications of species. Taxonomy is providing an increasingly 

important basis for managing species - the Borneo Broadfin Shark (Lamiopsis 

tephrodes Fowler, 1905) was recently resurrected from what was previously 

considered to be the monotypic genus Lamiopsis, as a result primarily of evidence 

of substantial DNA sequence divergence between sister species. Accurate 

taxonomy remains the product of painstaking effort from a committed group of 

experts using all contemporary tools available. 

Taxonomic studies of the fishes of Borneo started almost 200 years ago. In 1996, 

an international project funded by the Darwin Foundation provided data on 36 

chondrichthyan species from Sabah, and three years later a published checklist 

from Borneo included 82 species. During a National Science Foundation (NSF) 

study of the metazoan parasites of Borneo and their chondrichthyan hosts, this list 

was expanded to include 118 chondrichthyan species and provided the impetus 

for the international collaboration that has produced this book. This guide will aid 

the conservation and management of this most vulnerable group of marine fishes 

and provide much pleasure to people interested in the natural history of Borneo, 

particularly its highly diverse shark and ray fauna. 

Nic Bax 

Leader, Marine Biodiversity and Conservation Management Research, CSIRO 

Director, CERF Marine Biodiversity Hub. 

Australia

iv 

Contents 

Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii 

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 

How to use this book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 

Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 

Key to families and genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 

Hexanchiformes 

Hexanchidae (Cowsharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 

Squaliformes 

Squalidae (Dogfishes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 

Centrophoridae (Gulper Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 

Squatiniformes 

Squatinidae (Angelsharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 

Heterodontiformes 

Heterodontidae (Hornsharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 

Orectolobiformes 

Orectolobidae (Wobbegongs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 

Hemiscylliidae (Longtail Carpetsharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 

Ginglymostomatidae (Nurse Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 

Stegostomatidae (Zebra Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 

Rhincodontidae (Whale Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 

Lamniformes 

Alopiidae (Thresher Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 

Lamnidae (Mackerel Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 

Carcharhiniformes 

Scyliorhinidae (Catsharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 

Triakidae (Houndsharks). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 

Hemigaleidae (Weasel Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 

Carcharhinidae (Whaler Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 

Sphyrnidae (Hammerhead Sharks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

Rajiformes 

Pristoidei 

Pristidae (Sawfishes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 

Rhinoidei 

Rhinidae (Shark Rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146 

Rhynchobatoidei 

Rhynchobatidae (Wedgefishes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 

Rhinobatoidei 

Rhinobatidae (Shovelnose Rays). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154 

Torpedinoidei 

Narcinidae (Numbfishes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160 

Narkidae (Sleeper Rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166 

Rajoidei 

Rajidae (Skates) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 

Anacanthobatidae (Leg Skates) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178 

Myliobatoidei 

Plesiobatidae (Giant Stingarees) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 

Dasyatidae (Stingrays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182 

Gymnuridae (Butterfly Rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238 

Myliobatidae (Eagle Rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242 

Rhinopteridae (Cownose Rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252 

Mobulidae (Devilrays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256 

Chimaeriformes 

Chimaeridae (Shortnose Chimaeras) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268 

Images sourced outside Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270 

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271 

Checklist of Borneo sharks and rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277 

Scientific names index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283 

English common names index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 288 

Indonesian common names index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293 

Malaysian common names index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296

Introduction 

The region 

The island of Borneo lies at the western margin of the Coral Triangle of South-East 

Asia, a mega-diverse faunal region comprising the Philippines, Malaysia, Indonesia, 

Timor-Leste, Papua New Guinea and the Solomon Islands. Borneo is surrounded 

by four large marine basins, the Sulu, Celebes, Java and South China Seas. Divided 

administratively between the States of Brunei, Indonesia (Kalimantan), and Malaysia 

(Sabah and Sarawak), the 730,000 km 2 landmass supports a population of about 

18 million people. Considered to have unparalleled terrestrial biodiversity, its aquatic 

environments are equally complex and variable with a rich variety of habitats in both 

the sea and freshwater. Borneo has the longest rivers in Indonesia (Kapuas River, 

ca 1,140 km) and Malaysia (Rajang River, ca 560 km), as well as a few others longer 

than 800 km. Runoff from these rivers controls salinity in the nearby coastal zone, 

particularly in the south, forming expansive plumes of brackish water that create 

marginal estuaries that extend well into the open sea. Other parts of the island, such 

as the northern coast, are devoid of large rivers and here the habitats are dominated by 

freshwater intolerant marine species. There are also extensive areas of coral reef off the 

west coast. Very deep water habitats are restricted to parts of the Celebes and Sulu seas. 

Consequently, the intraregional distribution of the marine biota and its utilisation also 

vary considerably within the region. 

Biodiversity 

Borneo is considered to be a major centre of evolution and radiation of endemic 

terrestrial plants and animals. It is known to harbour 15,000 or so species of plants, 

more than 220 species of terrestrial mammals, and some 420 species of endemic birds. 

This high endemicity is largely due to extinction and speciation events during the 

Pleistocene ice ages and separation from continental Asia during the Holocene. A 

high level of endemism is shared by nearby aquatic systems, and of the 394 species of 

freshwater fishes, 149 are endemic to the island. Falling within the Coral Triangle, which 

covers only 2% of the world’s oceans but contains 76% of known coral species, the 

marine fish fauna is equally impressive. In a comprehensive biodiversity survey of coral 

reef fishes of the Indo Pacific, Allen (2008) recorded almost 1,500 species in the 100,000 

km 2 grid along the eastern border of Sabah and Kalimantan. 

Early work on Borneo fishes 

Borneo is historically important in ichthyology as it is the collection locality for some 

of the first taxonomic studies of the region’s fishes. Schlegel & Müller (1839) described 

the first marine fish from the region and soon after, Heckel (1843) described species 

from freshwater. Their work was followed by the unparalleled efforts of the Dutch 

field scientist, Pieter Bleeker, who between 1850 and 1876 described more than

S h a r k s a n d R a y s o f B o r n e o 

1,100 new fish species from the Indo-Malay archipelago, including a large number of 

chondrichthyans, and produced the historically important Atlas Ichthyologique des Indes 

Orientales Néerlandaises. Many of Bleeker’s 12,000 or so fish specimens are catalogued 

at the Natural History Museum in Leiden, Netherlands. Other famous 19 th C 

ichthyologists, including Richardson, Günther and Kaup, also made important contributions 

to our knowledge of this fascinating fauna. 

Cartilaginous fishes are relatively diverse in both marine and freshwater environments 

of Borneo. The region is the type locality for Pristis zijsron, Carcharhinus borneensis, 

C. sealei, Lamiopsis tephrodes and Sinobatis borneensis, as well as 10 new species 

discovered recently, i.e. Orectolobus leptolineatus, Cephaloscyllium sarawakensis, Glyphis 

fowlerae, Rhynchobatus springeri, Okamejei cairae, Okamejei jensenae, Himantura lobistoma, 

Pastinachus gracilicaudus, Pastinachus solocirostris and Pastinachus stellurostris. Until recently, 

this fauna was poorly known and accounts by field scientists in the mid-19 th C 

produced only patchy records of the region’s elasmobranchs. Cantor (1849) compiled 

a catalogue of Malaysian fishes that included 28 species of sharks and rays. Basic 

information, such as collection details, morphological traits, distribution within the 

region and basic biology, were not collected. In the early 20 th C, Henry Fowler 

and Albert Herre provided supplementary data on elasmobranchs from the region, 

especially from Sabah and Sarawak. Fowler, in particular, considered most of Bleeker’s 

new rays to be invalid. During a series of ad hoc surveys in the early 1990s, several of 

Bleeker’s species were rediscovered in fish markets of nearby Java rekindling interest in 

the taxonomy of the group. 

Environmental pressures 

About 120 million people, whose livelihoods depend directly on fishing, live along the 

coasts and islands of the Coral Triangle. This large population is placing huge pressures 

on the marine environment through a range of fishing activities, many of which are 

unreported and unregulated (but not illegal, particularly fishing activities of the numerous 

artisanal fishers, where marine fishes are considered as “free common-good resources”). 

The live fish trade, supplying dominant markets in Hong Kong and China, takes some 

30,000 t/yr of reef fish worth more than US$150/kg out of the Coral Triangle, with 

Kota Kinabalu the main base in Malaysia. As well as supplying restaurants, these fish also 

enter the ornamental fish trade and many are caught by illegal cyanide fishing. Due to 

dynamite fishing, illegal, unreported and unregulated (IUU) fishing by other South- 

East Asian countries, and over-harvesting of invertebrates such as triton, sea cucumber 

and giant clam, the area’s coral reefs and associated ecosystems are being increasingly 

threatened. While there are some Marine Parks in the region, the lack of enforcement 

reduces their effectiveness. However there are some recent encouraging initiatives. In 

March 2008, the Asia Development Bank provided a US$1.5 million grant for environmentalists 

and respective governments to protect the forests and seas of Brunei, Indonesia, 

Malaysia and the Philippines.

I n t r o d u c t i o n 

 

Shark fisheries 

Major environmental degradation in Borneo, along with a four-fold increase in 

worldwide fisheries since World War II, has coincided with a major increase in 

exploitation of cartilaginous fishes. Most species known from Borneo occur in coastal 

or freshwater habitats easily accessible to fisheries and other antropogenic activities 

impacting on these fishes. Indonesia traditionally has the largest annual global production of 

elasmobranchs but, despite the early focus on them, only recently has the composition of 

this fauna been comprehensively described. The declaration of the Indonesian Exclusive 

Economic Zone (EEZ) in 1983, which includes the waters of Kalimantan, gives Indonesia 

control of more than 5.4 million km 2 of sea. With a reported 118,000 and 121,000 

tonnes landed in 2003 and 2004 respectively, Indonesia has the largest chondrichthyan 

fishery in the world. Catches of Malaysian Borneo are smaller but far from insignificant. 

Chondrichthyans are caught locally by target fisheries, and as commercial and artisanal 

bycatch, using fishing methods that vary greatly geographically and between nations. In 

Indonesia, chondrichthyans are targeted using a variety of fishing methods, such as gill 

and tangle nets, longlines and harpoons. Fisheries that land substantial catches of elasmobranchs 

as bycatch include those operating small-scale bottom trawls, trammel and gill 

nets, and longlines and droplines. In Malaysia, demersal trawling and drift nets account for 

the bulk of the catch (DOFM, 2006). 

Darwin project 

In 1996, the first major investigation of the elasmobranchs of Sabah was initiated 

by an international research team led by Sarah Fowler and supported by funding 

from the UK Darwin Foundation. This project attempted to answer basic questions 

relating to the biodiversity, fisheries impact, biology, and conservation status of coastal 

marine and freshwater elasmobranchs of Sabah, and compile a preliminary checklist of 

chondrichthyans of greater Borneo. The survey work, which focused on both coastal 

marine and freshwater habitats, uncovered a relatively rich freshwater fauna. For 

example, Fowler & Payne (1995) recorded two species of rays, several species of rare 

sharks and a species of sawfish, from the Kinabatangan River in East Malaysia. A 

preliminary report on the project was published by Cook & Compagno (1996) providing 

data on 36 elasmobranch species in markets and freshwater habitats of Sabah. At the 

completion of the formal survey of elasmobranchs from fish markets, and river and 

coastal villages throughout Sabah, conducted by the IUCN Shark Specialist Group (see 

Fowler et al., 1999), 69 species of elasmobranchs (34 sharks and 35 rays) were recorded, 23 

of which were new records for the state. Of particular note was the listing of new species 

of the river shark, genus Glyphis, and new species of Mustelus, Orectolobus and Rhynchobatus. 

A checklist of chondrichthyan fishes of all of Borneo was compiled at the same time, 

which included 82 species (40 sharks, 41 rays and 1 chimaera). 

Following on from this project (1999-2004), a primarily Malaysian led study of the 

chondrichthyans of Brunei and Malaysia, including Sabah and Sarawak, led to the


collection of 760 specimens of 79 species, comprised of 43 sharks, 35 rays and one 

chimaera (Yano et al., 2005), but some of these species appear to be invalid. A new 

and valid species of catshark, Cephaloscyllium sarawakensis, was also described from this 

research. 

Figure 1 Map of Borneo showing the locations of landing sites and villages surveyed. The 

different landmass colours used correspond to the different countries making up Borneo: 

grey (Indonesia), green (Malaysia) and yellow (Brunei).


 

National Science Foundation project 

The Darwin project was followed by a more broad-based field project funded largely by 

the National Science Foundation (NSF) covering the whole of Borneo. In a multi-agency 

collaborative effort, involving parasite and elasmobranch taxonomists from institutions 

in the USA, Australia, Indonesia and Malaysia, this project focused on surveying and 

describing the chondrichthyans and their metazoan parasite faunas of inshore, estuarine 

and selected freshwater habitats. The project consisted of two phases: field surveys of 

Malaysian Borneo (2002–04) and Indonesian Kalimantan (2006–08) (Fig. 1). Earlier 

research suggested this region was home to well over 100 species of sharks and rays, many 

of which were poorly known, and some new to science. While records of metazoan 

parasites were virtually non-existent for elasmobranchs in this geographic region, based 

on studies elsewhere, it was predicted that the host fauna could yield over 500 parasite 

species with well over 70% new to science. In all, over 600 species of metazoan parasites, 

of which about 85% are new to science, were collected during the project. 

Several thousand elasmobranch specimens were examined from research and chartered 

fishing vessels, commercial trawlers, landing sites, and fish markets from around the 

island. In many cases, these specimens were documented with a series of digital photos. 

These images can be accessed via the elasmobranch host specimen database available 

at https://web2.uconn.edu/tapeworm/hosts.php. Tissue samples were taken from 

elasmobranch specimens whenever possible for subsequent DNA sequencing, 

comparative analysis and species validation. These sequences are being databased as part of 

a global elasmobranch sequence repository that will eventually serve as a baseline against 

which new elasmobranch records can be compared. The surveys led to the descriptions 

of an additional 9 new species, as well as specimens of endangered, rare or little known 

species, including the rediscovery of the Borneo Shark (Carcharhinus borneensis), once 

considered to be extinct. These baseline taxonomic data can be used to facilitate development 

of a resource management plan for elasmobranchs of the region. Several inshore 

species exhibit regional patterns in distribution that appear to be closely related to habitat 

availability. For example, several micro-endemic stingrays (genera Himantura and Pastinachus) 

occur in the environs of the large Kapuas River estuary in western Kalimantan, and 

the region has additional undocumented, extreme, intraspecific variability in some shark 

groups. 

Characteristics of the fauna 

The chondrichthyan fauna of Borneo presently contains 118 validated species, including 

52 sharks, 65 rays and 1 chimaera. The representation of the group at highest taxonomic 

levels (Table 1) is similar to regions adjacent (i.e. northwestern Australia, Philippines 

and Indonesia), but there are some differences in the relative importance of some 

subgroups. For example, ground sharks (Carcharhiniformes) and eagle rays and their allies 

(Myliobatoidei) are the dominant subgroups in each region comprising more than 20% 

of the species; the latter are more dominant in Borneo than any other region. Conversely, 

dogsharks (Squaliformes) and skates (Rajoidei) are relatively depauperate in Borneo and


Table 1 Proportional species richness (%) of shark and ray groups across the region. 

Higher taxa 

NW Philippines Indonesia Borneo 

Australia 

Heterodontiformes 1 1 - 1 

Lamniformes 5 6 6 2 

Orectolobiformes 7 6 6 7 

Carcharhiniformes 31 32 33 31 

Hexanchiformes 2 2 2 1 

Squaliformes 12 10 10 2 

Squatiniformes 1 1 1 1 

Pristiphoriformes - 1 - - 

Pristoidei 3 2 1 3 

Rhinoidei 1 1 1 1 

Rhynchobatoidei 1 2 2 2 

Rhinobatoidei 2 3 4 2 

Platyrhinoidei - 1 - - 

Torpedinoidei 3 4 1 4 

Rajoidei 8 7 1 4 

Myliobatoidei 20 21 33 38 

Chimaeriformes 3 2 1 1 

TOTAL 179 164 139 118 

this may be due to the comparative paucity of deepwater habitats near the coast. Clearly 

the richest chondrichthyan families in all regions, the whaler sharks (Carcharhinidae) 

and stingrays (Dasyatidae), both attained the highest relative proportions of any fauna in 

Borneo (Table 2). Also, eagle rays (Myliobatidae) and devil rays (Mobulidae) are more 

diverse in this region than off Australia. Once again, deepwater subgroups, such as the 

gulper sharks (Centrophoridae), spurdogs (Squalidae) and lantern sharks (Etmopteridae), 

were absent from the ten main families. 

Shelf demersal species, comprising about 70% of the fauna, are more dominant in Borneo 

than in other regions (Table 3); conversely, the proportions of outer shelf and slope species 

are comparatively low. Freshwater and estuarine species comprise almost 7% of the fauna; 

by far the highest proportion of any region and this may be linked to the diversity of 

these habitats in the region.


 

Table 2 Comparative proportional species richness (>2%) of the major shark and ray 

families across the region. 

Family NW Australia Philippines Indonesia Borneo 

Carcharhinidae 16 15 19 24 

Dasyatidae 12 11 19 25 

Scyliorhinidae 8 7 4 4 

Rajidae 8 7 4 

Triakidae 4 4 4 

Orectolobidae 3 

Centrophoridae 3 4 5 

Etmopteridae 3 

Squalidae 3 4 4 

Urolophidae 3 

Pristidae 3 3 

Myliobatidae 4 4 4 

Mobulidae 4 4 

Rhinobatidae 4 3 

Sphyrnidae 3 3 

Hemigaleidae 3 3 

Narcinidae 3 

Hemiscylliidae 4 

Rhynchobatidae 3 

Aims of this book 

This international collaborative survey, funded largely by the National Science 

Foundation, has provided a fuller understanding of the biodiversity of chondrichthyans 

and their parasite faunas in one of the most faunally diverse parts of the planet. This work 

complements related research on parasites and their host faunas conducted elsewhere 

over the past 25 years, as well as similar regional surveys of the chondrichthyans of 

the Philippines and Indonesia. In the late 1990s, an elasmobranch biodiversity and 

conservation project in the Philippines, defined the fauna and produced conservation 

and management strategies to ensure sustainability of chondrichthyan fisheries in the 

environs of the nearby Sulu Sea. Soon after, an ACIAR-funded investigation of the 

chondrichthyans of Indonesia led to the production of the first regional guide to the 

sharks and rays of this region. Collectively, these studies have contributed greatly to an 

improved knowledge of the composition and distribution of these fishes in the Indo- 

Malay Archipelago.


Table 3 Proportional species richness (%) for primary biomes across the region. 

Habitat type 

NW 

Australia 

Philippines Indonesia Borneo 

Freshwater/estuarine 2 - 1 7 

Continental/insular shelves 49 52 60 72 

Shelf to slope (demersal) 12 13 8 7 

Continental/insular slopes 24 18 14 3 

Shelf to oceanic (pelagic) 9 14 14 11 

Oceanic (pelagic) 2 1 3 1 

Wide ranging 1 1 1 -

How to use this book 

This book is designed to assist in the identification of sharks, rays and chimaeras that can 

be observed in fisheries catches or in domestic fish markets in the riverine, estuarine and 

marine regions of Borneo. This island includes both Indonesian and Malaysian landmass 

and contains the entire country of Brunei. Species included in this guide are mostly those 

observed in fish market surveys of Borneo, but include some additional species taken 

in other surveys (e.g. Yano et al., 2005) or species that have been observed by divers in 

the region (e.g. Silvertip Shark Carcharhinus albimarginatus, Tawny Nurse Shark Nebrius 

ferrugineus). 

A key is provided so that a specimen can be systematically identified to its family 

and genus group. Users can go straight to the relevant chapter when the family is 

already known. However, where users are in any doubt they should firstly refer to 

the key when making an identification. Species treatment sections follow the keys 

and these are organised in phylogenetic sequence of families, commencing with the 

cowsharks (Hexanchidae) and finishing with chimaeras (Chimaeridae). Each major 

group of sharks or chimaeras is colour coded at the level of order, while each major 

group of rays is colour coded at the level of suborder. Species within each family are 

arranged in alphabetical order of their scientific names and undescribed species are arranged 

prior to nominal members of the same genus. Species accounts have been kept relatively 

simple, but the use of technical ichthyological terms is sometimes unavoidable and users 

can consult the illustrated glossary for a definition of these terms. 

Species treatments 

Details of the identifying features, size, distribution, habitat and biology, and utilisation 

are provided for each species under appropriate subheadings. Additional information on 

other names, synonyms, conservation status and relevant literature, as well as images and 

maps, are also provided. 

Images and illustrations 

The primary image for each species is usually of a specimen from Borneo. However, in 

some circumstances, when local images were unavailable, a suitable image from a nearby 

region was used. A list of the species for which images were sourced from outside of 

Borneo is provided on page 270. 

Additional photographs were used, where possible, to highlight identifying characteristics 

of a species that are not apparent from the primary image alone. When photographs were 

unavailable or unsuitable, line drawings were used.

10 S h a r k s a n d R a y s o f B o r n e o 

Common and scientific names 

The recommended common names generally follow English names adopted 

by Compagno et al. (2005a) for sharks, and Last & Stevens (2009) and the relevant 

sections in Carpenter & Niem (1999) for rays, with the exception of some species that 

have a more commonly accepted regional name. Newly recognised species have been 

give new common names. Alternative English, Indonesian and Malaysian names used in 

the Indo–Malay Archipelago are also given under ‘Other names’. Local common names 

are based on information gathered during this project, and from Yano et al. (2005) and 

White et al. (2006b). 

The scientific name of each species consists of genus and species names, the name of 

the author(s) who named it, and the year in which it was named. Parentheses around 

the author(s) and date indicate that the species was originally placed in a different genus. 

Species of uncertain identity are referred to by their generic name followed by ‘sp.’. 

Identifying features 

These include the main features by which a species can be most easily identified in the 

field, such as body and tooth shapes, fin positions and dimensions, and colour pattern. Each 

character is ordered numerically and the corresponding number is usually positioned on 

either the primary or secondary images to illustrate the feature or at least its location. 

Size 

Maximum size, and the sizes at birth, hatching and sexual maturity, is given for each 

species when known. Unless stated otherwise, all measurements refer to total lengths of 

species. In the case of sharks, total length is measured as a straight line from the tip of the 

snout to the tip of the extended upper caudal-fin lobe. Ray sizes are also given as total 

lengths, except in the families Dasyatidae, Gymnuridae, Myliobatidae, Rhinopteridae and 

Mobulidae, where the tail is frequently absent or damaged. For these groups the main size 

measurement is disc width (DW), taken from the extremities of the left and right pectoral 

fins. The caudal filament at the tail tip of chimaeras is not included in the total length; 

instead, the length measurement is taken at the posterior edge of the caudal fin. 

Distribution 

Maps show the distribution of each species in Borneo with specimen records marked 

by circles; red is used for distributions recorded during the present study, blue for other 

distributional data, and orange for unpublished data collected in Kalimantan by one of 

us (Fahmi). An asterix indicates the type locality for those species originally described 

from Borneo. These maps reflect current knowledge, but many species are probably more 

widely distributed in the region. River drainages are not shown on the maps so some 

distributional appear to be on land. Global distributional ranges for species and their 

ranges within Borneo are provided separately in the text.

H o w t o u s e t h i s b o o k 

11 

Habitat and biology 

This section covers basic information on the habitats (pelagic and oceanic, demersal on 

continental shelf, etc.) and depth distributions of species, and also on aspects of their 

reproductive biology, age and growth, their prey and, in some cases, their potential 

danger to humans. 

Utilisation 

This section briefly details the way species are used in Borneo, including what parts of the 

animals are used. In some cases it relates to use in nearby regions when the full extent of 

use in Borneo is unclear or unknown. 

Conservation status 

The current (at time of writing) IUCN Red List Threatened Species Assessment for 

each species is included here, along with any other relevant conservation assessment 

(e.g. CITES). For the IUCN Red List category, species that are listed as Vulnerable, 

Endangered or Critically Endangered, are considered as ‘threatened’ species by the IUCN. 

Local synonyms 

Any other scientific names used for a particular species are listed in this section, including 

junior synonyms, temporary names and misidentifications (names followed by a colon). 

Only alternative names used within the Coral Triangle area are listed in this section, i.e. 

Borneo, Indonesia, Malaysian Peninsular, Philippines and New Guinea. The literature 

sources of any temporary names or misidentifications are also provided, as well as 

misidentified images in field guides. 

References 

This section includes relevant literature sources used for species in the general region. 

Primary regional references covering a large proportion of Borneo species, such as 

Carpenter & Niem (1999), Compagno et al. (2005a), Yano et al. (2005), White et al. 

(2006b) and Last & Stevens (2009), are not referred to repeatedly. Similarly, several major 

biological references relevant to the region were also not referred to repeatedly, i.e. White 

(2007c) and White & Dharmadi (2007). The full citations of references can be found in 

the Reference section beginning on page 271. 

Indexes 

Indexes of scientific names, English common names, and Indonesian and Malaysian 

local names are provided on pages 283–298, and a checklist of species known from 

Borneo is provided on pages 277–282. In the checklist, authorities are sometimes 

abbreviated to improve the layout, e.g. M. & H. for Müller & Henle, Bl. & Sch. for Bloch 

& Schneider.


Glossary 

acute – sharp or pointed. 

adelphophagy – method of embryonic nutrition where the embryo cannibalises other 

embryos within the uterus. 

adpressed – pressed flat against the body. 

alar thorns – paired patches of thorns on the outer disc of mature male skates. 

angular – forming a distinct angle. 

anterior – relating to front of or head end of an object. 

barbel – a slender, tentacle-like sensory structure on the head. 

benthic – living on the bottom of the ocean. 

benthopelagic – living above but near the bottom of the ocean. 

bilobate – having two lobes. 

blotch – an enlarged area or patch that is different in colour to adjacent areas. 

brackish – waters with a salinity between that of freshwater and saltwater. 

buccal – pertaining to the mouth cavity. 

bycatch – component of the catch excluding targeted commercial species. 

cartilage – a skeletal material consisting of a matrix of soft, white or translucent 

chondrin. 

caudal keel – a longitudinal fleshy ridge along side of the caudal peduncle. 

caudal peduncle – the posterior part of the body supporting the caudal fin. 

cephalic lobe – broad lobe on forehead of some rays. 

cephalopods – group of animals including cuttlefishes, nautili, squids and octopi. 

cetaceans – group of aquatic mammals including whales and dolphins. 

chevron – a V-shaped pattern of adjoining lines. 

chondrichthyans – group of fishes including sharks, rays and chimaeras. 

chondrin – a horny, gelatinous protein substance that forms the matrix of cartilage. 

circumglobal – distributed around the world within a certain latitudinal range. 

claspers – modified portions of pelvic fins in male chondrichthyans used for

G l o s s a r y 13 

transferring sperm to female during mating. 

cloaca – a common opening for digestive, urinary and reproductive tracts (also called 

vent). 

common name – the informal name for an animal, which often varies from place to 

place. 

compressed – flattened laterally from side to side. 

concave – curved inwards (opposite of convex) 

continental shelf – the shelf-like part of the seabed adjacent to the coast to a depth of 

about 200 m. 

continental slope – the typically steep, slope-like part of seabed bordering the 

continental shelf to a depth of about 2000 m. 

convex – arched, curved outwards (opposite of concave). 

cosmopolitan – having a worldwide distribution. 

crenulate – margin shaped into small rounded scallops. 

crescentic – shaped like the new moon (roughly C-shaped). 

crustaceans – group of invertebrate animals including crabs, shrimps, prawns, lobsters 

and crayfish. 

cusp – a projection on a tooth. 

cusplet – a small cusp. 

demersal – living on or near the bottom of the ocean. 

denticle – a small, tooth-like structure; plate-like scale of a cartilaginous fish. 

depressed – dorsoventrally flattened from top to bottom. 

depth – height of body or head from top to bottom; also distance from sea surface to 

the bottom. 

dermal – relating to the skin. 

dermal flaps – outgrowths of skin. 

disc – the combined head, trunk and enlarged pectoral fins of those cartilaginous fishes 

with depressed bodies, e.g. stingrays. 

distal – region, border or point remote from the site of attachment (opposite of 

proximal).


dorsal – relating to upper part or surface of back. 

dorsolateral – positioned or orientated between dorsal and lateral surfaces. 

dusky – slightly dark or greyish in colour. 

elasmobranchs – group of fishes including sharks and rays. 

electric organ – organ capable of delivering a mild electric shock. 

elongate – extended in length in relation to another object or drawn out. 

embedded – within surrounding tissue. 

endemic – native and restricted to a certain area. 

epipelagic – upper part of the oceanic zone from the surface to about 200 m. 

estuarine – living mainly in estuaries. 

falcate – curved like a sickle. 

family – a group term for classifying organisms, containing one or more closely related 

genera. 

fauna – the communities of animals in an area. 

fertilisation – the union of male and female cells to form a new individual. 

filter feeding – filtering food particles from the water by means of the gill rakers. 

fusiform – spindle-shaped, tapering at both ends. 

genus – a group term for classifying organisms, containing one or more related species. 

gestation – the period of development in the uterus from conception until birth. 

gillnet – a net used to tangle fishes. 

gill opening – an opening (usually slit-like in cartilaginous fishes) on head that 

connects the gill chamber to the exterior. 

granulations – fine denticles. 

habitat – the locality with its own particular environment in which an organism lives. 

hammer-shaped – shaped with paired lateral expansions, like the head of a mallet. 

head – specialised anterior part of an animal on which the mouth and major sensory 

organs are located; part other than body and tail. 

head clasper – small appendage on the forehead of mature male chimaeroid fishes.


histotrophy – form of embryonic nutrition where the developing embryos receive a 

lipid-rich histotroph, or uterine milk, usually delivered through extensions of the 

uterine wall called trophonemata. 

holotype – the specimen used in the original description of a species. 

hyomandibular pores – line of enlarged pores extending posteriorly from the mouth 

corners. 

infraorbital – area below the eye. 

interdorsal – space on dorsal surface between 1 st and 2 nd dorsal fins. 

interdorsal ridge – ridge of skin between dorsal fins. 

internarial space – distance between the nostrils. 

internasal flap – fleshy flap extending between nostrils, sometimes partly covering the 

mouth. 

interorbital space – area on top of head between eyes. 

jaws – part of mouth supporting teeth. 

juvenile – young fish, similar in form to adults but not yet sexually mature. 

keel – a fleshy ridge. 

lateral – referring to the sides. 

lateral-line canal – part of the sensory network of chimaeras; appearing as a distinct 

line. 

lateral ridges – fleshy expansions on sides of body. 

life cycle – the generalised history of a species from birth to death. 

lip – fleshy outer portion of jaws. 

longitudinal – lengthwise (opposite of transverse). 

longline – a fishing line bearing numerous baited hooks, typically set horizontally in 

water column. 

lunate – shaped like a crescent moon. 

margin – edge or rim. 

median – relating to the middle of an object. 

mesopelagic – living in open ocean at depths between 200 and 1000 m.


mouth – opening through which food enters the alimentary canal. 

mucous canal – part of the sensory network of the head of chimaeras. 

multicuspid – a tooth or denticle with several cusps. 

nape – region of head above and behind eyes. 

nasal curtain – see internasal flap. 

nasoral grooves – furrow connecting mouth to the nostrils. 

nektonic – swimming organisms capable of moving against a current and thus not 

only drifting. 

nictitating eyelid – a transparent, moveable membrane or inner eyelid that protects 

eye. 

nomenclature – the systematic naming of animals and plants. 

nostril – external opening of the nasal organs. 

nuchal – pertaining to the nape. 

obtuse – broadly rounded or having a blunt end. 

oceanic – living in the open ocean. 

ocellus (pl. ocelli) – an eye-like spot or marking with a marginal ring. 

ocular – relating to the eye. 

oophagy – method of matrotrophic embryonic nutrition in viviparous species where 

the embryo feeds on unfertilised eggs in the uterus. 

oral – relating to the mouth. 

orbit – bony cavity in skull where eyeball is housed. 

oviparous – producing eggs that hatch after being deposited from the body of a 

pregnant female. 

papillae – a small fleshy projection. 

parasitic – living and feeding in or on another organism to the detriment of that 

organism. 

pearl thorn – nuchal thorn of stingrays shaped like an embedded pearl. 

pedicel – small stalk, usually of a denticle. 

pelagic – free-swimming in the seas, oceans or open water and not associated with the 

bottom.


plain – uniformly coloured, without a contrasting colour pattern. 

placental - method of matrotrophic embryonic nutrition in viviparous species where 

nutrients are transferred across the mother’s uterine epithelium, which is intimately 

connected with foetal tissue (placenta). 

plankton – small animals or plants that drift in open water. 

population – a biological unit that represents the individuals of a species living in a 

certain area. 

pore – small secretory or sensory opening. 

posterior – relating to hind of or rear end of an object. 

precaudal pit – a transverse or longitudinal notch on caudal peduncle just anterior to 

origin of caudal fin in some sharks. 

predator – feeding on other animals. 

pre-pelvic clasper – small, retractable appendages on each side just forward of the 

pelvic fins in mature male chimaeroid fishes. 

preorbital – before (anterior to) the eye. 

purse seine – a fishing net used to encircle surface-dwelling fish and land onto a boat. 

quadrangular – shaped with four distinct edges or marigns. 

reticulated – divided into a network. 

reticulations – markings in a general form of a net. 

rhomboidal – diamond-shaped. 

rostral cartilage – a gristly structure supporting the snout. 

rostral teeth – tooth-like projections on the sides of the snout of sawfishes and sawsharks. 

rostrum (adj. rostral) – a projecting snout. 

rounded – margin evenly convex. 

saddle – a blotch extending across dorsal surface from one side to another. 

salinity – the concentration of salt in water. 

school – a close aggregation of fish swimming in association with each other. 

scientific name – the formal binomial name of an organism consisting of the genus 

and species names; only one valid scientific name exists per species.


scythe-like – shaped like a scythe; long and narrowly curved. 

seine – a fishing net that hangs vertically in water column and encircles fish when its 

ends are drawn together. 

serrate – saw-like. 

snout – part of head in front of eyes. 

species – actually or potentially inter-breeding populations that are reproductively 

isolated from other populations. 

spine – a sharp projecting point. 

spiracle – a respiratory opening behind the eye in sharks and rays. 

spiracular fold – fold of skin present on the hind margin of the spiracular opening in 

some ray species. 

spot – a regularly shaped or rounded area of a colour different to adjacent areas. 

squalene – oil produced from the liver of some sharks. 

stellate – star-shaped. 

stinging spine – large, serrated bony structure on the tail of some rays. 

stripe – a contrasting longitudinal pattern in the form of a line. 

subcircular – almost circular. 

subequal – almost equal. 

substrate – the substance forming the bottom of the sea or ocean. 

subterminal – positioned near but not at end of an object. 

supraorbital crests – enlarged ridges above the eyes. 

sympatric – living together in the same geographic area. 

symphysis (adj. symphysial) – relating to medial junction of either the upper or lower 

jaw. 

synonym (adj. synonymous) – each of two or more scientific names of the same rank 

used to denote the same taxon. 

tail – part of fish between cloaca and origin of the caudal fin. 

taxonomy – the science of classification of plants and animals. 

taxon – any formal taxonomic unit or category of organisms (genus, species, etc.)


teeth – hard outgrowths on jaws, used for biting and masticating food. 

teleost – large group containing most bony fishes. 

tendril – a slender, curling barbel. 

terminal – located at or forming the end of something. 

thorn – large denticles on surface of a ray or skate. 

tip – the extremity of a part of a fish. 

tooth rows – horizontal rows of teeth in the jaws. 

total length – longest length of a fish, from snout tip to upper caudal tip or tail tip. 

trammel net – a three layered fishing net used for entangling fish. 

transverse – directed crosswise, across width (opposite of longitudinal). 

trawl net – fishing net which is dragged behind a boat. 

tricuspid tooth – tooth with 3 cusps. 

trunk – part of fish between head and tail; between fifth gill slit and cloaca. 

ventral – relating to the lower part or surface. 

vertebrate – animal having a vertebral column or backbone. 

viviparous – producing live young from within the body of the parent female.


anterior 

eye 

spiracle 

snout tip 

gill opening 

dorsal-fin spine 

1 st dorsal fin 

2 nd dorsal fin 

precaudal pit 

posterior 

nostril 

mouth 

labial furrows 

pelvic fin anal fin 

caudal keel 

clasper (males) 

caudal fin 

structural features of sharks 

pectoral fin 

anterior margin 

apex 

spine 

posterior margin 

origin 

lobe 

free rear tip 

insertion 

inner margin 

shark dorsal fin 

barbel 

circumnarial groove 

nostril 

circumnarial fold 

oronasal groove 

upper labial furrow 

mouth 

symphysial groove 

lower labial furrow 

shark undersurface of head


upper origin 

dorsal margin 

upper lobe 

terminal lobe 

posterior tip 

terminal margin 

subterminal margin 

caudal fork 

subterminal notch 

upper postventral margin 

lower origin 

lower postventral margin 

shark caudal fin 

preventral margin 

lower lobe 

ventral tip 

ventral 

dorsal 

internasal flap 

mouth 

gill opening 

spiracle 

eye 

rostral cartilage 

shoulder 

pectoral fin 

cloaca 

pectoral fin insertion 

thorn 

pelvic fin 

caudal fin 

lateral tail fold 

clasper 

(male) 


1 st dorsal fin stinging spine 

structural features of rays


nostril 

lobes 

internasal flap 

fringe 

papillae 

mouth 

ray nostrils and mouth (oronasal) 

enlarged thorn 

stinging spine 

dorsal fold 

thorn 

granulations 

ventral fold 

ray tail 

head clasper (male) 

spine 

1 st dorsal fin 

lateral line 

mucous canal 

eye 


caudal-fin 

upper lobe 

mouth 

gill opening 

pre-pelvic 

clasper (male) 

clasper 

(male) 

anal fin 

caudal filament 

caudal-fin 

lower lobe 

pectoral fin 

pelvic fin 

structural features of chimaerids

R u n n i n g h e a d 23 

Key to families and genera 

1. Single gill opening on each side of head 

(fig. 1) .................................................. 

