Borneo Shark and Ray Guide

SHARKS and RAYS of
SHARKS AND RAYS OF BORNEO

Sharks and Rays of Borneo
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.
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
John J. Pogonoski 4
John D. Stevens 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, photo-
copying, 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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Collingwood VIC 3066
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Telephone: +61 3 9662 7666
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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 km2 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
km2 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
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 19th C
ichthyologists, including Richardson, Günther and Kaup, also made important contribu-
tions 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-19th 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 20th 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 environ-
mentalists and respective governments to protect the forests and seas of Brunei, Indonesia,
Malaysia and the Philippines.
Introduction
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 km2 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 elasmo-
branchs 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 Rhynchoba-
tus. 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).
Introduction
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 develop-
ment 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 Pastina-
chus) 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.
Introduction
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 Philippines Indonesia Borneo

Australia
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 Sharks and Rays of Borneo
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.
How to use this book 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
Glossary 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.
Glossary 15
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 1st and 2nd 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.
Glossary 17
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 saw-
sharks.
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.)
Glossary 19
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.
1st dorsal fin

anterior posterior
dorsal-fin spine 2nd dorsal fin
eye
gill opening precaudal pit
spiracle
snout tip
caudal fin
nostril
mouth
pelvic fin anal fin
labial furrows
caudal keel
clasper (males)
pectoral fin
structural features of sharks
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
lower labial furrow

symphysial groove
shark undersurface of head

Glossary 21
terminal lobe
posterior tip
upper lobe
terminal margin
dorsal margin
upper origin subterminal margin
subterminal notch
upper postventral margin
caudal fork
lower origin lower postventral margin
preventral margin
ventral tip
lower lobe
shark caudal fin
ventral dorsal
rostral cartilage
internasal flap mouth
eye
shoulder
gill opening
spiracle
pectoral fin
cloaca
pectoral fin insertion
thorn
pelvic fin
caudal fin
lateral tail fold
clasper
2nd dorsal fin
(male)
1st dorsal fin stinging spine
structural features of rays

nostril
lobes
internasal flap
fringe
papillae mouth
ray nostrils and mouth (oronasal)
stinging spine
enlarged thorn dorsal fold
thorn
granulations ventral fold
ray tail
1st dorsal fin

spine
head clasper (male) lateral line
eye 2nd dorsal fin

mucous canal caudal-fin
upper lobe
mouth
anal fin caudal-fin
gill opening pre-pelvic clasper
(male) lower lobe
clasper (male) caudal filament
pelvic fin
pectoral fin
structural features of chimaerids

Running head 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 fig. 2
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 fig. 3

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 fig. 4
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. 5

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

dorsal fin originating closer to insertion
of pelvic fins than to tail tip (fig. 6) ...... 6
0–2 dorsal fins; 1st dorsal fin originating fig. 6
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 fig. 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 ....... fig. 8
.... 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 fig. 9
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 fig. 10
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,
fig. 11 fig. 12
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. 13
Key to families and genera 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
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
fig. 15
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 fig. 16
......................................................... 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 fig. 17 fig. 18

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 fig. 19
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. 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 fig. 21 fig. 22
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 fig. 23
14. Caudal fin present (fig. 24); no thorns or

tubercles on disc or tail .........................
... Family Plesiobatidae, Giant Stingarees
fig. 24; 180
fig. 24
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
fig. 25 fig. 26
Skin folds present on undersurface of
tail, and sometimes on dorsal surface
after sting; base of tail relatively broad, fig. 27
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. 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 fig. 29
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
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 fig. 31
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 fig. 32
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
fig. 33
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 fig. 34
................. Family Mobulidae, Devilrays
Mouth terminal, located at end of snout

tip ............................................... Manta fig. 35
fig. 35; 256
Mouth subterminal, located on under-
surface of head .......................... Mobula
fig. 36; 258
No lateral, paddle-like processes on head,

instead with a pair of broadly rounded fig. 36
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 fig. 37

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 fig. 38
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
fig. 39 fig. 40
17. A single dorsal fin (fig. 42); 6 or 7 gill

openings on each side of head (fig. 42)
fig. 41
............ Family Hexanchidae, Cowsharks
fig. 42; 36
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 fig. 43
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) ................................................. fig. 44 fig. 45
... Family Centrophoridae, Gulper Sharks
fig. 46; 40
20. Head very broad, hammer-shaped, with

very widely separated eyes (fig. 47) ........ fig. 46
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
fig. 47
Lateral blades of head much shorter and
not wing-like; width of head less than
40% of total length .................... Sphyrna
fig. 48; 136
Head much narrower, not hammer-

shaped .............................................. 21
fig. 48
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
fig. 49
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. 50
....... Family Heterodontidae, Hornsharks

fig. 50; 44
Dorsal-fin spines absent; no crest over
fig. 51
eye ................................................... 23
23. Whole mouth forward of front margin

of eye (fig. 51) ................................... 24
Mouth partly beneath or behind front fig. 52
margin of eye (fig. 52) ....................... 28
24. Mouth very broad, terminal; caudal fin

forked (fig. 53) .......................................
... Family Rhincodontidae,Whale Sharks
fig. 53; 60
fig. 53
Mouth smaller, subterminal; caudal fin
not forked ...................................... 25
25. No fleshy lobe or groove on outer

margin of nostril (fig. 54) .................. 26
Fleshy lobe and groove present on outer
fig. 54 fig. 55
margin of nostril (fig. 55) ................ 27
26. Caudal fin very long, about equal to

trunk length (fig. 56); ridges present along fig. 56
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, fig. 57
Nurse Sharks
fig. 57; 56
27. Head and body strongly depressed

(fig. 58); skin flaps present on sides of
head (fig. 58) ......................................... 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
.......................... Family Hemiscylliidae,

Longtail Carpetsharks
fig. 59; 48
fig. 60
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
fig. 61
Caudal fin asymmetrical, heterocercal
(fig. 61); caudal-peduncle keel either low
or absent (fig. 61) ............................... 29
29. 1st dorsal-fin origin behind pelvic-fin

origin ... Family Scyliorhinidae, Catsharks
a. Upper labial furrows very long, more fig. 62 fig. 63
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 fig. 64
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
fig. 65 fig. 66
Anterior nasal flaps much shorter, smaller
than nostrils, not reaching mouth (fig. 66);
nasoral grooves absent ............. Apristurus
fig. 67; 66
c. 2nd dorsal fin much smaller than 1st fig. 67
dorsal fin (fig. 68); origin of 2nd dorsal
fin only slightly behind origin of anal fin
(fig. 68); stomach inflatable ....................
....................................... Cephaloscyllium
fig. 68; 70
fig. 68
Dorsal fins about equal in size (fig.
69); origin of 2nd dorsal fin behind
midbase of anal fin (fig. 69); stomach not
inflatable ................................ Halaelurus
fig. 69; 72 fig. 69
1st dorsal-fin origin well in front of

pelvic-fin base .................................. 30
30. Precaudal pits absent (fig. 71); anterior

fig. 70
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
fig. 71 fig. 72
31. Spiracles present; posterior margin of
2nd dorsal fin deeply concave; intestine
with spiral valves (fig. 73) ......................
...... Family Hemigaleidae, Weasel Sharks
fig. 73
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 fig. 74 fig. 75
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
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
fig. 77
c. Snout obtusely wedge-shaped in dorso-
ventral view; fins not falcate, posterior
margins of pelvic and pectoral fins
straight or slightly concave (fig. 78) .......
.......................................... Chaenogaleus
fig. 78; 78 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 fig. 79
Spiracles absent (except in Tiger Shark);

posterior margin of 2nd dorsal fin not
deeply concave; intestine with a scroll fig. 80
valve (fig. 80) ........................................
... Family Carcharhinidae, Whaler Sharks
a. Caudal peduncle with lateral keels

(fig. 81); spiracles present; upper labial
fig. 81
furrows, reaching forward to front of eyes; fig. 82
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 fig. 83
reaching forward to in front of eyes;
teeth not cockscomb-shaped .............. b
b. 2nd dorsal fin half or more height of 1st

dorsal fin ............................................ c
2nd dorsal fin less than half height of 1st fig. 84
dorsal fin ............................................ e
c. 1st 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 fig. 85 fig. 86
1stdorsal and upper caudal fin without
obvious white tips; teeth with a single
cusp and no lateral cusplets (fig. 86) ...... d
d. 2nd dorsal fin nearly as large as 1st dorsal

fin (fig. 87) ............................... Lamiopsis fig. 87
fig. 87; 120
2nd dorsal fin half to two thirds height of
1st dorsal fin (fig. 88) .................. Glyphis
fig. 88; 116
e. Head greatly depressed and trowel- fig. 88

shaped (fig. 89); pectoral fins triangular
(fig. 89); 1st dorsal-fin free rear tip

almost over mid-base of pelvic fin
(fig. 89) .................................. Scoliodon
fig. 89; 130
fig. 89
Head varying from conical to slightly
depressed; pectoral fins narrower; 1st
dorsal-fin free rear tip over or slightly
anterior to pelvic-fin origin ................. f
fig. 90
f. 2nd dorsal-fin origin well behind anal-fin
origin (usually over or just anterior to anal-
fin 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
fig. 91
2nd dorsal-fin origin usually about
opposite anal-fin origin (sometimes more
posterior but always well anterior of anal-
fin insertion) (fig. 91); posterior margin
of anal fin deeply concave or deeply fig. 92
notched (fig. 91); preanal ridges short,
about half anal-fin base or less .............. h
h. Notch present at posterior edge of eye