............ Family Chimaeridae, Chimaeras 

fig. 1; 266 

5–7 gill openings on each side of head 

........................................................... 2 

fig. 1 

2. Body flattened, ray-like; gill openings on 

undersurface of head (except in Angel 

Sharks) ..................................................... 3 

Body more or less fusiform, shark-like; 

gill openings on sides of head ........... 17 

3. Gill openings partly on sides of head 

(fig. 2); pectoral fins very large, clearly 

detached from head (fig. 2) ................. 

......... Family Squatinidae, Angel Sharks 

fig. 3; 42 

Gill openings entirely on undersurface 

of head; pectoral fins wholly or partly 

joined to head .................................... 4 

4. Snout saw-like, flattened, armed with 

lateral teeth (fig. 4) ................................. 

..................... Family Pristidae, Sawfishes 

Rostral teeth absent from basal quarter 

of saw (fig. 4); lower lobe of caudal fin 

large, more than half length of upper 

lobe (fig. 4) .......................... Anoxypristis 

fig. 4; 140 

Rostral teeth extending onto basal quarter 

of saw, almost to its base (fig. 5); lower 

lobe of caudal fin small, less than half 

length of upper lobe (fig. 5) .......... Pristis 

fig. 5; 142 

Snout not saw-like, no rostral teeth .... 5 

fig. 2 

fig. 3 

fig. 4 

fig. 5


5. Two prominent dorsal fins (fig. 6); 1 st 

dorsal fin originating closer to insertion 

of pelvic fins than to tail tip (fig. 6) ...... 6 

0–2 dorsal fins; 1 st dorsal fin originating 

closer to tail tip than to insertion of pelvic 

fins when 2 fins are present (fig. 7) ..... 10 

6. Caudal fin with a well developed 

ventral lobe (fig. 8); pectoral and pelvic 

fins separated slightly, not touching 

(fig. 8) .............................................. 7 

Ventral lobe of caudal fin not prominent 

(fig. 6); pectoral and pelvic fins touching 

or overlapping (fig. 6) ........................ 8 

7. Head triangular (fig. 8); upper disc with 

small thorns; two spiracular folds ....... 

.... Family Rhynchobatidae, Wedgefishes 

fig. 8; 148 

Head broadly rounded (fig. 9); upper 

disc with ridges lined with large thorns; 

no spiracular folds ................................. 

................ Family Rhinidae, Shark Rays 

fig. 9; 146 

8. Snout triangular, forming an acute angle 

at tip (fig. 10); body surface rough with 

thorns or fine denticles; no electric 

organs ................................................... 

... Family Rhinobatidae, Shovelnose Rays 

Anterior nasal aperture almost rectangular, 

very large (fig. 11); snout length more than 

5 times eye diameter .............Glaucostegus 

fig. 10; 154 

Anterior nasal aperture almost circular, 

relatively smaller (fig. 12); snout length less 

than 5 times eye diameter ........ Rhinobatos 

fig. 13; 158 

Snout broadly rounded (fig. 14); body 

surface entirely smooth; electric organs 

present ................................................ 9 

fig. 6 

fig. 7 

fig. 8 

fig. 9 

fig. 10 

fig. 11 fig. 12 

fig. 13

K e y t o f a m i l i e s a n d g e n e r a 25 

9. 2 dorsal fins present (fig. 14); a deep 

groove around mouth; teeth extending 

onto outer surfaces of upper and lower 

jaws in most species ............................... 

............. Family Narcinidae, Numbfishes 

fig. 14; 160 

0–1 dorsal fins present (figs 15 and 16); 

a shallow groove around mouth; teeth 

not extending onto outer surfaces of 

upper and lower jaws in most species 

............... Family Narkidae, Sleeper Rays 

No dorsal fins present (fig. 15) ....... Temera 

fig. 15; 168 

One dorsal fin present (fig. 16) ...... Narke 

fig. 16; 166 

10. Pelvic fin divided into two distinct lobes 

(fig. 17); no enlarged stinging spine on tail 

......................................................... 11 

Pelvic fin with only one lobe (fig. 18); 

usually with 1 or more enlarged, serrated 

stinging spines (scar visible when spine 

absent) .............................................. 12 

11. Entire dorsal surface smooth (except 

for alar thorns of male); preorbital snout 

more than 8 times eye diameter (fig. 19); 

tail very short and thin, without dorsal 

fins (fig. 19) .......................................... 

..... Family Anacanthobatidae, Leg Skates 

fig. 19; 178 

Thorns or fine denticles present on dorsal 

surface; preorbital snout much less than 8 

times eye diameter (fig. 20); tail slender 

(but not filamentous), with 2 dorsal fins 

when undamaged (fig. 20) ..................... 

........................... Family Rajidae, Skates 

fig. 20; 170 

12. Anterior part of head not extending 

forward beyond disc (fig. 21); eyes located 

fig. 14 

fig. 15 

fig. 16 

fig. 17 fig. 18 

fig. 19 

fig. 20


dorsally and well inward from disc margin 

(fig. 21) ............................................ 13 

Anterior part of head extending forward 

beyond disc (fig. 22); eyes located laterally 

on side of head (fig. 22) ...................... 15 

13. Disc very broad, width more than 1.5 

times length (fig. 23); tail very short and 

filamentous (fig. 23) .............................. 

...... Family Gymnuridae, Butterfly Rays 

fig. 23; 238 

Disc narrower, width less than 1.5 times 

length; tail much longer and thicker .... 

........................................................ 14 

14. Caudal fin present (fig. 24); no thorns or 

tubercles on disc or tail ......................... 

... Family Plesiobatidae, Giant Stingarees 

fig. 24; 180 

Caudal fin absent; central disc and dorsal 

surface of tail normally with some 

thorns or tubercles ................................. 

................... Family Dasyatidae, Stingrays 

a. No skin folds on tail; base of tail narrow 

and typically rounded to slightly 

compressed in cross-section (fig. 25) ..... b 

Skin folds present on undersurface of 

tail, and sometimes on dorsal surface 

after sting; base of tail relatively broad, 

distinctly depressed (fig. 26) .................. c 

b. No stinging spine; many long, sharp 

thorns over entire disc ............ Urogymnus 

fig. 27; 236 

One or more stinging spines on tail 

(if lost, scar visible); no long, sharp 

thorns present over sides of disc 

(sometimes with low thorns near 

midline of disc) .................... Himantura 

fig. 28; 190 

fig. 21 fig. 22 

fig. 23 

fig. 24 

fig. 25 fig. 26 

fig. 27 

fig. 28 

c. Disc oval or subcircular; ventral skin fold 

tall, extending to tail tip ..................... d


Disc more or less quadrangular; ventral 

skin fold terminating well before tail tip 

when undamaged ................................ e 

d. Disc oval (fig. 29); dorsal surface with 

numerous blue spots over a brownish 

background (fig. 29); tail with a blue 

stripe along each side ................ Taeniura 

fig. 29; 232 

Disc subcircular (fig. 30); dorsal surface 

with fine black and white mottling (fig. 

30); no blue spots or stripes on body or 

tail ......................................... Taeniurops 

fig. 30; 234 

e. Anterior margin of disc uniformly 

convex (fig. 31); ventral surface dark 

greyish ........................... Pteroplatytrygon 

fig. 31; 230 

Anterior margin of disc angular (fig. 32); 

ventral surface mostly pale .................... f 

f. Distance from cloaca to stinging spine 

exceeding half of disc width; ventral skin 

fold relatively tall, its maximum height 

equal to or exceeding spiracle length ...... 

............................................. Pastinachus 

fig. 32; 222 

Distance from cloaca to stinging spine 

less than half of disc width; ventral skin 

fold relatively low, its maximum height 

much less than spiracle length .............. g 

g. Tail banded black and white behind sting 

(fig. 33); blue-spotted with dark transverse 

band through eyes (fig. 33) ....... Neotrygon 

fig. 33; 220 

Tail not banded beyond sting (fig. 34); not 

blue-spotted and without dark transverse 

band through eyes (fig. 34) ......... Dasyatis 

fig. 34; 182 

15. A pair of long, paddle-like flaps 

situated laterally on front of head 

(fig. 35); teeth minute, in many rows 

................. Family Mobulidae, Devilrays 

fig. 30 

fig. 32 

fig. 34 

fig. 31 

fig. 29 

fig. 33


Mouth terminal, located at end of snout 

tip ............................................... Manta 

fig. 35; 256 

Mouth subterminal, located on undersurface 

of head .......................... Mobula 

fig. 36; 258 

fig. 35 

No lateral, paddle-like processes on head, 

instead with a pair of broadly rounded 

lobes (fig. 37) or a single, fleshy subrostral 

lobe (fig. 38); teeth large and plate-like, 

fewer than 10 rows in each jaw ......... 16 

16. Margin of subrostral lobe with a deep 

central notch (fig. 37); floor of mouth 

without papillae .................................... 

... Family Rhinopteridae, Cownose Rays 

fig. 37; 252 

Margin of subrostral lobe rounded (fig. 

38); floor of mouth with fleshy papillae 

.......... Family Myliobatidae, Eagle Rays 

Posterior margin of internasal flap 

deeply notched (inverted V-shape) (fig. 

39); teeth in a single row in each jaw 

............................................... Aetobatus 

fig. 38; 242 

Posterior margin of internasal flap almost 

straight or only undulating slightly 

(fig. 40); teeth in more than one row in 

each jaw ............................. Aetomylaeus 

fig. 41; 246 

17. A single dorsal fin (fig. 42); 6 or 7 gill 

openings on each side of head (fig. 42) 

............ Family Hexanchidae, Cowsharks 

fig. 42; 36 

fig. 36 

fig. 37 

fig. 38 

fig. 39 fig. 40 

fig. 41 

2 dorsal fins; 5 gill openings on each 

side of head ..................................... 18 

18. Anal fin absent ................................ 19 

Anal fin present, sometimes small or 

fig. 42


obscure (situated close to caudal fin) ... 20 

19. Teeth in both jaws similar in shape and 

size (fig. 44); no subterminal notch on 

caudal fin (fig. 43) ............................... 

................... Family Squalidae, Dogfishes 

fig. 43; 38 

Teeth in upper jaw differing in shape 

and/or size to those of lower jaw (fig. 45); 

strong subterminal notch present 

(fig. 46) ................................................. 

... Family Centrophoridae, Gulper Sharks 

fig. 46; 40 

20. Head very broad, hammer-shaped, with 

very widely separated eyes (fig. 47) ........ 

Family Sphyrnidae, Hammerhead Sharks 

Lateral blades of head very broad, narrow 

and wing-like; width of head close to 

half of total length ................... Eusphyra 

fig. 47; 134 

Lateral blades of head much shorter and 

not wing-like; width of head less than 

40% of total length .................... Sphyrna 

fig. 48; 136 

fig. 43 

fig. 44 fig. 45 

fig. 46 

fig. 47 

Head much narrower, not hammershaped 

.............................................. 21 

21. Upper caudal-fin lobe equal to or longer 

than half total length (fig. 49); body 

without spots or bands .......................... 

....... ..Family Alopiidae, Thresher Sharks 

fig. 49; 62 

Upper caudal-fin lobe much shorter 

than half total length (caudal fin also 

long in Zebra Sharks but their body 

spotted and/or banded) ................... 22 

22. Dorsal-fin spines present (fig. 50); medium 

to tall crest over eye (fig. 50) .................. 

fig. 48 

fig. 50 

fig. 49


....... Family Heterodontidae, Hornsharks 

fig. 50; 44 

Dorsal-fin spines absent; no crest over 

eye ................................................... 23 

23. Whole mouth forward of front margin 

of eye (fig. 51) ................................... 24 

Mouth partly beneath or behind front 

margin of eye (fig. 52) ....................... 28 

fig. 51 

fig. 52 

24. Mouth very broad, terminal; caudal fin 

forked (fig. 53) ....................................... 

... Family Rhincodontidae, Whale Sharks 

fig. 53; 60 

Mouth smaller, subterminal; caudal fin 

not forked ...................................... 25 

fig. 53 

25. No fleshy lobe or groove on outer 

margin of nostril (fig. 54) .................. 26 

Fleshy lobe and groove present on outer 

margin of nostril (fig. 55) ................ 27 

26. Caudal fin very long, about equal to 

trunk length (fig. 56); ridges present along 

sides of body (fig. 56) ........................... 

... Family Stegostomatidae, Zebra Sharks 

fig. 56; 58 

Caudal fin shorter, less than half trunk 

length (fig. 57); no ridges on body 

(fig. 57) ..... Family Ginglymostomatidae, 

Nurse Sharks 

fig. 57; 56 

fig. 57 

fig. 54 fig. 55 

fig. 56 

27. Head and body strongly depressed 

(fig. 58); skin flaps present on sides of 

head (fig. 58) ......................................... 

....... Family Orectolobidae, Wobbegongs 

fig. 58; 46 

Head and body cylindrical (fig. 59); no 

skin flaps on sides of head (fig. 59) ........ 

fig. 59 

fig. 58


.......................... Family Hemiscylliidae, 

Longtail Carpetsharks 

fig. 59; 48 

28. Caudal fin almost symmetrical, lunate 

(fig. 60); a strong keel on either side of 

caudal peduncle (fig. 60) ....................... 

........ Family Lamnidae, Mackerel Sharks 

fig. 60; 64 

Caudal fin asymmetrical, heterocercal 

(fig. 61); caudal-peduncle keel either low 

or absent (fig. 61) ............................... 29 

29. 1 st dorsal-fin origin behind pelvic-fin 

origin ... Family Scyliorhinidae, Catsharks 

fig. 60 

fig. 61 

a. Upper labial furrows very long, more 

than 3 times spiracle diameter, mostly 

extending to level of front of eye (fig. 62) 

........................................................... b 

Upper labial furrows much shorter or 

rudimentary, when present much less than 

3 times spiracle diameter, never extending 

to level of front of eye (fig. 63) ............ c 

b. Anterior nasal flaps greatly expanded, 

reaching mouth (fig. 65); nasoral grooves 

present ............................... Atelomycterus 

fig. 64; 68 

Anterior nasal flaps much shorter, smaller 

than nostrils, not reaching mouth (fig. 66); 

nasoral grooves absent ............. Apristurus 

fig. 67; 66 

c. 2 nd dorsal fin much smaller than 1 st 

dorsal fin (fig. 68); origin of 2 nd dorsal 

fin only slightly behind origin of anal fin 

(fig. 68); stomach inflatable .................... 

....................................... Cephaloscyllium 

fig. 68; 70 

Dorsal fins about equal in size (fig. 

69); origin of 2 nd dorsal fin behind 

midbase of anal fin (fig. 69); stomach not 

inflatable ................................ Halaelurus 

fig. 69; 72 

fig. 62 fig. 63 

fig. 64 

fig. 65 fig. 66 

fig. 67 

fig. 68 

fig. 69


1 st dorsal-fin origin well in front of 

pelvic-fin base .................................. 30 

30. Precaudal pits absent (fig. 71); anterior 

margin of caudal fin smooth (fig. 71) 

............ Family Triakidae, Hound Sharks 

fig. 70; 74 

Precaudal pits present (fig. 72); anterior 

margin of caudal fin usually rippled 

(fig. 72) ............................................ 31 

31. Spiracles present; posterior margin of 

2 nd dorsal fin deeply concave; intestine 

with spiral valves (fig. 73) ...................... 

...... Family Hemigaleidae, Weasel Sharks 

a. Lower teeth near symphysis with short, 

straight or weakly hooked cusps (fig. 74) 

that are concealed when mouth closed; 

gill slits small, less than twice eye length .. b 

Lower teeth near symphysis with long, 

strongly hooked cusps (fig. 75) that 

prominently protrude from mouth when 

closed; gill slits large, more than twice eye 

length ................................................. c 

b. Pelvic, dorsal and caudal fins not falcate 

(fig. 76); mouth long, narrowly arched; 

lower teeth near symphysis with erect 

cusps and slightly arched roots; cusplets 

present on lower teeth……….. Paragaleus 

fig. 76; 84 

Pelvic, dorsal and caudal fins strongly 

falcate (fig. 77); mouth very short, broadly 

arched; lower teeth near symphysis with 

erect cusps and highly arched roots; no 

cusplets on lower teeth ........... Hemigaleus 

fig. 77; 80 

c. Snout obtusely wedge-shaped in dorsoventral 

view; fins not falcate, posterior 

margins of pelvic and pectoral fins 

straight or slightly concave (fig. 78) ....... 

.......................................... Chaenogaleus 

fig. 78; 78 

fig. 70 

fig. 71 fig. 72 

fig. 73 

fig. 74 fig. 75 

fig. 76 

fig. 77 

fig. 78


Snout bluntly rounded in dorsoventral 

view; fins strongly falcate, posterior 

margins of pelvic and pectoral fins deeply 

concave (fig. 79) ................... Hemipristis 

fig. 79; 82 

Spiracles absent (except in Tiger Shark); 

posterior margin of 2 nd dorsal fin not 

deeply concave; intestine with a scroll 

valve (fig. 80) ........................................ 

... Family Carcharhinidae, Whaler Sharks 

a. Caudal peduncle with lateral keels 

(fig. 81); spiracles present; upper labial 

furrows, reaching forward to front of eyes; 

teeth cockscomb-shaped (fig. 82)........... 

.............................................. Galeocerdo 

fig. 83; 114 

Caudal peduncle without lateral keels 

(weak ones present in Prionace); spiracles 

absent; upper labial furrows short, never 

reaching forward to in front of eyes; 

teeth not cockscomb-shaped .............. b 

b. 2 nd dorsal fin half or more height of 1 st 

dorsal fin ............................................ c 

2 nd dorsal fin less than half height of 1 st 

dorsal fin ............................................ e 

c. 1 st dorsal and upper caudal fin with 

distinct white tips (fig. 84); teeth with a 

single cusp and lateral cusplets (fig. 85) ... 

.............................................. Triaenodon 

fig. 84; 132 

1 st dorsal and upper caudal fin without 

obvious white tips; teeth with a single 

cusp and no lateral cusplets (fig. 86) ...... d 

d. 2 nd dorsal fin nearly as large as 1 st dorsal 

fin (fig. 87) ............................... Lamiopsis 

fig. 87; 120 

2 nd dorsal fin half to two thirds height of 

1 st dorsal fin (fig. 88) .................. Glyphis 

fig. 88; 116 

e. Head greatly depressed and trowelshaped 

(fig. 89); pectoral fins triangular 

fig. 79 

fig. 80 

fig. 81 

fig. 83 

fig. 84 

fig. 87 

fig. 88 

fig. 82 

fig. 85 fig. 86


(fig. 89); 1 st dorsal-fin free rear tip 

almost over mid-base of pelvic fin 

(fig. 89) .................................. Scoliodon 

fig. 89; 130 

Head varying from conical to slightly 

depressed; pectoral fins narrower; 1 st 

dorsal-fin free rear tip over or slightly 

anterior to pelvic-fin origin ................. f 

f. 2 nd dorsal-fin origin well behind anal-fin 

origin (usually over or just anterior to analfin 

insertion) (fig. 90); posterior margin 

of anal fin nearly straight or shallowly 

concave (fig. 90); preanal ridges very 

long, subequal to anal-fin base length ... g 

2 nd dorsal-fin origin usually about 

opposite anal-fin origin (sometimes more 

posterior but always well anterior of analfin 

insertion) (fig. 91); posterior margin 

of anal fin deeply concave or deeply 

notched (fig. 91); preanal ridges short, 

about half anal-fin base or less .............. h 

h. Notch present at posterior edge of eye 

(fig. 93); 1 st dorsal-fin origin posterior 

to adpressed pectoral-fin free rear tips 

by a distance exceeding 4 th gill opening 

height (fig. 92) ......................... Loxodon 

fig. 92; 122 

Posterior rim of eye without a notch (fig. 

94); 1 st dorsal-fin origin over or only just 

posterior to adpressed pectoral-fin free 

rear tips (fig. 95) …….… Rhizoprionodon 

fig. 95; 126 

i. 1 st dorsal-fin base much closer to pelvicfin 

than pectoral-fin bases (fig. 96); 

colour brilliant dark blue above; weak keels 

present on caudal peduncle ......... Prionace 

fig. 96; 124 

1 st dorsal-fin base equidistant between 

pectoral and pelvic-fin bases (fig. 97); 

colour light to dark greyish, greyish brown 

or brownish above; no keels present on 

caudal peduncle .................. Carcharhinus 

fig. 97; 86 

fig. 89 

fig. 92 

fig. 93 

fig. 95 

fig. 96 

fig. 97 

fig. 90 

fig. 91 

fig. 94

SPECIES INFORMATION


Bluntnose Sixgill Shark 

Hexanchus griseus (Bonnaterre, 1788) 

2 1 

3 4 

Other names: Sixgill Shark, Bull Shark, Mud Shark, Cow Shark (English), Cucut 

Meong, Hiu Tahu Putih (Indonesian), Yu Insang Enam (Malaysian) 

Identifying features: 

1 6 gill slits 

2 small, greenish eye 

3 single dorsal fin 

4 dorsal fin usually plain (occasionally with narrow white posterior margin) 

5 snout broadly rounded (viewed ventrally) 

6 lower jaw with 6 rows of large, comb-like teeth 

Size: To at least 480 cm; males and females mature at 285–315 and 420 cm respectively; 

born at 65–70 cm. 

Distribution: Almost circumglobal in tropical and temperate waters. 

Habitat and biology: Adults normally found near the bottom in deepwater to at 

least 2500 m; young sometimes enter shallow coastal bays. Viviparous, with yolk-sac 

dependency; gives birth to litters of 47–108 pups. Diet includes bony fishes, elasmobranchs, 

cephalopods and crustaceans, and adults also feed on larger bony fishes (e.g. 

billfishes), cetaceans and seals. 

Utilisation: Caught rarely so of minimal commercial value. 

Conservation status: IUCN Red List: Near Threatened. 

Local synonyms: None. 

References: Ebert (2003); White & Dharmadi (2010).

H e x a n c h i f o r m e s 37 

5 

Ventral head 

Upper tooth 

In Borneo: Probably widespread, but 

known only from near Layang-layang, off 

western Sabah. 

Hexanchidae (Cowsharks) 

Lower tooth 

6


Western Highfin Spurdog 

Squalus altipinnis Last, White & Stevens, 2007 

4 

1 

5 4 3 

7 

Other names: None (English), none (Indonesian), Yu Minyak (Malaysian) 


1 pectoral-fin free rear tip not elongate 

2 snout narrowly pointed (viewed ventrally) 

3 2 nd dorsal fin relatively tall, upright 

4 dorsal fins with thick spines 

5 origin of 2 nd dorsal fin well posterior to pelvic-fin free rear tips 

6 dermal denticle crowns elevated on pedicels, broad, with 1 or 3 cusps 

7 posterior margin of caudal fin pale 

Size: To at least 71.5 cm; males probably smaller, mature at 54 cm; birth size unknown 

(late term embryos 19 cm long). 

Distribution: Possibly widespread in the Indo-Malay Archipelago, confirmed from 

northwestern Australia, Borneo, the Philippines and possibly Indonesia. 

Habitat and biology: Occurs near the bottom on the outer continental shelf and 

upper slope in 130–300 m. Viviparous, with yolk-sac dependency. Diet probably consists 

of small bony fishes, cephalopods and crustaceans. 

Utilisation: Caught occasionally by demersal longline fisheries and lobster pots operating 

in deepwater areas. Used for its meat when fresh, otherwise discarded. 

Conservation status: IUCN Red List: Data Deficient. 

Local synonyms: Squalus sp. 1: Compagno et al. (2005b). 

References: Last et al. (2007).

S q u a l i f o r m e s 39 

2 

Ventral head 

Late-term embryo (19 cm) 

In Borneo: Northern Borneo, from 

Sabah and northern Sarawak. 

7 

Squalidae (Dogfishes) 

6 

Flank denticle


Smallfin Gulper Shark 

Centrophorus moluccensis Bleeker, 1860 

3 

4 

3 

1 

Other names: Endeavour Dogfish (English), Hiu Botol Danten, Hiu Taji, Hiu Senget, 

Cucut Botol (Indonesian), Yu Minyak (Malaysian) 


1 pectoral-fin free rear tip greatly extended 

2 snout relatively short and broadly rounded (when viewed ventrally) 

3 2 nd dorsal fin relatively small, half or less height of 1 st dorsal; both fins with short 

spines 

4 origin of 2 nd dorsal fin well posterior to pelvic-fin free tips 

5 dermal denticles flat and sessile, not elevated on pedicels 

6 teeth of upper and lower jaws differing in shape 

Size: To at least 100 cm; males and females mature at about 70 and 85 cm respectively; 

born at about 33 cm. 

Distribution: Widespread in the Indo–West Pacific, from South Africa to Japan and 

Australia. 

Habitat and biology: Occurs near the bottom on the outer continental shelf and 

upper slope at depths of 125–820 m. Viviparous, with yolk-sac dependency; gives 

birth to litters of two pups every two years; gestation period unknown. Diet consists 

primarily of bony fishes and cephalopods, but also elasmobranchs and crustaceans. 

Utilisation: Caught infrequently by demersal longline in deepwater. Used for its meat 

and fins (low value), and liver oil (very high value). 



References: White & Dharmadi (2010).

S q u a l i f o r m e s 41 

2 

Ventral head 

Patch of flank denticles 

5 

In Borneo: Sparse records from off 

Sarawak and eastern Sabah. 

Upper teeth 

Lower teeth 

6 

Centrophoridae (Gulper Sharks)


Ocellated Angelshark 

Squatina tergocellatoides Chen, 1963 

1 

6 

3 

5 

8 8 

8 

7 

Other names: None (English), Hiu Kodok (Indonesian), Yu Pari (Malaysian) 


1 pectoral fins large, broad and angular 

2 interorbital space concave 

3 no enlarged predorsal thorns 

4 nasal barbels and anterior nasal flaps moderately expanded and finely fringed 

5 1 st dorsal-fin origin behind pelvic-fin free rear tips 

6 dorsal colour pale yellowish brown 

7 several pairs of dark blotches or ocelli on pectoral and pelvic fins, and on tail 

8 dorsal fins and upper caudal-fin lobe with dark bases 

Size: To at least 101 cm. 

Distribution: Known only from the Taiwan Straits and off Borneo. 

Habitat and biology: A poorly known, demersal shark found on the outer 

continental shelf (~160 m depth off Sarawak). Viviparous, with yolk-sac dependency 

(lecithotrophic); one pregnant female (~100 cm) contained at least 3 mid-term embryos 

(~14 cm). Diet unknown, probably consists of fishes and crustaceans. Angelsharks are 

swift ambush predators that use their flexible necks and protruding jaws to snatch prey.

S q u a t i n i f o r m e s 43 

Utilisation: Caught opportunistically rather than being targeted. Used for its meat. Fins 

of no commercial value. 

Conservation status: IUCN Red List: Vulnerable. 


References: None. 

Ventral head and abdomen 

2 

In Borneo: Off western Sabah and 

Sarawak. 

Squatinidae (Angelsharks) 

Front view of head 

4 

Anterior nasal flap (from front)


Zebra Hornshark 

Heterodontus zebra (Gray, 1831) 

2 

1 

3 

1 

Other names: Zebra Bullhead Shark, Zebra Port Jackson Shark, Bullhead Shark 

(English), Hiu (Indonesian), Yu Kebut (Malaysian) 


1 dorsal fins preceded by strong spines 

2 head large and blunt with low crests over eyes 

3 pale with numerous dark, narrow bands 

4 nostrils connected to mouth by a groove 

5 mouth small, near snout tip 

6 anterior teeth small, pointed, multicuspid 

7 posterior teeth molar-like with medial ridges 

Size: To at least 122 cm; males mature from about 61 cm; hatch at about 15 cm. 

Distribution: Western Pacific, from Japan southwards through the Philippines, Borneo 

and Indonesia to northern Australia. 

Habitat and biology: Common, but poorly known; continental and insular shelves 

mostly down to about 50 m, but deeper off Australia (150–200 m). Biology poorly 

known; oviparous, with large, screw-shaped egg cases that are presumably lodged into 

rocky crevices during embryonic development. Diet unknown, probably consists of 

benthic invertebrates such as sea urchins and crustaceans. 

Utilisation: Rarely caught and of little commercial value. Meat sold as fillets; fins have 

no value.

H e t e r o d o n t i f o r m e s 45 

Conservation status: IUCN Red List: Least Concern. 

Local synonyms: Cestracion amboinensis Regan, 1906. 


4 

5 

Ventral head 

6 7 

In Borneo: Northern Borneo, including 

Sarawak and Sabah. 

Heterodontidae (Hornsharks) 

Anterior upper tooth 

Posterior upper tooth


Indonesian Wobbegong 

Orectolobus leptolineatus Last, Pogonoski & White, 2010 

1 

2 

3 

4 

5 

Other names: Indo Wobbegong (English), Hiu Kodok, Hiu Lepang, Hiu Jenggot 

(Indonesian), Yu Misai (Malaysian) 


1 body strongly depressed 

2 nasal barbels branched 

3 dermal lobes present on side of head 

4 elaborate variegated pattern of dark saddles, and pale spots and fine reticulations 

5 anal fin present 

Size: To at least 120 cm; males mature at 85–88 cm and females by 94 cm. 

Distribution: Known from Indonesia, Borneo, the Philippines and Taiwan. 

Habitat and biology: Habitat and biology not well known; demersal on insular and 

continental shelves. Viviparous, with yolk-sac dependency; one pregnant female 

contained at least 4 mid-term embryos. Diet probably consists of demersal invertebrates 

and small fishes as with other wobbegong species. 

Utilisation: Not recorded during market surveys thus utilisation in the area not known 

but likely very minimal. 

Conservation status: IUCN Red List: Not Evaluated. 

Local synonyms: Orectolobus cf ornatus (De Vis): Compagno et al. (2005b), White et al. 

(2006b); Orectolobus maculatus (Bonnaterre): Pickell & Siagian (2000), Yano et al. (2005). 

References: Last et al. (2010c).

O r e c t o l o b i f o r m e s 47 

2 

3 

Ventral head 

Embryo (14 cm) 

In Borneo: Known from a few specimens 

collected off southeastern Sabah 

(Semporna) and Sarawak. 

Orectolobidae (Wobbegongs)


Indonesian Bambooshark 

Chiloscyllium hasseltii Bleeker, 1852 

1 

6 

3 

4 

4 

1 

5 

2 

Other names: Hasselt’s Bambooshark (English), Hiu Tekek (Indonesian), Yu-cicak 

Lampai, Yu Bodoh (Malaysian) 


1 body and tail relatively stout 

2 base of anal fin shorter than lower caudal-fin lobe 

3 no skin ridges on trunk 

4 posterior margins of dorsal fins straight to convex 

5 anal-fin origin close behind free rear tip of 2 nd dorsal fin 

6 adults lacking a colour pattern 

7 juveniles with black-edged saddle markings and blackish blotches on fins 

Size: To at least 78 cm; males mature at about 62 cm; hatch at about 9–12 cm. 

Distribution: Tropical Indo–West Pacific, from Myanmar and Vietnam to Indonesia 

and New Guinea. 

Habitat and biology: Little known, bottom-dwelling species, found inshore to 

depths to at least 12 m. Oviparous, depositing egg cases on benthic marine plants, with 

hatching occurring in December. Diet unknown, but presumably consists of bottomdwelling 

invertebrates, and possibly small fishes. 

Utilisation: Caught occasionally by gillnet, longline and trawl fisheries. Used for its 

meat. 


Local synonyms: Chiloscyllium obscurum Gray, 1851; Chiloscyllium dolganovi Kharin, 

1987; Chiloscyllium indicum var. obscura Günther, 1870; Chiloscyllium hasselti Bleeker: 

Compagno (2001), Compagno et al. (2005a), Yano et al. (2005). 

References: Compagno (2001).


Ventral head 

In Borneo: Central western Borneo, 

including northwestern Kalimantan and 

Sarawak. 

6 

Hemiscylliidae (Longtail Carpetsharks) 

Dorsal head and pectoral fins 

7 

Juvenile (34.5 cm)


Slender Bambooshark 

Chiloscyllium indicum (Gmelin, 1789) 

6 

1 

3 

4 

1 

4 

5 

2 

Other names: Ridgeback Bambooshark, Ridgeback Shark (English), Hiu Bongo, 

Cucut Dolok, Hiu Tekek (Indonesian), Yu-cicak Tembaga, Yu Bodoh, Yu Tokeh, Yu Pasir 

(Malaysian) 


1 body and tail very slender 

2 bases of anal fin and lower caudal-fin lobe about equal in length 

3 lateral skin ridges present on trunk 


5 anal-fin origin well behind free rear tip of 2 nd dorsal fin 

6 adults usually with numerous dark spots and blotches (no bands); spots barely evident 

in largest individuals 

7 juveniles more intensely spotted with black-edged saddles 


free-swimming at 13 cm. 

Distribution: Tropical Indo–West Pacific, from Sri Lanka to Indonesia; possibly from 

the Arabian Sea to the Solomon Islands, and north to Japan. 

Habitat and biology: A common, but little known, bottom-dwelling species 

occurring inshore; often enters freshwater and brackish areas. Oviparous, depositing 

small oval egg cases. Diet unknown but presumably dominated by small invertebrates. 

Three colour morphs exist in the environs of the Kapuas Estuary but, based on molecular 

analyses, these appear to be a single species. 

Utilisation: Caught often at some inshore localities using demersal gillnets and trawls. 

Used mainly for its meat, but of limited value due to its small size. 


Local synonyms: Chiloscyllium phymatodes Bleeker, 1852. 

References: Compagno (2001).


Ventral head 

6 

In Borneo: Patchy; recorded from 

southern and western Kalimantan and 

Sarawak. 

3 



7 

Juvenile (19 cm)


Whitespotted Bambooshark 

Chiloscyllium plagiosum (Bennett, 1830) 

1 

6 

3 

4 

1 

4 

5 

2 

Other names: None (English), Hiu Bongo, Cucut Dolok, Hiu Tekek (Indonesian), Yucicak 

Bintik, Yu Bodoh (Malaysian) 


1 body and tail relatively slender 

2 base of anal-fin much shorter than base of lower caudal-fin lobe 

3 weak or obscure lateral skin ridges on trunk 


5 anal-fin origin well behind free rear tip of 2 nd dorsal fin 

6 dark bands and spots interspersed with pale blotches 

Size: To about 95 cm; males mature at 50–63 cm; hatch at 10–13 cm. 

Distribution: Tropical Indo–West Pacific, from Madagascar through to Indonesia, and 

north to Japan. 

Habitat and biology: A common inshore, bottom-dwelling shark found on coral 

reefs, in tidal pools, and amongst rock crevices. Oviparous, depositing oval egg cases 

(~8 cm long); in the northwestern Pacific, hatching occurs from June to August. Both 

sexes mature at ~4 years of age in Taiwan populations. Nocturnal, diet consists primarily 

of bottom-dwelling invertebrates and small fishes. 

Utilisation: Caught rarely by demersal gillnet and trawl fisheries operating inshore. 

Used mainly for its meat, but of limited value due to its small size. The meat is processed 

(fish ball, fish cake), and now sold as fillets in Kota Kinabalu and Sandakan fish markets. 


Local synonyms: Chiloscyllium plagiosum var. interruptum Bleeker, 1852; Chiloscyllium 

margaritiferum Bleeker, 1863. 

References: Compagno (2001); Chen et al. (2007).


Ventral head 

In Borneo: South China Sea, from 

northwestern Kalimantan to northern 

Sabah. 

6 



6 

Juvenile (13 cm)


Brownbanded Bambooshark 

Chiloscyllium punctatum Müller & Henle, 1838 

1 

4 

4 

6 

3 

1 

5 

2 

Other names: Grey Carpetshark, Brownbanded Catshark, Spotted Catshark (English), 

Hiu Bongo, Hiu Batu, Hiu Gedok, Hiu Tekek (Indonesian), Yu-cicak Insang Putih,Yu 

Bodoh, Yu Tokeh, Yu Punai (Malaysian) 


1 body and tail moderately slender 

2 base of anal-fin much shorter than base of lower caudal-fin lobe 

3 no skin ridges on trunk 

4 posterior margins of dorsal fins straight or distinctly concave 

5 anal-fin origin over or just behind free rear tip of 2 nd dorsal fin 

6 adults plain coloured or with faint brownish bands 

7 juveniles with dark transverse bands usually with a scattering of small dark spots 

Size: To at least 132 cm (144 cm in captivity); males and females mature at 67–72 cm 

and by 68 cm respectively; hatch at 13–18 cm. 

Distribution: Tropical Indo–West Pacific, from the east coast of India to Australia and 

New Guinea, and north to Japan. 

Habitat and biology: A very common, inshore bottom-dwelling shark living in tidal 

pools on coral reefs, on muddy banks, and amongst mangroves and seagrasses; from the 

intertidal zone to a depth of 85 m or more. Oviparous, depositing rounded egg cases 

(11 by 5 cm); in captivity, hatching occurs between 90 and 153 days afterward. Diet 

consists primarily of bottom-dwelling invertebrates and small fishes. Can survive for 

many hours out of water. 

Utilisation: Caught often by demersal gillnet, longline and trawl fisheries operating in 

insular and continental shelf waters. Used for its meat. 

Conservation status: IUCN Red List: Near Threatened.



References: Compagno (2001). 

Ventral head 

In Borneo: Southern and western 

Kalimantan, Sarawak and Sabah. 



7 

Sub-adult (51 cm)


Tawny Nurse Shark 

Nebrius ferrugineus (Lesson, 1830) 

3 

5 

4 

4 

1 

Other names: Tawny Shark, Spitting Shark, Sleepy Shark, Madame X (English), Hiu 

Gedebong, Hiu Gedok, Hiu Bisu (Indonesian), Yu Semilang, Yu Bodoh (Malaysian) 


1 caudal fin elongate, asymmetrical, its length about one third of total length 

2 nostrils near tip of snout, with short barbels and nasoral grooves 

3 spiracle much smaller than eye 

4 dorsal fins similar in size, tips angular, close to tail 

5 body colour plain, yellowish to greyish brown 



Distribution: Widespread throughout the Indo–West and Central Pacific. 

Habitat and biology: A demersal shark living on coral and rocky reefs, and over 

sand flats, from the intertidal zone to at least 70 m depth. Viviparous, with yolk-sac 

dependency (in Japan, reported to be oophagous with small litters); retains up to 32 

egg cases that hatch inside the mother, giving birth to live pups. Sluggish and mostly 

nocturnal, usually resting in small groups near caves and rocky crevices during the day. 

Diet includes a variety of demersal invertebrates as well as small reef fishes; uses its 

pharynx as a powerful suction pump to extract prey from rocky crevices. 

Utilisation: Not recorded during market surveys thus utilisation in the area not known. 


Local synonyms: Ginglymostoma rueppellii Bleeker, 1852; Nebrius concolor Rüppell 1837. 

References: None.


2 

Upper tooth 

Lower tooth 

Ventral head 

4 

In Borneo: Uncommon, not observed in 

surveys, but known to occur on coral reefs 

in the Sulu Sea (Sipadan Islands) and Perak 

(Peninsular Malaysia). Unverified record 

from dive operator in Kota Kinabalu. 

4 

Ginglymostomatidae (Nurse Sharks) 

Lateral precaudal tail region


Zebra Shark 

Stegostoma fasciatum (Hermann, 1783) 

5 

1 

4 

3 

Other names: Leopard Shark (English), Hiu Belimbing, Kluyu Blimbingan 

(Indonesian), Yu Rimau, Yu Kebut, Yu Cicak, Yu Tokeh (Malaysian) 


1 colour pattern yellowish brown with numerous, dark brown spots in adults 

2 dark brown with vertical white bars and spots in juveniles (smaller than 70 cm) 

3 caudal fin very long, blade-like 

4 prominent ridges on dorsal surface and sides 

5 spiracle similar in size to eye 

Size: To at least 235 cm (reports of 354 cm need validation); males and females mature 

at about 145–185 and 170 cm respectively; hatch at about 20–36 cm. 

Distribution: Tropical Indo–West Pacific, from southeastern Africa to New Caledonia 

and Japan. 

Habitat and biology: A common inshore, demersal shark found over coral reefs and 

soft bottoms in shallow waters to at least 40 m. Oviparous; their large egg cases (13–17 

cm long, 8 cm wide), which are dark brown to purplish black with longitudinal striations, 

are anchored to the substrate by lateral masses of hair-like fibres. Diet consists 

primarily of gastropods and bivalves, but also crustaceans and small fishes. Active 

mostly at night, resting on the bottom during day. 

Utilisation: Used primarily for its meat and skin (exported to Thailand for leather). 

Protected under the Malaysian Fisheries Act, so catches are opportunistic and landings 

not reported. 


Local synonyms: Squalus pantherinus Kuhl & van Hasselt in Bleeker, 1852 (name


mentioned in synonmy with S. fasciatum); Scyllia quinquecornuatum van Hasselt, 1823; 

Squalus tigrinus Forster, 1781; Stegostoma varium Garman, 1913. 


Ventral head 

Lower tooth 

Upper tooth 

In Borneo: Western and northern 

Borneo, including Sarawak, Sabah and 

north-west and north-east Kalimantan. 

Stegostomatidae (Zebra Sharks) 

2 

Juvenile (49.5 cm)


Whale Shark 

Rhincodon typus Smith, 1828 

1 

6 

5 

4 

3 

Other names: None (English), Hiu Paus (Indonesian), Yu Paus (Malaysian) 


1 head very broad and flattened 

2 adults huge, attaining 12 m or more in length 

3 mouth very wide, almost terminal on head 

4 caudal peduncle depressed, with strong fleshy keels on sides 

5 prominent ridges on dorsal surface and sides 

6 dorsal and lateral surfaces with numerous creamy white spots between pale bars and 

stripes 

Size: To at least 12 m, but recent studies suggest lengths of up to 18 m; males and 

females mature at 300–400 and >760 cm respectively; born at about 40–64 cm. 

Distribution: Cosmopolitan in all tropical and warm temperate seas. 

Habitat and biology: The whale shark is the largest living fish. Epipelagic, oceanic 

and coastal, sometimes venturing into atoll lagoons; highly migratory, occurring 

singly or in large aggregations. Viviparous, with yolk-sac dependency, although was long 

thought to be oviparous as it retains egg cases in utero until pups hatch; a pregnant female 

caught off Taiwan had 300 pups nearing full term. Diet includes a variety of planktonic 

and nektonic prey. Ecotourism ventures have been established in many countries to take 

advantage of some of the predictable aggregations. 

Utilisation: Caught irregularly, and somewhat opportunistically, by traditional fishers 

using gillnets and harpoons. Used for its fins (extremely valuable as a display item) and 

meat. Population declines due to overfishing reported from the Philippines, Taiwan, 

Maldives and India. 

Conservation status: IUCN Red List: Vulnerable; Appendix II of CITES (to ensure 

that international trade is sustainable) and Appendix II of the Convention on Migratory 

Species (to encourage international management).


Local synonyms: Rhiniodon typus Smith, 1829. 


1 

3 

Upper tooth 

In Borneo: Recorded from northern 

and western Sabah and northwestern 

Kalimantan, but probably more widely 

distributed. Seasonal in Sabah, usually 

from February to April. 

Rhincodontidae (Whale Sharks) 

Lower tooth


Pelagic Thresher 

Alopias pelagicus Nakamura, 1935 

1 

2 

4 

5 

3 

Other names: None (English), Hiu Monyet, Hiu Tikus, Cucut Pedang, Tikusan 

(Indonesian), Yu Ekor Panjang (Malaysian) 


1 upper lobe of caudal fin nearly as long as rest of body 

2 head profile arched between eyes, no deep grooves on nape 

3 1 st dorsal fin closer to pectoral-fin rear tip than pelvic-fin base 

4 eye relatively large, almost central on side of head 

5 white part of belly not extending over pectoral-fin base 

Size: To at least 390 cm; males and females mature at about 245–270 and 265–290 cm 

respectively; born at about 130–160 cm. 

Distribution: Widespread in the tropical and subtropical Indo–Pacific. 

Habitat and biology: Oceanic, usually well offshore, from the surface to at least 

150 m depth. Viviparous, with oophagy, but no evidence of adelphophagy; gives birth to 

litters of two (one per uterus) after an unknown gestation period; no reproductive 

seasonality with females giving birth throughout the year. Males and females mature 

at 7–8 and 8–9 years respectively. Diet poorly known but presumably consists of small 

fishes and cephalopods. Possibly able to maintain a body temperature warmer than the 

surrounding seawater.

L a m n i f o r m e s 63 

Utilisation: Caught by the tuna and shark longline, and tuna drift net fisheries in deep 

oceanic waters. Used for its fins (high value), meat (salted), skins and cartilage. 



References: White (2007b). 

Ventral head 

In Borneo: Uncommon, South China 

Sea off Sabah and also the Natuna Islands 

off Sarawak; possibly more widespread. 

Alopiidae (Thresher Sharks) 

Upper tooth 

Lower tooth


Shortfin Mako 

Isurus oxyrinchus Rafinesque, 1810 

3 

1 

2 

Other names: Mako Shark, Blue Pointer, Mackerel Shark, Snapper Shark (English), 

Hiu Tenggiri, Hiu Anjing, Hiu Mako, Hiu Kakap (Indonesian), Jerung-mako Sirip 

Pendek, Yu Jerung (Malaysian) 


1 large lateral keel on either side of the caudal peduncle 

2 pectoral fins short, their length less than head length 

3 eye relatively small 

4 snout sharply pointed (viewed ventrally), its undersurface white 

5 long slender, pointed teeth (smooth edged and curved with bent tips) protruding 

from mouth 


respectively; born at about 60–70 cm. 

Distribution: Cosmopolitan in all tropical and temperate seas. 

Habitat and biology: Epipelagic and oceanic, from the surface to at least 650 m 

depth; occasionally found close inshore. Viviparous, with oophagy and possibly 

adelphophagy; gives birth to litters of 10–18 pups (maximum 25) after a 15–18 month 

gestation period, with a 3 year reproductive cycle. Males and females mature at 8 and 18 

years respectively. Diet consists primarily of small to very large fishes, including 

elasmobranchs, cephalopods and occasionally marine mammals. Potentially 

dangerous to humans, known to attack boats; considered to be the fastest swimming 

shark.

L a m n i f o r m e s 65 

Utilisation: Caught occasionally by tuna and shark longline fisheries. Used for its high 

quality meat, fins (high value in adults), jaws (highly prized), skin and cartilage. 


Local synonyms: Oxyrhina glauca Müller & Henle, 1839. 

References: White (2007b). 

4 

5 

Ventral head 

In Borneo: Only recorded off Sabah, but 

almost certainly more widespread. 

Lamnidae (Mackerel Sharks) 

Upper tooth 

5 

Lower tooth


Bigfin Catshark 

Apristurus platyrhynchus (Tanaka, 1909) 

6 

5 

4 

3 

2 1 

Other names: Spatulasnout Catshark (English), none (Indonesian), none (Malaysian) 


1 anal fin low and very long 

2 pectoral fins large, reaching almost to pelvic fins 

3 2 nd dorsal fin much larger than 1 st dorsal fin 

4 uniformly greyish to brownish 

5 gill slits and naked areas on fins black 

6 snout moderately long and flattened 

7 labial furrows very long 

8 anterior nasal flaps not greatly expanded, not reaching mouth and smaller than 

nostrils 

Size: To at least 71 cm; both sexes mature by about 60 cm. 

Distribution: Patchy in the Western Pacific, from Australia (including the Norfolk 

Ridge), Borneo, Philippines and Japan. 

Habitat and biology: A deepwater catshark living near the bottom on the continental 

slope at depths of 400–1080 m. Oviparous, but biology and diet unknown, presumably 

consisting of small invertebrates and bony fishes. 

Utilisation: Unknown; rarely encountered. 


Local synonyms: Pentanchus verweyi Fowler, 1934; Apristurus verweyi (Fowler). 

References: Kawauchi et al. (2008).

C a r c h a r h i n i f o r m e s 67 

8 

7 

Ventral head 

In Borneo: Known only from off 

Sipadan Island, in the Celebes Sea 

(eastern Sabah). 

Scyliorhinidae (Catsharks) 

Flank denticles


Coral Catshark 

Atelomycterus marmoratus (Bennett, 1830) 

1 

1 

3 

2 

Other names: None (English), Hiu Tokek, Cucut Tokek (Indonesian), Yu Cicak, Yu 

Tokeh, Yu Pendek (Malaysian) 


1 dorsal fins large, subequal in size, angled rearwards, with obvious white tips 

2 covered with numerous light grey and white spots; saddles obsolete 

3 prominent white stripe on side of head through gill slits 

4 labial furrows very long 

5 anterior nasal flaps greatly enlarged, extending to mouth 

Size: To at least 70 cm; both sexes mature at about 47–55 cm. 

Distribution: Tropical Indo–West Pacific, from Pakistan to New Guinea and southern 

China and Taiwan. 

Habitat and biology: A common, inshore catshark found in crevices and holes on 

rocky reefs. Oviparous, laying pairs of egg cases onto the substrate; egg cases elongate 

(6–8 cm long, 2 cm wide), brown, with two constricted waists and tendrils only at one 

end. Diet unknown, but presumably dominated by small invertebrates. 

Utilisation: Caught occasionally by fishers operating over coral reefs; probably 

adversely affected by dynamite fishing. Utilised for its meat but of limited value due to 

its small size; its fins have no value. 


Local synonyms: Scyllium marmoratum Bennett, 1830. 

References: Bor et al. (2003); White (2007a).


5 

4 

Ventral head 

In Borneo: Coastal waters of Sabah and 

Sarawak. 


Dorsal head and pectoral fins


Sarawak Swellshark 

Cephaloscyllium sarawakensis Yano, Ahmad & Gambang, 2005 

6 

2 

1 

3 

Other names: Sarawak Pygmy Swell Shark, Sarawak Pygmy Swellshark (English), none 

(Indonesian), Yu-buntal Sarawak, Yu Buncit, Yu Kembong, Yu Buntal (Malaysian) 


1 2 nd dorsal fin much smaller than 1 st dorsal fin, its origin only slightly behind analfin 

origin 

2 stomach inflatable 

3 no crest of enlarged denticles along dorsal caudal-fin margin 

4 no labial furrows 

5 anterior nasal flaps short, not reaching mouth 

6 colour pattern relatively simple, consisting of narrow dark saddles and large blotches 

on sides 

Size: To at least 48 cm; females and males appear to be mature by 32 and 35 cm 

respectively. 

Distribution: Tropical northwestern Pacific, from Borneo to Hainan Island, southern 

China. 

Habitat and biology: A recently described, demersal species found on the outer shelf 

at depths of about 120–165 m. Oviparous, depositing smooth, oval-shaped egg cases 

9–10 cm long by ~3 cm wide. Diet unknown, probably mainly small invertebrates and 

fishes. Stomach inflatable, filling with air or water if threatened. 

Utilisation: Bycatch of trawl fisheries but of limited value for food. 


Local synonyms: Cephaloscyllium circulopullum Yano, Ahmad & Gambang, 2005; 

Cephaloscyllium parvum Inoue & Nakaya, 2006. 

References: Schaaf-Da Silva & Ebert (2008).


5 

4 

Ventral head 

Egg case 

In Borneo: South China Sea, off Sarawak. 


Dorsal head and pectoral fins


Indonesian Speckled Catshark 

Halaelurus maculosus White, Last & Stevens, 2007 

2 

1 

1 

3 4 

Other names: None (English), Hiu Tokek (Indonesian), Yu Bodoh (Malaysian) 


1 dorsal fins of similar size 

2 clusters of small blackish spots on and between brownish saddles 

3 whitish ventrally 

4 pelvic-fin inner margins of adult males not joined above claspers 

5 upper labial furrows barely noticeable 

6 anterior nasal flaps short, not reaching mouth 

Size: To at least 53 cm; males mature at 36–40 cm. 

Distribution: Known from Indonesia, the Philippines and Malaysia. 

Habitat and biology: A recently described, demersal species; probably reef-dwelling 

on the outer continental shelf. Biology poorly known; oviparous, females depositing 6– 

12 oval-shaped egg cases. Diet unknown, but probably dominated by small invertebrates 

and fishes. 

Utilisation: Bycatch of trawl fisheries, used for its meat but of limited value due to its 

small size. 


Local synonyms: Halaelurus cf. buergeri (Müller & Henle): White et al. (2006b); 

Halaelurus buergeri (Müller & Henle): Yano et al. (2005); Halaelurus sp. 1: White (2007a). 

References: White (2007a); White et al. (2007).


6 

5 

Ventral head 

Egg case 


Sarawak. 


Dorsal head and anterior trunk


Starspotted Smoothhound 

Mustelus manazo Bleeker, 1854 

5 

4 

3 

1 

2 

Other names: Gummy Shark, Japanese Smoothhound (English), Hiu Kacang, Hiu Air, 

Cucut Londer (Indonesian), Yu-jaras Bintik, Yu Jaras (Malaysian) 


1 1 st dorsal fin plain or with a dusky tip 

2 caudal fin often with white posterior margin or tip 

3 sides with numerous small white spots (sometimes absent or only near lateral line) 

4 1 st dorsal-fin origin well behind pectoral-fin base 

5 eyes dorsolateral on head, with ridge beneath 

6 internarial width 1–2 times nostril width 

7 upper labial furrow much longer than lower furrow 

8 teeth in both jaws broad and blunt, not strongly compressed 

Size: To at least 117 cm; both sexes mature at 62–70 cm; born at about 30 cm. 

Distribution: Northwestern Pacific from Malaysia to southern Siberia; records from 

Kenya in the Western Indian Ocean require validation. 

Habitat and biology: Demersal on soft bottoms of the continental shelf and upper 

slope, including well inshore. Biology of Japanese and Taiwanese populations well 

studied. Viviparous, with histotrophy (no yolk sac-placenta as in some other members 

of the genus); gives birth to litters of 1–22 pups (average 5) after about a 10 month 

gestation. Fast growing with both sexes maturing at 3–4 years of age. Feeds 

predominantly on demersal invertebrates, mostly crustaceans. 

Utilisation: Caught rarely by trawlers. Used for its meat and fins, but of limited value. 

Conservation status: IUCN Red List: Data Deficient.


Local synonyms: Mustelus mosis Hemprich & Ehrenberg: Yano et al. (2005). 


6 

7 

8 

Ventral head 

In Borneo: Known from the South 

China Sea, off Sarawak and Sabah. 

Triakidae (Houndsharks)


Whitefin Smoothhound 

Mustelus widodoi White & Last, 2006 

1 

5 

4 

3 

2 

2 

Other names: None (English), Hiu Kacang, Hiu Air, Cucut Londer (Indonesian), Yu 

Jaras (Malaysian) 


1 1 st dorsal fin with a prominent white tip 

2 2 nd dorsal fin and caudal-fin upper lobe with distinct black tips 

3 sides lacking white spots 

4 1 st dorsal-fin origin well behind pectoral-fin base 

5 eyes dorsolateral on head, with ridge beneath 

6 internarial width 1–2 times nostril width 

7 lower labial furrow subequal to or slightly longer than upper furrow 

8 teeth in both jaws broad and blunt, not strongly compressed 

Size: To at least 110 cm; males mature at 83–89 cm. 

Distribution: Southern Indonesia (Java, Bali, Lombok and possibly Sumatra) and off 

Malaysian Borneo. 

Habitat and biology: Biology and habitat largely unknown. Demersal on the midcontinental 

shelf to upper slope in depths greater than 60 m. Reproductive strategy 

unknown; either viviparous, with histotrophy or viviparous, with a yolk sac-placenta as 

in other members of the genus. Diet unknown, presumably consisting of small fishes 

and invertebrates as in other members of the genus. 

Utilisation: Marketed infrequently, probably more abundant in the deeper parts of the 

South China Sea. Used for its meat and fins, but of limited use due to its rarity. 


Local synonyms: Mustelus sp. 1: White et al. (2006b). 

References: White & Last (2006); White (2007a).


6 

8 

7 

Ventral head 

In Borneo: Known only from northwestern 

Sabah. 

Triakidae (Houndsharks)


Hooktooth Shark 

Chaenogaleus macrostoma (Bleeker, 1852) 

3 

1 

2 

3 

3 3 

Other names: None (English), Hiu Kacang, Hiu Pilus (Indonesian), Yu Gigi Cangkuk 

(Malaysian) 


1 spiracles small 

2 gill slits large, more than twice eye length 

3 pelvic and dorsal fins, and ventral caudal-fin lobe, weakly falcate 

4 snout obtusely wedge-shaped (viewed from underneath) 

5 teeth protruding prominently when mouth closed 

6 front lower teeth with long, strongly hooked cusps 

Size: To at least 100 cm; males mature at about 68 cm; born at about 20 cm or more. 

Distribution: Tropical Indo–West Pacific, from the Gulf of Aden to southern Indonesia, 

and north to China. 

Habitat and biology: Occurs on or near the bottom near the coast to depths of at 

least 60 m. Viviparous, with a yolk-sac placenta; gives birth to litters of 4 pups after an 

unknown gestation period. Diet probably consists of small fishes, cephalopods and 

crustaceans. 

Utilisation: Caught rarely by inshore gillnet and longline fisheries. Used for its meat 

and fins but of limited value due to its small size. 



References: None.


4 

5 

Upper tooth 

Lower tooth 

Ventral head 

6 

In Borneo: Uncommon, recorded 

from Sarawak, Sabah and northeastern 

Kalimantan, but probably more widely 

distributed. 

Hemigaleidae (Weasel Sharks)


Sicklefin Weasel Shark 

Hemigaleus microstoma Bleeker, 1852 

3 

1 

2 

3 

3 

3 

Other names: Weasel Shark (English), Hiu Kacang, Hiu Pilus (Indonesian), Yu Bintik 

Putih, Yu Pasir (Malaysian) 



2 gill slits small, less than twice eye length 

3 pelvic and dorsal fins, and ventral caudal-fin lobe, strongly falcate 

4 mouth very short and broadly arched 

5 teeth concealed when mouth closed 

6 upper teeth with oblique, subtriangular cusps, lateral margins serrated 

7 front lower teeth with short, erect cusps (roots strongly arched) 


born at about 45 cm. 

Distribution: Tropical Indo–West Pacific, from the Red Sea to Indonesia and north to 

China, excluding Australia and New Guinea. 

Habitat and biology: Occurs on or near the bottom on the continental shelf. 

Viviparous, with a yolk-sac placenta; gives birth possibly twice a year to litters of 2–4 

pups after a gestation period of probably less than 6 months. Diet largely unknown, but 

a few stomachs examined contained cephalopods. 

Utilisation: Often caught by inshore gillnet, bottom trawl and, to a lesser extent, 

longline fisheries. Used for its meat and fins, but of limited value due to its small size. 

Conservation status: IUCN Red List: Vulnerable.


Local synonyms: Hemigaleus machlani Herre, 1929; Carcharias (Hypoprion) notatus 

Bleeker, 1850. 

References: White (2007a); Taylor & Bennett (2008). 

Upper tooth 

5 

4 

Ventral head 

6 

In Borneo: Widespread. 

Hemigaleidae (Weasel Sharks) 

7 

Lower tooth 

Pectoral fin - ventral


Fossil Shark 

Hemipristis elongata (Klunzinger, 1871) 

3 

1 

2 

3 

3 

3 

Other names: Snaggletooth Shark (English), Hiu Monas (Indonesian), Jerung Gigi 

Cerakah, Yu Putih (Malaysian) 



2 gill slits large, more than twice eye length 

3 pelvic and dorsal fins, and ventral caudal-fin lobe, strongly falcate 

4 snout bluntly rounded (viewed from underneath) 

5 teeth protruding prominently when mouth closed 

6 front lower teeth with long, strongly hooked cusps 

Size: To at least 230 cm; males and females mature at 110–136 and about 120 cm 

respectively; born at 45–52 cm. 

Distribution: Tropical (rarely warm temperate) waters of the Indo–West Pacific, from 

southeastern Africa to northern Australia, and north to China. 

Habitat and biology: Occurs on continental and insular shelves to depths of at least 

130 m. Viviparous, with a yolk-sac placenta; gives birth to litters of 2–11 pups after a 

7–8 month gestation period; possibly breeds every second year. Diet consists of cephalopods 

and fishes, including small elasmobranchs. 

Utilisation: Caught often by inshore gillnet, bottom trawl and longline fisheries. Used 

for its meat (good quality), fins and cartilage. Population declines have been reported in 

some regions. 


Local synonyms: Chaenogaleus macrostoma (Bleeker): White et al. (2006b); Hemipristis 

elongatus (Klunzinger): Yano et al. (2005).


References: White (2007a). 

4 

Upper tooth 

5 

Ventral head 

6 



Lower tooth 

Pectoral fin - ventral 

Juvenile (63 cm)


Straight-tooth Weasel Shark 

Paragaleus tengi (Chen, 1963) 

3 

3 

1 

2 

3 

3 

Other names: None (English), Hiu Kacang, Hiu Pasir (Indonesian), Yu Gigi Lurus 

(Malaysian) 



2 gill slits small, less than twice eye length 

3 pelvic and dorsal fins, and ventral caudal-fin lobe, not falcate 

4 snout pointed (viewed from underneath) 

5 mouth long, narrowly arched 

6 teeth concealed when mouth closed 

7 upper teeth with narrow, moderately long, semi-erect to oblique cusps, and distal 

cusplets 

8 anterior lower teeth with moderately long, erect cusps (roots slightly arched) 

Size: To at least 93 cm; free-swimming at 40 cm. 

Distribution: Tropical northwestern Pacific, from Indonesia to southern Japan. 

Habitat and biology: An inshore species, probably occurring on or near the bottom. 

Viviparous, with a yolk-sac placenta; biology and diet unknown. 

Utilisation: Caught occasionally by inshore gillnet, bottom trawl and longline fisheries. 

Used for its meat and fins, but of limited value due to its small size. 


Local synonyms: Negogaleus longicaudatus Bessednov, 1966. 

References: None.


4 

6 

5 

Upper tooth 

Ventral head 

7 

In Borneo: Off Sabah, Sarawak and 

eastern Kalimantan. 


8 

Lower tooth


Silvertip Shark 

Carcharhinus albimarginatus (Rüppell, 1837) 

3 

1 

2 

1 

1 

1 

1 

Other names: None (English), Cucut Lanjaman, Hiu Lanyam, Hiu Plen, Hiu Sonteng 

(Indonesian), none (Malaysian) 


1 1 st dorsal, pectoral, pelvic and caudal fins with prominent white tips 

2 interdorsal ridge present 

3 apex of 1 st dorsal fin pointed or narrowly rounded 

4 snout rather long and parabolic (viewed ventrally) 

5 upper teeth triangular, cusp oblique, edges distinctly notched and serrated 

6 lower teeth serrated but more slender and erect than uppers 



Distribution: Widespread in the tropical Indo–Pacific, from southeastern Africa to 

central America. 

Habitat and biology: An inshore and offshore pelagic shark, rather than oceanic; 

occurring throughout the water column to depths of at least 800 m. Viviparous, with a 

yolk-sac placenta; gives birth every second year to a litter of about 6 pups (maximum 

11) after a 12 month gestation period. Diet includes a variety of pelagic and demersal 

fishes. Known to be aggressive when provoked and is potentially dangerous to humans. 

Utilisation: Locally, rarely landed by longline and pelagic gillnet fisheries. Used elsewhere 

for its fins, meat, skin and cartilage. 





4 

Upper tooth 

Ventral head 

5 

In Borneo: Few confirmed local records, 

but observed by divers at the Spratly 

Islands in the South China Sea; probably 

also common around coral reefs off eastern 

Sabah. 

Carcharhinidae (Whaler Sharks) 

Lower tooth 

6


Graceful Shark 

Carcharhinus amblyrhynchoides (Whitley, 1934) 

1 

2 

1 

1 

1 

1 

Other names: Queensland Shark (English), Cucut Lanjaman, Hiu Bujit (Indonesian), 

Yu Jalur Putih (Malaysian) 


1 all fin tips usually black or dusky (except anal fin) 

2 interdorsal ridge absent 

3 snout rather short, narrowly rounded to pointed (viewed ventrally), internarial space 

1–1.2 times preoral snout length 

4 upper and lower teeth with a slender, erect cusp, no basal cusplets 

5 precaudal vertebrae usually less than 82 

Size: To at least 178 cm; both sexes mature at about 110–115 cm; born at 50–60 cm. 

Distribution: Tropical Indo–West Pacific, from the Gulf of Aden to northern Australia 

and the Philippines. 

Habitat and biology: Primarily pelagic in midwater over insular shelves, from close 

inshore to bottom depths of at least 50 m. Viviparous, with a yolk-sac placenta; gives 

birth to litters of an average of 3 pups (maximum 8) after a 9–10 month gestation 

period. Diet consists primarily of fish, with smaller amounts of crustaceans and cephalopods. 

Not dangerous to humans. 

Utilisation: Caught mainly as bycatch by longlining. Used for its fins and meat, but 

typically only small in size. 




References: White (2007c); Ahmad et al. (2008). 

3 

Upper tooth 

Ventral head 

4 



Lower tooth


Grey Reef Shark 

Carcharhinus amblyrhynchos (Bleeker, 1856) 

3 

2 

1 

Other names: Longnose Blacktail Shark (English), Cucut Lanjaman, Hiu Bujit, Merak 

Bulu, Hiu Lonjor (Indonesian), Yu Ekor Hitam (Malaysian) 


1 a wide, black posterior caudal-fin margin 

2 interdorsal ridge usually absent (sometimes weak) 

3 1 st dorsal fin relatively tall, its origin over pectoral-fin free rear tips 

4 snout broadly rounded (viewed ventrally) 

5 upper teeth long, narrowly triangular, with notch on one edge 

6 lower teeth narrow, more erect than uppers 

Size: To at least 255 cm, usually



Local synonyms: Galeolamna tufiensis Whitley, 1949; specimen figured as Carcharhinus 

galapagensis (Snodgrass & Heller) in Yano et al. (2005) is most likely a juvenile male of 

this species. 


4 

Ventral head 

In Borneo: Patchily distributed, but 

probably widespread. Recorded off 

Sabah, Sarawak, and eastern and western 

Kalimantan. 

1 



Upper tooth 

5 

6 

Lower tooth


Borneo Shark 

Carcharhinus borneensis (Bleeker, 1858) 

3 

2 

1 

Other names: None (English), Hiu (Indonesian), Yu Borneo (Malaysian) 


1 2 nd dorsal fin very low (height 2.2–2.5 in inner margin), its origin over midbase of 

anal fin 


3 1 st dorsal-fin inner margin relatively long, about half of fin base 

4 snout very long and pointed (viewed ventrally), not strongly calcified 

5 hyomandibular pores conspicuously enlarged alongside mouth corners 

6 upper teeth serrated with a narrow, oblique cusp and large basal cusplets 

7 lower teeth with an erect to oblique cusp and weak cusplets 

8 upper and lower tooth rows 11–12 

Size: To at least 65 cm; males mature at about 55–58 cm. 

Distribution: Known from off Malaysia, China, and possibly Indonesia and the 

Philippines but range probably now more restricted. 

Habitat and biology: A rarely caught coastal and inshore species. Biology and habitat 

virtually unknown. 

Utilisation: Presumably caught on line; used for its meat but of limited value due to its 

small size and restricted distribution. 

Conservation status: IUCN Red List: Endangered. 

Local synonyms: Specimen figured as Carcharhinus borneensis in Yano et al. (2005) is a 

juvenile Rhizoprionodon acutus (Rüppell). 

References: White et al. (2010a).


4 

Upper tooth 

Lower tooth 

Ventral head 

5 

6 

7 

In Borneo: Presently known only 

from Mukah (Sarawak); type collected 

in the 19 th century from northwestern 

Kalimantan. 

Carcharhinidae (Whaler Sharks)


Spinner Shark 

Carcharhinus brevipinna (Müller & Henle, 1839) 

1 

2 

Other names: Longnose Grey Whaler, Inkytail Shark, Smoothfang Shark (English), 

Hiu Bujit, Hiu Plen, Merak Bulu, Cucut Lanjaman (Indonesian), Yu Muncung Panjang 

(Malaysian) 


1 1 st dorsal-fin usually slightly behind pectoral-fin free rear tips 


3 snout long and pointed (viewed ventrally) 

4 all fins, except pelvics, with conspicuous black tips in specimens over 1 m 

5 upper labial furrows conspicuous 

6 upper and lower teeth similar, nearly symmetrical, with a very low and narrow, erect 

cusp 

Size: To about 300 cm; both sexes mature at about 190–200 cm; born at 60–81 cm. 

Distribution: Tropical and warm temperate waters of the Indian, western Pacific and 

Atlantic Oceans. 

Habitat and biology: Generally found nearshore to at least 75 m depth on the 

continental shelf. Viviparous, with a yolk-sac placenta; gives birth every second year to 

litters of 3–15 pups after a 10–12 month gestation period. Both sexes mature at 8–10 

years. Diet consists primarily of small pelagic fishes, and smaller quantities of 

cephalopods. An active schooling species capable of spinning out of the water during 

feeding runs through fish schools. 

Utilisation: Caught by longlining, with juveniles a very abundant component of 

inshore gillnet fisheries in parts of Indonesia. Used for its fins, meat, skin and cartilage. 



Local synonyms: Carcharias (Prionodon) brevipinna Bleeker, 1853. 


3 

5 In Borneo: Probably widespread. 

Ventral head 

Upper tooth 

6 


Lower tooth 

Lateral head


Whitecheek Shark 

Carcharhinus dussumieri (Müller & Henle, 1839) 

1 

2 

3 

Other names: Widemouth Blackspot Shark (English), Cucut Lanjaman, Hiu Bujit 

(Indonesian), Yu Pasir (Malaysian) 


1 1 st dorsal fin moderately tall, broad, not falcate 

2 low interdorsal ridge usually present 

3 only 2 nd dorsal fin with a black tip (usually covering about half of fin) 

4 snout rather long, broadly parabolic (viewed ventrally) 

5 upper teeth with strongly oblique cusp, flanked on one side by strong, serrated 

cusplets 

6 lower teeth narrow, upright, no cusplets 

Size: To at least 94 cm; both sexes mature at about 70–75 cm; born at 34–40 cm. 

Distribution: Tropical waters of the Indo–West Pacific, from the Red Sea to northern 

Australia, New Guinea and Japan. 

Habitat and biology: A common demersal inshore species from close inshore to 

about 170 m depth. Viviparous, with a yolk-sac placenta; gives birth each year to litters 

of 2 (maximum 4) pups after an unknown gestation period; no reproductive 

seasonality. Diet consists primarily of fish, and to a lesser extent crustaceans and 

cephalopods. Molecular analysis has identified the existence of several forms in the 

Indo–Pacific that could be separate species. 

Utilisation: Commonly caught by bottom trawl and demersal gillnet fisheries. Used for 

its fins and meat but of limited value due to its small size. 



Local synonyms: Carcharias (Prionodon) tjutjot Bleeker, 1852; Carcharias (Prionodon) 

javanicus Bleeker, 1852. 


4 

Upper tooth 

Ventral head 

5 



Lower tooth 

6


Silky Shark 

Carcharhinus falciformis (Müller & Henle, 1839) 

1 

2 

3 

Other names: None (English), Mungsing, Hiu Lonjor, Cucut Lanjaman, Hiu Bujit, 

Hiu Lanyam (Indonesian), Yu Jereh, Yu Pasir (Malaysian) 


1 1 st dorsal-fin origin well behind pectoral-fin free rear tips 


3 2 nd dorsal-fin low with very long inner margin, 1.6–3.0 times its height 

4 snout rather long, narrowly rounded (viewed ventrally) 

5 upper teeth narrow with one edge prominently notched 

6 lower teeth narrow, upright 

Size: To about 350 cm, but usually



Local synonyms: Aprionodon sitankaiensis Herre, 1934. 


4 

Upper tooth 

Ventral head 

5 

In Borneo: Eastern Sabah, and northwestern 

and eastern Kalimantan; probably 

widespread. 


Lower tooth 

6


Bull Shark 

Carcharhinus leucas (Müller & Henle, 1839) 

2 

1 

3 

Other names: River Whaler, Freshwater Whaler (English), Cucut Bekeman, Hiu Buas, 

Hiu Bujit (Indonesian), Jerung Sapi, Yu Jerung (Malaysian) 


1 2 nd dorsal fin relatively large (about a third height of 1 st dorsal fin) 


3 notch on posterior margin of anal fin not acute (usually 90º or more) 

4 snout very short and broadly rounded (viewed ventrally), preoral length less than 

internarial space 

5 upper teeth triangular, edges heavily serrated 

6 lower teeth narrowly triangular, erect 

7 usually 12 tooth rows on each side of lower jaw 

Size: To at least 400 cm; males and females mature at 197–226 and 180–230 cm 


Distribution: Cosmopolitan in most tropical and warm temperate waters, including 

freshwater and brackish rivers and lakes. 

Habitat and biology: A coastal, estuarine and freshwater shark, occurring near the 

bottom to depths of at least 150 m. Viviparous, with a yolk-sac placenta; gives birth to 

litters of 1–13 pups after a 10–11 month gestation period. Both sexes mature at about 

20 years and attain 50 years, in South African populations. Omnivorous, diet includes 

turtles, crocodiles, birds, dolphins, terrestrial mammals and crustaceans, but prefers bony 

fishes and elasmobranchs. An extremely aggressive shark that has frequently attacked 

humans. Molecular analysis has identified slight population differences between Indo– 

Pacific and Atlantic forms.