(fig. 93); 1st dorsal-fin origin posterior
fig. 93
to adpressed pectoral-fin free rear tips
by a distance exceeding 4th gill opening fig. 94
height (fig. 92) ......................... Loxodon
fig. 92; 122
Posterior rim of eye without a notch (fig.
94); 1st dorsal-fin origin over or only just
posterior to adpressed pectoral-fin free fig. 95
rear tips (fig. 95) …….… Rhizoprionodon
fig. 95; 126
i. 1st dorsal-fin base much closer to pelvic-

fin than pectoral-fin bases (fig. 96);
colour brilliant dark blue above;weak keels
present on caudal peduncle ......... Prionace fig. 96
fig. 96; 124
1st 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
fig. 97; 86
SPECIES INFORMATION
Bluntnose Sixgill Shark

Hexanchus griseus (Bonnaterre, 1788)
3 4
2 1
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, elasmo-
branchs, 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).
Hexanchiformes 37
In Borneo: Probably widespread, but

known only from near Layang-layang, off
Hexanchidae (Cowsharks)
western Sabah.
Ventral head
Upper tooth
Lower tooth
6
Western Highfin Spurdog

Squalus altipinnis Last, White & Stevens, 2007
4
4 3
5
1
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 2nd dorsal fin relatively tall, upright
4 dorsal fins with thick spines
5 origin of 2nd 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).
Squaliformes 39
In Borneo: Northern Borneo, from

Sabah and northern Sarawak.
Ventral head
Squalidae (Dogfishes)
7
Late-term embryo (19 cm)
Flank denticle
Smallfin Gulper Shark

Centrophorus moluccensis Bleeker, 1860
3
4
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 2nd dorsal fin relatively small, half or less height of 1st dorsal; both fins with short
spines
4 origin of 2nd 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).
Squaliformes 41
Centrophoridae (Gulper Sharks)

In Borneo: Sparse records from off
Sarawak and eastern Sabah.
Ventral head
Upper teeth
6
5
Lower teeth
Patch of flank denticles
Ocellated Angelshark
Squatina tergocellatoides Chen, 1963
6
3
5
8
8 8
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 1st 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.
Squatiniformes 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.
Squatinidae (Angelsharks)
Ventral head and abdomen In Borneo: Off western Sabah and
Sarawak.
4
Front view of head Anterior nasal flap (from front)
Zebra Hornshark
Heterodontus zebra (Gray, 1831)
1
1
2
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.
Heterodontiformes 45
Conservation status: IUCN Red List: Least Concern.

Local synonyms: Cestracion amboinensis Regan, 1906.
References: None.
Heterodontidae (Hornsharks)
Ventral head
In Borneo: Northern Borneo, includ-

ing Sarawak and Sabah.
6 7
Anterior upper tooth Posterior upper tooth

Indonesian Wobbegong
Orectolobus leptolineatus Last, Pogonoski & White, 2010
4
5
2
3
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).
Orectolobiformes 47
Ventral head In Borneo: Known from a few speci-
Orectolobidae (Wobbegongs)
mens collected off southeastern Sabah
(Semporna) and Sarawak.
Embryo (14 cm)

Indonesian Bambooshark
Chiloscyllium hasseltii Bleeker, 1852
1 4
4
6 3
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 2nd 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 bottom-
dwelling 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).
Orectolobiformes 49
Hemiscylliidae (Longtail Carpetsharks)

Ventral head In Borneo: Central western Borneo,
including northwestern Kalimantan and
Sarawak.
Dorsal head and pectoral fins
Juvenile (34.5 cm)

Slender Bambooshark
Chiloscyllium indicum (Gmelin, 1789)
1 4 4
6 3
5 2
1
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 2nd 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.
Orectolobiformes 51

Ventral head In Borneo: Patchy; recorded from
southern and western Kalimantan and
Sarawak.
3
6
Juvenile (19 cm)

Whitespotted Bambooshark
Chiloscyllium plagiosum (Bennett, 1830)
4
1 4
6 3
1 5 2
Other names: None (English), Hiu Bongo, Cucut Dolok, Hiu Tekek (Indonesian),Yu-
cicak 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 2nd 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).
Orectolobiformes 53

Ventral head
In Borneo: South China Sea, from
northwestern Kalimantan to northern
Sabah.
Juvenile (13 cm)

Brownbanded Bambooshark
Chiloscyllium punctatum Müller & Henle, 1838
1 4 4
6 3
5
1 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 2nd 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.
Orectolobiformes 55


Ventral head
In Borneo: Southern and western
Kalimantan, Sarawak and Sabah.
Sub-adult (51 cm)

Tawny Nurse Shark

Nebrius ferrugineus (Lesson, 1830)
4 1
4
3 5
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
born at 40–60 cm.
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.
Orectolobiformes 57
Ginglymostomatidae (Nurse Sharks)

Ventral head
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
Upper tooth from dive operator in Kota Kinabalu.
Lower tooth
Lateral precaudal tail region

Zebra Shark
Stegostoma fasciatum (Hermann, 1783)
3
5
4
1
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 stria-
tions, 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
Orectolobiformes 59
mentioned in synonmy with S. fasciatum); Scyllia quinquecornuatum van Hasselt, 1823;

Squalus tigrinus Forster, 1781; Stegostoma varium Garman, 1913.
References: None.
Stegostomatidae (Zebra Sharks)

Ventral head
In Borneo: Western and northern

Upper tooth Borneo, including Sarawak, Sabah and
north-west and north-east Kalimantan.
Lower tooth
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).
Orectolobiformes 61
Local synonyms: Rhiniodon typus Smith, 1829.

References: None.
1
3
Rhincodontidae (Whale Sharks)

In Borneo: Recorded from northern
and western Sabah and northwestern
Kalimantan, but probably more widely
distributed. Seasonal in Sabah, usually
from February to April.
Upper tooth
Lower tooth
Pelagic Thresher
Alopias pelagicus Nakamura, 1935
3
2
4
5
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 1st 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.
Lamniformes 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).
Alopiidae (Thresher Sharks)

In Borneo: Uncommon, South China
Sea off Sabah and also the Natuna Islands
off Sarawak; possibly more widespread.
Ventral head
Upper tooth
Lower tooth
Shortfin Mako
Isurus oxyrinchus Rafinesque, 1810
3 1
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.
Lamniformes 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).
Lamnidae (Mackerel Sharks)

5
In Borneo: Only recorded off Sabah, but

almost certainly more widespread.
Ventral head
Upper tooth
Lower tooth
Bigfin Catshark
Apristurus platyrhynchus (Tanaka, 1909)
3
6 4
5
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 2nd dorsal fin much larger than 1st 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).
Carcharhiniformes 67
In Borneo: Known only from off
Scyliorhinidae (Catsharks)
Sipadan Island, in the Celebes Sea
(eastern Sabah).
Ventral head
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).
Ventral head In Borneo: Coastal waters of Sabah and
Sarawak.

Sarawak Swellshark
Cephaloscyllium sarawakensis Yano, Ahmad & Gambang, 2005
6 1
3
2
Other names: Sarawak Pygmy Swell Shark, Sarawak Pygmy Swellshark (English), none
(Indonesian),Yu-buntal Sarawak,Yu Buncit,Yu Kembong,Yu Buntal (Malaysian)
1 2nd dorsal fin much smaller than 1st dorsal fin, its origin only slightly behind anal-
fin 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).
Ventral head
In Borneo: South China Sea, off Sarawak.
Egg case

Indonesian Speckled Catshark

Halaelurus maculosus White, Last & Stevens, 2007
1
1
2
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).
Ventral head
In Borneo: Known only from off
Sarawak.
Egg case
Dorsal head and anterior trunk

Starspotted Smoothhound
Mustelus manazo Bleeker, 1854
1
4
5 2
3
Other names: Gummy Shark, Japanese Smoothhound (English), Hiu Kacang, Hiu Air,
Cucut Londer (Indonesian),Yu-jaras Bintik,Yu Jaras (Malaysian)
1 1st 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 1st 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.
Local synonyms: Mustelus mosis Hemprich & Ehrenberg: Yano et al. (2005).
References: None.
Triakidae (Houndsharks)
8
7
In Borneo: Known from the South

China Sea, off Sarawak and Sabah.
Ventral head
Whitefin Smoothhound
Mustelus widodoi White & Last, 2006
1
4 2 2
5 3
Other names: None (English), Hiu Kacang, Hiu Air, Cucut Londer (Indonesian),Yu
Jaras (Malaysian)
1 1st dorsal fin with a prominent white tip
2 2nd dorsal fin and caudal-fin upper lobe with distinct black tips
3 sides lacking white spots
4 1st 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 mid-
continental 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).
In Borneo: Known only from north-

western Sabah.
Triakidae (Houndsharks)
Ventral head
Hooktooth Shark
Chaenogaleus macrostoma (Bleeker, 1852)
3
3
1
2
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.
Hemigaleidae (Weasel Sharks)

In Borneo: Uncommon, recorded
from Sarawak, Sabah and northeastern
Kalimantan, but probably more widely
distributed.
Ventral head
Upper tooth
6
Lower tooth
Sicklefin Weasel Shark

Hemigaleus microstoma Bleeker, 1852
3
1
2
3
3
Other names: Weasel Shark (English), Hiu Kacang, Hiu Pilus (Indonesian),Yu Bintik
Putih,Yu Pasir (Malaysian)
1 spiracles small
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.
Local synonyms: Hemigaleus machlani Herre, 1929; Carcharias (Hypoprion) notatus

Bleeker, 1850.
References: White (2007a); Taylor & Bennett (2008).

4
5
In Borneo: Widespread.
Ventral head
Upper tooth
6
7
Lower tooth
Pectoral fin - ventral
Fossil Shark
Hemipristis elongata (Klunzinger, 1871)
3
1
2
3
3
Other names: Snaggletooth Shark (English), Hiu Monas (Indonesian), Jerung Gigi
Cerakah,Yu Putih (Malaysian)
1 spiracles small
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 cephalo-
pods 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).