Utilisation: Landed occasionally by coastal longline and gillnet fisheries. Used for its 

fins, meat, skin, jaws and cartilage. 




4 

Ventral head 

In Borneo: Widespread, concentrated 

near major river outflows. 


Juvenile (86 cm) 

Upper tooth 

6 

5 

Lower tooth


Common Blacktip Shark 

Carcharhinus limbatus (Müller & Henle, 1839) 

4 

1 

2 

4 

4 

4 

4 

Other names: Blacktip Whaler (English), Hiu Kejen, Merak Bulu, Cucut Lanjaman, 

Hiu Lanyam, Hiu Bujit (Indonesian), Yu Trihitam, Yu Jereh, Yu Kepak Hitam (Malaysian) 


1 1 st dorsal-fin origin above or just behind pectoral-fin insertions 


3 snout long and pointed (viewed ventrally), internarial space 1.3–1.7 times preoral 

snout 

4 juveniles with obvious black fin tips (dorsal, pectoral and lower lobe of caudal fin 

plain in specimens >130 cm) 

5 upper labial furrows short, barely noticeable 

6 upper and lower teeth similar, nearly symmetrical, with an erect, narrow cusp 

Size: To at least 250 cm; males and females mature at 165–180 and 183–194 cm 


Distribution: Cosmopolitan in tropical and warm temperate waters. 

Habitat and biology: Mainly pelagic over continental and insular shelves, usually 

inshore, but occasionally found well offshore. Viviparous, with a yolk-sac placenta; gives 

birth every two years to litters of about 4–7 pups (maximum 10) after a 10–12 month 

gestation period. Males and females mature at 6–7 and 5–6 years respectively. Diet 

consists primarily of bony fishes and elasmobranchs, as well as crustaceans and cephalopods. 

Harmless to humans, unless stimulated by food. Molecular analysis has identified 

differences between Indo–Pacific and Atlantic forms that could be separate species.


Utilisation: Caught by longline and inshore gillnet fisheries. Used for its fins, meat, skin 

and cartilage. 


Local synonyms: Carcharias (Prionodon) pleurotaenia Bleeker, 1852. 


3 

Ventral head 

5 


Upper tooth 


6 

Lateral head 

Lower tooth


Hardnose Shark 

Carcharhinus macloti (Müller & Henle, 1839) 

3 

2 

1 

Other names: None (English), Hiu Aron (Indonesian), Yu Muncung Keras, Yu Jereh, 

Yu Pasir (Malaysian) 


1 2 nd dorsal-fin origin about over midbase of anal fin 


3 1 st dorsal-fin inner margin extremely long, about two thirds of fin base 

4 snout long and pointed (viewed ventrally); rostrum very hard and strongly calcified 

5 upper teeth with narrow, oblique or nearly erect central cusp, bordered by strong 

cusplets 

6 lower teeth tall, narrow, upright, smooth edged 

Size: To at least 110 cm; males and females mature at 70–75 cm; born at 40–45 cm. 

Distribution: Tropical Indo–West Pacific, from eastern Africa to northern Australia and 

New Guinea. 

Habitat and biology: Occurs from close inshore to at least 170 m depth, sometimes in 

large aggregations. Viviparous, with a yolk-sac placenta; gives birth every second year to 

litters of usually 2 pups after a gestation period of about 12 months. Diet consists 

primarily of small fishes, but also cephalopods and crustaceans. 

Utilisation: Caught occasionally by inshore demersal gillnet fisheries. Used for its fins 

and meat, but of limited value due to its small size. 



References: None.


4 

Upper tooth 

Ventral head 

5 

In Borneo: Patchy, off Sarawak, Sabah 

and eastern Kalimantan. 


Lower tooth 

6


Blacktip Reef Shark 

Carcharhinus melanopterus (Quoy & Gaimard, 1824) 

1 

2 

3 

7 

Other names: Blacktip Shark, Guliman (English), Hiu Mada, Kluyu Karang, Hiu 

Bujit (Indonesian), Yu Sirip Hitam, Yu Kepak Hitam, Yu Nipah (Malaysian) 


1 1 st dorsal fin with a thick black tip (its inner edge sharply defined and often 

bordered by white area) 


3 distinct pale stripe along each flank 

4 snout very short, broadly rounded (viewed ventrally), preoral length subequal to 

internarial space 

5 upper teeth with narrow, oblique central cusp and low basal cusplets 

6 lower teeth narrow, upright to oblique, edges finely serrated 

7 caudal fin lower lobe with thick black tip (other fins often with smaller black tips) 

Size: To at least 142 cm; both sexes mature at 95–110 cm; born at 35–50 cm. 

Distribution: Tropical Indo–West Pacific and Central Pacific, from southeastern Africa 

to the Central Pacific Islands, and eastern Mediterranean. 

Habitat and biology: One of the most common reef sharks, usually living in lagoons 

and near the fringes of reefs. Viviparous, with a yolk-sac placenta; gives birth each year 

to litters of 2–4 pups after an 8–9 month gestation period (breeds every second year in 

Aldabra, Western Indian Ocean). Diet consists primarily of fishes, but also cephalopods, 

crustaceans and other molluscs. Not regarded as dangerous but has been known to 

attack waders and spearfishers.


Utilisation: Caught by shallow-set lines and nets, and probably adversely affected by 

dynamite fishing in the region. Used for its fins and meat. 




4 

Ventral head 



Upper tooth 

5 

Lower tooth 

6


Sandbar Shark 

Carcharhinus plumbeus (Nardo, 1827) 

3 

1 

2 

Other names: Thickskin Shark, Brown Shark (English), Hiu Teteri, Cucut Lanjaman 

(Indonesian), Yu Kulit Tebal (Malaysian) 


1 1 st dorsal fin very broad and tall, more than half of predorsal length (smaller in 

newborns) 


3 origin of 1 st dorsal-fin over or just anterior to pectoral-fin insertion 

4 snout rather long and broadly rounded (viewed ventrally) 

5 upper teeth broadly triangular, cusp erect to slightly oblique 

6 lower teeth narrow, upright, edges finely serrated 

Size: To at least 240 cm; males and females mature at 130–180 cm and 145–185 cm 


Distribution: Cosmopolitan, but patchy, in tropical and warm temperate waters. 

Habitat and biology: Occurs over continental and insular shelves, and adjacent deep 

water, from the intertidal to depths of at least 280 m, usually near the bottom in 20– 

60 m depth. Viviparous, with a yolk-sac placenta; gives birth every two or three years 

to litters of 5–12 (maximum 14) pups after a 12 month gestation period. Males and 

females mature at 14 and 16 years respectively, in Australian populations. Diet consists 

primarily of fishes, but also cephalopods and crustaceans. Not reported to be dangerous 

to humans. Molecular analysis has identified differences between Indo–Pacific and 

Atlantic forms that could be separate species.


Utilisation: Occasionally caught in the shark longline and tuna gillnet fisheries. Used 

for its fins (high value in adults), meat, skin and cartilage. 




4 

Upper tooth 

Ventral head 

5 

In Borneo: Off Sabah, Sarawak and 

Brunei, but probably more widespread. 


Lower tooth 

6


Blackspot Shark 

Carcharhinus sealei (Pietschmann, 1913) 

1 

2 

3 

Other names: None (English), Cucut Lanjaman (Indonesian), Yu Pasir (Malaysian) 


1 1 st dorsal fin moderately tall, moderately falcate 

2 low interdorsal ridge usually present 

3 only 2 nd dorsal fin with a black tip (covering more than half of fin, often extending 

onto body) 

4 snout rather long, narrowly parabolic (viewed ventrally) 

5 upper teeth with strongly oblique cusp, flanked on one side by strong, smooth-edged 

cusplets 

6 lower teeth narrow, upright, usually with conspicuous cusplets 

Size: To at least 95 cm; males and females mature at >80 cm and 68–75 cm respectively; 


Distribution: Tropical Indo–West Pacific, from southeastern Africa to New Guinea 

and China (not found in Australian waters). 

Habitat and biology: A common coastal species, demersal in inshore waters from the 

surf zone down to at least 40 m depth. Viviparous, with a yolk-sac placenta; gives birth 

to litters of 1 or 2 pups after a gestation period of ~9 months; fast growing, matures 

after 1 year and maximum age about 5 years. Diet consists of small fishes, crustaceans 

and cephalopods. Not dangerous to humans. 

Utilisation: Caught frequently off Sarawak and northwestern Kalimantan by trawl and 

gillnet fisheries. Used for its fins and meat but of limited value due to its small size. 



Local synonyms: Carcharias borneensis Seale, 1910. 


4 

Upper tooth 

Ventral head 

5 

In Borneo: Widespread, but no current 

records from southwestern Kalimantan. 


Lower tooth 

6


Spot-tail Shark 

Carcharhinus sorrah (Müller & Henle, 1839) 

2 

1 3 

1 

1 

Other names: Sorrah Shark, School Shark (English), Cucut Lanjaman, Hiu Bujit, 

Lanyam, Merak Bulu, Mungsing (Indonesian), Yu Kepak Hitam (Malaysian) 


1 2 nd dorsal, pectoral and ventral tip of caudal fin strikingly black-tipped 


3 2 nd dorsal fin very low, inner margin extremely long (2.0–2.6 times fin height) 

4 snout long and moderately pointed (viewed ventrally) 

5 upper teeth with oblique cusp, flanked on one side by strong cusplets 

6 lower teeth narrow, oblique, without cusplets 

Size: To at least 160 cm; males and females mature at 90–115 cm and 95–118 cm 


Distribution: Tropical Indo–West Pacific, from southeastern Africa to northern 

Australia, Solomon Islands and Japan. 

Habitat and biology: A common shark, particularly off western Borneo; occurring 

over continental and insular shelves, including around coral reefs, from the intertidal to 

at least 80 m depth. Viviparous, with a yolk-sac placenta; giving birth each year to 

litters of about 3–8 pups after a 10 month gestation period. Growth rapid with both 

sexes maturing at about 2–3 years. Diet consists primarily of bony fishes, but also 

cephalopods and crustaceans. Not dangerous to humans. 

Utilisation: Common catch of the shark longline and inshore gillnet fisheries. Used 

widely for its fins, meat, skin and cartilage. 



Local synonyms: Specimen figured as Carcharhinus hemiodon (Müller & Henle) in Yano 

et al. (2005) appears to be an adult male of this species. 


4 

Upper tooth 

Ventral head 

5 



Lower tooth 

6


Tiger Shark 

Galeocerdo cuvier (Péron & Lesueur, 1822) 

2 1 

3 

Other names: None (English), Mungsing Jara, Hiu Macan, Hiu Omas (Indonesian), Yu 

Tenggiri, Yu Giling, Jerung Tenggiri (Malaysian) 


1 dark, vertical bars on sides (less distinct in adults) 

2 spiracle present, small and slit-like 

3 caudal peduncle with a low, rounded, lateral keel 

4 snout very short and bluntly rounded (viewed ventrally) 

5 upper labial furrows very long, subequal to preoral length 

6 teeth in both jaws heavily serrated, cockscomb-shaped, one edge deeply notched, the 

other convex 

Size: To at least 600 cm (one record of 740 cm); males and females mature at 300–305 

and 250–350 cm respectively; born at about 51–76 cm. 

Distribution: Cosmopolitan in all tropical seas, making seasonal incursions into warm 

temperate areas. 

Habitat and biology: Occurs close inshore to the outer continental shelf, including the 

intertidal zone and surface, to at least 150 m depth. The only non-placental whaler shark; 

viviparous, with histotrophy; gives birth to litters of an average of 33 pups (maximum 82) 

after a 12–16 month gestation period. Males and females mature at 7–8 and 7–12 years 

respectively. Omnivorous, feeding on a broad variety of prey including fishes, elasmobranchs, 

dugongs, birds, turtles, snakes, dolphins, birds, crustaceans and cephalopods. 

Potentially very dangerous to humans given its large size and presence close inshore, but 

normally not aggressive. Probably belongs to its own family of sharks.


Utilisation: Caught occasionally by the shark longline, tangle net and bottom trawl 

fisheries. Valuable for its fins, meat, skin, jaws and cartilage. 


Local synonyms: Carcharias (Prionodon) fasciatus Bleeker, 1852; Galeocerdo fasciatus van 

Kampen, 1907. 


4 

5 

Ventral head 

In Borneo: Almost certainly widespread; 

current records are from off Sarawak, 

Sabah and eastern Kalimantan. 


Upper tooth 

6 

Lower tooth


Mukah River Shark 

Glyphis sp. 

5 

4 

2 

6 

1 

5 

5 

Other names: None (English), none (Indonesian), Yu Sungai (Malaysian) 


1 precaudal pits longitudinal (not crescentic) 

2 2 nd dorsal fin only moderately tall, slightly less than half height of 1 st dorsal fin 

3 eyes moderately large 

4 1 st dorsal-fin free rear tip just anterior to pelvic-fin origins 

5 fins plain 

6 2 nd dorsal-fin posterior margin moderately concave 

7 waterline (demarcation of dark dorsal and pale ventral colours) sharp, extending 

through level of mid-eye on head 

Size: Known from two immature males of 610 and 658 mm. An adult male Glyphis 

(1660 mm) caught in Sampit Bay (South Kalimantan) is possibly conspecific. 

Distribution: Possibly endemic to waters of Borneo. 

Habitat and biology: Very rare; probably occurs in turbid rivers, brackish backwaters 

and marine areas adjacent to major river outflows. Biology unknown; presumably viviparous, 

with yolk-sac placenta. Diet also unknown, presumably primarily bony fishes. 

Conservation status needs investigating, likely to be highly threatened. 

Utilisation: Caught rarely. Probably utilised for its meat and fins where caught. 


Local synonyms: None.


References: Fahmi & Adrim (2009); Compagno et al. (2010). 

3 

7 

Lateral head 

In Borneo: Known from only two 

specimens collected from Mukah in 

Sarawak. A specimen collected from 

Sampit Bay in south Kalimantan may 

also represent this species. 



Borneo River Shark 

Glyphis fowlerae Compagno, White & Cavanagh, 2010 

3 

7 

4 

2 

6 

1 

5 

5 

Other names: None (English), none (Indonesian), Yu Sungai (Malaysian) 



2 2 nd dorsal fin tall, half to three fifths of 1 st dorsal-fin height 

3 eyes noticeably small 

4 1 st dorsal-fin free rear tip just anterior to pelvic-fin origins 

5 fins mainly plain, except for dark patch on pectoral-fin bases and dusky tip on 

ventral caudal-fin lobe 

6 2 nd dorsal-fin posterior margin nearly straight or shallowly concave 

7 waterline (demarcation of dark dorsal and pale ventral colours) diffuse, extending 

below eye on head 

8 lower anterior teeth with an entirely serrated cusp, no spear-like tips; total tooth row 

counts 60–63 

9 total vertebral centra 196–205, precaudal centra 108–114 

Size: To at least 200 cm; born at about 50 cm. 

Distribution: Endemic to waters of Borneo. 

Habitat and biology: Very rare; occurs in turbid rivers and brackish backwaters. 

Biology essentially unknown; presumably viviparous, with yolk-sac placenta. Diet also 

unknown, presumably primarily bony fishes. Conservation status needs investigating, 

likely to be highly threatened. 

Utilisation: Caught infrequently, and only in specific locations, by artisanal fishers using


lines and gill nets in the Kinabatangan River. Utilised for its meat. 


Local synonyms: Glyphis sp. B: Compagno & Niem (1998); Compagno et al. (2005); 

Glyphis sp.: Yano et al. (2005). 

References: Compagno et al. (2010). 

7 

Ventral head 

3 

In Borneo: Recorded only from the 

Kinabatangan River, northeastern Sabah. 

Upper tooth 


Lateral head 

Lower tooth 

8


Borneo Broadfin Shark 

Lamiopsis tephrodes (Fowler, 1905) 

3 

2 

1 

4 

Other names: None (English), Hiu Bujit (Indonesian), Yu Sirip Lebar (Malaysian) 



2 2 nd dorsal fin very tall, similar to height of 1 st dorsal fin 


4 pectoral fin long with a very broad base 

5 snout rather long, parabolic (viewed ventrally) 

6 upper teeth with a high, broadly triangular, erect to weakly oblique cusp 

7 lower teeth with an erect, high, hooked, smooth-edged cusp 



Distribution: Not well defined; tropical waters of the Indo–Malay Archipelago, north 

to China. 

Habitat and biology: Probably a range-restricted shark occurring in shallow coastal 

and inshore waters, usually with high turbidity. Viviparous, with a yolk-sac placenta; 

gives birth to litters of 4–8 (usually 8) pups after a gestation period of about 8 months. 

Dietary composition not known, but probably consists of small fishes, crustaceans and 

cephalopods. Harmless to humans. 

Utilisation: Caught irregularly by inshore gillnets at locations influenced by runoff 

from rivers. Used for its fins and meat. 

Conservation status: IUCN Red List: Not Evaluated.


Local synonyms: Lamiopsis temmincki (Müller & Henle): Yano et al. (2005), White et al. 

(2006b); Carcharhinus microphthalmus Chu, 1960. 

References: White et al. (2010c). 

5 

Upper tooth 

Ventral head 

6 

In Borneo: Occurs around the Borneo 

coastline, but more common at particular 

localities, such as Mukah (Sarawak). 


Lower tooth 

7


Sliteye Shark 

Loxodon macrorhinus Müller & Henle, 1839 

1 

3 

2 

5 

Other names: Jordan’s Blue Dogshark, Slender Dogshark (English), Hiu Kejen 

(Indonesian), Yu Mata Lekuk (Malaysian) 


1 eye large with distinct notch on its posterior edge 

2 2 nd dorsal-fin origin over anal-fin insertion 

3 1 st dorsal-fin origin well posterior to pectoral-fin free rear tips 

4 small and inconspicuous labial furrows 

5 preanal ridges very long, equal to anal-fin base length 

6 snout very long, parabolic (viewed ventrally) 

7 teeth in both jaws similar, with a strongly oblique, narrowly triangular, smoothedged 

cusp 



Distribution: Tropical waters of the Indo–West Pacific, from southeastern Africa to 

Australia and southern Japan. 

Habitat and biology: A common species occurring near the bottom in shallow, clear, 

inshore waters to at least 100 m depth. Viviparous, with a yolk-sac placenta; gives birth 

each year to litters of 2 pups (maximum 4); no reproductive seasonality. Diet consists 

primarily of small fishes and crustaceans, as well as cephalopods. Harmless to humans. 

Utilisation: Caught frequently by inshore demersal gillnet fisheries. Used for its fins 

and meat, but of limited value due to its small size. 


Local synonyms: Carcharias (Scoliodon) dumerilii Bleeker, 1856. 

References: None.


6 

4 In Borneo: Mainly northern Borneo; off 

Sarawak, Sabah, and eastern and western 

Kalimantan. 

Ventral head 

Upper tooth 

Lower tooth 

7 

Eye 

1 



Blue Shark 

Prionace glauca (Linnaeus, 1758) 

4 

1 3 

2 

Other names: Blue Whaler, Great Blue Shark (English), Hiu Aer, Hiu Lalaek, Hiu 

Karet, Cucut Selendang (Indonesian), Jerung Biru (Malaysian) 


1 1 st dorsal-fin base closer to pelvic-fin base than pectoral-fin base 

2 pectoral fins very long and scythe-like 

3 caudal peduncle with weak lateral keels 

4 dorsal surface indigo blue, ventral surface white 

5 snout very long and narrowly rounded (viewed ventrally) 

6 upper teeth with a narrow, triangular, oblique and finely serrated cusp 

7 lower teeth with a slender, erect, finely serrated cusp 

Size: To at least 383 cm; both sexes mature at about 210–220 cm (somewhat variable); 


Distribution: Cosmopolitan in all tropical and temperate seas. 

Habitat and biology: The most wide-ranging of all sharks; typically oceanic and 

pelagic, from the surface to at least 600 m depth; capable of long migrations. Viviparous, 

with a yolk-sac placenta; gives birth annually or every second year to litters of usually 

30–40 pups (maximum 135) after a 9–12 month gestation period. Growth is fairly 

rapid with males and females maturing at 4–6 and 5–7 years respectively. Diet consists 

primarily of small pelagic fishes and cephalopods, as well as demersal fishes, small sharks 

and seabirds. Potentially dangerous to humans, but not particularly aggressive. 

Utilisation: Common bycatch of tuna and shark longline fisheries in Indonesia but 

infrequently landed by Borneo fishers. Valued for its fins, meat, skin, jaws and cartilage.





5 

Ventral head 

In Borneo: Known from a single record 

off Sabah; probably more widespread. 


Upper tooth 

6 

Lower tooth 

7


Milk Shark 

Rhizoprionodon acutus (Rüppell, 1837) 

1 

2 

Other names: Longman’s Dogshark, Fish Shark, White-eye Shark (English), Hiu Pilus, 

Hiu Plen, Mungsing, Hiu Pisang (Indonesian), Yu Muncung Susu (Malaysian) 


1 2 nd dorsal fin smaller than anal fin, its origin about opposite anal-fin insertion 

2 preanal ridges very long, about equal to anal-fin base length 

3 snout long and narrowly rounded (viewed ventrally) 

4 upper labial furrows long and prominent, 1.4–2.0% of total length 

5 hyomandibular pores alongside mouth corners enlarged in a distinct series, usually 

more than 16 in total for both sides 

6 teeth in both jaws with an oblique, narrowly triangular cusp 

Size: To about 110 cm (a 178 cm individual recorded off Africa); males and females 

mature at about 75–79 and 75–83 cm respectively; born at 30–40 cm. 

Distribution: Tropical Indo–West Pacific, from southeastern Africa to northern 

Australia and southern Japan, and Eastern Atlantic. 

Habitat and biology: A common species occurring mainly near the bottom, over 

continental and insular shelves, from the intertidal to about 200 m depth. Viviparous, 

with a yolk-sac placenta; gives birth each year to litters of usually 2–5 pups (maximum 8) 

after a gestation period of about 12 months; no seasonality in its reproductive cycle; size 

at birth varies regionally. Diet consists primarily of small fishes, and to a lesser extent 

cephalopods and crustaceans. Molecular analysis has identified differences between 

Indo-Malay and Australian populations that could be separate species. 

Utilisation: Commonly caught by inshore demersal gillnet fisheries. Used for its fins 

and meat, but of limited value due to its small size.



Local synonyms: Carcharias (Scoliodon) walbeehmi Bleeker, 1856; Carcharias (Prionodon) 

sorrahkowah Bleeker, 1854; Carcharhinus borneensis (Bleeker): Yano et al. (2005). 


3 

4 

5 In Borneo: Mainly northern Borneo; 

from Sabah, Sarawak, and eastern and 

western Kalimantan. 

Ventral head 


Upper tooth 

6 

Lower tooth


Grey Sharpnose Shark 

Rhizoprionodon oligolinx Springer, 1964 

1 

2 

Other names: None (English), Hiu Pilus, Hiu Pisang (Indonesian), Yu Muncung 

Minyak (Malaysian) 


1 2 nd dorsal fin smaller than anal fin, its origin opposite anal-fin insertion 

2 preanal ridges very long, about equal to anal-fin base length 

3 snout long and narrowly rounded (viewed ventrally) 

4 upper labial furrows short, 0.2–1.3% of total length 

5 hyomandibular pores alongside mouth corners enlarged in a distinct series, usually 

less than 14 in total for both sides 

6 teeth in both jaws with an oblique, narrowly triangular cusp 

7 small gap often visible at symphysis of upper and lower jaws when mouth closed 

(viewed ventrally) 

Size: To about 70 cm; males and females mature at about 29–45 and 32–41 cm 


Distribution: Tropical Indo–West Pacific, from the Red Sea to Indonesia and southern 

Japan (only a single validated record from the Gulf of Carpentaria in northern Australia). 

Habitat and biology: A small, common, inshore shark, schooling close inshore to at 

least 36 m depth. Viviparous, with a yolk-sac placenta; gives birth to litters of 3–5 pups 

after an unknown gestation period. Diet probably consists primarily of small fishes. 

Utilisation: Caught by inshore demersal gillnet fisheries. Used for its fins and meat, but 

of limited value due to its small size. 

Conservation status: IUCN Red List: Least Concern.


Local synonyms: Scoliodon intermedius Garman, 1913; specimen figured as Carcharhinus 

sp. in Yano et al. (2005) appears to be of a small individual of this species. 


3 

4 

7 

Ventral head 

5 

In Borneo: Widely distributed. 


Upper tooth 

6 

Lower tooth


Pacific Spadenose Shark 

Scoliodon macrorhynchos (Bleeker, 1852) 

2 

1 3 

5 

4 

Other names: None (English), Mungsing, Hiu Kejen (Indonesian), Yu Padi, Yu Pasir, 

Yu Jereh (Malaysian) 


1 head and snout strongly depressed, trowel-shaped 

2 1 st dorsal fin large, closer to pelvic-fin base than to pectoral-fin base 

3 2 nd dorsal fin very small, its origin well behind origin of a much larger anal fin 

4 pectoral fin very small and triangular (not falcate) 

5 caudal fork shallowly concave 

6 teeth small with a strongly oblique, smooth-edged, blade-like cusp 

Size: To about 74 cm; both sexes mature at 25–36 cm; born at 12–15 cm. 

Distribution: Indo–Malayan region eastwards to Borneo, China and Taiwan; extremities 

of range need investigating. 

Habitat and biology: A common species that is very abundant in coastal waters, 

particularly near major freshwater outflows; often forming large schools near the 

bottom in rocky habitats. Viviparous, with a yolk-sac placenta; eggs released by females 

are very small with small yolk reserves so an unusual placenta and umbilical cord 

nourish the pups from a very early stage of the gestation period. Diet presumably 

consists primarily of crustaceans and small fishes. 

Utilisation: Common bycatch of the inshore demersal trawl and gillnet fisheries, 

particularly off Sarawak and Kalimantan. Used for its meat, but of limited value due to 

its small size. 

Conservation status: IUCN Red List: Not Evaluated.


Local synonyms: Scoliodon laticaudus Müller & Henle: Yano et al. (2005), White et al. 

(2006b). 

References: White et al. (2010b). 

1 

Ventral head 

In Borneo: Mainly off western Borneo, 

including Sarawak, and western and eastern 

Kalimantan. 


Upper tooth 

6 

Lower tooth


Whitetip Reef Shark 

Triaenodon obesus (Rüppell, 1837) 

1 

2 

3 

1 

Other names: Whitetip Shark, Blunthead Shark (English), Hiu Bokem, Hiu Karang 

Buas, Hiu Coklat (Indonesian), Yu Sirip Putih (Malaysian) 


1 1 st dorsal and upper lobe of caudal fin with distinctive white tips 


3 2 nd dorsal fin large, about half to three quarters height of 1 st dorsal fin 

4 snout very short, broadly rounded, tip blunt (viewed ventrally) 

5 upper and lower teeth smooth-edged with a long, narrow central cusp, flanked either 

side by a strong cusplet 

Size: To about 200 cm; males and females mature at 112–118 and 114–122 cm respectively; 


Distribution: Tropical Indo–Pacific from southeastern Africa to Central America, 

including the Pacific Islands. 

Habitat and biology: A reef-associated shark resting on or near the bottom in caves 

and crevices on coral reefs during the day, usually in clear shallow water in depths of 

8–40 m, but has also been reported from a depth of 330 m. Viviparous, with a yolk-sac 

placenta; gives birth to litters of about 2 or 3 pups (maximum 5) after a 10–12 month 

gestation period. Curious but rarely aggressive species. Known to be ciguatoxic in rare 

circumstances. 

Utilisation: Caught by inshore line and net fisheries, and probably adversely affected by 

dynamite fishing. Used for its fins and meat. 





4 

Upper tooth 

Ventral head 

5 

In Borneo: Mainly northern Borneo; 

from Sabah, Sarawak, and northwestern 

and eastern Kalimantan. 


Lower tooth


Winghead Shark 

Eusphyra blochii (Cuvier, 1816) 

1 

2 

3 

5 

Other names: Slender Hammerhead (English), Hiu Capil, Hiu Bingkoh, Hiu Caping, 

Hiu Martil (Indonesian), Yu-tukul Palang, Yu Tanduk, Yu Palang, Yu Mata Jauh, Yu 

Bengkong, Yu Sanggul (Malaysian) 


1 head extremely broad, wing-shaped, its width about half of total length 

2 1 st dorsal fin very tall, strongly falcate 

3 origin of 1 st dorsal fin over pectoral-fin bases 

4 midline of head with a shallow indentation 

5 upper precaudal pit forming a narrow longitudinal groove (not crescentic) 

6 teeth similar in both jaws, relatively small, smooth-edged and oblique 



Distribution: Widepread in the tropical Indo–West Pacific, from the Persian Gulf to 

northern Australia and New Guinea, north to Taiwan. 

Habitat and biology: Occurs in shallow areas of the continental and insular shelves. 

Viviparous, with a yolk-sac placenta; gives birth to litters of 6–25 pups after a gestation 

period of 10–11 months. Diet consists primarily of small fishes, but also crustaceans and 

some cephalopods.


Utilisation: Used for its meat but less important than other hammerheads due to its 

small average size. 


Local synonyms: Zygaena latycephala van Hasselt, 1823. 

References: Stevens & Lyle (1989). 

4 

1 

In Borneo: A few records off Sabah, 

Sarawak and northwestern Kalimantan 

but probably widespread. 

Sphyrnidae (Hammerhead Sharks) 

Ventral head 

Upper tooth 

6 

Lower tooth


Scalloped Hammerhead 

Sphyrna lewini (Griffith & Smith, 1834) 

1 

3 

4 

6 

5 

Other names: Kidney-headed Shark (English), Hiu Capil, Hiu Bingkoh, Hiu Caping, 

Hiu Martil, Hiu Parang (Indonesian), Yu-tukul Sanggul, Yu Tukul, Yu Palang, Yu Mata 

Jauh, Yu Bengkong, Yu Sanggul, Yu Parang (Malaysian) 


1 head broad, its width less than a third of total length 

2 anterior margin of head arched, barely indented at midline 

3 1 st dorsal fin tall, moderately falcate 

4 2 nd dorsal fin short with long rear tip and weakly concave posterior margin 

5 anal-fin base barely longer than 2 nd dorsal-fin base 

6 upper precaudal pit forming a crescentic groove 

7 upper teeth narrowly triangular (erect anteriorly becoming oblique posteriorly) 

Size: To at least 350 cm, possibly to 420 cm; males mature at 140–180 cm and females 

at 200–230 cm; born at 40–50 cm. 


Habitat and biology: The most abundant tropical hammerhead; occurs over continental 

and insular shelves and adjacent deep water, from the surface to at least 275 m depth; 

juveniles usually close inshore. Viviparous, with a yolk-sac placenta; gives birth to litters 

of 13–41 (mean 25) pups in October to January after a gestation of 9–10 months. Males 

and females mature at 7–10 and about 15 years respectively. Diet consists primarily of 

bony fishes and cephalopods, but also sharks and rays. Potentially dangerous to humans. 

Often form large diurnal aggregations that disperse to deepwater at night to feed. Two 

distinct forms have been identified from molecular analysis of Borneo specimens but,


although there appear to be differences in the shape of the dark marking on the lower 

caudal lobe, these forms have not yet been clearly distinguished based on their morphology. 

Utilisation: Retained for their fins, meat and cartilage. Population declines recorded in 

some parts of the Indo–West Pacific. 


Local synonyms: Zygaena indica van Hasselt, 1823. 

References: Stevens & Lyle (1989); White et al. (2008). 

2 

1 

Ventral head 

In Borneo: Probably widely distributed, 

but few records from southern Kalimantan. 


Upper tooth 

7 

Lower tooth


Great Hammerhead 

Sphyrna mokarran (Rüppell, 1837) 

3 

4 

6 

1 

5 

Other names: None (English), Hiu Capil, Hiu Bingkoh, Hiu Caping, Hiu Martil, Hiu 

Parang (Indonesian), Yu Tukul, Yu Parang (Malaysian) 


1 head broad, its width less than a third of total length 

2 anterior margin of head nearly straight, shallowly indented at midline 

3 1 st dorsal fin very tall, strongly falcate in adults 

4 2 nd dorsal fin tall with short rear tip and strongly concave posterior margin 

5 anal-fin base longer than 2 nd dorsal-fin base 

6 upper precaudal pit forming a crescentic groove 

7 upper teeth triangular and oblique, with serrations 

Size: To at least 450 cm (although some records to 600 cm); in Australia, males and 

females mature at about 225 and 210–228 cm (mature at a much larger size in South 

African populations); born at 50–70 cm. 


Habitat and biology: Continental and insular shelves and semi-oceanic, from the 

surface, and from very shallow water, to at least 80 m depth. Viviparous, with a yolksac 

placenta; gives birth to litters of 6–42 pups after a gestation period of 11 months. 

Diet consists primarily of bony fishes and elasmobranchs, but also cephalopods and 

crustaceans. Potentially dangerous to humans and known to intimidate spear fishers 

occasionally, but most often timid when observed. 

Utilisation: Retained for their fins, meat and cartilage. Large population declines


recorded in some areas. 



References: Stevens & Lyle (1989). 

2 

1 

Ventral head 

In Borneo: Recorded from Sabah and 

Sarawak, but probably widely distributed. 


Upper tooth 

7 

Lower tooth


Narrow Sawfish 

Anoxypristis cuspidata (Latham, 1794) 

3 

1 

2 

Other names: Pointed Sawfish (English), Cucut Gergaji, Hiu Parang, Pamprang, 

Parangpang (Indonesian), Yu Gergaji Jarang, Yu Parangan, Beroi, Kan Sua, Yu Gergaji, Yu 

Parang, Yu Todak (Malaysian) 


1 1 st dorsal-fin origin well posterior to pelvic-fin origins 

2 ventral lobe of caudal fin well developed 

3 rostral saw very long and narrow 

4 18–23 pairs of rostral teeth 

5 rostral teeth absent from base of saw 

6 rostral teeth slightly closer together near tip than at middle of saw 

Size: To at least 350 cm, records to 600 cm very doubtful; males and females mature at 

about 200 and 225 cm respectively; born at about 70 cm. 

Distribution: Tropical Indo–West Pacific, from the Red Sea to New Guinea, including 

tropical eastern and western Australia, and southern Japan. 

Habitat and biology: Demersal in coastal embayments to well offshore (to at least 

40 m depth). Juveniles mainly coastal, shallower than 10 m. Viviparous, with yolk-sac 

dependency; gives birth to litters of about 15 pups after a 4–5 month gestation. Females 

and males mature at 4 and 5 years respectively. Diet probably consists of small fishes and 

invertebrates. 

Utilisation: Still caught occasionally by demersal tangle net and trawl fisheries in the 

Arafura Sea but possibly extinct in parts of the Indo–Pacific. Presumably once used for 

its meat, skin and cartilage. The rostral saw has been traditionally kept as a curio.

P r i s t o i d e i 141 

Conservation status: IUCN Red List: Critically Endangered; Appendix I of CITES. 


References: Manjaji (2002a, b). 

Caudal fin 

2 

In Borneo: Single records from southern 

Kalimantan and near Sandakan, northeastern 

Sabah. 

Pristidae (Sawfishes) 

6 

4 5 

Ventral head


Freshwater Sawfish 

Pristis microdon Latham, 1794 

3 

1 

2 

Other names: Great-tooth Sawfish, Wide Sawfish (English), Hiu Gergaji, Hiu 

Parang (Indonesian), Yu-gergaji Seragam, Yu Parangan, Yu Gergaji, Yu Parang, Yu Todak 

(Malaysian) 


1 1 st dorsal-fin origin well forward of pelvic-fin origins 

2 ventral lobe of caudal-fin short, but conspicuous 

3 rostral saw short and broad 


5 rostral teeth present near base of saw 

6 rostral teeth near tip and at middle of saw a similar distance apart 

Size: Reported to reach about 700 cm, but probably closer to 600 cm; both sexes 

maturing at 240–300 cm; born at 70–90 cm. 

Distribution: Tropical Indo–West Pacific, from India to New Guinea, including 

Australia; poorly defined, possibly more widespread. 

Habitat and biology: Lives on or near the bottom in estuaries, rivers and lakes, as well 

as coastal marine waters, to depths of at least 60 m; elsewhere, juveniles are most common 

in freshwater with large adults in estuaries or the sea. Probably breeds in estuaries. 

Viviparous, with yolk-sac dependency; females, which take almost a decade to reach 

maturity, have litters of up to 12 pups after a 5 month gestation. Diet probably changes 

during its life, consisting mainly of small fishes and demersal invertebrates. Uses saw to 

stun prey, and can also inflict serious injury to humans unless handled with care. 

Utilisation: Occasionally caught in the Kinabatangan River using demersal gillnets by 

artisanal fishers. Highly susceptible to tangle nets and extant populations are likely to be


seriously depleted. This form of fishing may have been responsible for local extinctions 

in other rivers of the region. Once used widely for its fins and meat (both of very high 

value), and skin and cartilage. The rostral saw has been traditionally kept as a curio. 