5
Ventral head
Upper tooth
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)
1 spiracles small
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.
5
6

In Borneo: Off Sabah, Sarawak and
eastern Kalimantan.
Ventral head
Upper tooth
8
Lower tooth
Silvertip Shark
Carcharhinus albimarginatus (Rüppell, 1837)
3
1
1
2
1
1
Other names: None (English), Cucut Lanjaman, Hiu Lanyam, Hiu Plen, Hiu Sonteng
(Indonesian), none (Malaysian)
1 1st dorsal, pectoral, pelvic and caudal fins with prominent white tips
2 interdorsal ridge present
3 apex of 1st 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
born at 70–80 cm.
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.

References: None.
Carcharhinidae (Whaler Sharks)

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 east-
Ventral head ern Sabah.
Upper tooth
5
Lower tooth 6
Graceful Shark
Carcharhinus amblyrhynchoides (Whitley, 1934)
1
2 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 cephalo-
pods. 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).

Ventral head
Upper tooth
Lower tooth
Grey Reef Shark

Carcharhinus amblyrhynchos (Bleeker, 1856)
3
2
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 1st 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 <180 cm; both sexes mature at about 130–140 cm;
born at 50–60 cm.
Distribution: Widespread throughout the tropical Indo–Pacific, from eastern Africa to
the Galapagos Islands.
Habitat and biology: Among the most common sharks inhabiting coral reefs; lives
mainly near deep drop-offs or in lagoon passages, from the surface to about 280 m
depth.Viviparous, with a yolk-sac placenta; gives birth to litters of 1–6 pups after 12–14
month gestation period. Males and females mature at 9 and 11 years respectively, in
Australian populations. Diet consists primarily of small fishes, and to a lesser extent
crustaceans and cephalopods. Can become aggressive when provoked and considered
potentially dangerous to humans.
Utilisation: Caught by longline and inshore gillnet fisheries, and possibly adversely
affected by dynamite fishing in the region. Used for fins, meat, skin and cartilage.

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.
References: None.

In Borneo: Patchily distributed, but
probably widespread. Recorded off
Sabah, Sarawak, and eastern and western
Ventral head Kalimantan.
Embryo (67 cm)
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 2nd dorsal fin very low (height 2.2–2.5 in inner margin), its origin over midbase of
anal fin
3 1st 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).

In Borneo: Presently known only
5 from Mukah (Sarawak); type collected
in the 19th century from northwestern
Kalimantan.
Ventral head
Upper tooth
6
Lower tooth 7
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 1st 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.

References: None.

5 In Borneo: Probably widespread.
Ventral head
Upper tooth
Lower tooth
Lateral head
Whitecheek Shark
Carcharhinus dussumieri (Müller & Henle, 1839)
2 3
Other names: Widemouth Blackspot Shark (English), Cucut Lanjaman, Hiu Bujit
(Indonesian),Yu Pasir (Malaysian)
1 1st dorsal fin moderately tall, broad, not falcate
2 low interdorsal ridge usually present
3 only 2nd 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.
References: None.

Ventral head
Upper tooth
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 1st dorsal-fin origin well behind pectoral-fin free rear tips
3 2nd 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 <250 cm; both sexes mature at about 200–210 cm;
born at 53–73 cm off Indonesia.
Distribution: Circumtropical, occasionally making seasonal incursions into warm
temperate zones.
Habitat and biology: Oceanic and pelagic, but most abundant in offshore waters
close to land masses; usually near surface, but also to depths of at least 500 m.Vivipa-
rous, with a yolk-sac placenta; gives birth to litters of about 8 pups (maximum 16)
after an unknown gestation period; no reproductive seasonality with females breeding
throughout the year. Males and females mature at 11–13 and 15 years respectively, in
Indonesian populations. Diet consists primarily of fishes, as well as cephalopods and
pelagic crabs. Not considered dangerous to humans. Molecular analysis has identified
differences between Indo–Pacific and Atlantic forms that could be separate species.
Utilisation: Common component of the catch of shark and tuna longline, and tuna
gillnet fisheries in Indonesia but not landed in large quantities in Borneo. Used for its
fins, meat, skin and cartilage.

Local synonyms: Aprionodon sitankaiensis Herre, 1934.
References: None.

In Borneo: Eastern Sabah, and north-
western and eastern Kalimantan; probably
Ventral head widespread.
Upper tooth
5
Lower tooth 6
Bull Shark
Carcharhinus leucas (Müller & Henle, 1839)
2
1
Other names: River Whaler, Freshwater Whaler (English), Cucut Bekeman, Hiu Buas,
Hiu Bujit (Indonesian), Jerung Sapi,Yu Jerung (Malaysian)
1 2nd dorsal fin relatively large (about a third height of 1st 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.
References: None.

In Borneo: Widespread, concentrated
Ventral head near major river outflows.
Juvenile (86 cm)
Upper tooth
6
5 Lower tooth
Common Blacktip Shark

Carcharhinus limbatus (Müller & Henle, 1839)
1
2
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 1st 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 cepha-
lopods. 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.
References: None.

Ventral head
Upper tooth
Lower tooth
Lateral head
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 2nd dorsal-fin origin about over midbase of anal fin
3 1st 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.

In Borneo: Patchy, off Sarawak, Sabah
and eastern Kalimantan.
Ventral head
Upper tooth
5
Lower tooth 6
Blacktip Reef Shark

Carcharhinus melanopterus (Quoy & Gaimard, 1824)
1
2
Other names: Blacktip Shark, Guliman (English), Hiu Mada, Kluyu Karang, Hiu
Bujit (Indonesian),Yu Sirip Hitam,Yu Kepak Hitam,Yu Nipah (Malaysian)
1 1st 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.
References: None.

4
Ventral head
Upper tooth
5
Lower tooth 6
Sandbar Shark
Carcharhinus plumbeus (Nardo, 1827)
1
3
2
Other names: Thickskin Shark, Brown Shark (English), Hiu Teteri, Cucut Lanjaman
(Indonesian),Yu Kulit Tebal (Malaysian)
1 1st dorsal fin very broad and tall, more than half of predorsal length (smaller in
newborns)
3 origin of 1st 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.
References: None.

In Borneo: Off Sabah, Sarawak and
Brunei, but probably more widespread.
Ventral head
Upper tooth 5
Lower tooth 6
Blackspot Shark
Carcharhinus sealei (Pietschmann, 1913)
1
2 3
Other names: None (English), Cucut Lanjaman (Indonesian),Yu Pasir (Malaysian)

1 1st dorsal fin moderately tall, moderately falcate
2 low interdorsal ridge usually present
3 only 2nd 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;
born at 33–45 cm.
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.

References: None.

In Borneo: Widespread, but no current
records from southwestern Kalimantan.
Ventral head
Upper tooth
5
6
Lower tooth
Spot-tail Shark
Carcharhinus sorrah (Müller & Henle, 1839)
2
13
1
1
Other names: Sorrah Shark, School Shark (English), Cucut Lanjaman, Hiu Bujit,
Lanyam, Merak Bulu, Mungsing (Indonesian),Yu Kepak Hitam (Malaysian)
1 2nd dorsal, pectoral and ventral tip of caudal fin strikingly black-tipped
3 2nd 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.
References: None.

Ventral head
Upper tooth
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, elasmo-
branchs, 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.
References: None.

4
In Borneo: Almost certainly widespread;

current records are from off Sarawak,
Ventral head Sabah and eastern Kalimantan.
Upper tooth
Lower tooth
Mukah River Shark

Glyphis sp.
4 2 6
1
Other names: None (English), none (Indonesian),Yu Sungai (Malaysian)

1 precaudal pits longitudinal (not crescentic)
2 2nd dorsal fin only moderately tall, slightly less than half height of 1st dorsal fin
3 eyes moderately large
4 1st dorsal-fin free rear tip just anterior to pelvic-fin origins
5 fins plain
6 2nd 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 vivip-
arous, 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.
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
4 2
3 6
1
7
5
5
Other names: None (English), none (Indonesian),Yu Sungai (Malaysian)

2 2nd dorsal fin tall, half to three fifths of 1st dorsal-fin height
3 eyes noticeably small
4 1st 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 2nd 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).

In Borneo: Recorded only from the
Kinabatangan River, northeastern Sabah.
Ventral head
3
7 Upper tooth
Lateral head 8
Lower tooth
Borneo Broadfin Shark

Lamiopsis tephrodes (Fowler, 1905)
2
3
1
Other names: None (English), Hiu Bujit (Indonesian),Yu Sirip Lebar (Malaysian)
2 2nd dorsal fin very tall, similar to height of 1st 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
born at 40–60 cm.
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.
Local synonyms: Lamiopsis temmincki (Müller & Henle):Yano et al. (2005), White et al.
(2006b); Carcharhinus microphthalmus Chu, 1960.
References: White et al. (2010c).

In Borneo: Occurs around the Borneo
coastline, but more common at particular
localities, such as Mukah (Sarawak).
Ventral head
Upper tooth
6
Lower tooth 7
Sliteye Shark
Loxodon macrorhinus Müller & Henle, 1839
3
1 2
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 2nd dorsal-fin origin over anal-fin insertion
3 1st 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, smooth-
edged 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
Local synonyms: Carcharias (Scoliodon) dumerilii Bleeker, 1856.
References: None.

4 In Borneo: Mainly northern Borneo; off
Sarawak, Sabah, and eastern and western
Kalimantan.
Ventral head
Upper tooth 1
7
Eye
Lower tooth
Blue Shark
Prionace glauca (Linnaeus, 1758)
1 3
4
Other names: Blue Whaler, Great Blue Shark (English), Hiu Aer, Hiu Lalaek, Hiu
Karet, Cucut Selendang (Indonesian), Jerung Biru (Malaysian)
1 1st 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);
born at 35–50 cm.
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.

References: None.

In Borneo: Known from a single record
off Sabah; probably more widespread.
Ventral head
Upper tooth 6
Lower tooth 7
Milk Shark
Rhizoprionodon acutus (Rüppell, 1837)
Other names: Longman’s Dogshark, Fish Shark, White-eye Shark (English), Hiu Pilus,
Hiu Plen, Mungsing, Hiu Pisang (Indonesian),Yu Muncung Susu (Malaysian)
1 2nd 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

Local synonyms: Carcharias (Scoliodon) walbeehmi Bleeker, 1856; Carcharias (Prionodon)
sorrahkowah Bleeker, 1854; Carcharhinus borneensis (Bleeker):Yano et al. (2005).
References: None.

4
5 In Borneo: Mainly northern Borneo;
from Sabah, Sarawak, and eastern and
western Kalimantan.
Ventral head
Upper tooth
Lower tooth
Grey Sharpnose Shark

Rhizoprionodon oligolinx Springer, 1964
Other names: None (English), Hiu Pilus, Hiu Pisang (Indonesian),Yu Muncung
Minyak (Malaysian)
1 2nd 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.
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.
References: None.

7
In Borneo: Widely distributed.
5
4
Ventral head
Upper tooth
Lower tooth
Pacific Spadenose Shark

Scoliodon macrorhynchos (Bleeker, 1852)
2
1 3
Other names: None (English), Mungsing, Hiu Kejen (Indonesian),Yu Padi,Yu Pasir,
Yu Jereh (Malaysian)
1 head and snout strongly depressed, trowel-shaped
2 1st dorsal fin large, closer to pelvic-fin base than to pectoral-fin base
3 2nd 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.
Local synonyms: Scoliodon laticaudus Müller & Henle: Yano et al. (2005), White et al.
(2006b).
References: White et al. (2010b).

In Borneo: Mainly off western Borneo,
including Sarawak, and western and eastern
Kalimantan.
Ventral head
Upper tooth
Lower tooth
Whitetip Reef Shark

Triaenodon obesus (Rüppell, 1837)
1
1
3
2
Other names: Whitetip Shark, Blunthead Shark (English), Hiu Bokem, Hiu Karang
Buas, Hiu Coklat (Indonesian),Yu Sirip Putih (Malaysian)
1 1st dorsal and upper lobe of caudal fin with distinctive white tips
3 2nd dorsal fin large, about half to three quarters height of 1st 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 respec-
tively; born at 52–60 cm.
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.

References: None.

Ventral head
In Borneo: Mainly northern Borneo;
from Sabah, Sarawak, and northwestern
and eastern Kalimantan.
Upper tooth
Lower tooth
Winghead Shark
Eusphyra blochii (Cuvier, 1816)
2
1
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 1st dorsal fin very tall, strongly falcate
3 origin of 1st 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
born at 32–47 cm.
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).
Sphyrnidae (Hammerhead Sharks)

4
In Borneo: A few records off Sabah,

Sarawak and northwestern Kalimantan
but probably widespread.
Ventral head Upper tooth
Lower tooth
Scalloped Hammerhead
Sphyrna lewini (Griffith & Smith, 1834)
1
4
6
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 1st dorsal fin tall, moderately falcate
4 2nd dorsal fin short with long rear tip and weakly concave posterior margin
5 anal-fin base barely longer than 2nd 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).

1
In Borneo: Probably widely distributed,

but few records from southern Kalimantan.
Ventral head
Upper tooth
7
Lower tooth
Great Hammerhead
Sphyrna mokarran (Rüppell, 1837)
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 1st dorsal fin very tall, strongly falcate in adults
4 2nd dorsal fin tall with short rear tip and strongly concave posterior margin
5 anal-fin base longer than 2nd 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 yolk-
sac 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
In Borneo: Recorded from Sabah and

Sarawak, but probably widely distributed.
Ventral head
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 1st 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.
Pristoidei 141
Conservation status: IUCN Red List: Critically Endangered; Appendix I of CITES.

References: Manjaji (2002a, b).
Pristidae (Sawfishes)
2
In Borneo: Single records from southern

Caudal fin Kalimantan and near Sandakan, north-
eastern Sabah.
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 1st 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
Pristoidei 143
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.
2
In Borneo: Occurs in the Kinabatangan
Caudal fin River; possibly once more widespread in
larger rivers and adjacent marine bays of
the region.
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 1st 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.Vivipa-
rous, 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
invertebrates.
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.
Pristoidei 145
2
Caudal fin
In Borneo: Recent records from central
Sarawak (near Bintulu), and northeastern
(Labuk River) and eastern Sabah (Kinaba-
tangan River); holotype collected in the
19th century from southern Kalimantan.
4 5
Ventral head
Shark Ray
Rhina ancylostoma Bloch & Schneider, 1801
5 1
2
Other names: Bowmouth Guitarfish, Mud Skate (English), Hiu Barong, Kupu-
kupu Brangkas,Yunbun Karang (Indonesian), Pari-kemejan Kepala Keras,Yu Kemejan
(Malaysian)
1 1st 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 depend-
ency; 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.
Rhinoidei, Rhynchobatoidei & Rhinobatoidei 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).
Rhinidae (Shark Rays)

4
Lateral head
In Borneo: Known from coastal areas of
Sabah, Sarawak, and western and southern
Kalimantan.
Ventral head
Whitespotted Wedgefish
Rhynchobatus australiae Whitley, 1939
6 4 4
2
5 1
3
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 1st 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 1st dorsal fin less than 1.5 times height of 2nd (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
born at 46–50 cm.
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.
References: None.
Rhynchobatidae (Wedgefishes)
6
In Borneo: Mainly western and north-

ern Borneo.
Ventral head
Dorsal view of eyes

and spiracles
Smoothnose Wedgefish
Rhynchobatus laevis (Bloch & Schneider, 1801)
6 2 4
4
5 1 8
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 1st 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 1st dorsal fin exceeding 1.5 times height of 2nd (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.
References: None.
In Borneo: Recorded from off north-
eastern Sabah.
Ventral head
7
Dorsal view of eyes
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 1st 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 1st dorsal fin less than 1.5 times height of 2nd (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).
In Borneo: A few records from brackish
habitats of southern Sarawak.
Ventral head
7
Dorsal view of eyes
and spiracles
Clubnose Guitarfish
Glaucostegus thouin (Anonymous, 1798)
2
1
3 6
4
Other names: None (English), Cucut Mandrong, Liongbun (Indonesian), Pari-

kemejan 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.
Local synonyms: Rhinobatos thouin (Anonymous [Lacepède]): Yano et al. (2005), White
et al. (2006b), also misspelled as Rhinobatos thouini.
References: None.
Rhinobatidae (Shovelnose Rays)

5
In Borneo: Known from Sarawak,

Sabah and southeastern and western
Kalimantan.
Ventral head
6
Lateral tail and dorsal fins
Giant Guitarfish
Glaucostegus typus (Bennett, 1830)
2
1
3
4
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).
References: None.

5
In Borneo: Primarily from eastern

Borneo, also known from Sarawak and
western Kalimantan.
Ventral head
6
Lateral tail and dorsal fins
Taiwan Guitarfish
Rhinobatos formosensis Norman, 1926
2
1
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).

In Borneo: Off Sarawak and Sabah.
Ventral head
Embryo (19 cm)
1st dorsal fin 2nd dorsal fin

Shortlip Numbfish
Narcine brevilabiata Bessednov, 1966
3
2
4
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.
To 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).
Narcinidae (Numbfishes)
7 6
Mouth and nostrils

In Borneo: Off Sarawak and eastern
Sabah.
Juvenile (14 cm)

Rough Numbfish
Narcine lingula Richardson, 1846
5
2 3
4
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 deliver-
ing 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
Mouth and nostrils
In Borneo: Uncommon locally, known

from southern Sarawak and eastern
Sabah.
Darkspotted Numbfish
Narcine maculata (Shaw, 1804)
5
3
2
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).
6
7
Mouth and nostrils
In Borneo: Recorded locally from

southern Sarawak; another specimen
recorded in literature from unspecified
locality off Sabah.
Spot-tail Sleeper Ray

Narke dipterygia (Bloch & Schneider, 1801)
1 5
4
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 yolk-
sac 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).
Narkidae (Sleeper Rays)

7
Mouth and nostrils
In Borneo: Off central Sarawak.
Ventral
Finless Sleeper Ray

Temera hardwickii Gray, 1831
4
5
2 1
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).
Narkidae (Sleeper Rays)

7
Mouth and nostrils
In Borneo: Single record from the South

China Sea off southwestern Sarawak.
Ventral
Kwangtung Skate
Dipturus kwangtungensis (Chu 1960)
3 1
5
2 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
Rajoidei 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).
Rajidae (Skates)
8
1 In Borneo: Off Sarawak.
Ventral disc (female)
Lateral posterior tail

Borneo Sand Skate

Okamejei cairae Last, Fahmi & Ishihara, 2010
3 1
7
2
6
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.
Rajoidei 173
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).
Rajidae (Skates)
8
Ventral disc (female) In Borneo: Off western Borneo, including

Sabah, Sarawak and northwestern Kali-
mantan.
7
6
Lateral tail
Yellowspotted Skate
Okamejei hollandi (Jordan & Richardson, 1909)
3 1
2
6 7
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 well-
defined 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
Rajoidei 175
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).
References: None.
Rajidae (Skates)
8
1
In Borneo: Off western Borneo, including
Ventral disc (female) Sarawak and Kalimantan.
6 7
Lateral tail
Philippine Ocellate Skate

Okamejei jensenae Last & Lim, 2010
1
3
7
2
6
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.
Rajoidei 177
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).
Rajidae (Skates)
8
In Borneo: Off Lahad Datu, eastern

Ventral disc (female) Sabah.
Lateral posterior tail

Borneo Leg Skate

Sinobatis borneensis (Chan, 1965)
3
2
4
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.
Rajoidei 179

Local synonyms: Previously as Anacanthobatis borneensis Chan.
References: Last & Compagno (1999).
Anacanthobatidae (Leg Skates)

Mouth and nostrils
In Borneo: Known from the holotype

collected in the South China Sea, off
Sarawak.
Ventral
Giant Stingaree
Plesiobatis daviesi (Wallace, 1967)
4
3
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
Myliobatoidei 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.
References: None.
Plesiobatidae (Giant Stingarees)

Mouth and nostrils
In Borneo: Known only from off Sarawak.
1
Lateral posterior tail with caudal fin
Smalleye Stingray
Dasyatis microps (Annandale, 1908)
1
4
6
5
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
Myliobatoidei 183
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).
In Borneo: Northeastern Sabah.