Conservation status: IUCN Red List: Critically Endangered; Appendix II of CITES. 


References: Manjaji (2002a, b). 

Caudal fin 

2 

In Borneo: Occurs in the Kinabatangan 

River; possibly once more widespread in 

larger rivers and adjacent marine bays of 

the region. 


6 

4 5 

Ventral head


Green Sawfish 

Pristis zijsron Bleeker, 1851 

3 

1 

2 

Other names: Narrowsnout Sawfish, Sawfish (English), Hiu Gergaji, Hiu Parang 

(Indonesian), Yu-gergaji Silih, Yu Parangan, Yu Gergaji, Yu Parang, Yu Todak (Malaysian) 


1 1 st dorsal-fin origin slightly posterior to pelvic-fin origins 

2 no obvious ventral caudal-fin lobe 

3 rostral saw moderately long and slender 


5 rostral teeth present near base of saw 

6 rostral teeth much closer together near tip than at mid-length of saw 

Size: To at least 540 cm, possibly 730 cm; both sexes mature at about 300 cm; born at 

about 80 cm. 

Distribution: Tropical Indo–West Pacific, from southeastern Africa to New Guinea 

and Australia, north in the Pacific to Vietnam. 

Habitat and biology: Demersal in coastal bays and estuaries, also well offshore. Viviparous, 

with yolk-sac dependency; gives birth to litters of about 12 pups. Matures at about 

9 years in Australian populations. Diet consists of small schooling fishes and demersal 


Utilisation: Now caught rarely by demersal gill net and trawlers in the Indonesian 

Archipelago. Apparently extinct through much of its range. Once, used widely for its 

meat, fins, skin and cartilage. The rostral saw has been traditionally kept as a curio. 

Conservation status: IUCN Red List: Critically Endangered; Appendix I of CITES. 

Local synonyms: Pristis dubius Bleeker, 1852. 

References: Manjaji (2002a, b).


6 

Caudal fin 

2 

In Borneo: Recent records from central 

Sarawak (near Bintulu), and northeastern 

(Labuk River) and eastern Sabah (Kinabatangan 

River); holotype collected in the 

19 th century from southern Kalimantan. 


4 5 

Ventral head


Shark Ray 

Rhina ancylostoma Bloch & Schneider, 1801 

3 

5 

1 

2 

Other names: Bowmouth Guitarfish, Mud Skate (English), Hiu Barong, Kupukupu 

Brangkas, Yunbun Karang (Indonesian), Pari-kemejan Kepala Keras, Yu Kemejan 

(Malaysian) 


1 1 st dorsal-fin origin slightly anterior to pelvic-fin origin 

2 caudal fin lunate, upper and lower lobes almost symmetrical 

3 head and snout thick, margin broadly rounded 

4 spiracles without skin folds on posterior margin 

5 ridges on orbits, mid-body and shoulders bearing clusters of strong thorns 

Size: To at least 270 cm; males mature at about 150–175 cm; a 51 cm individual was a 

newborn. 

Distribution: Widespread in the Indo–West Pacific, from southern Africa to New 

Guinea and Australia, and north to southern Japan. 

Habitat and biology: Demersal on soft substrates near coastal areas and reefs; to at 

least 70 m, but depth distribution is poorly defined. Viviparous, with yolk-sac dependency; 

biology poorly known, one pregnant female contained 9 mid-term embryos 

(27–31 cm). Diet consists primarily of bottom-dwelling crustaceans and molluscs. 

Handling large adults can be hazardous. They create a nuisance to trawl fishers as they 

are difficult to remove and their roughness can damage smaller species in the catch.

R h i n o i d e i , R h y n c h o b a t o i d e i & R h i n o b a t o i d e i 147 

Utilisation: Caught occasionally by demersal tangle net, and possibly by trawl and 

longline fisheries. Used throughout the region for fresh and dried-salted meat, and their 

high-value fins. 



References: Devadoss & Batcha (1995); Frimodt (1995). 

5 

4 

Lateral head 

In Borneo: Known from coastal areas of 

Sabah, Sarawak, and western and southern 

Kalimantan. 

Rhinidae (Shark Rays) 

3 

Ventral head


Whitespotted Wedgefish 

Rhynchobatus australiae Whitley, 1939 

6 

5 

2 

3 

1 

4 

4 

8 

Other names: Whitespotted Guitarfish, Whitespot Ray, Whitespot Shovelnose Ray 

(English), Paredung, Pangrum, Hiu Lontar, Liongbun, Petong, Hiu Kemenyan, Hiu 

Minsong (Indonesian), Pari-kemejan Tompok Putih, Yu Kia-kia (Malaysian) 


1 origin of 1 st dorsal-fin slightly posterior to pelvic-fin origin 

2 a diffuse-edged, black spot on each pectoral fin of juveniles (less than 60 cm), 

becoming faint or absent in large adults 

3 diagonal row of three equidistant white spots usually above black pectoral spot 

4 1 st dorsal fin less than 1.5 times height of 2 nd (in adults) 

5 no large dark spots over or behind eyes 

6 snout bottle-shaped, constricted slightly near tip 

7 spiracles with two skin folds on posterior margin 

8 caudal fin with a well-developed lower lobe 



Distribution: Tropical Indo–West Pacific from Taiwan to northern Australia, including the 

Philippines; probably wider ranging in the Indian Ocean but currently not well defined 

and possibly includes additional unidentified species. 

Habitat and biology: Demersal on soft bottoms near the coast and sand patches on 

coral reefs, inshore to depths of at least 60 m. Viviparous, with yolk-sac dependency; 

gives birth to litters of 7–19 pups after an unknown gestation period; no apparent 

reproductive synchronicity. Diet consists primarily of large demersal crustaceans and 

molluscs. In the past, has been confused with another large wedgefish, R. djiddensis, from


the Red Sea and western Indian Ocean. 

Utilisation: Caught most commonly by demersal tangle net, and occasionally trawl and 

longline fisheries. Sought after for its fins and meat, which are both highly valued. 

Conservation status: IUCN Red List: Vulnerable; of long-term conservation concern. 



6 

Ventral head 

In Borneo: Mainly western and northern 

Borneo. 

Rhynchobatidae (Wedgefishes) 

7 

Dorsal view of eyes 

and spiracles


Smoothnose Wedgefish 

Rhynchobatus laevis (Bloch & Schneider, 1801) 

6 

5 

2 

1 

4 

4 

8 

3 

Other names: Giant Guitarfish, Sandshark, Shovelnose Shark (English), Paredung, 

Pangrum, Hiu Lontar, Liongbun, Petong, Hiu Kemenyan, Hiu Minsong (Indonesian), 

Pari Kemejan, Yu Kia-kia (Malaysian) 


1 origin of 1 st dorsal-fin almost over pelvic-fin origin 

2 a sharp-edged, black spot on each pectoral fin of juveniles (less than 60 cm), often 

forming a ring in large adults 

3 black pectoral spot usually closely surrounded by 3 white spots (no diagonal upper 

row of spots) 

4 1 st dorsal fin exceeding 1.5 times height of 2 nd (in large adults) 

5 no large dark spots around eyes 

6 snout not obviously constricted near tip 



Size: Possibly to at least 250 cm, maybe larger; size of maturity and birth presently 

unknown. 

Distribution: Considered to be widespread in the tropical Indo–West Pacific, from 

southeastern Africa to Indonesia, and north to southern Japan. Populations across this 

broad region may represent different species; probably not occurring in Australian seas. 

Habitat and biology: Demersal on soft bottoms and near coral reefs, as well as inshore 

and offshore on the continental shelf. Viviparous, with yolk-sac dependency; 

reproduction not well known. Diet probably consists of demersal invertebrates and 

small fishes.


Utilisation: Caught occasionally by demersal tangle net, trawl and longline fisheries. 

Sought after for its fins and meat, which are both highly valued. 

Conservation status: IUCN Red List: Vulnerable; of long-term conservation concern. 



6 

Ventral head 

In Borneo: Recorded from off northeastern 

Sabah. 


7 


and spiracles


Broadnose Wedgefish 

Rhynchobatus springeri Compagno & Last, 2010 

6 

2 

4 

5 1 

4 

3 

8 

Other names: None (English), Paredung, Pangrum, Hiu Lontar, Liongbun, Petong, 

Hiu Kemenyan, Hiu Minsong (Indonesian), Pari Kemejan, Yu Kia-kia (Malaysian) 


1 origin of 1 st dorsal-fin above or slightly behind pelvic-fin origin 

2 a large, sharp-edged, black spot on each pectoral fin of juveniles and adults 

3 black pectoral spot usually closely surrounded by 4–5 white spots (no diagonal upper 

row of spots) 

4 1 st dorsal fin less than 1.5 times height of 2 nd (in adults) 

5 usually with dark markings on and/or behind eyes 

6 snout broad with a convex margin 



Size: Adult males to at least 130 cm, females possibly larger. 

Distribution: Poorly defined, probably Thailand, Malaysia, Philippines, Singapore and 

Indonesia. 

Habitat and biology: Demersal on soft bottoms in coastal bays and estuarine 

habitats, otherwise poorly known. Probably viviparous, with yolk-sac dependency. Diet 

unknown, but most wedgefishes feed primarily on demersal crustaceans, molluscs and 

small fishes. 

Utilisation: Caught occasionally by tangle net fishers. Utilised for its meat, fins 

(extremely high value) and cartilage.


Conservation status: IUCN Red List: Not Evaluated; of long-term conservation 

concern. 


References: Compagno & Last (2010). 

6 

Ventral head 

In Borneo: A few records from brackish 

habitats of southern Sarawak. 


7 


and spiracles


Clubnose Guitarfish 

Glaucostegus thouin (Anonymous, 1798) 

1 

4 

2 

3 

6 

Other names: None (English), Cucut Mandrong, Liongbun (Indonesian), Parikemejan 

Muncung Panjang, Yu Kemejan, Yu Kia-kia (Malaysian) 


1 snout tip enlarged to form an elongate knob 

2 dorsal surface plain, without blotches or spots 

3 denticles along midline of dorsal surface distinct 

4 clear part of snout sharply demarcated from rest of head near eyes 

5 nostrils broad and slit-like with narrow nasal flaps 

6 lower lobe of caudal fin short 

Size: To about 300 cm; largest observed from the region 240 cm. 

Distribution: Widespread in the Indo–West Pacific from the Red Sea through Thailand, 

Vietnam, Malaysia, Indonesia, New Guinea and Japan; apparently not known from 

Peninsula Malaysia or nearby Philippines. 

Habitat and biology: Demersal on insular and continental shelves to about 100 m 

depth; local depth information not reliable, reported from inshore but probably more 

abundant offshore on the continental shelf. Biology poorly known, presumably viviparous, 

with yolk-sac dependency. Diet consists of benthic invertebrates and small fishes. 

Utilisation: Caught infrequently by local demersal tangle net and trawl fisheries. Used 

opportunistically for its meat and fins (both very high value), as well as skin and cartilage; 

an esteemed food fish in India. Likely to become a major conservation concern if fished 

at existing levels. 

Conservation status: IUCN Red List: Vulnerable.


Local synonyms: Rhinobatos thouin (Anonymous [Lacepède]): Yano et al. (2005), White 

et al. (2006b), also misspelled as Rhinobatos thouini. 


5 

Ventral head 

In Borneo: Known from Sarawak, 

Sabah and southeastern and western 

Kalimantan. 

Rhinobatidae (Shovelnose Rays) 

Lateral tail and dorsal fins 

6


Giant Guitarfish 

Glaucostegus typus (Bennett, 1830) 

1 

4 

2 

3 

6 

Other names: Common Shovelnose Ray, Giant Shovelnose Ray, Shovelnose Ray, 

Shovelnose Shark (English), Hiu Cermin, Paitpait, Pari Bandrong, Pari Gitar, Pari 

Kekeh, Petong (Indonesian), Pari-kemejan Muncung Pendek, Yu Pari (Malaysian) 


1 snout tip not forming a knob 

2 dorsal surface plain, without blotches or spots 

3 denticles along midline of dorsal surface distinct 

4 clear part of snout sharply demarcated from rest of head near eyes 

5 nostrils broad and slit-like with narrow nasal flaps 


Size: To about 270 cm, reports to 400 cm are likely to be erroneous; maturing at 150– 

180 cm; born at 38–40 cm. 

Distribution: Widespread in the Indo–West Pacific from India through Indonesia to 

northern Australia and possibly Melanesia; north in the Pacific to Thailand but now 

apparently absent in the Philippines. 

Habitat and biology: Demersal on insular and continental shelves, also known from 

estuaries and sometimes in freshwater; juveniles occur inshore around mangroves and 

atolls, adults occur further offshore to depths of about 100 m. Viviparous, with yolk-sac 

dependency; biology poorly known; reportedly able to breed in freshwater. Diet consists 

largely of shellfish (prawns and crabs), as well as small fishes and cephalopods. 

Utilisation: Caught often by demersal tangle net fisheries, and sometimes by trawling.


Marketed for its meat and fins (both very high value), and skins and cartilage; sold fresh 

and possibly dried and salted. 


Local synonyms: Rhinobatos typus Bennett: Yano et al. (2005), White et al. (2006b). 


5 

Ventral head 

In Borneo: Primarily from eastern 

Borneo, also known from Sarawak and 



Lateral tail and dorsal fins 

6


Taiwan Guitarfish 

Rhinobatos formosensis Norman, 1926 

1 

2 

3 

4 6 

Other names: None (English), none (Indonesian), Yu Kemejan (Malaysian) 


1 snout tip not forming a knob 

2 dorsal surface usually plain (often with faint orange blotches) 

3 denticles along midline of dorsal surface very small 

4 clear part of snout not sharply demarcated from rest of head near eyes 

5 nostrils with broad apertures and wide nasal flaps 


7 young covered with small ocelli 

Size: To at least 84 cm; males mature by 64 cm; size at birth unknown (late term 

embryos up to 20 cm in length). 

Distribution: Known from off Malaysian Borneo, the Philippines and Taiwan. 

Habitat and biology: Demersal inshore; biology and depth information largely 

unknown. Presumably viviparous, with yolk-sac dependency as with other members 

of the family. Diet probably consists mainly of bottom-dwelling invertebrates such as 

marine worms and shellfishes. 

Utilisation: Taken occasionally as bycatch of inshore trawl fisheries. Too rarely caught 

to be of value; probably used for its meat when landed. 


Local synonyms: Rhinobatos cf. schlegelii Müller & Henle: Compagno et al. (2005b). 

References: Compagno et al. (2005b).


5 

Ventral head 

7 


In Borneo: Off Sarawak and Sabah. 


1 st dorsal fin 2 nd dorsal fin


Shortlip Numbfish 

Narcine brevilabiata Bessednov, 1966 

1 

3 

4 

2 

5 

Other names: Shortlip Electric Ray (English), Pari Listrik, Pari Kontak (Indonesian), 

Pari Karan Tompok Besar (Malaysian) 


1 disc subcircular, thick and flabby 

2 eyes raised, not embedded in skin 

3 two similar-sized dorsal fins 

4 dark spots on upper snout numerous, distinctly smaller than eye 

5 3–5 pairs of very large, dark blotches on disc 

6 exposed part of tooth bands in jaws relatively broad 

7 mouth subequal to or only slightly narrower than width between lateral edges of 

nostrils 

8 ventral surfaces of pelvic and pectoral fins often broadly dark edged 

Size: To at least 32 cm. 

Distribution: Known from the South China Sea in the Northwest Pacific, from 

Malaysia to China; possibly extends north to the East China and Yellow Seas. 

Habitat and biology: Demersal on the inner continental shelf, known from depths of 

41–70 m. Biology largely unknown, presumably viviparous, with yolk-sac dependency.

T o r p e d i n o i d e i 161 

Diet presumably consists of small fishes and benthic invertebrates; mouth can protrude 

as a tube to extract prey from the substrate. Like other members of the genus, the 

electric organs in the disc are capable of delivering a moderate shock on direct contact. 

Utilisation: Caught regularly as bycatch of bottom trawling. A low-grade food species 

and used for fish meal. 



References: Carvalho (1999). 

7 

6 

Mouth and nostrils 

In Borneo: Off Sarawak and eastern 

Sabah. 

Narcinidae (Numbfishes) 

Juvenile (14 cm)


Rough Numbfish 

Narcine lingula Richardson, 1846 

1 

4 

2 

5 

3 

Other names: Chinese Numbfish, Rough Electric Ray (English), Pari Listrik, Pari 

Kontak (Indonesian), none (Malaysian) 


1 disc subcircular to oval, thick and flabby 

2 eyes not embedded in skin 


4 dark spots on upper snout usually sparse, size similar to eye (atypically smaller) 

5 upper disc with greatly enlarged, symmetrically arranged blotches 

6 exposed part of tooth bands in jaws relatively narrow 

7 mouth subequal to or only slightly narrower than width between lateral edges of 

nostrils 

8 ventral surfaces of pelvic and pectoral fins not dark edged 

Size: To at least 38 cm; males mature at about 24 cm. 

Distribution: Patchily distributed in the Indo–West Pacific, possibly from eastern India 

to Hong Kong, and south to Java; may consist of more than a single species each with 

more restricted ranges. 

Habitat and biology: Demersal in shallow inshore waters on the continental shelf. 

Biology unknown, presumably viviparous, with yolk-sac dependency. Diet is likely to 

consist of small fishes and invertebrates. Numbfishes of the region are not well known 

and some names currently used may change as more information becomes available.


Like other members of the genus, the electric organs in the disc are capable of delivering 

a moderate shock on direct contact. 

Utilisation: Caught occasionally in the region by bottom trawl. Not considered for 

food and used for fish meal. 



References: Carvalho (1999); Carvalho et al. (1999). 

7 

6 


In Borneo: Uncommon locally, known 

from southern Sarawak and eastern 

Sabah. 

Narcinidae (Numbfishes)


Darkspotted Numbfish 

Narcine maculata (Shaw, 1804) 

1 

2 

5 

3 

4 

Other names: Darkfin Numbfish, Darkspotted Electric Ray (English), Pari Listrik, 

Pari Kontak (Indonesian), Pari-karan Hitam, Pari Karan, Pari Letrik, Pari Serban 

(Malaysian) 


1 disc subcircular, thick and flabby 

2 eyes not embedded in skin 


4 dark spots on upper snout similar in size to eye 

5 upper disc without greatly enlarged, symmetrically arranged blotches 

6 exposed part of tooth bands in jaws relatively narrow 

7 mouth usually narrower than width between lateral edges of nostrils 

8 ventral surfaces of pelvic and pectoral fins not broadly dark edged 


Distribution: Widespread in the Indo–West Pacific, from western India to China, 

south to Java; possibly wider ranging. 

Habitat and biology: Demersal inshore and offshore on the continental shelf, known 

from depths to about 70 m. Biology largely unknown, presumably viviparous, with


yolk-sac dependency. Diet probably consists of small fishes and benthic invertebrates; 

mouth can protrude as a tube to extract prey from the substrate. Like other members of 

the genus, the electric organs in the disc are capable of delivering a moderate shock on 

direct contact. 

Utilisation: Only caught rarely by bottom trawl. Considered as low grade for food and 

used for fish meal. 



References: Carvalho (1999); Carvalho et al. (1999). 

7 

6 


In Borneo: Recorded locally from 

southern Sarawak; another specimen 

recorded in literature from unspecified 

locality off Sabah. 

Narcinidae (Numbfishes)


Spot-tail Sleeper Ray 

Narke dipterygia (Bloch & Schneider, 1801) 

2 

4 

3 

1 

5 

Other names: Numbray, Spot-tail Electric Ray (English), Pari Listrik, Pari Kontak 

(Indonesian), Pari Karan (Malaysian) 


1 one dorsal fin 

2 eyes very small, partly embedded in skin 

3 disc moderately enlarged, its length (measured to cloaca) slightly longer than tail 

4 spiracle lateral to eye 

5 caudal fin longer than deep 

6 nostrils slit-like 

7 mouth very small and not strongly arched 

8 undersurfaces of disc and pelvic fin uniformly pale 

Size: To at least 18 cm; males mature by about 15 cm. 

Distribution: Tropical Indo–West Pacific, from Oman to Malaysia and north to Japan; 

may occur off Sumatra. 

Habitat and biology: Demersal on the continental shelf, both inshore and offshore; 

locally in 70–95 m depth. Biology largely unknown, presumably viviparous, with yolksac 

dependency. Diet also unknown but probably consists of small invertebrates, such as 

polychaetes that live in mud and sand.


Utilisation: Infrequent bycatch of both inshore and offshore bottom trawl fisheries 

operating over soft bottoms on the continental shelf. Not used for food. 



References: Compagno & Last (1999b). 

6 

7 


In Borneo: Off central Sarawak. 

Narkidae (Sleeper Rays) 

8 

8 

Ventral


Finless Sleeper Ray 

Temera hardwickii Gray, 1831 

4 

3 

2 1 

5 

Other names: Finless Electric Ray (English), Pari Listrik, Pari Kontak (Indonesian), 

Pari Karan Belangkas, Pari Karas (Malaysian) 


1 no dorsal fins 

2 eyes bulging, not embedded in skin 

3 disc greatly enlarged, its length (measured to cloaca) much longer than tail 

4 spiracle posterior to eye 

5 caudal fin about as long as deep 

6 nostrils almost circular 

7 mouth small and not strongly arched 

8 undersurfaces of disc and pelvic fin with broad brownish margins 

Size: Dubiously reported to about 46 cm, but mostly less than 15 cm; both sexes 

mature at about 11–12 cm. 

Distribution: Tropical Indo–West Pacific, from the Andaman Sea off Myanmar to 

Singapore, and Vietnam. 

Habitat and biology: Demersal on the continental shelf. Biology poorly known, 

viviparous, with yolk-sac dependency; one pregnant female had 4 pups. Diet presumably 

consists of small invertebrates. Paired electric organs in the disc are capable of delivering


a moderate shock on direct contact. 

Utilisation: Caught rarely by bottom trawl on the continental shelf. Not used for food. 



References: Compagno & Last (1999b). 

6 

7 


In Borneo: Single record from the South 

China Sea off southwestern Sarawak. 

Narkidae (Sleeper Rays) 

8 

8 

Ventral


Kwangtung Skate 

Dipturus kwangtungensis (Chu 1960) 

3 

2 

5 

1 

6 

7 

4 

Other names: None (English), none (Indonesian), Pari Belanda (Malaysian) 


1 pelvic fin divided into two distinct lobes 

2 snout supported by a firm cartilage 

3 snout narrowly triangular, long (preorbital length 4–5 times orbit diameter) 

4 dorsal surface dark brown with cloudy pattern of pale blotches surrounded by spots 

5 axil of pectoral fin with similar pattern to rest of disc 

6 thorns on tail confined to midline 

7 dorsal fins not widely separated 

8 most of ventral disc dark brown 

Size: To at least 50 cm; size of maturity unknown. 

Distribution: Found in the western North Pacific, from Japan, the Korean Peninsula, 

Taiwan to Borneo. 

Habitat and biology: Demersal on the continental shelf, probably on silty or muddy

R a j o i d e i 171 

bottoms. Oviparous, females likely to produce small egg cases with horny extensions at 

each corner. Diet probably consists of small crustaceans. 

Utilisation: Caught rarely as a bycatch of trawl fisheries. 


Local synonyms: Raja (Okamejei) hollandi Jordan & Richardson: Yano et al. (2005). 

References: Ishihara (1987). 

8 

1 

In Borneo: Off Sarawak. 

Rajidae (Skates) 

Ventral disc (female) 

7 

Lateral posterior tail


Borneo Sand Skate 

Okamejei cairae Last, Fahmi & Ishihara, 2010 

4 

3 

2 

1 

6 

7 

5 

Other names: None (English), Pari, Pari Luncur (Indonesian), Pari-raja Tompok 

(Malaysian) 




3 snout broadly triangular, quite long (preorbital length ~3 times orbit diameter) 

4 dorsal surface yellowish brown with clusters of minute black flecks 

5 dark ring-like marking near axil of pectoral fin 

6 thorns on tail minute 

7 dorsal fins widely separated 

8 ventral disc margin distinctly paler than central part 

Size: To at least 39 cm; males maturing at about 34 cm. 

Distribution: Possibly endemic to the South China Sea. 

Habitat and biology: Demersal over sandy and muddy bottoms on the continental 

shelf. Oviparous, producing small, rectangular egg cases with horny extensions at each 

corner. Diet unknown, but probably consists of small crustaceans, worms and molluscs.


Similar forms occur in the Western Central Pacific, from the East China Sea to Java 

(eastern Indonesia). Depth range not well defined, but probably mainly on the mid 

continental shelf (~70-150 m). 

Utilisation: Caught as occasional bycatch of trawl fisheries and utilised solely for its 

meat. 


Local synonyms: ?Okamejei boesemani (Ishihara): Compagno et al. (2005b); Raja 

(Okamejei) boesemani Ishihara: Yano et al. (2005). 

References: Last et al. (2010a). 

8 

1 



In Borneo: Off western Borneo, including 

Sabah, Sarawak and northwestern Kalimantan. 

6 

7 

Lateral tail


Yellowspotted Skate 

Okamejei hollandi (Jordan & Richardson, 1909) 

4 

3 

1 

2 

6 7 

5 

Other names: Holland Skate, Thornback Ray (English), Pari Luncur (Indonesian), Pari 

Belanda (Malaysian) 





4 dorsal surface brownish with pale blotches, black flecks (when present) not in welldefined 

clusters 

5 axil of pectoral fin with a pale-edged brown blotch 

6 thorns on tail not minute 

7 dorsal fins very widely separated 

8 most of ventral disc medium to dark brown 

Size: To at least 45 cm; males maturing at about 30 cm. 

Distribution: Western Pacific, from southern Japan, East China Sea, Taiwan, and the 

South China Sea. 

Habitat and biology: Demersal over sandy and muddy bottoms on the continental


shelf. Oviparous, females produce small, egg cases (up to 7 x 4 cm) with horny 

extensions at each corner. Diet consists of small crustaceans and bony fishes. Occurs with 

O. cairae but appears to be less common. Depth range not well defined, but probably 

mainly on the mid continental shelf (~70–150 m). 

Utilisation: Caught as an occasional bycatch of trawl fisheries and utilised solely for its 

meat. 


Local synonyms: Figured (Plate 211) as Raja (Okamejei) boesemani Ishihara: Yano et al. 

(2005). 


8 


1 


In Borneo: Off western Borneo, including 

Sarawak and Kalimantan. 

6 7 

Lateral tail


Philippine Ocellate Skate 

Okamejei jensenae Last & Lim, 2010 

4 

3 

2 

1 

6 

7 

5 

Other names: Yellow-spotted Skate (English), none (Indonesian), Pari Belanda 

(Malaysian) 





4 dorsal surface brownish with scattered yellowish spots 

5 pair of large dark spots with white-spotted borders (removed with mucous when 

washed) 

6 thorns on tail small 

7 dorsal fins relatively close together 

8 outer half of ventral disc distinctly darker than central part 

Size: To at least 37 cm; male size unknown but immature at 24 cm. 

Distribution: Probably endemic to the Indo-Malay Archipelago, in the Sulu and 

Celebes seas.


Habitat and biology: Demersal over soft bottoms on the continental shelf. Like all 

skates, presumably oviparous, producing small, rectangular egg cases. Diet also unknown, 

but probably consists of small benthic invertebrates. 

Utilisation: Caught occasionally as bycatch by trawlers and sold opportunistically. 


Local synonyms: Okamejei sp. 1: Compagno et al. (2005b). 

References: Compagno et al. (2005b); Last & Lim (2010). 

8 

1 


In Borneo: Off Lahad Datu, eastern 

Sabah. 


7 

Lateral posterior tail


Borneo Leg Skate 

Sinobatis borneensis (Chan, 1965) 

3 

2 

4 

5 

Other names: None (English), none (Indonesian), none (Malaysian) 


1 pelvic fin divided into two leg-like lobes 

2 snout floppy, supported by a thin, flexible cartilage 

3 snout very long, pointed (preorbital length more than 5 times orbit diameter) 

4 tail very slender, lacking dorsal fins 

5 dorsal surface of disc distinctly paler than ventral surface 


Distribution: Narrow-ranging in the tropical Indo-West Pacific, off Taiwan, the 

Philippines and in the South China Sea, off Malaysian Borneo. 

Habitat and biology: Occurs demersally on the continental slope in 475–835 m 

depth. Probably oviparous, biology poorly known. 

Utilisation: Caught rarely and not used for food.



Local synonyms: Previously as Anacanthobatis borneensis Chan. 

References: Last & Compagno (1999). 


In Borneo: Known from the holotype 

collected in the South China Sea, off 

Sarawak. 

Anacanthobatidae (Leg Skates) 

1 

Ventral


Giant Stingaree 

Plesiobatis daviesi (Wallace, 1967) 

1 

2 

3 

4 

5 

Other names: Deepwater Stingray (English), Pari (Indonesian), Pari Lembik Merah, 

Pari Merah (Malaysian) 


1 tail with a long, thin, lobe-like caudal fin 

2 disc upper surface with small granular denticles 

3 snout very long, >6 times orbit diameter 

4 no dorsal fin, or skin folds on side or undersurface of tail 

5 flesh flabby 


respectively; born at about 50 cm. 

Distribution: Tropical Indo–West Pacific, from southeastern Africa to New Caledonia, 

and from the Central Pacific (Hawaii). 

Habitat and biology: Mainly soft silty and muddy bottoms on continental slopes at 

depths of 275–680 m. Presumably viviparous, with histotrophy; very little known of its

M y l i o b a t o i d e i 181 

biology. Diet consists primarily of small fishes, cephalopods and crustaceans. Its long, 

flexible snout enables it to feed in soft sediments; occasionally feeds on mesopelagic 

species suggesting that it might also catch prey well off the bottom. Should be handled 

carefully because of its long, sharp sting. 

Utilisation: Caught infrequently by demersal longline fisheries in deepwater. Used for 

its meat, but not highly regarded. 





In Borneo: Known only from off Sarawak. 

Plesiobatidae (Giant Stingarees) 

1 

Lateral posterior tail with caudal fin


Smalleye Stingray 

Dasyatis microps (Annandale, 1908) 

2 

6 

1 

5 

4 

3 

Other names: Thickspine Giant Stingray (English), Pari Sapi, Pari Babi (Indonesian), 

Pari Gergasi (Malaysian) 


1 no enlarged thorns along central disc or tail (covered instead with minute spiny 

denticles) 

2 disc very broad and strongly rhomboidal with very angular apices 

3 pinkish brown above with rows of white spots on each side of pectoral fins 

4 skin folds on tail reduced to low ridges (occasionally indistinct) 

5 tail very broad based, tapering markedly near sting 

6 very small eyes and large spiracles 

Size: Attains at least 220 cm DW (>300 cm TL). 

Distribution: Patchy in the Indo–West Pacific, mainly from India to northern


Australia, but also from southeastern Africa. 

Habitat and biology: Mainly benthopelagic on the inner continental shelf to at least 

50 m depth; sometimes entering estuaries and river mouths. Biology unknown; 

presumably viviparous, with histotrophy, and diet probably consists of large crustaceans 

and small fishes. 

Utilisation: Caught very rarely in tangle nets. Used mainly for its meat, but uncommon 

in markets. 



References: Pierce et al. (2008). 

Dasyatidae (Stingrays) 

In Borneo: Northeastern Sabah.


Dwarf Black Stingray 

Dasyatis parvonigra Last & White, 2008 

2 

5 

1 

6 

Other names: None (English), Ikan Pari (Indonesian), Pari (Malaysian) 


1 thorns confined to central disc (single row on midline with shorter row either side) 

2 disc rhomboidal with somewhat angular apices 

3 short, low dorsal skin fold on tail beyond sting 

4 long, low ventral skin fold on tail 

5 tail moderately long, becoming much more slender beyond sting, but without banding 

6 no thorns on tail before sting 

7 ventral disc dark edged (not yellowish) 

Size: To at least 51 cm DW (~110 cm TL); males mature at about 35 cm DW. 

Distribution: Known only from northwestern Australia, Indonesia and Malaysia;


possibly also West Papua and the Philippines. 

Habitat and biology: Demersal on insular and continental shelves in 60–125 m. Diet 

and reproductive biology unknown; presumably viviparous, with histotrophy. 

Utilisation: Caught incidentally by the demersal trawl and trammel-net fisheries 

operating in shelf waters. Used for its meat. 


Local synonyms: Dasyatis cf. longa (Garman): White et al. (2006b); Dasyatis cf. longus 

(Garman): White & Dharmadi (2007). 

References: Last & White (2008). 

7 


Ventral disc 

In Borneo: Known from Sabah, including 

the eastern border with Kalimantan. 

1 

Nuchal area 

3 

4 

Lateral tail


Cow Stingray 

Dasyatis ushiei (Jordan & Hubbs, 1925) 

2 

5 

1 

6 

Other names: None (English), Pari Boro, Pari Jantung (Indonesian), Pari Ekor Duri 

(Malaysian) 


1 thorns extending along midline of disc and onto tail 

2 disc rhomboidal with angular apices 

3 low ridge-like dorsal skin fold on tail beyond sting 

4 long, low ventral skin fold on tail 

5 tail long, tapering in thickness beyond sting, but without banding 

6 cluster of large thorns on tail before sting in adults 

7 ventral disc dark edged (sometimes faint) 

Size: To at least 202 cm DW (~400 cm TL); males mature at about 117 cm DW; probably 

born at about 30–35 cm DW. 

Distribution: Not well defined, possibly western North Pacific to eastern Indonesia 

on insular and continental shelves.


Habitat and biology: Demersal on insular and continental shelves to at least 190 m 

(possibly to 360 m). Biology largely unknown; presumably viviparous, with histotrophy. 

Diet probably consists of small benthic fishes and invertebrates. 

Utilisation: Caught occasionally by demersal tangle net and longline fisheries 

operating in shelf waters. Used for its meat, cartilage and thorny tail. 


Local synonyms: Dasyatis cf. ushiei (Jordan & Hubbs): White et al. (2006b), White & 

Dharmadi (2007); Dasyatis cf thetidis Ogilby: Yano et al. (2005). 


Ventral disc 

7 

In Borneo: Known from off Sarawak 

and Sabah. 


1 

Nuchal area 

6 

3 

Lateral tail 

4


Sharpnose Stingray 

Dasyatis zugei (Müller & Henle, 1841) 

2 

7 

3 

1 

8 

Other names: Pale-edged Stingray (English), Pari Biasa, Toka-toka, Tuka-tuka 

(Indonesian), Pari Ketuka, Pari Nyiru, Pari Tanjung (Malaysian) 


1 a few small thorns along midline of disc 

2 disc with broadly rounded apices 

3 snout very elongate, pointed 

4 no oral papillae in mouth 

5 weak dorsal skin fold on tail beyond sting 

6 long, low ventral skin fold 

7 tail relatively short, tapering in thickness beyond sting and without banding 

8 row of small thorns on tail before sting in adults 

9 ventral disc pale or dark edged 

Size: To at least 29 cm DW (75 cm TL); males and females mature at about 16 and 

19 cm DW respectively; born at 7–10 cm DW.


Distribution: Indo–West Pacific, from India to eastern Indonesia (Bali) and northwards 

to southern Japan. 

Habitat and biology: Demersal on insular and continental shelves, to a depth of 

at least 40 m. Viviparous, with histotrophy; gives birth to litters of 1–4 pups after an 

unknown gestation period; no reproductive synchronicity. Diet consists primarily of 

shrimps, but includes other crustaceans and small fishes. 

Utilisation: Locally, landed in large quantities as bycatch of bottom trawling. One of 

the most commercially important rays in Borneo; used for fresh and dried meat. 





9 

Ventral disc 

In Borneo: Probably widespread, although 

currently few records from eastern 

Kalimantan. 

1 

Nuchal area 

5 

Lateral tail 

6


Pink Whipray 

Himantura fai Jordan & Seale, 1906 

5 

4 

3 

2 

6 

1 

7 

Other names: None (English), Pari Minyak, Pari Tembaga (Indonesian), Pari Pucat 

(Malaysian) 


1 no obvious band of denticles on central disc 

2 disc profile quadrangular with a short, broad snout 

3 upper surface uniformly yellowish or greyish pink (lacking an obvious pattern) 

4 no skin folds on tail 

5 tail long, whip-like and uniformly dark beyond sting (not banded) 

6 sting situated anteriorly on tail 

7 midline of tail before sting without enlarged thorny denticles 

Size: To at least 184 cm DW (500 cm TL); males mature at 108–115 cm DW; embryos 

of 68 cm DW have been recorded. 

Distribution: Poorly known, but probably widespread in the Indian Ocean and 

Western Pacific from southern Africa to Micronesia, including Australia.


Habitat and biology: Lives over soft substrates near the coast and on the inner 

continental shelf, from the intertidal zone to about 70 m depth, occasionally deeper 

to 200 m; schools near coral reefs and sand cays in some areas. Biology poorly known; 

viviparous, with histotrophy, and diet probably consists of small fishes and bottomdwelling 


Utilisation: Occasional catch of the demersal tangle net and bottom trawl fisheries. 

Used mainly for its meat. 


Local synonyms: Image of MSR659 in Yano et al. (2005) is of H. pastinacoides. 