Dasyatidae (Stingrays)
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;
Myliobatoidei 185
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
In Borneo: Known from Sabah, including

Ventral disc the eastern border with Kalimantan.
Nuchal area
4
Lateral tail
Cow Stingray
Dasyatis ushiei (Jordan & Hubbs, 1925)
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.
Myliobatoidei 187
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).
References: None.
7
Ventral disc In Borneo: Known from off Sarawak

and Sabah.
Nuchal area
6 3
4
Lateral tail
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.
Myliobatoidei 189
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.
References: None.
9
In Borneo: Probably widespread, al-
though currently few records from eastern
Ventral disc Kalimantan.
Nuchal area
Lateral tail 6
Pink Whipray
Himantura fai Jordan & Seale, 1906
5
4
6
2
7
1
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.
Myliobatoidei 191
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 bottom-
dwelling invertebrates.
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.
References: None.
Ventral disc In Borneo: Widespread.
Nuchal area
Whitespotted Whipray
Himantura gerrardi (Gray, 1851)
5
4
8
7 1 6
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
Myliobatoidei 193
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.
References: None.
Ventral disc
Nuchal area
Mangrove Whipray
Himantura granulata (Macleay, 1883)
6
5
2
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.
Myliobatoidei 195
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.
References: None.
In Borneo: Known from off Sabah and
northeastern Kalimantan.
Ventral disc
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.
Myliobatoidei 197
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.
References: None.
In Borneo: Mainly known from Sabah
Ventral disc and Sarawak; rarely caught off eastern,
northwestern and southern Kalimantan.
Dorsal tail thorns

1
Nuchal area
Roughback Whipray
Himantura kittipongi Vidthayanon & Roberts, 2005
6
5
1
9
8
7
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.
Myliobatoidei 199
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.
References: None.
4
In Borneo: Known locally from the

Pawan and Kapuas rivers, western
Kalimantan.
Ventral disc
Nuchal area
Leopard Whipray
Himantura leoparda Manjaji-Matsumoto & Last, 2008
5
4
3
6
1
8
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.
Myliobatoidei 201
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).
In Borneo: Off Sabah, and northwestern
Ventral disc and northeastern Kalimantan.
Nuchal area 3
5
Tail section - dorsal
Tail section - ventral Juvenile (200 mm DW)

Tubemouth Whipray
Himantura lobistoma Manjaji-Matsumoto & Last, 2006
5
2
3
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.
Myliobatoidei 203
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.
In Borneo: Known mainly from western
Ventral disc Borneo; occasionally off southern and
eastern Kalimantan.
7
Nuchal area
Protrusible mouth Late-term embryo (18 cm DW)

Longnose Marbled Whipray

Himantura oxyrhyncha (Sauvage, 1878)
6
5
3
2
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.
Myliobatoidei 205
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.
References: None.
4
In Borneo: Known from north-west

Kalimantan in the environs of the
Kapuas River.
Ventral disc
7
Nuchal area
Round Whipray
Himantura pastinacoides (Bleeker, 1852)
5
6
2
7
3
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.
Myliobatoidei 207
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.
References: None.
4
In Borneo: Widespread, concentrated

near large river estuaries.
Ventral disc
Nuchal area
Giant Freshwater Whipray

Himantura polylepis (Bleeker, 1852)
6
5
2
1 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.
Myliobatoidei 209
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).
In Borneo: Known from Sabah and

eastern Kalimantan.
Ventral disc
Nuchal area
White-edge Freshwater Whipray

Himantura signifer Compagno & Roberts, 1982
5
3
2
9 7
1 8
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
Myliobatoidei 211
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.
References: None.
In Borneo: Kapuas, Sampi and Rungan

rivers, western and southern Kalimantan.
Ventral disc
Nuchal area
Bleeker’s Whipray
Himantura uarnacoides (Bleeker, 1852)
5
6
9
7
2
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.
Myliobatoidei 213
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.
References: None.
4
In Borneo: Widespread, less common off

western Sabah.
Ventral disc
Nuchal area
Juvenile (277 mm TL)

Reticulate Whipray
Himantura uarnak (Forsskål, 1775)
3
7
1 9
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.
Myliobatoidei 215
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).
References: None.
In Borneo: Widespread, but patchy.
Ventral disc
Nuchal area
Juvenile (262 mm DW)

Honeycomb Whipray
Himantura undulata (Bleeker, 1852)
6 5
1 7
9
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.
Myliobatoidei 217
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).
In Borneo: Sarawak, northeastern
Sabah, and northwestern and eastern
Kalimantan; probably more widespread.
Ventral disc
8 4
Nuchal area
Juvenile (350 mm DW)

Dwarf Whipray
Himantura walga (Müller & Henle, 1841)
2
3
6
1
5
7
Other names: None (English), Pari Kikir, Toka Toka (Indonesian), Pari Ketuka, Tuka-
tuka, 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
Myliobatoidei 219
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.
References: None.
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)
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.
Myliobatoidei 221
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).
References: None.
Ventral disc
Nuchal area
3
Lateral tail
Eastern Cowtail Stingray

Pastinachus atrus (Macleay, 1883)
5
2
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.
Myliobatoidei 223
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).
References: None.
Ventral disc
Nuchal area
Lateral tail - at midlength of ventral fold

Narrowtail Stingray
Pastinachus gracilicaudus Last & Manjaji-Matsumoto, 2010
4
3 1 6 8
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
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.
Myliobatoidei 225
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).
In Borneo: Patchy, in western and
northern Borneo; not observed from most
of southern and eastern Kalimantan.
Ventral disc
Nuchal area
Roughnose Stingray
Pastinachus solocirostris Last, Manjaji & Yearsley, 2005
4
5
1 7
3
6
8
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 <3 times height of tail above)
6 tail relatively broad-based
Size: To at least 69 cm DW (>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
Myliobatoidei 227
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).
In Borneo: Widespread off western
Borneo; isolated records from southern
Ventral disc and southeastern Kalimantan.
3
Nuchal area
5 Denticles at apex of snout

Starynose Stingray
Pastinachus stellurostris Last, Fahmi & Naylor, 2010
4 5
1
3 6 8
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 <3 times height of tail above)
6 tail relatively broad-based
8 row of small thorns along middle of tail
Size: To at least 43 cm DW (134 cm TL).
Distribution: Possibly endemic to Borneo.
Habitat and biology: Habitat and biology unknown; demersal and probably associated
with the large river estuaries of the region. Presumably viviparous, with histotrophy. Diet
Myliobatoidei 229
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.
References: None.
In Borneo: Restricted, known only from
northwestern Kalimantan.
Ventral disc
Nuchal area 3
Denticles at apex of snout

5

Pelagic Stingray
Pteroplatytrygon violacea (Bonaparte, 1832)
4
5
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
Myliobatoidei 231
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.
References: None.
5
Ventral disc In Borneo: Known from off Sarawak

and Sabah, but likely to be more widely
distributed.
4
Lateral tail
Bluespotted Fantail Ray

Taeniura lymma (Forsskål, 1775)
2
5
4
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.
Myliobatoidei 233
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.
References: None.
In Borneo: Probably widespread,
although less common in Sarawak and
parts of Kalimantan.
Ventral disc
3
Lateral tail
Blotched Fantail Stingray

Taeniurops meyeni (Müller & Henle, 1841)
3
2
4
6
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.
Myliobatoidei 235
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).
References: None.
In Borneo: Known only from Sabah and
Ventral disc off Sarawak.
3
Lateral tail
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.
Myliobatoidei 237
Utilisation: Caught rarely by demersal tangle net and trawl fisheries. Probably used for
its meat, but may be discarded.
References: None.
In Borneo: Uncommon, known from
off Sabah and eastern Kalimantan;
Ventral disc possibly more widely distributed in
appropriate habitat.
Lateral head Mouth and nostrils

Longtail Butterfly Ray

Gymnura poecilura (Shaw, 1804)
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.
Myliobatoidei 239
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)
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 inter-
spersed 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.
Myliobatoidei 241
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).
Gymnuridae (Butterfly Rays)

1
2
Lateral anterior tail
In Borneo: Sarawak, Sabah, and eastern

and western Kalimantan.
Ventral disc
Longhead Eagle Ray

Aetobatus flagellum (Bloch & Schneider, 1801)
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
Myliobatoidei 243
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.
References: None.
Myliobatidae (Eagle Rays)

3
Ventral head
In Borneo: Uncommon, known from
western Kalimantan and northeastern
Sabah.
Upper tooth plate
Lower tooth plate

Whitespotted Eagle Ray

Aetobatus ocellatus (Kuhl, 1823)
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.
Myliobatoidei 245
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).
Ventral head
Upper tooth plate
Juvenile - dorsal disc (465 mm DW) Lower tooth plate

Mottled Eagle Ray

Aetomylaeus maculatus (Gray, 1834)
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.
Myliobatoidei 247
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 publi-
cation date listed on Eschmeyer’s Catalog of Fishes as 1 October 1834.
References: None.

5
Ventral head
Upper tooth plate
Juvenile - dorsal disc (~250 mm DW) Lower tooth plate

Banded Eagle Ray

Aetomylaeus nichofii (Bloch & Schneider, 1801)
1
6
Other names: Barbless Eagle Ray, Nieuhof ’s Eagle Ray (English), Pari Burung, Pe
Lamping (Indonesian), Pari-lang Jalur (Malaysian)
2 sting absent
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.
Myliobatoidei 249
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.
References: None.

5
Ventral head
Upper tooth plate
Lower tooth plate

Ornate Eagle Ray

Aetomylaeus vespertilio (Bleeker, 1852)
1
6
Other names: Reticulate Eagle Ray (English), Pari Burung (Indonesian), Pari Lang
Bunga (Malaysian)
2 sting absent
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-
Myliobatoidei 251
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.
References: None.

5
Ventral head
In Borneo: Known locally from a
few individuals collected off Sabah and
Kalimantan.
Upper tooth plate
Lower tooth plate

Javanese Cownose Ray

Rhinoptera javanica Müller & Henle, 1841
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.
Myliobatoidei 253
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 bottom-
dwelling invertebrates.
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.
Rhinopteridae (Cownose Rays)

References: None.
In Borneo: Apparently uncommon,

Ventral head known only from eastern Sabah and
western Kalimantan.
5 6
7
3
Lateral dorsal fin and anterior tail
Lateral head
Short-tail Cownose Ray

Rhinoptera jayakari Boulenger, 1895
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 1st 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.
Myliobatoidei 255
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).
Rhinopteridae (Cownose Rays)

References: None.
Ventral head In Borneo: Known off eastern Sabah,

southern Sarawak and northwestern
Kalimantan.
5 6

Lateral head
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 5th 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.
temperate waters.
Habitat and biology: Wide-ranging, pelagic species found near the surface in tropical
Myliobatoidei 257
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 5th 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.
Mobulidae (Devilrays)
References: White et al. (2006a); Marshall et al. (2009).
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.
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
Myliobatoidei 259
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.
References: None.
2
3
7
In Borneo: Known from off Kuching,

Sarawak.
Ventral head
Dorsolateral head
Japanese Devilray
Mobula japanica (Müller & Henle, 1841)
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.
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
Myliobatoidei 261
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).
Late-term embryo (498 mm DW)
2 In Borneo: Recorded from Semporna in

east Sabah.; probably widespread.
3
Lateral dorsal fin and sting
Dorsolateral head Ventral head

Lesser Devilray
Mobula kuhlii (Müller & Henle, 1841)
1
2
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.
Myliobatoidei 263
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
3
7
Ventral head In Borneo: Northeastern Borneo, off

Sabah.
4
6
Dorsolateral head
Bentfin Devilray
Mobula thurstoni (Lloyd, 1908)
1
2
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
Myliobatoidei 265
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.
2
Ventral head In Borneo: Single record from the

Sandakan fish market, northeastern
Sabah.
7
6

Dorsolateral head
Silver Chimaera
Chimaera phantasma Jordan & Snyder, 1900
6 2 5
7
3
1
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 1st dorsal-fin spine long, extending to near or beyond fin apex
5 2nd 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.
Chimaeriformes 267

Local synonyms: Chimaera sp.: Yano et al. (2005).
References: Masuda et al. (1975); Dagit (2006).
Chimaeridae (Shortnose Chimaeras)

8
Lateral head In Borneo: Known only from off

Sarawak.
1
Lateral tail
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
Acknowledgements 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
References
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Azman, Z. (2008) Panduan mengenali spesies ikan yu di Malaysia, Brunei Darussalam,
Indonesia dan Thailand. Jabatan Perikanan Malaysia. 209 pp.
Allen, G.R. (2008). Conservation hotspots of biodiversity and endemism for Indo–
Pacific coral reef fishes. Aquatic Conservation: Marine & Freshwater Ecosystems 18:
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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 Western Highfin Spurdog
Family Centrophoridae: Gulper Sharks
 Centrophorus moluccensis Bleeker, 1860 Smallfin Gulper Shark
Order Squatiniformes: Angelsharks

Family Squatinidae: Angelsharks
 Squatina tergocellatoides Chen, 1963 Ocellated Angelshark
Order Heterodontiformes: Hornsharks

Family Heterodontidae: Hornsharks
 Heterodontus zebra (Gray, 1831) Zebra Hornshark
Order Orectolobiformes: Carpet Sharks

Family Orectolobidae: Wobbegongs
 Orectolobus leptolineatus Last et al., 2010c Indonesian Wobbegong
Family Hemiscylliidae: Longtail Carpetsharks
 Chiloscyllium hasseltii Bleeker, 1852 Indonesian Bambooshark
 Chiloscyllium indicum (Gmelin, 1789) Slender Bambooshark
 Chiloscyllium plagiosum (Bennett, 1830) Whitespotted Bambooshark
 Chiloscyllium punctatum M. & H., 1838 Brownbanded Bambooshark
Family Ginglymostomatidae: Nurse Sharks
 Nebrius ferrugineus (Lesson, 1830) Tawny Nurse Shark
Family Stegostomatidae: Zebra Sharks
 Stegostoma fasciatum (Hermann, 1783) Zebra Shark
Family Rhincodontidae: Whale Sharks

 Rhincodon typus Smith, 1828 Whale Shark
Order Lamniformes: Mackerel Sharks

Family Alopiidae: Thresher Sharks
 Alopias pelagicus Nakamura, 1935 Pelagic Thresher
Family Lamnidae: Mackerel Sharks

 Isurus oxyrinchus Rafinesque, 1810 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
Checklist of species 279
 Carcharhinus leucas (M. & H., 1839) Bull Shark

 Carcharhinus limbatus (M. & H., 1839) Common Blacktip Shark
 Carcharhinus macloti (M. & H., 1839) Hardnose Shark
 Carcharhinus melanopterus (Q. & G., 1824) Blacktip Reef Shark
 Carcharhinus plumbeus (Nardo, 1827) Sandbar Shark
 Carcharhinus sealei (Pietschmann, 1913) Blackspot Shark
 Carcharhinus sorrah (M. & H., 1839) Spot-tail Shark
 Galeocerdo cuvier (Péron & Lesueur, 1822) Tiger shark
 Glyphis sp. Mukah River Shark
 Glyphis fowlerae Compagno et al., 2010 Borneo River Shark
 Lamiopsis tephrodes (Fowler, 1905) Borneo Broadfin Shark
 Loxodon macrorhinus M. & H., 1839 Sliteye Shark
 Prionace glauca (Linnaeus, 1758) Blue Shark
 Rhizoprionodon acutus (Rüppell, 1837) Milk Shark
 Rhizoprionodon oligolinx Springer, 1964 Grey Sharpnose Shark
 Scoliodon macrorhynchos (Bleeker, 1852) Pacific Spadenose Shark
 Triaenodon obesus (Rüppell, 1837) Whitetip Reef Shark
Family Sphyrnidae: Hammerhead Sharks
 Eusphyra blochii (Cuvier, 1816) Winghead Shark
 Sphyrna lewini (Griffith & Smith, 1834) Scalloped Hammerhead
 Sphyrna mokarran (Rüppell, 1837) Great Hammerhead
Order Rajiformes: Rays (Batoids)

Suborder Pristoidei: Sawfishes
Family Pristidae: Sawfishes
 Anoxypristis cuspidata (Latham, 1794) Narrow Sawfish
 Pristis microdon Latham, 1794 Freshwater Sawfish
 Pristis zijsron Bleeker, 1851 Green Sawfish
Suborder Rhinoidei: Shark Rays

Family Rhinidae: Shark Rays
 Rhina ancylostoma Bl. & Sch., 1801 Shark Ray
Suborder Rhynchobatoidei: Wedgefishes

Family Rhynchobatidae: Wedgefishes
 Rhynchobatus australiae Whitley, 1939 Whitespotted Wedgefish
 Rhynchobatus laevis (Bl. & Sch., 1801) Smoothnose Wedgefish
 Rhynchobatus springeri Comp. & Last, 2010 Broadnose Wedgefish
Suborder Rhinobatoidei: Shovelnose Rays

Family Rhinobatidae: Shovelnose Rays
 Glaucostegus thouin (Anonymous, 1798) Clubnose Guitarfish
 Glaucostegus typus (Bennett, 1830) Giant Guitarfish
 Rhinobatos formosensis Norman, 1926 Taiwan Guitarfish
Suborder Torpedinoidei: Electric Rays

Family Narcinidae: Numbfishes
 Narcine brevilabiata Bessednov, 1966 Shortlip Numbfish
 Narcine lingula Richardson, 1846 Rough Numbfish
 Narcine maculata (Shaw, 1804) Darkspotted Numbfish
Family Narkidae: Sleeper Rays
 Narke dipterygia (Bl. & Sch., 1801) Spot-tail Sleeper Ray
 Temera hardwickii Gray, 1831 Finless Sleeper Ray
Suborder Rajoidei: Skates

Family Rajidae: Skates
 Dipturus kwangtungensis (Chu 1960) Kwangtung Skate
 Okamejei cairae Last et al., 2010a Borneo Sand Skate
 Okamejei hollandi (Jordan & Rich., 1909) Yellowspotted Skate
 Okamejei jensenae Last & Lim, 2010 Philippine Ocellate Skate

Checklist of species 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 Narrowtail Stingray