Ventral disc 


Nuchal area


Whitespotted Whipray 

Himantura gerrardi (Gray, 1851) 

5 

4 

2 

3 

7 

1 

6 

8 

Other names: Sharpnose Stingray, Bluntnose Whiptail Ray, Banded Whiptail Ray 

(English), Pari Bintang, Pari Super, Pari Batu (Indonesian), Pari Bintik Putih, Pari Batik 

(Malaysian) 


1 widely spaced, granular denticles on central disc (absent in small juveniles) 

2 disc profile quadrangular 

3 upper surface of disc usually with numerous white spots (rarely plain greyish brown) 


5 tail long, whip-like, with alternating light and dark bands (rarely faint) 


7 central disc usually with 1–5 small thorns 

8 tail lacking thorns 

Size: Possibly to about 100 cm DW, rarely exceeding 85 cm DW (~ 220 cm TL); males


and females mature at 46–48 cm DW and about 64 cm DW respectively; born at 18– 

21 cm DW. 

Distribution: Widespread in the Indo–West Pacific from India to eastern Indonesia, 

north to Taiwan; possibly New Guinea but not Australia. 

Habitat and biology: Demersal on soft bottoms, from inshore to a depth of at least 

60 m. Viviparous, with histotrophy; gives birth to litters of 1–4 pups after an unknown 

gestation period; small individuals present in trawl catches in all months indicating no 

reproductive seasonality. Diet unknown, but presumably consists of bivalves, crustaceans 

and small fishes. Two distinct forms, detected by genetic methods, occur in Borneo but 

these need further resolution. 

Utilisation: Very important commercial species, caught in large quantities in the 

demersal tangle net, bottom trawl and trammel net fisheries. Used mainly for its highly 

regarded fresh meat and skin (valuable as leather). 


Local synonyms: Trygon macrurus Bleeker, 1852. 



Ventral disc 


1 

7 

Nuchal area


Mangrove Whipray 

Himantura granulata (Macleay, 1883) 

2 

6 

5 

7 

4 

9 

3 

1 

8 

Other names: Coachwhip Ray, Macleay’s Coachwhip Ray, Mangrove Ray, 

Mangrove Stingray, Whitetail Stingray, Whitetail Whipray (English), Pari Sapi 

(Indonesian), Pari Bakau (Malaysian) 


1 band of flat denticles on central disc in adults 

2 disc almost circular in profile 

3 snout broadly triangular, short 

4 upper surface greyish with small white flecks (often also with dark mucous) 


6 tail moderately long, whip-like, whitish posterior to sting 


8 central disc usually without enlarged thorns 

9 tail lacking thorns 

Size: To at least 141 cm DW (>350 cm TL); males mature at about 55–65 cm DW; 

born at about 14 cm DW.


Distribution: Possibly widespread in the Indo–West Pacific, from the Red Sea and 

Maldives to Micronesia and northern Australia. 

Habitat and biology: Demersal, in coastal waters but also inshore on the continental 

shelf to a depth of at least 85 m; juveniles occur mainly in mangroves and estuaries, 

adults on coral reefs. Viviparous, with histotrophy; biology little known. Diet consists 

primarily of crabs and prawns. 

Utilisation: Caught occasionally in the dryer northeastern part of Borneo by demersal 

tangle net and hook-and-line fisheries. Used fresh for its meat. 





In Borneo: Known from off Sabah and 

northeastern Kalimantan. 

Ventral disc 

1 

8 

Nuchal area


Jenkins’ Whipray 

Himantura jenkinsii (Annandale, 1909) 

5 

2 

4 

3 

6 

1 

7 

Other names: Rough-back Stingray (English), Pari Aer, Pari Duri, Pari Kikir 

(Indonesian), Pari Pasir, Pari Naga (Malaysian) 


1 band of flat denticles on central disc (poorly developed in young) 

2 disc profile quadrangular with a short, broad snout 

3 upper surface uniformly yellowish brown (rarely with small dark spots on disc near 

tail base) 


5 tail moderately long, whip-like, uniformly dark (not banded) 


7 central disc and tail with row of upright thorns 

Size: To at least 150 cm DW (about 300 cm TL); males mature at 70–85 cm DW; born 

at about 23 cm DW. 

Distribution: Widespread in the Indo–West Pacific, from southern Africa, Madagascar and 

the Red Sea through Pakistan to eastern Indonesia, the Philippines and northern Australia.


Habitat and biology: Demersal on sandy substrates, often in caves and under ledges, 

inshore to depths of at least 90 m. Viviparous, with histotrophy; biology little known. 

Diet presumably consists of crustaceans and small fishes. 

Utilisation: Occasional catch of the tangle net, bottom trawl and line fisheries. Used 

for fresh meat. 


Local synonyms: Himantura draco Compagno & Heemstra, 1984. 


Ventral disc 

In Borneo: Mainly known from Sabah 

and Sarawak; rarely caught off eastern, 

northwestern and southern Kalimantan. 


7 

Dorsal tail thorns 

1 

7 

Nuchal area


Roughback Whipray 

Himantura kittipongi Vidthayanon & Roberts, 2005 

6 

5 

3 

9 

1 

7 

8 

2 

Other names: None (English), Pari Sungai (Indonesian), Pari Sungai (Malaysian) 


1 broad denticle band on disc of adults (rough, often with scattered thorns) 

2 disc profile subcircular to oval with short, sharply pointed tip 

3 uniformly brownish above (without pattern but often spotted or blotched) 

4 whitish below with a broad dark outer margin 


6 tail long, narrow-based, whip-like, not banded 

7 single pearl thorn on central disc 

8 a row of slightly enlarged thorns on midline of tail 

9 eyes small 

Size: To at least 37 cm DW; males mature at about 25 cm DW. 

Distribution: Endemic to the Indo–Malay Archipelago; off Thailand, peninsula 

Malaysia and Borneo.


Habitat and biology: Demersal in fresh and brackish water, mainly on silty bottoms. 

Biology poorly known; viviparous, with histotrophy. Diet unknown; given its unusually 

small mouth, probably consists of small bottom-dwelling invertebrates, particularly small 

crustaceans. 

Utilisation: Caught incidentally by demersal tangle nets, traps, and hook and line. 

Possibly used for its meat when fresh; may be collected for the aquarium trade. 




4 


In Borneo: Known locally from the 

Pawan and Kapuas rivers, western 

Kalimantan. 

Ventral disc 

7 

Nuchal area


Leopard Whipray 

Himantura leoparda Manjaji-Matsumoto & Last, 2008 

5 

4 

3 

1 

8 

6 

2 

Other names: Undulate Whipray (English), Pari Macan (Indonesian), Pari Rimau 

(Malaysian) 


1 broad band of flat denticles on central disc (sparse or absent in small juveniles) 

2 disc profile somewhat quadrangular 

3 in adults, upper surface of disc with leopard-like markings (juveniles with large dark 

spots) 


5 tail long, whip-like, variably banded (rarely faint) 


7 central disc with row of up to 15 heart-shaped thorns 

8 midline of tail before sting without thorns 

Size: To at least 140 cm DW (about 410 cm TL); males mature at 70–80 cm DW; born 

at about 20 cm DW.


Distribution: Eastern Indian and western Pacific Oceans, from Myanmar to Papua 

New Guinea, including islands of the Japanese Archipelago and northern Australia. 

Habitat and biology: Demersal on soft substrates inshore to at least 70 m depth. 

Viviparous, with histotrophy; biology little known due to confusion with other strongly 

patterned whiprays. Diet presumably consists of crustaceans and small fishes. 

Utilisation: Caught occasionally in the demersal tangle net, trawl and line fisheries. 

Commercially important and used mainly for fresh meat. 


Local synonyms: Himantura undulata (Bleeker, 1852): Yano et al. (2005), White et al. 

(2006b). 

References: Manjaji-Matsumoto & Last (2008). 


Ventral disc 

In Borneo: Off Sabah, and northwestern 

and northeastern Kalimantan. 

1 

7 

Nuchal area 

Tail section - dorsal 

5 

3 

Tail section - ventral 

Juvenile (200 mm DW)


Tubemouth Whipray 

Himantura lobistoma Manjaji-Matsumoto & Last, 2006 

6 

2 

3 

5 

1 

9 

8 

Other names: None (English), Pari Lumpur, Pari Tanjung (Indonesian), Pari Mulut 

Tajam (Malaysian) 


1 denticle band very broad, margin pointed on snout 

2 disc oval, snout extremely elongate and broadly triangular 

3 brownish to greyish above (without a pattern) 

4 mouth strongly protrusible 


6 tail long, whip-like and plain (not banded) 

7 central disc usually 2–4 small pearl thorns 

8 no other enlarged thorns on disc or tail 

9 eyes small 

Size: To about 100 cm DW; males and females mature by 49 and 70 cm DW respectively; 

born at >18 cm DW.


Distribution: Endemic to Borneo and eastern Sumatra. 

Habitat and biology: Demersal on soft substrates inshore; biology little known. 

Viviparous, with histotrophy. Diet unknown, probably consists of crustaceans and 

small fishes. 

Utilisation: Caught commonly inshore off western Borneo, mainly by bottom trawl 

and line fisheries. Used mainly for fresh meat. 





Ventral disc 

In Borneo: Known mainly from western 

Borneo; occasionally off southern and 


7 

Nuchal area 

4 

Protrusible mouth 

Late-term embryo (18 cm DW)


Longnose Marbled Whipray 

Himantura oxyrhyncha (Sauvage, 1878) 

6 

5 

2 

3 

9 

1 

8 

Other names: Marbled Freshwater Stingray, Marbled Whipray (English), Pari 

(Indonesian), none (Malaysian) 


1 band of denticles on central disc narrow in adults 

2 disc profile oval with long, sharply pointed tip 

3 brownish with an ornate reticulate pattern 

4 uniformly whitish below, occasionally greyish or blackish marginally 


6 tail long, broad-based, whip-like and spotted 

7 central disc with large pearl thorn 

8 row of small heart-shaped thorns on mid-anterior tail of adults 

9 eyes very small 

Size: To about 37 cm DW (126 cm TL); free-swimming by 9 cm DW. 

Distribution: Borneo, Cambodia and Thailand.


Habitat and biology: Demersal in freshwater and estuaries; localised and common in 

lowland rivers and streams on soft substrates. Viviparous, with histotrophy; biology little 

known. Diet probably consists of small crustaceans and molluscs. 

Utilisation: Caught in demersal tangle nets, bottom trap and by line. Used mainly in 

the aquarium trade. 


Local synonyms: Dasybatus krempfi Chabanaud, 1923. 


Ventral disc 

4 

In Borneo: Known from north-west 

Kalimantan in the environs of the 

Kapuas River. 


7 

Nuchal area


Round Whipray 

Himantura pastinacoides (Bleeker, 1852) 

5 

6 

2 

3 

7 

1 

9 

Other names: None (English), Pari Aer, Pari Pasir (Indonesian), Pari Bulat (Malaysian) 


1 denticle band very broad, margin rounded on snout 

2 profile of disc almost oval; snout short, broadly triangular 

3 brownish or greyish above (without a pattern) 

4 whitish below, sometimes with a broad greyish posterior margin 


6 tail long, whip-like and dark posteriorly (not banded) 


8 usually one large, greenish pearl thorn on central disc 


Size: To at least 86 cm DW (186 cm TL), possibly to about 100 cm DW; males mature 

about 43–46 cm DW and females mature by 58 cm DW; birth size about 15–16 cm 

DW.


Distribution: Indo–Malay Archipelago, including Borneo and Sumatra; records from 

Javanese fish markets were probably caught elsewhere. 

Habitat and biology: Demersal, inshore, mostly in coastal embayments, estuaries and 

near large river mouths. Presumably viviparous, with histotrophy; biology not known. 

Diet probably consists of crustaceans and small fishes. 

Utilisation: Common bycatch of bottom trawl, line and beach seine fisheries off 

southern Borneo. Used for its meat (fresh), skin and cartilage. 


Local synonyms: Trygon pareh Bleeker, 1852. 


4 


Ventral disc 

In Borneo: Widespread, concentrated 

near large river estuaries. 

8 

Nuchal area


Giant Freshwater Whipray 

Himantura polylepis (Bleeker, 1852) 

6 

2 

5 

3 

1 

9 

7 

Other names: Freshwater Whipray, Giant Freshwater Stingray (English), Pari 

(Indonesian), Pari Air Tawar, Pari Sungai (Malaysian) 


1 no obvious band of flat denticles on central disc 

2 disc profile almost circular with long, sharp pointed tip 

3 greyish or brownish above (without a pattern) 

4 white below with very broad dark margin 


6 tail long, thin-based, whip-like and blackish (not banded) 


8 central disc usually with 2 heart-shaped thorns (no preceding row of slightly smaller 

thorns) 

9 midline of tail before sting lacking enlarged thorns 

Size: To at least 192 cm DW (~500 cm TL), probably larger; males mature at about 110 

cm DW.


Distribution: Widespread but patchy in the Indo–West Pacific, from India to eastern 

Indonesia; a similar form off Australia, and possibly New Guinea, is a separate species 

(H. dalyensis Last & Manjaji-Matsumoto, 2008). 

Habitat and biology: A very large coastal ray living primarily in rivers and estuaries, 

often well upstream. Probably common in the Kinabatangan River but rarely caught; 

sometimes in coastal marine and brackish habitats in other parts of Indonesia. Viviparous, 

with histotrophy; biology not well known. Feeds mainly on small fishes and crustaceans. 

Utilisation: Caught accidentally by local fishers, mainly by line and occasionally in 

gillnets and traps. Used for food and possibly its cartilage. 

Conservation status: IUCN Red List: Vulnerable; Critically Endangered in Thailand 

(as Himantura chaophraya Monkolprasit & Roberts, 1990). 

Local synonyms: Himantura chaophraya Monkolprasit & Roberts, 1990. 

References: Last & Manjaji-Matsumoto (2008). 


4 

In Borneo: Known from Sabah and 


Ventral disc 

8 

Nuchal area


White-edge Freshwater Whipray 

Himantura signifer Compagno & Roberts, 1982 

6 

2 

3 

5 

9 

1 

8 

7 

Other names: Freshwater Stingray, Pale Whipray, White-rim Stingray (English), Pari 

Sungai (Indonesian), Pari Air Tawar, Pari Sungai (Malaysian) 


1 band of denticles on central disc absent or narrow in adults 

2 disc profile almost circular with short, sharply pointed tip 

3 yellowish or greyish brown above (often with a faint reticulate pattern) with an 

obvious white edge 

4 uniformly whitish below 


6 tail long, broad-based, whip-like and whitish (not banded) 


8 central disc and tail lacking thorns 

9 eyes very small 

Size: To at least 37 cm DW (~134 cm TL); males maturing at 21–23 cm DW, females at


25–26 cm DW; born at about 11–12 cm DW. 

Distribution: Thailand, Indonesia and Malaysia, including central Sumatra, mainly in 

freshwater. 

Habitat and biology: Demersal on soft substrates in rivers, streams, and occasionally 

estuaries; ventures several hundred kilometres upstream. Viviparous, with histotrophy; 

biology little known. Diet consists of small, benthic invertebrates, such as crustaceans, 

shellfish and insect larvae. 

Utilisation: Caught mainly by trap, spear and bottom-set lines. Used for fresh meat and 

in the aquarium trade. 




4 


In Borneo: Kapuas, Sampi and Rungan 

rivers, western and southern Kalimantan. 

Ventral disc 

1 

Nuchal area


Bleeker’s Whipray 

Himantura uarnacoides (Bleeker, 1852) 

6 

5 

3 

1 

2 

9 

7 

Other names: Whitenose Whipray (English), Pari Tanjung, Pe Pasung, Pari Lumpur 

(Indonesian), Pari Pasir (Malaysian) 


1 denticle band very broad, margin rounded on snout 

2 profile of disc weakly rhomboidal, snout narrowly triangular 


4 whitish below, sometimes with a narrow dark posterior margin 


6 tail long, whip-like and pale (not banded) 


8 usually 1–3 large pearl thorns on central disc 


Size: Reported to reach 119 cm DW (300 cm TL), but rarely larger than 95 cm DW; 

males maturing at 50–60 cm DW; born about 18 cm DW. 

Distribution: Indo–West Pacific, from India to eastern Indonesia, including Malaysia.


Habitat and biology: Demersal on soft substrates near the coast to a depth of at least 

30 m. Viviparous, with histotrophy; biology little known. Diet unknown, probably 

consists of crustaceans and small fishes. 

Utilisation: Common catch component of demersal trawl and line fisheries. Used 

mainly for its meat and skin (exported to Thailand). 


Local synonyms: Trygon bleekeri Blyth, 1860. 


4 

In Borneo: Widespread, less common off 

western Sabah. 


Ventral disc 

8 

Nuchal area 

Juvenile (277 mm TL)


Reticulate Whipray 

Himantura uarnak (Forsskål, 1775) 

5 

3 

7 

1 

9 

6 

2 

Other names: Reticulate Whipray, Coachwhip Ray, Longtail Ray, Leopard Stingray 

(English), Pari Macan, Pari Merica (Indonesian), Pari Rimau (Malaysian) 



2 disc profile somewhat quadrangular 

3 in adults upper surface with fine dark spots and reticulations 

4 in juveniles upper surface with relatively small dark spots 


6 tail long, whip-like, variably banded (often faint in adults) 


8 central disc usually with 2 heart-shaped thorns (not preceded by row of slightly 

smaller thorns) 


Size: To at least 160 cm DW (450 cm TL); males mature at 82–84 cm DW; born at 

21–28 cm DW.


Distribution: Widespread in the Indo–West Pacific, from southern Africa and the eastern 

Mediterranean, through to the Philippines, Caroline Islands and northern Australia. 

Habitat and biology: Demersal on soft substrates, commonly in intertidal regions, but 

also to depths of at least 50 m. Viviparous, with histotrophy; biology little known. Diet 

probably consists of crustaceans and small fishes. Two distinct genetic forms exist off 

Borneo that may represent more than a single species. 

Utilisation: Commonly caught in small quantities by bottom trawlers; also taken by gill 

net and longline. Used fresh for its meat; also harvested for its skin and cartilage. 


Local synonyms: Trygon punctata Günther, 1870; Himantura toshi Whitley: Yano et al. 

(2005). 



In Borneo: Widespread, but patchy. 

Ventral disc 

4 

8 

Nuchal area 



Honeycomb Whipray 

Himantura undulata (Bleeker, 1852) 

6 

5 

3 

1 

9 

7 

2 

Other names: Bleeker’s Variegate Ray, Ocellate Whipray (English), Pari Macan 

(Indonesian), Pari Batik, Pari Rimau (Malaysian) 



2 disc profile weakly quadrangular (pectoral-fin apex broadly rounded) 

3 in adults upper surface with pattern of large ocelli and honeycomb-like reticulations 

4 in juveniles upper surface with relatively large dark spots 


6 tail long, whip-like, with dark pattern or bands 


8 central disc usually with dominant pearl-shaped thorn followed by 2–3 slightly 

smaller thorns 


Size: To at least 130 cm DW; males maturing at 60–70 cm DW; born at about 26– 

27 cm DW. 

Distribution: Indo–West Pacific, poorly defined; from India to eastern Indonesia; no 

confirmed records from Australia.


Habitat and biology: Demersal inshore on soft substrates, but little other information 

available as often confused with other similar species. Viviparous, with histotrophy; biology 

little known. Diet presumably consists of crustaceans and small fishes. 

Utilisation: Caught occasionally in demersal trawl, longline and tangle nets. Used 

mainly for its meat, possibly also skin and cartilage. 


Local synonyms: Trygon favus Annandale, 1909; Himantura fava Annandale: White et 

al. (2006b); Himantura sp. B: Yano et al. (2005). 


Ventral disc 

In Borneo: Sarawak, northeastern 

Sabah, and northwestern and eastern 

Kalimantan; probably more widespread. 


8 

Nuchal area 

4 



Dwarf Whipray 

Himantura walga (Müller & Henle, 1841) 

2 

3 

6 

1 

7 

5 

Other names: None (English), Pari Kikir, Toka Toka (Indonesian), Pari Ketuka, Tukatuka, 

Pari Tanjung (Malaysian) 


1 narrow band of flat denticles on central disc in adults 

2 profile of disc almost oval, snout broadly triangular 


4 whitish below, sometimes with a yellowish grey posterior margin 


6 tail short, not whip-like (end bulbous in adult females) 


8 mid-disc thorns absent or rudimentary 

9 tail thorns very elongate, bases nearly half eye diameter in length 

Size: To at least 24 cm DW (~38 cm TL); males and females mature at 16–18 cm DW; 

born at 8–10 cm DW. 

Distribution: Off Thailand, Malaysia and Indonesia, but possibly also occurs further 

west to India. 

Habitat and biology: Demersal on insular and continental shelves, and occasionally in 

coastal embayments. Viviparous, with histotrophy; gives birth to litters of 1–2 pups after


an unknown gestation period; no reproductive synchronicity. Diet consists primarily of 

small benthic crustaceans, but also small fishes. 

Utilisation: Landed in large quantities by bottom trawl fisheries. Used for its meat, 

fresh and dried, and is in demand locally. 


Local synonyms: Trygon heterurus Bleeker, 1852; images of Himantura imbricata (Bloch 

& Schneider) in Yano et al. (2005) appear to be this species. 


4 

In Borneo: Widespread, but not recorded 

from eastern Kalimantan. 


Ventral disc 

8 

9 

Nuchal area 

Anterior tail - dorsal view 

6 

Dorsal post-sting tail - mature female


Bluespotted Maskray 

Neotrygon kuhlii (Müller & Henle, 1841) 

4 

6 

5 

1 

7 

Other names: Bluespotted Stingaree, Bluespotted Stingray, Kuhl’s Stingray (English), 

Pari Blentik, Pari Kodok (Indonesian), Pari Lalat, Pari Macan, Pari Riman, Pari Rimau, 

Pari Tanjung, Kiampau (Malaysian) 


1 a few short thorns confined to midline of disc 

2 short, low skin fold on dorsal tail just posterior to sting tip 

3 long, low skin fold on ventral surface of tail 

4 tail not whip-like, its tip with broad black and white bands 

5 snout short with black bar through eyes 

6 large, bright, blue spots on upper disc 

7 usually no thorns on tail before sting 

Size: To at least 42 cm DW; males are mature by about 26 cm DW; born at about 

14 cm DW. 

Distribution: Indo–West Pacific, from southern Africa and India through southeast 

Asia and northern Australia, north to Japan and east to Micronesia and Melanesia; 

probably consists of a complex of species with narrow ranges.


Habitat and biology: Demersal on insular and continental shelves to a depth of 

90 m. Viviparous, with histotrophy; gives birth to litters of 1–2 pups after an unknown 

gestation period; no reproductive synchronicity. Diet consists primarily of benthic 

crustaceans and small fishes. 

Utilisation: One of the most commercially important rays in the region; caught in 

large quantities by bottom trawl, trammel and fish trap fisheries. Used mainly for fresh 

meat. 


Local synonyms: Dasyatis kuhlii (Müller & Henle): White et al. (2006b), White & 

Dharmadi (2007). 



Ventral disc 


1 

Nuchal area 

2 

Lateral tail 

3


Eastern Cowtail Stingray 

Pastinachus atrus (Macleay, 1883) 

4 

2 

5 

7 

3 

1 

6 

8 

Other names: Banana-tail Ray, Bull Ray, Fantail Ray, Feathertail Ray, Feathertail 

Stingray (English), Pari Bendera (Indonesian), Pari Bendera, Pari Daun, Pari Nyonya, 

Pari Tanjung (Malaysian) 


1 broad band of flat denticles on central disc 

2 disc profile quadrangular with a short, rounded snout 

3 snout apex mostly lacking denticles 

4 ventral skin fold terminating well before tail tip (when undamaged) 

5 fold blackish, very deep (maximum depth >3.5 times height of tail above, more than 

5 times in adults) 

6 tail relatively broad based 

7 sting situated posteriorly on tail (its distance from cloaca more than half disc width) 

8 no thorn row along middle of tail 

Size: To almost 200 cm DW (>300 cm TL); males mature at about 96–98 cm DW; 

born at about 18 cm DW.


Distribution: Tropical Indo–West Pacific including Malaysia, Indonesia, Australia, New 

Guinea, and possibly Melanesia and Micronesia; also known from the Western Indian 

Ocean (Madagascar). 

Habitat and biology: Demersal, coastal to inner continental shelf to a depth of at least 

60 m; occurs in coral lagoons, intertidal mangrove estuaries and also known from rivers 

in freshwater. Viviparous, with histotrophy; biology little known. Diet of adults presumably 

consists of large crustaceans and small fishes. 

Utilisation: Caught frequently by bottom trawl, with smaller quantities by longline and 

beach seine. Good food fish, used locally mainly for fresh meat. 


Local synonyms: Pastinachus sephen (Forsskål): Yano et al. (2005), White et al. (2006b). 



Ventral disc 


5 

Nuchal area 

Lateral tail - at midlength of ventral fold


Narrowtail Stingray 

Pastinachus gracilicaudus Last & Manjaji-Matsumoto, 2010 

4 

5 

7 

3 

1 

6 

8 

2 

Other names: None (English), Pari Bendera (Indonesian), none (Malaysian) 



2 disc profile quadrangular with a short, rounded snout 

3 snout apex mostly lacking denticles 


5 fold typically pale, relatively slender (maximum depth usually < 3 times height of tail 

above) 

6 tail relatively broad based 

7 sting situated posteriorly on tail (its distance from cloaca more than half disc width) 

8 no thorn row along middle of tail 

Size: To almost 83 cm DW (>200 cm TL); males mature by 67 cm DW; free-swimming 

by 30 cm DW. 

Distribution: Not well defined; known from the Indo–Malay Archipelago.


Habitat and biology: Demersal on continental shelf, mainly inshore. Viviparous, with 

histotrophy; biology little known. Diet probably consists of crustaceans and small fishes. 

Utilisation: Caught occasionally by demersal tangle net, bottom trawl and longline 

fisheries. Used mainly for its meat, and probably for its skin. 


Local synonyms: Confused with Pastinachus sephen (Forsskål). 

References: Last & Manjaji-Matsumoto (2010). 

Ventral disc 

In Borneo: Patchy, in western and 

northern Borneo; not observed from most 

of southern and eastern Kalimantan. 


5 

Nuchal area 



Roughnose Stingray 

Pastinachus solocirostris Last, Manjaji & Yearsley, 2005 

2 

4 

5 

3 

1 

6 

8 

7 

Other names: None (English), Pari Gunung, Pari Bendera (Indonesian), Pari Mulut 

Kasar (Malaysian) 



2 disc profile strongly quadrangular with an acute, triangular snout 

3 snout apex covered with enlarged, spear-shaped or fork-like denticles 


5 fold relatively slender (maximum depth usually 150 cm TL); males mature at about 36–39 cm DW; born 

at about 20 cm DW. 

Distribution: Confined to Indonesia and Malaysia (Borneo and Sumatra). 

Habitat and biology: Habitat and biology largely unknown; demersal inshore on


continental shelf, mostly near large estuaries. Presumably viviparous, with histotrophy. 

Diet probably consists of small crustaceans and fishes. 

Utilisation: Caught occasionally by bottom trawl, line and demersal gillnet fisheries 

operating off Sumatra, Sarawak and Kalimantan. Good food fish, used locally for its meat 

and probably its skin. 


Local synonyms: Pastinachus cf. sephen (Forsskål): Yano et al. (2005). 

References: Last et al. (2005). 

Ventral disc 

In Borneo: Widespread off western 

Borneo; isolated records from southern 

and southeastern Kalimantan. 


Nuchal area 

3 

5 

Denticles at apex of snout 



Starynose Stingray 

Pastinachus stellurostris Last, Fahmi & Naylor, 2010 

4 5 

7 

3 

1 

6 

8 

2 

Other names: None (English), Pari Bendera (Indonesian), none (Malaysian) 



2 disc profile strongly quadrangular with an acute, triangular snout 

3 snout apex covered with enlarged, star-shaped denticles 


5 fold relatively slender (maximum depth


probably consists of invertebrates and small fishes. 

Utilisation: Caught rarely by line fisheries; rare in markets, probably used for its meat. 


Local synonyms: Probably confused with Pastinachus solocirostris Last, Manjaji & 

Yearsley, 2005. 


Ventral disc 

In Borneo: Restricted, known only from 

northwestern Kalimantan. 


Nuchal area 

3 

5 

Denticles at apex of snout 



Pelagic Stingray 

Pteroplatytrygon violacea (Bonaparte, 1832) 

2 

5 

4 

1 

3 

Other names: Blue Stingray, Violet Stingray (English), Pari Lampin, Pari Lemer, 

Payubek (Indonesian), Pari Hitam (Malaysian) 


1 row of small thorns along midline of disc 

2 anterior profile of disc strongly convex and obtuse 

3 outer edge of disc angular 

4 ventral skin fold on tail very low, terminating well before tail tip 

5 both dorsal and ventral surfaces dark brownish to black (often with a purplish 

tinge) 

Size: To at least 59 cm DW (at least 130 cm TL), elsewhere reported to 80 cm DW; 

males and females mature at 35–38 and 45–50 cm DW respectively; born at 16–20 cm 

DW. 

Distribution: Cosmopolitan in tropical and temperate waters. 

Habitat and biology: Fully pelagic stingray found in the open ocean, usually in upper


100 m and rarely found inshore. Viviparous, with histotrophy; gives birth to litters of 

2–9 pups after a 4 month gestation period. Diet consists primarily of jellyfishes, squids, 

crustaceans and fishes. 

Utilisation: Occasional bycatch of tuna longline and gillnet fisheries operating 

throughout the region. Used for fresh meat when caught. 




5 

Ventral disc 

In Borneo: Known from off Sarawak 

and Sabah, but likely to be more widely 

distributed. 


Lateral tail 

4


Bluespotted Fantail Ray 

Taeniura lymma (Forsskål, 1775) 

3 

2 

4 

5 

1 

6 

Other names: Blue-spotted Lagoon Ray, Blue-spotted Ribbontail Ray, Lagoon Ray, 

Lesser Fantail Ray, Reef Ray, Ribbontail Stingray (English), Ikan Pari, Pari Kembang, 

Pari Totol (Indonesian), Pari Karang, Pari Reben, Kiampau (Malaysian) 


1 dorsal surface almost smooth to granular (denticles very small) 

2 profile of disc oval 

3 ventral skin fold on tail relatively deep, extending to tail tip 

4 upper surface with numerous bright blue spots 

5 stings present (usually 2), located near end of tail 

6 tail with blue stripe on each side before sting 

Size: To at least 35 cm DW (at least 75 cm TL); males and females mature at 20–22 and 

20–24 cm DW respectively; born at 13–14 cm DW. 

Distribution: Widespread in Indo–West Pacific, from southern Africa to the Solomon 

Islands, south to tropical Australia and north to the Philippines and Vietnam.


Habitat and biology: Dominant ray in coral reef habitats; occurs inshore to depths of 

at least 20 m. Viviparous, with histotrophy; biology little known. Diet consists primarily 

of molluscs, worms and small crustaceans. 

Utilisation: Caught over shallow coral reefs; common in small quantities in markets. 

Used widely for its meat. 


Local synonyms: Trygon ornatus Gray, 1830. 


In Borneo: Probably widespread, 

although less common in Sarawak and 

parts of Kalimantan. 


Ventral disc 

Lateral tail 

3


Blotched Fantail Stingray 

Taeniurops meyeni (Müller & Henle, 1841) 

2 

4 

3 

6 

1 

5 

Other names: Blackblotched Stingray, Blackspotted Stingray, Bull Ray, Fantail Stingray, 

Giant Reef Ray, Marble Ribbontail Ray, Round Ribbontail Ray, Speckle Stingray 

(English), Pari Babi, Pari Merica (Indonesian), Pari Kelabu, Pari Reben (Malaysian) 


1 dorsal surface almost smooth (denticles very small) 

2 profile of disc circular 

3 ventral skin fold on tail deep, extending to tail tip 

4 upper surface dark with white blotches and mottling (often faint) 

5 large sting present (usually 1) 

6 tail uniformly black posterior to sting 

Size: To at least 180 cm DW (330 cm TL); males mature at 100–110 cm DW; born at 

30–35 cm DW. 

Distribution: Widespread in the Indo–West Pacific from South Africa to Japan, south 

to Micronesia, including northern Australia.


Habitat and biology: Demersal on soft bottoms near coral reefs; also reported offshore 

to depths exceeding 400 m. Viviparous, with histotrophy; biology little known. Diet 

consists primarily of bivalves, crustaceans and small fishes. 

Utilisation: Caught occasionally by demersal tangle net and longline, rarely by bottom 

trawl fisheries. Used for its meat and cartilage in small quantities. 


Local synonyms: Taeniura melanospilus Bleeker, 1853; Taeniura meyeni Müller & Henle, 

1841: Yano et al. (2005), White et al. (2006b), White & Dharmadi (2007). 


Ventral disc 

In Borneo: Known only from Sabah and 

off Sarawak. 


Lateral tail 

3


Porcupine Ray 

Urogymnus asperrimus (Bloch & Schneider, 1801) 

2 

1 

3 

4 

Other names: Roughskin Stingaree, Solander’s Ray, Thorny Ray (English), Pari Duren 

(Indonesian), Pari Dedap, Pari Duri, Pari Durian (Malaysian) 


1 dorsal surface very rough, covered with long, sharp thorns 

2 profile of disc almost circular 

3 base of tail narrow, rounded in cross-section rather than depressed 

4 no skin folds or stinging spines on tail 

Size: Possibly to 147 cm DW; males and females mature at 90 and 100 cm DW 

respectively. 

Distribution: Widespread in the Indo–West Pacific; east Africa and the Red Sea to 

northern Australia and Fiji, north to Japan. Also reported off central Africa in the tropical 

eastern Atlantic. 

Habitat and biology: Demersal on insular and continental shelves; found on rubble and 

sand near coral reefs. Viviparous, with histotrophy; biology little known. Diet presumably 

consists mainly of bivalves and crustaceans which it extracts from the sediments. Its thorns 

are sharp and can inflict a painful injury.


Utilisation: Caught rarely by demersal tangle net and trawl fisheries. Probably used for 

its meat, but may be discarded. 




Ventral disc 

In Borneo: Uncommon, known from 

off Sabah and eastern Kalimantan; 

possibly more widely distributed in 

appropriate habitat. 


1 

Lateral head 

Mouth and nostrils


Longtail Butterfly Ray 

Gymnura poecilura (Shaw, 1804) 

4 

1 

2 

3 

Other names: Long-tailed Butterfly Ray (English), Pari Lowo, Pari Kelelawar 

(Indonesian), Pari Kelawar Bintik, Pari Tembikar (Malaysian) 


1 no dorsal fin 

2 tail lacking a stinging spine and with ~9 dark bands 

3 tail length about equal to snout-vent length (when undamaged) 

4 dorsal surface with faint whitish spots, sometimes plain 

Size: To at least 99 cm DW; males mature at about 45 cm DW; born at about 20–26 cm 

DW. 

Distribution: Tropical Indo–West Pacific; from the Red Sea eastward to southern 

Japan and eastern Indonesia. 

Habitat and biology: Demersal on the continental shelf to a depth of at least 30 m. 

Appears to prefer sandy habitats, often in shallow inshore waters and on sand banks.


Viviparous, with histotrophy; litters of up to 7 pups after an unknown gestation period. 

Feeds primarily on small fishes, crustaceans and molluscs. 

Utilisation: Caught frequently by demersal trawlers, and taken occasionally by trammel 

net. Utilised widely for its meat, but of limited value due to its small size. 



References: Jacobsen & Bennett (2009). 


Gymnuridae (Butterfly Rays) 

Ventral disc


Zonetail Butterfly Ray 

Gymnura zonura (Bleeker, 1852) 

4 

3 

Other names: Bleeker’s Butterfly Ray (English), Pari Lowo, Pari Kelelawar, Pari 

Payung (Indonesian), Pari Kelawar, Pari Tembikar (Malaysian) 


1 small dorsal fin often present at base of tail 

2 small stinging spine present 

3 tail length about 2/3 snout-vent length (when undamaged) 

4 dorsal colour pattern variable, complex, usually with numerous dark spots interspersed 

with larger yellowish spots 

Size: To at least 108 cm DW; males mature by 46–50 cm DW; born at about 20–21 cm 

DW. 

Distribution: Tropical Indo–West Pacific, from India to eastern Indonesia, including 

the Philippines. 

Habitat and biology: Demersal inshore, in shallow water to depths of at least 37 

m. Viviparous, with histotrophy; litters of 2–4 pups, gestation period unknown. Diet 

consists primarily of small fishes and crustaceans.


Utilisation: Frequently caught as bycatch of demersal trawl, trammel and gillnet 

fisheries, and occasionally by tangle nets. Utilised for its meat throughout the region, 

but of limited value due to its small size. 


Local synonyms: Gymnura japonica (Temminck & Schlegel): White et al. (2006b), 

White & Dharmadi (2007); Aetoplatea zonura Bleeker: Yano et al. (2005). 

References: Jacobsen & Bennett (2009). 

1 

Lateral anterior tail 

2 

In Borneo: Sarawak, Sabah, and eastern 

and western Kalimantan. 