 Pastinachus solocirostris Last et al., 2005 Roughnose Stingray
 Pastinachus stellurostris Last et al., 2010b Starynose Stingray
 Pteroplatytrygon violacea (Bonaparte, 1832) Pelagic Stingray
 Taeniura lymma (Forsskål, 1775) Bluespotted Fantail Ray
 Taeniurops meyeni (M. & H., 1841) Blotched Fantail Stingray
 Urogymnus asperrimus (Bl. & Sch., 1801) Porcupine Ray
Family Gymnuridae: Butterfly Rays
 Gymnura poecilura (Shaw, 1804) Longtail Butterfly Ray
 Gymnura zonura (Bleeker, 1852) Zonetail Butterfly Ray
Family Myliobatidae: Eagle Rays
 Aetobatus flagellum (Bl. & Sch., 1801) Longhead Eagle Ray
 Aetobatus ocellatus (Kuhl, 1823) Whitespotted Eagle Ray
 Aetomylaeus maculatus (Gray, 1834) Mottled Eagle Ray
 Aetomylaeus nichofii (Bl. & Sch., 1801) Banded Eagle Ray
 Aetomylaeus vespertilio (Bleeker, 1852) Ornate Eagle Ray
Family Rhinopteridae: Cownose Rays
 Rhinoptera javanica M. & H., 1841 Javanese Cownose Ray
 Rhinoptera jayakari Boulenger, 1895 Short-tail Cownose Ray
Family Mobulidae: Devilrays
 Manta birostris (Walbaum, 1792) Manta Ray
 Mobula sp. Borneo Devilray
 Mobula japanica (M. & H., 1841) Japanese Devilray
 Mobula kuhlii (M. & H., 1841) Lesser Devilray
 Mobula thurstoni (Lloyd, 1908) 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 Carcharhinus brevipinna 94
Aetobatus flagellum 242 Carcharhinus dussumieri 96
Aetobatus ocellatus 244 Carcharhinus falciformis 98
Aetomylaeus maculatus 246 Carcharhinus leucas 100
Aetomylaeus nichofii 248 Carcharhinus limbatus 102
Aetomylaeus vespertilio 250 Carcharhinus macloti 104
albimarginatus, Carcharhinus 86 Carcharhinus melanopterus 106
Alopias pelagicus 62 Carcharhinus plumbeus 108
Alopiidae 29, 62–63 Carcharhinus sealei 110
altipinnis, Squalus 38 Carcharhinus sorrah 112
amblyrhynchoides, Carcharhinus 88 Centrophoridae 29, 40–41
amblyrhynchos, Carcharhinus 90 Centrophorus moluccensis 40
Anacanthobatidae 25, 178–179 Cephaloscyllium sarawakensis 70
ancylostoma, Rhina 146 Chaenogaleus macrostoma 78
Anoxypristis cuspidata 140 Chiloscyllium hasseltii 48
Apristurus platyrhynchus 66 Chiloscyllium indicum 50
asperrimus, Urogymnus 236 Chiloscyllium plagiosum 52
Atelomycterus marmoratus 68 Chiloscyllium punctatum 54
atrus, Pastinachus 222 Chimaera phantasma 266
australiae, Rhynchobatus 148 Chimaeridae 23, 266–267
Chimaeriformes 266–267
B cuspidata, Anoxypristis 140
birostris, Manta 256 cuvier, Galeocerdo 114
blochii, Eusphyra 134
borneensis, Carcharhinus 92 D
borneensis, Sinobatis 178 Dasyatidae 26, 182–237
brevilabiata, Narcine 160 Dasyatis microps 182
brevipinna, Carcharhinus 94 Dasyatis parvonigra 184
Dasyatis ushiei 186
C Dasyatis zugei 188
cairae, Okamejei 172 daviesi, Plesiobatis 180
Carcharhinidae 33, 86–133 dipterygia, Narke 166
Carcharhiniformes 66–139 Dipturus kwangtungensis 170
Carcharhinus albimarginatus 86 dussumieri, Carcharhinus 96
Carcharhinus amblyrhynchoides 88
Carcharhinus amblyrhynchos 90 E
Carcharhinus borneensis 92 elongata, Hemipristis 82
Eusphyra blochii 134 Hexanchus griseus 36

Himantura fai 190
F Himantura gerrardi 192
fai, Himantura 190 Himantura granulata 194
falciformis, Carcharhinus 98 Himantura jenkinsii 196
fasciatum, Stegostoma 58 Himantura kittipongi 198
ferrugineus, Nebrius 56 Himantura leoparda 200
flagellum, Aetobatus 242 Himantura lobistoma 202
formosensis, Rhinobatos 158 Himantura oxyrhyncha 204
fowlerae, Glyphis 118 Himantura pastinacoides 206
Himantura polylepis 208
G
Himantura signifer 210
Galeocerdo cuvier 114
Himantura uarnacoides 212
gerrardi, Himantura 192
Himantura uarnak 214
Ginglymostomatidae 30, 56–57
Himantura undulata 216
glauca, Prionace 124
Himantura walga 218
Glaucostegus thouin 154
hollandi, Okamejei 174
Glaucostegus typus 156
Glyphis fowlerae 118 I
Glyphis sp. 116 indicum, Chiloscyllium 50
gracilicaudus, Pastinachus 224 Isurus oxyrinchus 64
granulata, Himantura 194
griseus, Hexanchus 36 J
Gymnura poecilura 238 japanica, Mobula 260
Gymnura zonura 240 javanica, Rhinoptera 252
Gymnuridae 26, 238–241 jayakari, Rhinoptera 254
jenkinsii, Himantura 196
H jensenae, Okamejei 176
Halaelurus maculosus 72
hardwickii,Temera 168 K
hasseltii, Chiloscyllium 48 kittipongi, Himantura 198
Hemigaleidae 32, 78–85 kuhlii, Mobula 262
Hemigaleus microstoma 80 kuhlii, Neotrygon 220
Hemipristis elongata 82 kwangtungensis, Dipturus 170
Hemiscylliidae 31, 48-55
Heterodontidae 30, 44–45 L
Heterodontiformes 44–45 laevis, Rhynchobatus 150
Heterodontus zebra 44 Lamiopsis tephrodes 120
Hexanchidae 28, 36–37 Lamnidae 31, 64–65
Hexanchiformes 36–37 Lamniformes 62–63
SR cuinennitni gf i ch enaadm e s i n d e x 285
leoparda, Himantura 200 Narcine maculata 164

leptolineatus, Orectolobus 46 Narcinidae 25, 160–165
leucas, Carcharhinus 100 Narke dipterygia 166
lewini, Sphyrna 136 Narkidae 25, 166–169
limbatus, Carcharhinus 102 Nebrius ferrugineus 56
lingula, Narcine 162 Neotrygon kuhlii 220
lobistoma, Himantura 202 nichofii, Aetomylaeus 248
Loxodon macrorhinus 122
lymma,Taeniura 232
O
obesus,Triaenodon 132
M ocellatus, Aetobatus 244
macloti, Carcharhinus 104 Okamejei cairae 172
macrorhinus, Loxodon 122 Okamejei hollandi 174
macrorhynchos, Scoliodon 130 Okamejei jensenae 176
macrostoma, Chaenogaleus 78 oligolinx, Rhizoprionodon 128
maculata, Narcine 164 Orectolobidae 30, 46–47
maculatus, Aetomylaeus 246 Orectolobiformes 46–61
maculosus, Halaelurus 72 Orectolobus leptolineatus 46
manazo, Mustelus 74 oxyrhyncha, Himantura 204
Manta birostris 256 oxyrinchus, Isurus 64
marmoratus, Atelomycterus 68
melanopterus, Carcharhinus 106
P
meyeni,Taeniurops 234 Paragaleus tengi 84
microps, Dasyatis 182 parvonigra, Dasyatis 184
microstoma, Hemigaleus 80 Pastinachus atrus 222
Mobula japanica 260 Pastinachus gracilicaudus 224
Mobula kuhlii 262 Pastinachus solocirostris 226
Mobula sp. 258 Pastinachus stellurostris 228
Mobula thurstoni 264 pastinacoides, Himantura 206
Mobulidae 27, 256–265 pelagicus, Alopias 62
mokarran, Sphyrna 138 phantasma, Chimaera 266
moluccensis, Centrophorus 40 plagiosum, Chiloscyllium 52
Mustelus manazo 74 platyrhynchus, Apristurus 66
Mustelus widodoi 76 Plesiobatidae 26, 180–181
Myliobatidae 28, 242–251 Plesiobatis daviesi 180
Myliobatoidei 180–265 plumbeus, Carcharhinus 108
poecilura, Gymnura 238
N polylepis, Himantura 208
Narcine brevilabiata 160 Prionace glauca 124
Narcine lingula 162 Pristidae 23, 140–145
Pristis microdon 142 Sphyrna mokarran 138

Pristis zijsron 144 Sphyrnidae 29, 134–139
Pristoidei 140–145 springeri, Rhynchobatus 152
Pteroplatytrygon violacea 230 Squalidae 29, 38–39
punctatum, Chiloscyllium 54 Squaliformes 38–41
Squalus altipinnis 38
R Squatina tergocellatoides 42
Rajidae 25, 172–179 Squatinidae 23, 42–43
Rajoidei 172–179 Squatiniformes 42–43
Rhina ancylostoma 146 Stegostoma fasciatum 58
Rhincodontidae 30, 60–61 Stegostomatidae 30, 58–59
Rhincodon typus 60 stellurostris, Pastinachus 228
Rhinidae 24, 146–147
Rhinobatidae 24, 154–159 T
Rhinobatoidei 154–159 Taeniura lymma 232
Rhinobatos formosensis 158 Taeniurops meyeni 234
Rhinoidei 146–147 Temera hardwickii 168
Rhinoptera javanica 252 tengi, Paragaleus 84
Rhinoptera jayakari 254 tephrodes, Lamiopsis 120
Rhinopteridae 28, 252–255 tergocellatoides, Squatina 42
Rhizoprionodon acutus 126 thouin, Glaucostegus 154
Rhizoprionodon oligolinx 128 thurstoni, Mobula 264
Rhynchobatidae 24, 148–153 Torpedinoidei 160–169
Rhynchobatoidei 147 Triaenodon obesus 132
Rhynchobatus australiae 148 Triakidae 32, 74–77
Rhynchobatus laevis 150 typus, Glaucostegus 156
Rhynchobatus springeri 152 typus, Rhincodon 60
S U
sarawakensis, Cephaloscyllium 70 uarnacoides, Himantura 212
Scoliodon macrorhynchos 130 uarnak, Himantura 214
Scyliorhinidae 31, 66–73 undulata, Himantura 216
sealei, Carcharhinus 110 Urogymnus asperrimus 236
signifer, Himantura 210 ushiei, Dasyatis 186
Sinobatis borneensis 178 V
solocirostris, Pastinachus 226
vespertilio, Aetomylaeus 250
sorrah, Carcharhinus 112
violacea, Pteroplatytrygon 230
sp., Glyphis 116
sp., Mobula 258 W
Sphyrna lewini 136 walga, Himantura 218
Scientific names index 287
widodoi, Mustelus 76
Z
zebra, Heterodontus 44
zonura, Gymnura 240
zugei, Dasyatis 188
English common names index
A Borneo Leg Skate 178