Gymnuridae (Butterfly Rays) 

Ventral disc


Longhead Eagle Ray 

Aetobatus flagellum (Bloch & Schneider, 1801) 

2 

5 

1 

Other names: None (English), Pari Burung (Indonesian), Pari Lang (Malaysian) 


1 dorsal disc surface plain, unspotted 

2 snout very long, narrowly rounded 

3 nasal curtain V-shaped 

4 teeth in a single row in both jaws, chevron-shaped 

5 spiracles dorsolateral on head 

Size: To at least 72 cm DW; males mature at >44 cm DW. 

Distribution: Tropical Indo–West Pacific, from the Red Sea to Indonesia and Malaysia; 

Japanese populations appear to belong to a larger, distinct species. 

Habitat and biology: Habitat and biology poorly known; locally, appears to occur 

offshore. Presumably viviparous, with histotrophy. Diet unknown but likely to consist of


bivalves and other hard-shelled invertebrates. 

Utilisation: Caught infrequently, usually by bottom trawl and demersal gillnet fisheries. 

Used for its meat, but of limited value due to its rarity. 




Upper tooth plate 

3 

Ventral head 

In Borneo: Uncommon, known from 

western Kalimantan and northeastern 

Sabah. 

Myliobatidae (Eagle Rays) 

4 

Lower tooth plate


Whitespotted Eagle Ray 

Aetobatus ocellatus (Kuhl, 1823) 

2 

5 

1 

Other names: Bonnet Skate, Duckbill Ray, Spotted Eagle Ray (English), Pari Burung, 

Pari Ayam (Indonesian), Pari Lang Bintik Putih, Pari Lang, Pari Helang (Malaysian) 


1 dorsal disc surface usually with numerous white spots (sometimes sparse and 

confined to posterior disc) 

2 snout moderately long, broadly rounded 

3 nasal curtain V-shaped 

4 teeth in a single row in both jaws, chevron-shaped 

5 spiracles dorsolateral on head 

Size: To at least 300 cm DW (possibly to 330 cm DW), more commonly to about 

160 cm DW; males mature at 100–110 cm DW; born at 33–36 cm DW. 

Distribution: Tropical and warm temperate seas of the Indo–West Pacific, from the 

Red Sea and South Africa to Hawaii, north to Japan, and south to Australia.


Habitat and biology: Inshore and benthopelagic on continental shelves, occasionally 

found in estuaries and near atolls, but also well offshore in open water. Viviparous, with 

histotrophy; gives birth to litters of 2–4 pups after an unknown gestation period. Diet 

consists primarily of bivalves, as well as gastropods, polychaetes, small fishes, crustaceans 

and cephalopods. Belongs to a group of very similar, white-spotted eagle rays. 

Utilisation: Commonly taken by demersal tangle net, bottom trawl, inshore gillnet 

and, to a lesser extent, demersal longline. Caught for its meat and cartilage. 


Local synonyms: Aetobatus narinari (Euphrasen, 1790): Yano et al. (2005), White et al. 

(2006b); Aetobatus cf. narinari (Euphrasen, 1790): Compagno et al. (2005b); Aetobatus 

guttatus (Shaw, 1804): Compagno & Last (1999a); Raia quinqueaculeata Quoy & Gaimard, 

1824. 

References: White et al. (2010d). 

3 

Ventral head 




4 

Juvenile - dorsal disc (465 mm DW) 

Lower tooth plate


Mottled Eagle Ray 

Aetomylaeus maculatus (Gray, 1834) 

1 

6 

4 

3 

2 

Other names: Ocellate Eagle Ray, Ornate Eagle Ray (English), Pari Burung 

(Indonesian), Pari-lang Tompok Putih (Malaysian) 


1 fleshy ridge on side of head not connected to edge of disc 

2 sting absent 

3 dorsal-fin posterior margin upright, its origin slightly posterior to pelvic-fin 

insertions 

4 upper surface brown with whitish spots (with weak ocelli on posterior half of disc in 

young) 

5 edge of nasal curtain near mouth almost straight 

6 spiracles lateral on head 

7 small eagle ray, attaining about 100 cm DW 

Size: To at least 100 cm DW; males mature by 71 cm DW; born at about 29 cm DW. 

Distribution: Eastern Indian Ocean to western Pacific including Malaysia, Thailand, 

Indonesia and southern China; possibly west to India and excluding Australia.


Habitat and biology: Largely unknown, but typically found inshore. Presumably 

viviparous, with histotrophy. Diet unknown, but possibly consists of bivalves and other 

bottom-dwelling invertebrates. A similar form in the western Indian Ocean appears to 

be another species. 

Utilisation: Caught occasionally in the region by bottom trawl, tangle net and inshore 

gillnet fisheries. Used for its meat, but of limited value as it is not frequently landed. 


Local synonyms: None. Authority previously considered to be Gray, 1832 but publication 

date listed on Eschmeyer’s Catalog of Fishes as 1 October 1834. 


5 

Ventral head 




4 

Juvenile - dorsal disc (~250 mm DW) 

Lower tooth plate


Banded Eagle Ray 

Aetomylaeus nichofii (Bloch & Schneider, 1801) 

1 

6 

4 

3 

2 

Other names: Barbless Eagle Ray, Nieuhof’s Eagle Ray (English), Pari Burung, Pe 

Lamping (Indonesian), Pari-lang Jalur (Malaysian) 




3 dorsal-fin posterior margin strongly angled, its origin slightly anterior to or opposite 

pelvic-fin insertions 

4 upper surface yellowish brown with about 5 bluish bands (often faint) 



Size: To at least 72 cm DW; males mature at 39–42 cm DW; born at about 17 cm DW. 

Distribution: Indo–West Pacific from India (possibly westwards to the Arabian Gulf) 

through Indonesia and northward to Japan; excluding Australia.


Habitat and biology: Occurs on or near the bottom intertidally, but mainly offshore 

to depths of at least 70 m. Little known of its biology; viviparous, with histotrophy; gives 

birth to litters of up to 4 pups after an unknown gestation period. Diet presumably consists 

of bivalves and other small invertebrates. A similar form occurring in Australian seas 

appears to be a distinct species. 

Utilisation: Frequently caught by bottom trawl and inshore gillnets but in low numbers. 

Used for its meat, but of limited value due to its small size. 




5 

Ventral head 




Lower tooth plate


Ornate Eagle Ray 

Aetomylaeus vespertilio (Bleeker, 1852) 

1 

6 

4 

3 

2 

Other names: Reticulate Eagle Ray (English), Pari Burung (Indonesian), Pari Lang 

Bunga (Malaysian) 




3 dorsal-fin posterior margin upright, its origin posterior to pelvic-fin insertions 

4 upper surface greenish brown with thin dark transverse lines anteriorly and whitish 

rings and dark reticulations posteriorly 



7 large eagle ray, attaining 300 cm or more DW 

Size: To at least 300 cm DW (possibly to 350 cm DW); males mature by 170 cm DW. 

Distribution: Indo–West Pacific but patchy; southern Africa to China (including 

Taiwan), and south to northern Australia. 

Habitat and biology: Habitat and biology little known, typically found on the continen-


tal shelf but rarely near the coast. Presumably viviparous, with histotrophy. Diet probably 

consists of bivalves and other bottom-dwelling invertebrates. Juveniles of the smaller 

Aetomylaeus maculatus also have an ornate colour pattern but lack distinctive transverse 

black lines on the anterior disc. 

Utilisation: Caught rarely by bottom trawl and tangle net fisheries. Used primarily for 

its meat, but of limited value as it is not frequently landed. 




5 

Ventral head 

In Borneo: Known locally from a 

few individuals collected off Sabah and 

Kalimantan. 



Lower tooth plate


Javanese Cownose Ray 

Rhinoptera javanica Müller & Henle, 1841 

1 

2 

5 

7 

4 

Other names: Cow Ray, Flapnose Ray (English), Pari Burung, Pari Elang 

(Indonesian), Pari-kelawar Jawa, Pari Ayam, Pari Lang, Pari Susun (Malaysian) 


1 snout strongly notched medially to form two lobes 

2 head relatively narrow, its width at spiracle 14.5–16% of disc width 

3 rostral flap short, its fleshy posterior edge not (or just) reaching mouth 

4 tail long, about 2.6–3.4 times disc length (when undamaged) 

5 dorsal-fin origin distinctly behind pectoral-fin insertions 

6 posterior margin of dorsal fin strongly concave 

7 sting base (when present) behind dorsal-fin free rear tip 

8 large species, adults possibly exceeding 150 cm DW 

Size: To at least 150 cm DW; males still immature at 90 cm DW, newborns observed at 

38 cm DW.


Distribution: Central western Pacific, from Thailand to Australia and north to the East 

China Sea; poorly defined due to confusion with other species. 

Habitat and biology: Preferred habitat and biology poorly known; patchy, but most 

common inshore in bays, estuaries and near coral reefs. Viviparous, with histotrophy; one 

large pregnant female in Indonesia (~130 cm DW) contained a single, late-term embryo 

of about 60 cm DW. Diet presumably consists mainly of bivalves and other bottomdwelling 


Utilisation: Caught occasionally by bottom trawl, demersal inshore gill and tangle net 

fisheries. Sold mainly for its meat, but not of high value. 


Local synonyms: Rhinoptera adspersa Müller & Henle, 1841. 


1 

3 

Rhinopteridae (Cownose Rays) 

Ventral head 

In Borneo: Apparently uncommon, 

known only from eastern Sabah and 


3 

5 6 

7 

Lateral dorsal fin and anterior tail 

Lateral head


Short-tail Cownose Ray 

Rhinoptera jayakari Boulenger, 1895 

1 

2 

5 

7 

4 

Other names: None (English), Pari Elang (Indonesian), Pari Lang (Malaysian) 


1 snout strongly notched medially to form two lobes 

2 head relatively broad, its width at spiracle 16.5–17.5% of disc width 

3 rostral flap long, its fleshy posterior edge extending past mouth (almost to 1 st gill 

slit) 

4 tail short, about 1.4–1.8 times disc length (when undamaged) 

5 dorsal-fin origin over or slightly forward of pectoral-fin insertions 

6 dorsal-fin posterior margin almost upright 

7 sting base (when present) beneath dorsal-fin inner margin 

8 smaller species, adults attaining at least 90 cm DW 

Size: To at least 90 cm DW, possibly to 110 cm DW in the western Indian Ocean; male 

holotype sexually mature at about 73 cm DW.


Distribution: Widespread in the tropical Indo–West Pacific, from southern Africa to 

eastern Indonesia, north to Japan and possibly Australia; poorly defined and often 

confused with R. javanica. 

Habitat and biology: Preferred habitat and biology poorly known. Often pelagic in 

large schools, near the coast and well offshore; the more abundant of the two cownose 

rays in the region. Presumably viviparous, with histotrophy. Diet unknown but probably 

consisting mainly of bottom-dwelling invertebrates, as well as small pelagic animals. 

Utilisation: Caught occasionally by bottom trawl, demersal inshore gill and tangle net 

fisheries. Sold mainly for its meat, but of limited value. 


Local synonyms: Rhinoptera sewelli Misra, 1946; Rhinoptera sp. 1: White et al. (2006b). 


1 

3 

Rhinopteridae (Cownose Rays) 

Ventral head 

In Borneo: Known off eastern Sabah, 

southern Sarawak and northwestern 

Kalimantan. 

5 6 

7 

3 

Lateral head 

Lateral dorsal fin and anterior tail


Manta Ray 

Manta birostris (Walbaum, 1792) 

2 

1 

Other names: Devilfish, Manta, Giant Manta (English), Pari Kerbua, Plampangan, 

Cawang Kalung (Indonesian), Pari Kelawar, Pari Paus (Malaysian) 


1 head very broad 

2 mouth blackish, at end of snout 

3 teeth absent from upper jaw 

4 calcified lump posterior to dorsal fin 

5 large triangular blotch abutting 5 th gill slit 

6 attains a very large size, often well over 4 m wide 

Size: To at least 670 cm DW, unconfirmed reports to 910 cm DW need verifying; 

males and females mature at about 375–380 and 380–410 cm DW respectively; born at 

122–149 cm DW. 

Distribution: Circumtropical, occasionally making seasonal incursions into warm 

temperate waters. 

Habitat and biology: Wide-ranging, pelagic species found near the surface in tropical


seas. Common near coral reefs but also occurs in the open ocean and inshore in muddy 

bays and river mouths. Swims continuously and rarely rests on the bottom. Viviparous, 

with histotrophy; gives birth to a single pup after an unknown gestation period. Feeds 

on zooplankton and small fishes; appears frequently over coral reefs during seasonal 

plankton blooms. Tends to be inquisitive and is easily approached by divers. A related 

species, M. alfredi (Krefft), has recently been recognised and has been recorded from 

north of Sabah. It lacks a caudal spine and calcified mass behind the dorsal fin, and has a 

paler mouth with a narrow streak at the end of the 5 th gill slit. 

Utilisation: Considered to be very vulnerable to overfishing but not observed in local 

markets. Elsewhere, caught occasionally by gillnet and harpoon fisheries and utilised for 

its gill filter plates (very high value), meat, cartilage and skin. Has considerable value for 

dive ecotourism. 



References: White et al. (2006a); Marshall et al. (2009). 

3 

Mobulidae (Devilrays) 

5 

Ventral head 

In Borneo: Single record from a fish 

market in northwestern Kalimantan 

presumably this species; observed regularly 

by divers at Derawan and Sipadan islands 

off eastern Borneo. 

4 

Embryo 

Lateral dorsal fin and anterior tail


Borneo Devilray 

Mobula sp. 

1 

2 

4 

5 

Other names: None (English), none (Indonesian), Pari Paus (Malaysian) 


1 cephalic lobes relatively long, about 15% DW 

2 anterior margin of snout deeply concave 

3 mouth width much less than twice distance from its angle to junction of head lobe 

4 dorsal fin white tipped 

5 base of tail almost quadrangular in cross section 

6 spiracles located underneath disc edge 

7 mouth subterminal, located on underside of head 

Size: To at least 80 cm DW; nothing known of size at maturity or birth. 

Distribution: Possibly confined to the Indo–West Pacific, but its range not well 

defined. 

Habitat and biology: An uncommon, presumably inshore pelagic species. Biology 

unknown, presumably viviparous, with histotrophy. Diet probably consists of planktonic


crustaceans, and possibly small fishes and cephalopods. Known from few specimens and 

its relationship to M. kuhlii needs further evaluation. 

Utilisation: Limited and not commonly observed in markets. 




1 

2 

3 

7 

Ventral head 

In Borneo: Known from off Kuching, 

Sarawak. 


6 

Dorsolateral head


Japanese Devilray 

Mobula japanica (Müller & Henle, 1841) 

1 

Other names: Spinetail Mobula, Spinetail Devilray (English), Cawang Genul, 

Lampengan, Pari Lambingan, Plampangan (Indonesian), Pari Paus (Malaysian) 


1 disc broad and slightly falcate 

2 dorsal fin conspicuously white tipped 

3 small sting present on tail base 

4 spiracles slit-like and elliptical, located above disc edge 


Size: To at least 310 cm DW (usually less than 250 cm); males mature at 198–205 cm; 

late-term embryos observed up to 113 cm DW. 

Distribution: Circumtropical, occasionally making seasonal incursions into warm 

temperate waters. 

Habitat and biology: A large pelagic inshore and offshore (possibly oceanic) devilray. 

Viviparous, with histotrophy; gives birth to a single pup after an unknown gestation


period. Feeds mainly on zooplankton and small fishes. Occurs in large schools and is 

occasionally observed by divers. Often confused with other devilrays. 

Utilisation: Rarely landed in Borneo but common bycatch of gillnet fisheries in other 

parts of the archipelago. Used for its gill filter plates (very high value), meat, cartilage 

and skin where caught in larger numbers. 


Local synonyms: Mobula japonica (Müller & Henle): Yano et al. (2005). 

References: White et al. (2006a). 

Late-term embryo (498 mm DW) 

2 

3 

In Borneo: Recorded from Semporna in 

east Sabah.; probably widespread. 


Lateral dorsal fin and sting 

4 

5 

Dorsolateral head 

Ventral head


Lesser Devilray 

Mobula kuhlii (Müller & Henle, 1841) 

2 

1 

5 

Other names: Shortfin Devilray, Shorthorn Devilray (English), Pari Lambingan, Pari 

Lampengan (Indonesian), Pari Kelawar, Pari Paus, Tanpa Duri (Malaysian) 


1 cephalic lobes relatively short, 12–14% DW 

2 anterior margin of snout slightly concave 

3 mouth width much more than twice distance from its angle to junction of head lobe 

4 dorsal fin not white tipped 

5 base of tail quadrangular in cross section 

6 spiracles subcircular, located underneath disc edge 


Size: To at least 120 cm DW; males mature at 115–119 cm DW; born at about 31 cm 

DW. 

Distribution: Indo–West Pacific from eastern Africa to Indonesia, but actual range not 

well defined.


Habitat and biology: A locally uncommon, small, inshore devilray. Viviparous, with 

histotrophy; probably giving birth to only a single pup. Diet unknown but presumably 

consists of planktonic crustaceans, and possibly small fishes and cephalopods. Other 

Mobula species probably occur in the deep oceanic waters off northern and eastern 

Borneo. 

Utilisation: Caught occasionally by sea gypsies off eastern Sabah and marketed mainly 

for its meat. 


Local synonyms: Dicerobatis draco Günther, 1872. 


1 

2 

3 

7 


Ventral head 

In Borneo: Northeastern Borneo, off 

Sabah. 

4 

6 

5 


Dorsolateral head


Bentfin Devilray 

Mobula thurstoni (Lloyd, 1908) 

1 

2 

5 

6 

Other names: Smoothtail Mobula (English), Pari Lempengan, Pari Lambingan, 

Plampangan (Indonesian), Pari Paus, Sirip Lekuk (Malaysian) 


1 cephalic lobes relatively short, 12–14% DW 

2 anterior margin of snout slightly concave 

3 mouth width much more than twice distance from its angle to junction of head lobe 

4 dorsal fin with prominent white tip 

5 outer anterior margin of pectoral fin with distinct concavity 

6 tail base depressed 

7 spiracles subcircular, located underneath disc edge 


Size: To at least 189 cm DW; males mature at 150–154 cm DW; born at 65–85 cm DW. 

Distribution: Probably circumtropical, but only known from scattered localities. 

Habitat and biology: Elsewhere in the archipelago, this medium-size devilray is a


common coastal inshore and offshore pelagic species. Viviparous, with histotrophy; 

probably giving birth to a single pup. Feeds almost exclusively on planktonic 

crustaceans. 

Utilisation: Probably caught only occasionally and used for its meat, cartilage and skin. 




1 

2 

3 

8 

Ventral head 

In Borneo: Single record from the 

Sandakan fish market, northeastern 

Sabah. 


4 

6 

7 


Dorsolateral head


Silver Chimaera 

Chimaera phantasma Jordan & Snyder, 1900 

4 

6 

2 

3 

5 

1 

7 

Other names: Ghost Shark (English), Hiu Hantu, Hiu Pogak (Indonesian), Chimara 

Perak, Ikan Tikus (Malaysian) 


1 anal fin present, separated from caudal fin by a narrow notch 

2 trunk lateral line canal with tight undulations along entire length 

3 body silver, occasionally with poorly defined dark longitudinal stripes 

4 1 st dorsal-fin spine long, extending to near or beyond fin apex 

5 2 nd dorsal fin dusky with a darker outer margin 

6 snout blunt, fleshy and short 

7 tail whip-like 

8 preopercular and oral lateral-line canals branch separately from the infraorbital canal 

Size: To at least 110 cm; males and females mature at >65 cm. 

Distribution: Western North Pacific, from Japan south to the Philippines and Borneo. 

Habitat and biology: Demersal on continental shelves and upper slopes in depths of 

20–550 m (150–200 m off Sarawak). Oviparous, depositing elliptical egg capsules (20 

cm in length) which have numerous longitudinal lines on surface. Diet consists of small 

benthic animals. 

Utilisation: Rarely caught in deeper waters by longlines and trawlers. Probably used for 

its meat as the flesh is good quality.

C h i m a e r i f o r m e s 267 


Local synonyms: Chimaera sp.: Yano et al. (2005). 

References: Masuda et al. (1975); Dagit (2006). 

6 

Lateral head 

1 

8 

Lateral tail 


Sarawak. 

7 

Chimaeridae (Shortnose Chimaeras) 

2 

Trunk lateral line canal


Acknowledgements 

Many people have helped in one way or another in both the preparation of the book 

and in the project itself. This ground-breaking, joint Indonesian–Malaysian–American– 

Australian project was made possible primarily by the financial support of the National 

Science Foundation (NSF) since 2002 (grants NSF BS&I Nos. DEB 0103640, DEB 

0542941, DEB 0542846). Funding support was also made available from a Lerner-Gray 

post-doctoral award to KJ in 2006 and also the Judith Humphry Shaw fund. 

The project could not have succeeded without the assistance and advice of numerous 

colleagues. We would particularly like to thank: Dato’ Junaidi bin Che Ayub (Director 

General of Fisheries Malaysia), Raja Mohammad Noordin bin Raja Omar (Director 

of Fisheries Research Institute Malaysia), Albert Chuan Gambang (Head of Fisheries 

Research Institute Sarawak), Saleem Mustafa (Director, Borneo Marine Research Institute, 

Universiti Malaysia Sabah), Ridzwan Abdul Rahman (former Director, Borneo Marine 

Research Institute, Universiti Malaysia Sabah), Edward Rooney Biusing (former Deputy 

Director Fisheries Sabah) and Mohammed Adrim (LIPI). Colleagues participating in the 

UK Darwin project, particularly Sarah Fowler, Leonard Compagno, Rachael Cavanagh, 

Scott Mycock, Sid Cook and Michael Sugden, helped set the scene for this project. 

The genetics component of the project was made possible by the help of a number of 

colleagues, particularly Kerri Matthes, Clemens Lakner, Neil Aschliman (Florida State 

University) and Shannon Corrigan (CSIRO). 

Projects such as these are not possible without assistance during the numerous surveys 

and we would like to thank the following people for their help in the field: Loren Caira, 

Claire Healy (Royal Ontario Museum, Toronto), Hamri bin Hgi-Kating (Mukah trawler, 

Sarawak), Latip Sait (Sematan trawler, Sarawak), Asni Etin, Masni Etin and Salik Etin 

(Kampung Abai, Kinabatangan), staff of the Sabah Fisheries Department, Semporna 

district (for arranging use of their boat), Anchain and Roslie (Kampung Tetabuan, 

Sabah), the village head and family of Kampung Tetabuan at Labuk Bay, staff of UMS 

Seaweed Project, Pulau Banggi, Kudat district (for arranging use of their boat), Justin 

Tampuling (Universiti Malaysia Sabah), Mengke (Sungai Kakap), Junardi and Mugi 

Utomo (University of Tanjungpura), and the numerous people at each of the landing sites 

visited. Thanks also to Zainal Abidin and family (Kuching Fisheries, Sarawak), Rayner 

Galid (Director of Fisheries, Kota Kinabalu), Charles Greenwald (Jakarta), Isnol (fisheries 

officer, Sungai Kakap, Kalimantan), Alang (Jungkat ray collector, Kalimantan), Albert 

Gambang (Kuching Fisheries, Sarawak) for allowing us to participate on a cruise on their 

vessel KK Manchong. 

Survey trips would not be possible without acquisition of collecting permits and we 

acknowledge the following for their assistance: Annadel Cabanban (Universiti Malaysia 

Sabah), Margarita Naming and Madam Eileen Yen (Sarawak Biodiversity Center), George

A c k n o w l e d g e m e n t s 269 

Chong (Kuching Fisheries, Sarawak), Suharsono (Director of Research Centre for 

Oceanography LIPI, Jakarta), Ono Kurnaen Sumadhinarga (LIPI, Jakarta), Dedy Darnaedi 

(LIPI, Bogor), and Ruben Silitonga and Sri Wahyono (LIPI, Jakarta). 

The collections in Malaysian Borneo were conducted under the following permits 

and research agreements: UPE:40/200/19SJ.924 and UPE:40/200/19SJ.925 from the 

Economic Planning Unit in Kuala Lumpur; No. JKM 100-24/13/1/223(59) from the 

Chief Minister’s Department, Kota Kinabalu, Sabah, Malaysia; No. SBC-RA-0050-JNC 

from the Sarawak Biodiversity Center in Kuching; permit Nos. 06252/SU.3/KS/2006 and 

3861/SU.3/KS/2007 from LIPI in Jakarta, and 1586/FRP/SM/VII/2008 from RISTEK 

in Jakarta. Specimens collected during the project were deposited at various ichthyological 

collections and we thank the staff at these facilities, particularly Renny Kurnia Hadiaty 

and Ahmad Jauhar Arief (Museum Zoologicum Bogoriense, MZB, Cibinong), Albert Lo 

and George Jonis (Sabah Museum, Kota Kinabalu); Alastair Graham (CSIRO Marine & 

Atmospheric Research), Dave Catania (California Academy of Sciences), and Morice 

Rowan, Stefanus Simon and Syuhaime Ahmat Ali (Borneo Marine Research Institute, 

Universiti Malaysia Sabah, Kota Kinabalu). A number of taxonomic issues arose during 

this project and we would like to thank the following people for providing material for 

comparison and assistance during museum visits: Martien van Oijen and Ronald de Ruiter 

(Rijksmuseum van Natuurlijke Histoire, RMNH, Leiden); Romain Causse, Bernard Séret, 

Guy Duhamel, Patrice Pruvost and Zora Gabsi (Muséum national d’Histoire naturelle, 

MNHN, Paris); Patrick Campbell and Oliver Crimmen (British Museum of Natural 

History, BMNH, London); Peter Bartsch (Museum fur Naturkunde, ZMB, Berlin); Ronald 

Vonk and Hielke Praagman (Zoologisch Museum, ZMA, Amsterdam); Gento Shinohara 

(National Science Museum, NSMT, Tokyo); Toshio Kawai (Hokkaido University, HUMZ, 

Hokkaido); Masato Moteki (Tokyo University Marine Science, MTUF, Tokyo); Kelvin 

Lim (National University of Singapore, NUS); Renny Kurnia Hadiaty and Ahmad Jauhar 

Arief (Museum Zoologicum Bogoriense, MZB, Jakarta); and Aaron Henderson (Sultan 

Qaboos University, Oman) and Alec Moore (RSK Group, UK). 

The following CSIRO Marine & Atmospheric Research (Wealth from Oceans Flagship) 

staff are also thanked: David Smith for project support, Louise Bell for cover page design, 

Louise Conboy for image preparation and acquistion, Melody Puckridge and Bob Ward 

for associated Barcode of Life input for some of the difficult taxa, Dan Gledhill for 

technical assistance, Joel MacKeen and Meredith Hepburn for acquisition of references, 

Tim Fountain for acquisition of meristic data, John Manger and Briana Elwood (CSIRO 

Publishing) for access to illustrations and map templates as well as assistance with aspects of 

book publishing. Thanks also to Georgina Davis, Stephanie Barnes and Lindsay Marshall 

(www.stickfigurefish.com.au) for providing high quality illustrations for the book. 

Some images were sourced from external localities and those that provided these images 

are thanked: Alec Moore (RSK Group, UK), Dr O. Gadig (Universidade Estadual Paulista, 

Brazil), Mike Severns (Hawaii) and Dr J. Randall (Bishop Museum, Hawaii).


Images sourced outside Borneo 

Hexanchus griseus: lateral (Brazil). Centrophorus moluccensis: lateral (Lombok, Indonesia). 

Orectolobus leptolineatus: ventral head, embryo (Lombok, Indonesia). Chiloscyllium 

plagiosum: juvenile (Hong Kong). Nebrius ferrugineus: lateral, ventral head (Western 

Australia); dorsal fins (Singapore). Stegostoma fasciatum: lateral, ventral head (Queensland, 

Australia). Rhincodon typus: lateral (Japan). Alopias pelagicus: lateral, ventral head (Bali, 

Indonesia). Isurus oxyrinchus: lateral, ventral head (Lombok, Indonesia). Apristurus 

platyrhynchus: lateral, ventral head (New South Wales, Australia); flank denticles (Western 

Australia). Halaelurus maculosus: lateral, ventral head, dorsal head and anterior trunk (Bali, 

Indonesia); egg case (Lombok, Indonesia). Chaenogaleus macrostoma: lateral (Qatar); ventral 

head (West Java, Indonesia). Carcharhinus albimarginatus: lateral, ventral head (Western 

Australia). Carcharhinus amblyrhynchos: embryo (Lombok, Indonesia). Carcharhinus 

dussumieri: ventral head (West Java, Indonesia). Carcharhinus falciformis: lateral, ventral 

head (Lombok, Indonesia). Carcharhinus plumbeus: lateral (Bali, Indonesia). Galeocerdo 

cuvier: lateral (Bali, Indonesia); ventral head (Queensland, Australia). Prionace glauca: lateral 

(Tasmania, Australia); ventral head (Queensland, Australia).Triaenodon obesus: lateral 

(Bali, Indonesia). Eusphyra blochii: lateral (West Java, Indonesia); ventral head (northern 

Australia). Anoxypristis cuspidata: dorsal, ventral head, caudal fin (Queensland, Australia). 

Pristis microdon: dorsal, ventral head, caudal fin (Queensland, Australia). Pristis zijsron: dorsal, 

ventral head, caudal fin (Queensland, Australia). Rhina ancylostoma: dorsal (Bali, Indonesia); 

lateral head (East Java, Indonesia). Glaucostegus typus: ventral head (Queensland, Australia); 

lateral tail (West Papua, Indonesia). Plesiobatis daviesi: dorsal (West Java, Indonesia); oronasal 

(New South Wales, Australia). Dasyatis microps: dorsal (Gulf of Carpentaria, Australia). 

Dasyatis ushiei: dorsal, ventral disc, nuchal area (Lombok, Indonesia). Himantura fai: nuchal 

area (Queensland, Australia). Himantura granulata: dorsal (East Java, Indonesia); nuchal 

area (Queensland, Australia). Himantura jenkinsii: dorsal tail thorns, nuchal area (West 

Java, Indonesia). Himantura leoparda: ventral disc, nuchal area (Queensland, Australia), tail 

section - dorsal and ventral (Manila, Philippines). Himantura uarnacoides: nuchal area (West 

Java, Indonesia). Pteroplatytrygon violacea: dorsal (Lombok, Indonesia); ventral disc (Bali, 

Indonesia). Taeniurops meyeni: dorsal, ventral disc (Bali, Indonesia). Urogymnus asperrimus: 

dorsal (Marshall Islands). Gymnura zonura: lateral anterior tail (Bali, Indonesia). Aetobatus 

flagellum: dorsal, ventral head (West Java, Indonesia). Aetomylaeus vespertilio: dorsal, ventral 

head (Queensland, Australia). Rhinoptera javanica: ventral head, lateral head, lateral dorsal 

fin and anterior tail (Queensland, Australia). Rhinoptera jayakari: lateral head, lateral dorsal 

fin and anterior tail (Lombok, Indonesia). Manta birostris: dorsal, lateral dorsal fin and 

anterior tail (Lombok, Indonesia); embryo (West Java, Indonesia). Mobula japanica: dorsal, 

dorsolateral head (Lombok, Indonesia); lateral dorsal fin and sting (East Java, Indonesia); 

ventral head (Bali, Indonesia). Mobula thurstoni: dorsal, ventral head, dorsolateral head, 

lateral dorsal fin and anterior tail (Lombok, Indonesia).

271 

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Checklist of Borneo sharks and rays 

Order Hexanchiformes: Cow and Frilled Sharks 

Family Hexanchidae: Sixgill and Sevengill Sharks 

Hexanchus griseus (Bonnaterre, 1788) Bluntnose Sixgill Shark 

Order Squaliformes: Dogfish Sharks 

Family Squalidae: Dogfishes 

Squalus altipinnis Last et al., 2007 

Family Centrophoridae: Gulper Sharks 

Centrophorus moluccensis Bleeker, 1860 

Order Squatiniformes: Angelsharks 

Family Squatinidae: Angelsharks 

Squatina tergocellatoides Chen, 1963 

Order Heterodontiformes: Hornsharks 

Family Heterodontidae: Hornsharks 

Heterodontus zebra (Gray, 1831) 

Order Orectolobiformes: Carpet Sharks 

Family Orectolobidae: Wobbegongs 

Orectolobus leptolineatus Last et al., 2010c 

Family Hemiscylliidae: Longtail Carpetsharks 

Chiloscyllium hasseltii Bleeker, 1852 

Chiloscyllium indicum (Gmelin, 1789) 

Chiloscyllium plagiosum (Bennett, 1830) 

Chiloscyllium punctatum M. & H., 1838 

Family Ginglymostomatidae: Nurse Sharks 

Nebrius ferrugineus (Lesson, 1830) 

Family Stegostomatidae: Zebra Sharks 

Stegostoma fasciatum (Hermann, 1783) 

Western Highfin Spurdog 

Smallfin Gulper Shark 

Ocellated Angelshark 

Zebra Hornshark 

Indonesian Wobbegong 

Indonesian Bambooshark 

Slender Bambooshark 

Whitespotted Bambooshark 

Brownbanded Bambooshark 

Tawny Nurse Shark 

Zebra Shark


Family Rhincodontidae: Whale Sharks 

Rhincodon typus Smith, 1828 

Order Lamniformes: Mackerel Sharks 

Family Alopiidae: Thresher Sharks 

Alopias pelagicus Nakamura, 1935 

Family Lamnidae: Mackerel Sharks 

Isurus oxyrinchus Rafinesque, 1810 

Whale Shark 

Pelagic Thresher 

Shortfin Mako 

Order Carcharhiniformes: Ground Sharks 

Family Scyliorhinidae: Catsharks 

Apristurus platyrhynchus (Tanaka, 1909) Bigfin Catshark 

Atelomycterus marmoratus (Bennett, 1830) Coral Catshark 

Cephaloscyllium sarawakensis Yano et al., 2005 Sarawak Swellshark 

Halaelurus maculosus White et al., 2007 Indonesian Speckled Catshark 

Family Triakidae: Houndsharks 

Mustelus manazo Bleeker, 1854 

Starspotted Smoothhound 

Mustelus widodoi White & Last, 2006 Whitefin Smoothhound 

Family Hemigaleidae: Weasel Sharks 

Chaenogaleus macrostoma (Bleeker, 1852) Hooktooth Shark 

Hemigaleus microstoma Bleeker, 1852 Sicklefin Weasel Shark 

Hemipristis elongata (Klunzinger, 1871) Fossil Shark 

Paragaleus tengi (Chen, 1963) 

Straight-tooth Weasel Shark 

Family Carcharhinidae: Whaler Sharks 

Carcharhinus albimarginatus (Rüppell, 1837) Silvertip Shark 

Carcharhinus amblyrhynchoides (Whitley, 1934) Graceful Shark 

Carcharhinus amblyrhynchos (Bleeker, 1856) Grey Reef Shark 

Carcharhinus borneensis (Bleeker, 1858) Borneo Shark 

Carcharhinus brevipinna (M. & H., 1839) Spinner Shark 

Carcharhinus dussumieri (M. & H., 1839) Whitecheek Shark 

Carcharhinus falciformis (M. & H., 1839) Silky Shark

C h e c k l i s t o f s p e c i e s 279 

Carcharhinus leucas (M. & H., 1839) 

Carcharhinus limbatus (M. & H., 1839) 

Carcharhinus macloti (M. & H., 1839) 

Carcharhinus melanopterus (Q. & G., 1824) 

Carcharhinus plumbeus (Nardo, 1827) 

Carcharhinus sealei (Pietschmann, 1913) 

Carcharhinus sorrah (M. & H., 1839) 

Galeocerdo cuvier (Péron & Lesueur, 1822) 

Glyphis sp. 

Glyphis fowlerae Compagno et al., 2010 

Lamiopsis tephrodes (Fowler, 1905) 

Loxodon macrorhinus M. & H., 1839 

Prionace glauca (Linnaeus, 1758) 

Rhizoprionodon acutus (Rüppell, 1837) 

Rhizoprionodon oligolinx Springer, 1964 

Scoliodon macrorhynchos (Bleeker, 1852) 

Triaenodon obesus (Rüppell, 1837) 

Family Sphyrnidae: Hammerhead Sharks 

Eusphyra blochii (Cuvier, 1816) 

Sphyrna lewini (Griffith & Smith, 1834) 

Sphyrna mokarran (Rüppell, 1837) 

Bull Shark 

Common Blacktip Shark 

Hardnose Shark 

Blacktip Reef Shark 

Sandbar Shark 

Blackspot Shark 

Spot-tail Shark 

Tiger shark 

Mukah River Shark 

Borneo River Shark 

Borneo Broadfin Shark 

Sliteye Shark 

Blue Shark 

Milk Shark 

Grey Sharpnose Shark 

Pacific Spadenose Shark 

Whitetip Reef Shark 

Winghead Shark 

Scalloped Hammerhead 

Great Hammerhead 

Order Rajiformes: Rays (Batoids) 

Suborder Pristoidei: Sawfishes 

Family Pristidae: Sawfishes 

Anoxypristis cuspidata (Latham, 1794) 

Pristis microdon Latham, 1794 

Pristis zijsron Bleeker, 1851 

Narrow Sawfish 

Freshwater Sawfish 

Green Sawfish


Suborder Rhinoidei: Shark Rays 

Family Rhinidae: Shark Rays 

Rhina ancylostoma Bl. & Sch., 1801 

Suborder Rhynchobatoidei: Wedgefishes 

Family Rhynchobatidae: Wedgefishes 

Rhynchobatus australiae Whitley, 1939 

Rhynchobatus laevis (Bl. & Sch., 1801) 

Rhynchobatus springeri Comp. & Last, 2010 

Suborder Rhinobatoidei: Shovelnose Rays 

Family Rhinobatidae: Shovelnose Rays 

Glaucostegus thouin (Anonymous, 1798) 

Glaucostegus typus (Bennett, 1830) 

Rhinobatos formosensis Norman, 1926 

Suborder Torpedinoidei: Electric Rays 

Family Narcinidae: Numbfishes 

Narcine brevilabiata Bessednov, 1966 

Narcine lingula Richardson, 1846 

Narcine maculata (Shaw, 1804) 

Family Narkidae: Sleeper Rays 

Narke dipterygia (Bl. & Sch., 1801) 

Temera hardwickii Gray, 1831 

Suborder Rajoidei: Skates 

Family Rajidae: Skates 

Dipturus kwangtungensis (Chu 1960) 

Okamejei cairae Last et al., 2010a 

Okamejei hollandi (Jordan & Rich., 1909) 

Okamejei jensenae Last & Lim, 2010 

Shark Ray 

Whitespotted Wedgefish 

Smoothnose Wedgefish 

Broadnose Wedgefish 

Clubnose Guitarfish 

Giant Guitarfish 

Taiwan Guitarfish 

Shortlip Numbfish 

Rough Numbfish 

Darkspotted Numbfish 

Spot-tail Sleeper Ray 

Finless Sleeper Ray 

Kwangtung Skate 

Borneo Sand Skate 

Yellowspotted Skate 

Philippine Ocellate Skate

C h e c k l i s t o f s p e c i e s 281 

Family Anacanthobatidae: Leg Skates 

Sinobatis borneensis (Chan, 1965) 

Borneo Leg Skate 

Suborder Myliobatoidei: Stingrays 

Family Plesiobatidae: Giant Stingarees 

Plesiobatis daviesi (Wallace, 1967) 

Giant Stingaree 

Family Dasyatidae: Stingrays 

Dasyatis microps (Annandale, 1908) Smalleye Stingray 

Dasyatis parvonigra Last & White, 2008 Dwarf Black Stingray 

Dasyatis ushiei (Jordan & Hubbs, 1925) Cow Stingray 

Dasyatis zugei (M. & H., 1841) 

Sharpnose Stingray 

Himantura fai Jordan & Seale, 1906 Pink Whipray 

Himantura gerrardi (Gray, 1851) 

Whitespotted Whipray 

Himantura cf gerrardi * 

Borneo Whitespotted Whipray 

Himantura granulata (Macleay, 1883) Mangrove Whipray 

Himantura jenkinsii (Annandale, 1909) Jenkins’ Whipray 

Himantura kittipongi Vidth. & Rob., 2005 Roughback Whipray 

Himantura leoparda Manjaji-M. & Last, 2008 Leopard Whipray 

Himantura lobistoma Manjaji-M. & Last, 2006 Tubemouth Whipray 

Himantura oxyrhyncha (Sauvage, 1878) Longnose Marbled Whipray 

Himantura pastinacoides (Bleeker, 1852) Round Whipray 

Himantura polylepis (Bleeker, 1852) Giant Freshwater Whipray 

Himantura signifer Compagno & Rob., 1982 White-edge Freshwater Whipray 

Himantura uarnacoides (Bleeker, 1852) Bleeker’s Whipray 

Himantura uarnak (Forsskål, 1775) Reticulate Whipray 

Himantura cf uarnak * 

Finespot Whipray 

Himantura undulata (Bleeker, 1852) Honeycomb Whipray 

Himantura walga (M. & H., 1841) Dwarf Whipray 

Neotrygon kuhlii (M. & H. 1841) 

Bluespotted Maskray 

Pastinachus atrus (Macleay, 1883) 

Eastern Cowtail Stingray


Pastinachus gracilicaudus Last & M.-M., 2010 

Pastinachus solocirostris Last et al., 2005 

Pastinachus stellurostris Last et al., 2010b 

Pteroplatytrygon violacea (Bonaparte, 1832) 

Taeniura lymma (Forsskål, 1775) 

Taeniurops meyeni (M. & H., 1841) 

Urogymnus asperrimus (Bl. & Sch., 1801) 

Family Gymnuridae: Butterfly Rays 

Gymnura poecilura (Shaw, 1804) 

Gymnura zonura (Bleeker, 1852) 

Family Myliobatidae: Eagle Rays 

Aetobatus flagellum (Bl. & Sch., 1801) 

Aetobatus ocellatus (Kuhl, 1823) 

Aetomylaeus maculatus (Gray, 1834) 

Aetomylaeus nichofii (Bl. & Sch., 1801) 

Aetomylaeus vespertilio (Bleeker, 1852) 

Family Rhinopteridae: Cownose Rays 

Rhinoptera javanica M. & H., 1841 

Rhinoptera jayakari Boulenger, 1895 

Family Mobulidae: Devilrays 

Manta birostris (Walbaum, 1792) 

Mobula sp. 

Mobula japanica (M. & H., 1841) 

Mobula kuhlii (M. & H., 1841) 

Mobula thurstoni (Lloyd, 1908) 

Narrowtail Stingray 

Roughnose Stingray 

Starynose Stingray 

Pelagic Stingray 

Bluespotted Fantail Ray 

Blotched Fantail Stingray 

Porcupine Ray 

Longtail Butterfly Ray 

Zonetail Butterfly Ray 

Longhead Eagle Ray 

Whitespotted Eagle Ray 

Mottled Eagle Ray 

Banded Eagle Ray 

Ornate Eagle Ray 

Javanese Cownose Ray 

Short-tail Cownose Ray 

Manta Ray 

Borneo Devilray 

Japanese Devilray 

Lesser Devilray 

Bentfin Devilray 

Order Chimaeriformes: Chimaeras 

Family Chimaeridae: Shortnose Chimaeras 

Chimaera phantasma Jordan & Snyder, 1900 Silver Chimaera 

* denotes species for which full treatments are not provided in the guide.

283 

Scientific names index 

A 

acutus, Rhizoprionodon 126 

Aetobatus flagellum 242 

Aetobatus ocellatus 244 

Aetomylaeus maculatus 246 

Aetomylaeus nichofii 248 

Aetomylaeus vespertilio 250 

albimarginatus, Carcharhinus 86 

Alopias pelagicus 62 

Alopiidae 29, 62–63 

altipinnis, Squalus 38 

amblyrhynchoides, Carcharhinus 88 

amblyrhynchos, Carcharhinus 90 

Anacanthobatidae 25, 178–179 

ancylostoma, Rhina 146 

Anoxypristis cuspidata 140 

Apristurus platyrhynchus 66 

asperrimus, Urogymnus 236 

Atelomycterus marmoratus 68 

atrus, Pastinachus 222 

australiae, Rhynchobatus 148 

B 

birostris, Manta 256 

blochii, Eusphyra 134 

borneensis, Carcharhinus 92 

borneensis, Sinobatis 178 

brevilabiata, Narcine 160 

brevipinna, Carcharhinus 94 

C 

cairae, Okamejei 172 

Carcharhinidae 33, 86–133 

Carcharhiniformes 66–139 

Carcharhinus albimarginatus 86 

Carcharhinus amblyrhynchoides 88 

Carcharhinus amblyrhynchos 90 

Carcharhinus borneensis 92 

Carcharhinus brevipinna 94 

Carcharhinus dussumieri 96 

Carcharhinus falciformis 98 

Carcharhinus leucas 100 

Carcharhinus limbatus 102 

Carcharhinus macloti 104 

Carcharhinus melanopterus 106 

Carcharhinus plumbeus 108 

Carcharhinus sealei 110 

Carcharhinus sorrah 112 

Centrophoridae 29, 40–41 

Centrophorus moluccensis 40 

Cephaloscyllium sarawakensis 70 

Chaenogaleus macrostoma 78 

Chiloscyllium hasseltii 48 

Chiloscyllium indicum 50 

Chiloscyllium plagiosum 52 

Chiloscyllium punctatum 54 

Chimaera phantasma 266 

Chimaeridae 23, 266–267 

Chimaeriformes 266–267 

cuspidata, Anoxypristis 140 

cuvier, Galeocerdo 114 

D 

Dasyatidae 26, 182–237 

Dasyatis microps 182 

Dasyatis parvonigra 184 

Dasyatis ushiei 186 

Dasyatis zugei 188 

daviesi, Plesiobatis 180 

dipterygia, Narke 166 

Dipturus kwangtungensis 170 

dussumieri, Carcharhinus 96 

E 

elongata, Hemipristis 82


Eusphyra blochii 134 

F 

fai, Himantura 190 

falciformis, Carcharhinus 98 

fasciatum, Stegostoma 58 

ferrugineus, Nebrius 56 

flagellum, Aetobatus 242 

formosensis, Rhinobatos 158 

fowlerae, Glyphis 118 

G 

Galeocerdo cuvier 114 

gerrardi, Himantura 192 

Ginglymostomatidae 30, 56–57 

glauca, Prionace 124 

Glaucostegus thouin 154 

Glaucostegus typus 156 

Glyphis fowlerae 118 

Glyphis sp. 116 

gracilicaudus, Pastinachus 224 

granulata, Himantura 194 

griseus, Hexanchus 36 

Gymnura poecilura 238 

Gymnura zonura 240 

Gymnuridae 26, 238–241 

H 

Halaelurus maculosus 72 

hardwickii, Temera 168 

hasseltii, Chiloscyllium 48 

Hemigaleidae 32, 78–85 

Hemigaleus microstoma 80 

Hemipristis elongata 82 

Hemiscylliidae 31, 48-55 

Heterodontidae 30, 44–45 

Heterodontiformes 44–45 

Heterodontus zebra 44 

Hexanchidae 28, 36–37 

Hexanchiformes 36–37 

Hexanchus griseus 36 

Himantura fai 190 

Himantura gerrardi 192 

Himantura granulata 194 

Himantura jenkinsii 196 

Himantura kittipongi 198 

Himantura leoparda 200 

Himantura lobistoma 202 

Himantura oxyrhyncha 204 

Himantura pastinacoides 206 

Himantura polylepis 208 

Himantura signifer 210 

Himantura uarnacoides 212 

Himantura uarnak 214 

Himantura undulata 216 

Himantura walga 218 

hollandi, Okamejei 174 

I 

indicum, Chiloscyllium 50 

Isurus oxyrinchus 64 

J 

japanica, Mobula 260 

javanica, Rhinoptera 252 

jayakari, Rhinoptera 254 

jenkinsii, Himantura 196 

jensenae, Okamejei 176 

K 

kittipongi, Himantura 198 

kuhlii, Mobula 262 

kuhlii, Neotrygon 220 

kwangtungensis, Dipturus 170 

L 

laevis, Rhynchobatus 150 

Lamiopsis tephrodes 120 

Lamnidae 31, 64–65 

Lamniformes 62–63

RS cu ine nitn i g f i c h en adm e s i n d e x 285 

leoparda, Himantura 200 

leptolineatus, Orectolobus 46 

leucas, Carcharhinus 100 

lewini, Sphyrna 136 

limbatus, Carcharhinus 102 

lingula, Narcine 162 

lobistoma, Himantura 202 

Loxodon macrorhinus 122 

lymma, Taeniura 232 

M 

macloti, Carcharhinus 104 

macrorhinus, Loxodon 122 

macrorhynchos, Scoliodon 130 

macrostoma, Chaenogaleus 78 

maculata, Narcine 164 

maculatus, Aetomylaeus 246 

maculosus, Halaelurus 72 

manazo, Mustelus 74 

Manta birostris 256 

marmoratus, Atelomycterus 68 

melanopterus, Carcharhinus 106 

meyeni, Taeniurops 234 

microps, Dasyatis 182 

microstoma, Hemigaleus 80 

Mobula japanica 260 

Mobula kuhlii 262 

Mobula sp. 258 

Mobula thurstoni 264 

Mobulidae 27, 256–265 

mokarran, Sphyrna 138 

moluccensis, Centrophorus 40 

Mustelus manazo 74 

Mustelus widodoi 76 

Myliobatidae 28, 242–251 

Myliobatoidei 180–265 

N 

Narcine brevilabiata 160 

Narcine lingula 162 

Narcine maculata 164 

Narcinidae 25, 160–165 

Narke dipterygia 166 

Narkidae 25, 166–169 

Nebrius ferrugineus 56 

Neotrygon kuhlii 220 

nichofii, Aetomylaeus 248 

O 

obesus, Triaenodon 132 

ocellatus, Aetobatus 244 

Okamejei cairae 172 

Okamejei hollandi 174 

Okamejei jensenae 176 

oligolinx, Rhizoprionodon 128 

Orectolobidae 30, 46–47 

Orectolobiformes 46–61 

Orectolobus leptolineatus 46 

oxyrhyncha, Himantura 204 

oxyrinchus, Isurus 64 

P 

Paragaleus tengi 84 

parvonigra, Dasyatis 184 

Pastinachus atrus 222 

Pastinachus gracilicaudus 224 

Pastinachus solocirostris 226 

Pastinachus stellurostris 228 

pastinacoides, Himantura 206 

pelagicus, Alopias 62 

phantasma, Chimaera 266 

plagiosum, Chiloscyllium 52 

platyrhynchus, Apristurus 66 

Plesiobatidae 26, 180–181 

Plesiobatis daviesi 180 

plumbeus, Carcharhinus 108 

poecilura, Gymnura 238 

polylepis, Himantura 208 

Prionace glauca 124 

Pristidae 23, 140–145


Pristis microdon 142 

Pristis zijsron 144 

Pristoidei 140–145 

Pteroplatytrygon violacea 230 

punctatum, Chiloscyllium 54 

R 

Rajidae 25, 172–179 

Rajoidei 172–179 

Rhina ancylostoma 146 

Rhincodontidae 30, 60–61 

Rhincodon typus 60 

Rhinidae 24, 146–147 

Rhinobatidae 24, 154–159 

Rhinobatoidei 154–159 

Rhinobatos formosensis 158 

Rhinoidei 146–147 

Rhinoptera javanica 252 

Rhinoptera jayakari 254 

Rhinopteridae 28, 252–255 

Rhizoprionodon acutus 126 

Rhizoprionodon oligolinx 128 

Rhynchobatidae 24, 148–153 

Rhynchobatoidei 147 

Rhynchobatus australiae 148 

Rhynchobatus laevis 150 

Rhynchobatus springeri 152 

S 

sarawakensis, Cephaloscyllium 70 

Scoliodon macrorhynchos 130 

Scyliorhinidae 31, 66–73 

sealei, Carcharhinus 110 

signifer, Himantura 210 

Sinobatis borneensis 178 

solocirostris, Pastinachus 226 

sorrah, Carcharhinus 112 

sp., Glyphis 116 

sp., Mobula 258 

Sphyrna lewini 136 

Sphyrna mokarran 138 

Sphyrnidae 29, 134–139 

springeri, Rhynchobatus 152 

Squalidae 29, 38–39 

Squaliformes 38–41 

Squalus altipinnis 38 

Squatina tergocellatoides 42 

Squatinidae 23, 42–43 

Squatiniformes 42–43 

Stegostoma fasciatum 58 

Stegostomatidae 30, 58–59 

stellurostris, Pastinachus 228 

T 

Taeniura lymma 232 

Taeniurops meyeni 234 

Temera hardwickii 168 

tengi, Paragaleus 84 

tephrodes, Lamiopsis 120 

tergocellatoides, Squatina 42 

thouin, Glaucostegus 154 

thurstoni, Mobula 264 

Torpedinoidei 160–169 

Triaenodon obesus 132 

Triakidae 32, 74–77 

typus, Glaucostegus 156 

typus, Rhincodon 60 

U 

uarnacoides, Himantura 212 

uarnak, Himantura 214 

undulata, Himantura 216 

Urogymnus asperrimus 236 

ushiei, Dasyatis 186 

V 

vespertilio, Aetomylaeus 250 

violacea, Pteroplatytrygon 230 

W 

walga, Himantura 218

S c i e n t i f i c n a m e s i n d e x 287 

widodoi, Mustelus 76 

Z 

zebra, Heterodontus 44 

zonura, Gymnura 240 

zugei, Dasyatis 188


English common names index 

A 

Angelsharks 42–43 

B 

Banana-tail Ray 222 

Banded Eagle Ray 248 

Banded Whiptail Ray 192 

Barbless Eagle Ray 248 

Bentfin Devilray 264 

Bigfin Catshark 66 

Blackblotched Stingray 234 

Blackspot Shark 110 

Blackspotted Stingray 234 

Blacktip Reef Shark 106 

Blacktip Shark 106 

Blacktip Whaler 102 

Bleeker’s Butterfly Ray 240 

Bleeker’s Variegate Ray 216 

Bleeker’s Whipray 212 

Blotched Fantail Stingray 234 

Blue-spotted Lagoon Ray 232 

Blue-spotted Ribbontail Ray 232 

Blue Pointer 64 

Blue Shark 124 

Bluespotted Fantail Ray 232 

Bluespotted Maskray 220 

Bluespotted Stingaree 220 

Bluespotted Stingray 220 

Blue Stingray 230 

Blue Whaler 124 

Blunthead Shark 132 

Bluntnose Sixgill Shark 36 

Bluntnose Whiptail Ray 192 

Bonnet Skate 244 

Borneo Broadfin Shark 120 

Borneo Devilray 258 

Borneo Leg Skate 178 

Borneo River Shark 118 

Borneo Sand Skate 172 

Borneo Shark 92 

Bowmouth Guitarfish 146 

Broadnose Wedgefish 152 

Brownbanded Bambooshark 54 

Brownbanded Catshark 54 

Brown Shark 108 

Bullhead Shark 44 

Bull Ray 222, 234 

Bull Shark 36, 100 

Butterfly Rays 238–241 

C 

Catsharks 66–73 

Chinese Numbfish 162 

Clubnose Guitarfish 154 

Coachwhip Ray 194, 214 

Common Blacktip Shark 102 

Common Shovelnose Ray 156 

Coral Catshark 68 

Cownose Rays 252–255 

Cow Ray 252 

Cow Shark 36 

Cowsharks 36–37 

Cow Stingray 186 

D 

Darkfin Numbfish 164 

Darkspotted Electric Ray 164 

Darkspotted Numbfish 164 

Deepwater Stingray 180 

Devilfish 256 

Devilrays 256–265 

Dogfishes 38–39 

Duckbill Ray 244

E n g l i s h c o m m o n n a m e s i n d e x 289 

Dwarf Black Stingray 184 

Dwarf Whipray 218 

E 

Eagle Rays 242–251 

Eastern Cowtail Stingray 222 

Endeavour Dogfish 40 

F 

Fantail Ray 222 

Fantail Stingray 234 

Feathertail Ray 222 

Feathertail Stingray 222 

Finless Electric Ray 168 

Finless Sleeper Ray 168 

Fish Shark 126 

Flapnose Ray 252 

Fossil Shark 82 

Freshwater Sawfish 142 

Freshwater Stingray 210 

Freshwater Whaler 100 

Freshwater Whipray 208 

G 

Ghost Shark 266 

Giant Freshwater Stingray 208 

Giant Freshwater Whipray 208 

Giant Guitarfish 150, 156 

Giant Manta 256 

Giant Reef Ray 234 

Giant Shovelnose Ray 156 

Giant Stingaree 180 

Giant Stingarees 180–181 

Graceful Shark 88 

Great-tooth Sawfish 142 

Great Blue Shark 124 

Great Hammerhead 138 

Green Sawfish 144 

Grey Carpetshark 54 

Grey Reef Shark 90 

Grey Sharpnose Shark 128 

Guliman 106 

Gulper Sharks 40–41 

Gummy Shark 74 

H 

Hammerhead Sharks 134–139 

Hardnose Shark 104 

Hasselt’s Bambooshark 48 

Holland Skate 174 

Honeycomb Whipray 216 

Hooktooth Shark 78 

Hornsharks 44–45 

Houndsharks 74–77 

I 

Indonesian Bambooshark 48 

Indonesian Speckled Catshark 72 

Indonesian Wobbegong 46 

Indo Wobbegong 46 

Inkytail Shark 94 

J 

Japanese Devilray 260 

Japanese Smoothhound 74 

Javanese Cownose Ray 252 

Jenkins’ Whipray 196 

Jordan’s Blue Dogshark 122 

K 

Kidney-headed Shark 136 

Kuhl’s Stingray 220 

Kwangtung Skate 170 

L 

Lagoon Ray 232 

Leg Skates 178–179 

Leopard Shark 58 

Leopard Stingray 214 

Leopard Whipray 200 

Lesser Devilray 262


Lesser Fantail Ray 232 

Long-tailed Butterfly Ray 238 

Longhead Eagle Ray 242 

Longman’s Dogshark 126 

Longnose Blacktail Shark 90 

Longnose Grey Whaler 94 

Longnose Marbled Whipray 204 

Longtail Butterfly Ray 238 

Longtail Carpetsharks 48–55 

Longtail Ray 214 

M 

Mackerel Shark 64 

Mackerel Sharks 64–65 

Macleay’s Coachwhip Ray 194 

Madame X 56 

Mako Shark 64 

Mangrove Ray 194 

Mangrove Stingray 194 

Mangrove Whipray 194 

Manta 256 

Manta Ray 256 

Marbled Freshwater Stingray 204 

Marbled Whipray 204 

Marble Ribbontail Ray 234 

Milk Shark 126 

Mottled Eagle Ray 246 

Mud Shark 36 

Mud Skate 146 

Mukah River Shark 116 

N 

Narrow Sawfish 140 

Narrowsnout Sawfish 144 

Narrowtail Stingray 224 

Nieuhof ’s Eagle Ray 248 

Numbfishes 160–165 

Numbray 166 

Nurse Sharks 56–57 

O 

Ocellated Angelshark 42 

Ocellate Eagle Ray 246 

Ocellate Whipray 216 

Ornate Eagle Ray 246, 250 

P 

Pacific Spadenose Shark 130 

Pale-edged Stingray 188 

Pale Whipray 210 

Pelagic Stingray 230 

Pelagic Thresher 62 

Philippine Ocellate Skate 176 

Pink Whipray 190 

Pointed Sawfish 140 

Porcupine Ray 236 

Q 

Queensland Shark 88 

R 

Reef Ray 232 

Reticulate Eagle Ray 250 

Reticulate Whipray 214 

Ribbontail Stingray 232 

Ridgeback Bambooshark 50 

Ridgeback Shark 50 

River Whaler 100 

Rough-back Stingray 196 

Roughback Whipray 198 

Rough Electric Ray 162 

Roughnose Stingray 226 

Rough Numbfish 162 

Roughskin Stingaree 236 

Round Ribbontail Ray 234 

Round Whipray 206 

S 

Sandbar Shark 108 

Sandshark 150

E n g l i s h c o m m o n n a m e s i n d e x 291 

Sarawak Pygmy Swell Shark 70 

Sarawak Pygmy Swellshark 70 

Sarawak Swellshark 70 

Sawfish 144 

Sawfishes 140–145 

Scalloped Hammerhead 136 

School Shark 112 

Shark Ray 146 

Shark Rays 146–147 

Sharpnose Stingray 188, 192 

Short-tail Cownose Ray 254 

Shortfin Devilray 262 

Shortfin Mako 64 

Shorthorn Devilray 262 

Shortlip Electric Ray 160 

Shortlip Numbfish 160 

Shortnose Chimaeras 266–267 

Shovelnose Ray 156 

Shovelnose Rays 154–159 

Shovelnose Shark 150, 152, 156 

Sicklefin Weasel Shark 80 

Silky Shark 98 

Silver Chimaera 266 

Silvertip Shark 86 

Sixgill Shark 36 

Skates 170–177 

Sleeper Rays 166–169 

Sleepy Shark 56 

Slender Bambooshark 50 

Slender Dogshark 122 

Slender Hammerhead 134 

Sliteye Shark 122 

Smalleye Stingray 182 

Smallfin Gulper Shark 40 

Smoothfang Shark 94 

Smoothnose Wedgefish 150 

Smoothtail Mobula 264 

Snaggletooth Shark 82 

Snapper Shark 64 

Solander’s Ray 236 

Sorrah Shark 112 

Spatulasnout Catshark 66 

Speckle Stingray 234 

Spinetail Devilray 260 

Spinetail Mobula 260 

Spinner Shark 94 

Spitting Shark 56 

Spot-tail Electric Ray 166 

Spot-tail Shark 112 

Spot-tail Sleeper Ray 166 

Spotted Catshark 54 

Spotted Eagle Ray 244 

Starspotted Smoothhound 74 

Starynose Stingray 228 

Stingrays 182–237 

Straight-tooth Weasel Shark 84 

T 

Taiwan Guitarfish 158 

Tawny Nurse Shark 56 

Tawny Shark 56 

Thickskin Shark 108 

Thickspine Giant Stingray 182 

Thornback Ray 174 

Thorny Ray 236 

Thresher Sharks 62–63 

Tiger Shark 114 

Tubemouth Whipray 202 

U 

Undulate Whipray 200 

V 

Violet Stingray 230 

W 

Weasel Shark 80 

Weasel Sharks 78–85 

Wedgefishes 148–153


Western Highfin Spurdog 38 

Whaler Sharks 88–133 

Whale Shark 60 

Whale Sharks 60–61 

White-edge Freshwater Whipray 210 

White-eye Shark 126 

White-rim Stingray 210 

Whitecheek Shark 96 

Whitefin Smoothhound 76 

Whitenose Whipray 212 

Whitespot Ray 148 

Whitespot Shovelnose Ray 148 

Whitespotted Bambooshark 52 

Whitespotted Eagle Ray 244 

Whitespotted Guitarfish 148 

Whitespotted Wedgefish 148 

Whitespotted Whipray 192 

Whitetail Stingray 194 

Whitetail Whipray 194 

Whitetip Reef Shark 132 

Whitetip Shark 132 

Widemouth Blackspot Shark 96 

Wide Sawfish 142 

Winghead Shark 134 

Wobbegongs 46–47 

Y 

Yellow-spotted Skate 176 

Yellowspotted Skate 174 

Z 

Zebra Bullhead Shark 44 

Zebra Hornshark 44 

Zebra Port Jackson Shark 44 

Zebra Shark 58 

Zebra Sharks 58–59 

Zonetail Butterfly Ray 240

293 

Indonesian common names index 

C 

Cawang Genul 260 

Cawang Kalung 256 

Cucut Bekeman 100 

Cucut Botol 40 

Cucut Dolok 50, 52 

Cucut Gergaji 140 

Cucut Lanjaman 86, 88, 90, 94, 96, 98, 

102, 108, 110, 112 

Cucut Londer 74, 76 

Cucut Mandrong 154 

Cucut Meong 36 

Cucut Pedang 62 

Cucut Selendang 124 

Cucut Tokek 68 

H 

Hiu 44, 92, 148 

Hiu Aer 124 

Hiu Air 74, 76 

Hiu Anjing 64 

Hiu Aron 104 

Hiu Barong 146 

Hiu Batu 54 

Hiu Belimbing 58 

Hiu Bingkoh 134, 136, 138 

Hiu Bisu 56 

Hiu Bokem 132 

Hiu Bongo 52, 54 

Hiu Botol Danten 40 

Hiu Buas 100 

Hiu Bujit 88, 90, 94, 96, 98, 100, 

102, 106, 112, 120 

Hiu Capil 134, 136, 138 

Hiu Caping 134, 136, 138 

Hiu Cermin 156 

Hiu Coklat 132 

Hiu Gedebong 56 

Hiu Gedok 54, 56 

Hiu Gergaji 142, 144 

Hiu Hantu 266 

Hiu Jenggot 46 

Hiu Kacang 74, 76, 78, 80, 84 

Hiu Kakap 64 

Hiu Karang Buas 132 

Hiu Karet 124 

Hiu Kejen 102, 122, 130 

Hiu Kemenyan 148, 150, 152 

Hiu Kodok 42, 46 

Hiu Lalaek 124 

Hiu Lanyam 86, 98, 102 

Hiu Lepang 46 

Hiu Lonjor 90, 98 

Hiu Lontar 148, 150, 152 

Hiu Macan 114 

Hiu Mada 106 

Hiu Mako 64 

Hiu Martil 134, 136, 138 

Hiu Minsong 150, 152 

Hiu Monas 82 

Hiu Monyet 62 

Hiu Omas 114 

Hiu Parang 136, 138, 140, 142, 144 

Hiu Pasir 84 

Hiu Paus 60 

Hiu Pilus 78, 80, 126, 128 

Hiu Pisang 126, 128 

Hiu Plen 86, 94, 126 

Hiu Pogak 266 

Hiu Senget 40 

Hiu Sonteng 86 

Hiu Tahu Putih 36 

Hiu Taji 40 

Hiu Tekek 48, 50, 52, 54


Hiu Tenggiri 64 

Hiu Teteri 108 

Hiu Tikus 62 

Hiu Tokek 68, 72 

I 

Ikan Pari 184, 232 

K 

Kluyu Blimbingan 58 

Kluyu Karang 106 

Kupu-kupu Brangkas 146 

L 

Lampengan 260 

Lanyam 112 

Liongbun 148, 150, 152, 154 

M 

Merak Bulu 90, 94, 102, 112 

Minsong 148 

Mungsing 98, 112, 126, 130 

Mungsing Jara 114 

P 

Paitpait 156 

Pamprang 140 

Pangrum 148, 150, 152 

Parangpang 140 

Paredung 148, 150, 152 

Pari 172, 180, 204, 208 

Pari Aer 196, 206 

Pari Ayam 244 

Pari Babi 182, 234 

Pari Bandrong 156 

Pari Batu 192 

Pari Bendera 222, 224, 226, 228 

Pari Biasa 188 

Pari Bintang 192 

Pari Blentik 220 

Pari Boro 186 

Pari Burung 242, 244, 246, 248, 250, 252 

Pari Duren 236 

Pari Duri 196 

Pari Elang 252, 254 

Pari Gitar 156 

Pari Gunung 226 

Pari Jantung 186 

Pari Kekeh 156 

Pari Kelelawar 238, 240 

Pari Kembang 232 

Pari Kerbua 256 

Pari Kikir 196, 218 

Pari Kodok 220 

Pari Kontak 160, 162, 164, 166, 168 

Pari Lambingan 260, 262, 264 

Pari Lampengan 262 

Pari Lampin 230 

Pari Lemer 230 

Pari Lempengan 264 

Pari Listrik 160, 162, 164, 166, 168 

Pari Lowo 238, 240 

Pari Lumpur 202, 212 

Pari Luncur 172, 174 

Pari Macan 200, 214, 216 

Pari Merica 214, 234 

Pari Minyak 190 

Pari Pasir 206 

Pari Payung 240 

Pari Sapi 182, 194 

Pari Sungai 198, 210 

Pari Super 192 

Pari Tanjung 202, 212 

Pari Tembaga 190 

Pari Totol 232 

Payubek 230 

Pe Lamping 248 

Pe Pasung 212 

Petong 148, 150, 152, 156

I n d o n e s i a n c o m m o n n a m e s i n d e x 295 

Plampangan 256, 260, 264 

T 

Tikusan 62 

Toka-toka 188 

Toka Toka 218 

Tuka-tuka 188 

Y 

Yunbun Karang 146


Malaysian common names index 

B 

Bengkong 134 

Beroi 140 

C 

Chimara Perak 266 

I 

Ikan Tikus 266 

J 

Jerung-mako Sirip 64 

Jerung Biru 124 

Jerung Gigi Cerakah 82 

Jerung Sapi 100 

Jerung Tenggiri 114 

K 

Kan Sua 140 

Kemejan 152 

Kiampau 220, 232 

P 

Pari 152, 184 

Pari-karan Hitam 164 

Pari-kelawar Jawa 252 

Pari-kemejan Kepala Keras 146 

Pari-kemejan Muncung Panjang 154 

Pari-kemejan Muncung Pendek 156 

Pari-kemejan Tompok Putih 148 

Pari-lang Jalur 248 

Pari-lang Tompok Putih 246 

Pari-raja Tompok 172 

Pari Air Tawar 208, 210 

Pari Ayam 252 

Pari Bakau 194 

Pari Batik 192, 216 

Pari Belanda 170, 174, 176 

Pari Bendera 222 

Pari Bintik Putih 192 

Pari Bulat 206 

Pari Daun 222 

Pari Dedap 236 

Pari Duri 236 

Pari Durian 236 

Pari Ekor Duri 186 

Pari Gergasi 182 

Pari Helang 244 

Pari Hitam 230 

Pari Karan 164, 166 

Pari Karan Belangkas 168 

Pari Karang 232 

Pari Karan Tompok Besar 160 

Pari Karas 168 

Pari Kelabu 234 

Pari Kelawar 240, 256, 262 

Pari Kelawar Bintik 238 

Pari Kemejan 150 

Pari Ketuka 188, 218 

Pari Lalat 220 

Pari Lang 242, 244, 252, 254 

Pari Lang Bintik Putih 244 

Pari Lang Bunga 250 

Pari Lembik Merah 180 

Pari Letrik 164 

Pari Macan 220 

Pari Merah 180 

Pari Mulut Kasar 226 

Pari Mulut Tajam 202 

Pari Naga 196 

Pari Nyiru 188 

Pari Nyonya 222 

Pari Pasir 196, 212 

Pari Paus 256, 258, 260, 262, 264 

Pari Pucat 190

M a l a y s i a n c o m m o n n a m e s i n d e x 297 

Pari Reben 232, 234 

Pari Riman 220 

Pari Rimau 200, 214, 216, 220 

Pari Serban 164 

Pari Sungai 198, 208, 210 

Pari Susun 252 

Pari Tanjung 188, 218, 220, 222 

Pari Tembikar 238, 240 

Pendek 64 

S 

Sirip Lekuk 264 

T 

Tanpa Duri 262 

Tuka-tuka 218 

Y 

Yu 134 

Yu-buntal Sarawak 70 

Yu-cicak Bintik 52 

Yu-cicak Insang Puih 54 

Yu-cicak Lampai 48 

Yu-cicak Tembaga 50 

Yu-gergaji Seragam 142 

Yu-gergaji Silih 144 

Yu-jaras Bintik 74 

Yu-tukul Palang 134 

Yu-tukul Sanggul 136 

Yu Bengkong 136 

Yu Bintik Putih 80 

Yu Bodoh 48, 50, 52, 54, 56, 72 

Yu Borneo 92 

Yu Buncit 70 

Yu Buntal 70 

Yu Cicak 58, 68 

Yu Ekor Hitam 90 

Yu Ekor Panjang 62 

Yu Gergaji 140, 142, 144 

Yu Gergaji Jarang 140 

Yu Gigi Cangkuk 78 

Yu Gigi Lurus 84 

Yu Giling 114 

Yu Insang Enam 36 

Yu Jalur Putih 88 

Yu Jaras 74, 76 

Yu Jereh 98, 102, 104, 130 

Yu Jerung 64, 100 

Yu Kebut 44, 58 

Yu Kembong 70 

Yu Kemejan 146, 154, 158 

Yu Kepak Hitam 102, 106, 112 

Yu Kia-kia 148, 150, 152, 154 

Yu Kulit Tebal 108 

Yu Mata Jauh 134, 136 

Yu Mata Lekuk 122 

Yu Minyak 38, 40 

Yu Misai 46 

Yu Muncung Keras 104 

Yu Muncung Minyak 128 

Yu Muncung Panjang 94 

Yu Muncung Susu 126 

Yu Nipah 106 

Yu Padi 130 

Yu Palang 134, 136 

Yu Parang 136, 138, 140, 142, 144 

Yu Parangan 140, 142, 144 

Yu Pari 42, 156 

Yu Pasir 50, 80, 96, 98, 104, 110, 130 

Yu Paus 60 

Yu Pendek 68 

Yu Punai 54 

Yu Putih 82 

Yu Rimau 58 

Yu Sanggul 134, 136 

Yu Semilang 56 

Yu Sirip Hitam 106 

Yu Sirip Lebar 120 

Yu Sirip Putih 132


Yu Sungai 116, 118 

Yu Tanduk 134 

Yu Tenggiri 114 

Yu Todak 140, 142, 144 

Yu Tokeh 50, 54, 58, 68 

Yu Trihitam 102 

Yu Tukul 136, 138

Sharks and Rays of Borneo 

This full-colour field guide complements a similar book on the economically important 

sharks and rays of Indonesia. It is the result of a collaborative project between the 

governments of the United States, Malaysia, Indonesia and Australia, and funded by 

the National Science Foundation. 

The first comprehensive reference on the sharks and rays of Borneo, it contains 

everything you need to know about recognising and identifying the sharks, rays 

and chimaeras caught and marketed in Indonesia. Its user-friendly layout contains 

information on identifying features, size, distribution, local common names, habitat, 

biology and conservation status of 118 species. It is an essential reference for all 

shark and ray enthusiasts – including local fishers and consumers, fisheries and 

conservation officers and scientists.

Whaler Sharks - seafdec.org.my

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