Borneo River Shark 118
Angelsharks 42–43
Borneo Sand Skate 172
B Borneo Shark 92
Bowmouth Guitarfish 146
Banana-tail Ray 222
Broadnose Wedgefish 152
Banded Eagle Ray 248
Brownbanded Bambooshark 54
Banded Whiptail Ray 192
Brownbanded Catshark 54
Barbless Eagle Ray 248
Brown Shark 108
Bentfin Devilray 264
Bullhead Shark 44
Bigfin Catshark 66
Bull Ray 222, 234
Blackblotched Stingray 234
Bull Shark 36, 100
Blackspot Shark 110
Butterfly Rays 238–241
Blackspotted Stingray 234
Blacktip Reef Shark 106 C
Blacktip Shark 106 Catsharks 66–73
Blacktip Whaler 102 Chinese Numbfish 162
Bleeker’s Butterfly Ray 240 Clubnose Guitarfish 154
Bleeker’s Variegate Ray 216 Coachwhip Ray 194, 214
Bleeker’s Whipray 212 Common Blacktip Shark 102
Blotched Fantail Stingray 234 Common Shovelnose Ray 156
Blue-spotted Lagoon Ray 232 Coral Catshark 68
Blue-spotted Ribbontail Ray 232 Cownose Rays 252–255
Blue Pointer 64 Cow Ray 252
Blue Shark 124 Cow Shark 36
Bluespotted Fantail Ray 232 Cowsharks 36–37
Bluespotted Maskray 220 Cow Stingray 186
Bluespotted Stingaree 220
Bluespotted Stingray 220 D
Blue Stingray 230 Darkfin Numbfish 164
Blue Whaler 124 Darkspotted Electric Ray 164
Blunthead Shark 132 Darkspotted Numbfish 164
Bluntnose Sixgill Shark 36 Deepwater Stingray 180
Bluntnose Whiptail Ray 192 Devilfish 256
Bonnet Skate 244 Devilrays 256–265
Borneo Broadfin Shark 120 Dogfishes 38–39
Borneo Devilray 258 Duckbill Ray 244
English common names index 289
Dwarf Black Stingray 184 Grey Sharpnose Shark 128

Dwarf Whipray 218 Guliman 106
Gulper Sharks 40–41
E Gummy Shark 74
Eagle Rays 242–251
Eastern Cowtail Stingray 222 H
Endeavour Dogfish 40 Hammerhead Sharks 134–139
Hardnose Shark 104
F Hasselt’s Bambooshark 48
Fantail Ray 222 Holland Skate 174
Fantail Stingray 234 Honeycomb Whipray 216
Feathertail Ray 222 Hooktooth Shark 78
Feathertail Stingray 222 Hornsharks 44–45
Finless Electric Ray 168 Houndsharks 74–77
Finless Sleeper Ray 168
I
Fish Shark 126
Indonesian Bambooshark 48
Flapnose Ray 252
Indonesian Speckled Catshark 72
Fossil Shark 82
Indonesian Wobbegong 46
Freshwater Sawfish 142
Indo Wobbegong 46
Freshwater Stingray 210
Inkytail Shark 94
Freshwater Whaler 100
Freshwater Whipray 208 J
G Japanese Devilray 260
Japanese Smoothhound 74
Ghost Shark 266
Javanese Cownose Ray 252
Giant Freshwater Stingray 208
Jenkins’ Whipray 196
Giant Freshwater Whipray 208
Jordan’s Blue Dogshark 122
Giant Guitarfish 150, 156
Giant Manta 256 K
Giant Reef Ray 234 Kidney-headed Shark 136
Giant Shovelnose Ray 156 Kuhl’s Stingray 220
Giant Stingaree 180 Kwangtung Skate 170
Giant Stingarees 180–181
Graceful Shark 88 L
Great-tooth Sawfish 142 Lagoon Ray 232
Great Blue Shark 124 Leg Skates 178–179
Great Hammerhead 138 Leopard Shark 58
Green Sawfish 144 Leopard Stingray 214
Grey Carpetshark 54 Leopard Whipray 200
Grey Reef Shark 90 Lesser Devilray 262
Lesser Fantail Ray 232 O

Long-tailed Butterfly Ray 238 Ocellated Angelshark 42
Longhead Eagle Ray 242 Ocellate Eagle Ray 246
Longman’s Dogshark 126 Ocellate Whipray 216
Longnose Blacktail Shark 90 Ornate Eagle Ray 246, 250
Longnose Grey Whaler 94
Longnose Marbled Whipray 204 P
Longtail Butterfly Ray 238 Pacific Spadenose Shark 130
Longtail Carpetsharks 48–55 Pale-edged Stingray 188
Longtail Ray 214 Pale Whipray 210
Pelagic Stingray 230
M Pelagic Thresher 62
Mackerel Shark 64 Philippine Ocellate Skate 176
Mackerel Sharks 64–65 Pink Whipray 190
Macleay’s Coachwhip Ray 194 Pointed Sawfish 140
Madame X 56 Porcupine Ray 236
Mako Shark 64
Mangrove Ray 194 Q
Mangrove Stingray 194 Queensland Shark 88
Mangrove Whipray 194
Manta 256 R
Manta Ray 256 Reef Ray 232
Marbled Freshwater Stingray 204 Reticulate Eagle Ray 250
Marbled Whipray 204 Reticulate Whipray 214
Marble Ribbontail Ray 234 Ribbontail Stingray 232
Milk Shark 126 Ridgeback Bambooshark 50
Mottled Eagle Ray 246 Ridgeback Shark 50
Mud Shark 36 River Whaler 100
Mud Skate 146 Rough-back Stingray 196
Mukah River Shark 116 Roughback Whipray 198
Rough Electric Ray 162
N Roughnose Stingray 226
Narrow Sawfish 140 Rough Numbfish 162
Narrowsnout Sawfish 144 Roughskin Stingaree 236
Narrowtail Stingray 224 Round Ribbontail Ray 234
Nieuhof ’s Eagle Ray 248 Round Whipray 206
Numbfishes 160–165
Numbray 166 S
Nurse Sharks 56–57 Sandbar Shark 108
Sandshark 150
English common names index 291
Sarawak Pygmy Swell Shark 70 Solander’s Ray 236

Sarawak Pygmy Swellshark 70 Sorrah Shark 112
Sarawak Swellshark 70 Spatulasnout Catshark 66
Sawfish 144 Speckle Stingray 234
Sawfishes 140–145 Spinetail Devilray 260
Scalloped Hammerhead 136 Spinetail Mobula 260
School Shark 112 Spinner Shark 94
Shark Ray 146 Spitting Shark 56
Shark Rays 146–147 Spot-tail Electric Ray 166
Sharpnose Stingray 188, 192 Spot-tail Shark 112
Short-tail Cownose Ray 254 Spot-tail Sleeper Ray 166
Shortfin Devilray 262 Spotted Catshark 54
Shortfin Mako 64 Spotted Eagle Ray 244
Shorthorn Devilray 262 Starspotted Smoothhound 74
Shortlip Electric Ray 160 Starynose Stingray 228
Shortlip Numbfish 160 Stingrays 182–237
Shortnose Chimaeras 266–267 Straight-tooth Weasel Shark 84
Shovelnose Ray 156
Shovelnose Rays 154–159 T
Shovelnose Shark 150, 152, 156 Taiwan Guitarfish 158
Sicklefin Weasel Shark 80 Tawny Nurse Shark 56
Silky Shark 98 Tawny Shark 56
Silver Chimaera 266 Thickskin Shark 108
Silvertip Shark 86 Thickspine Giant Stingray 182
Sixgill Shark 36 Thornback Ray 174
Skates 170–177 Thorny Ray 236
Sleeper Rays 166–169 Thresher Sharks 62–63
Sleepy Shark 56 Tiger Shark 114
Slender Bambooshark 50 Tubemouth Whipray 202
Slender Dogshark 122
U
Slender Hammerhead 134
Undulate Whipray 200
Sliteye Shark 122
Smalleye Stingray 182 V
Smallfin Gulper Shark 40 Violet Stingray 230
Smoothfang Shark 94
Smoothnose Wedgefish 150 W
Smoothtail Mobula 264 Weasel Shark 80
Snaggletooth Shark 82 Weasel Sharks 78–85
Snapper Shark 64 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 Hiu Gedebong 56
Hiu Gedok 54, 56
Cawang Genul 260
Hiu Gergaji 142, 144
Cawang Kalung 256
Hiu Hantu 266
Cucut Bekeman 100
Hiu Jenggot 46
Cucut Botol 40
Hiu Kacang 74, 76, 78, 80, 84
Cucut Dolok 50, 52
Hiu Kakap 64
Cucut Gergaji 140
Hiu Karang Buas 132
Cucut Lanjaman 86, 88, 90, 94, 96, 98,
102, 108, 110, 112 Hiu Karet 124
Cucut Londer 74, 76 Hiu Kejen 102, 122, 130
Cucut Mandrong 154 Hiu Kemenyan 148, 150, 152
Cucut Meong 36 Hiu Kodok 42, 46
Cucut Pedang 62 Hiu Lalaek 124
Cucut Selendang 124 Hiu Lanyam 86, 98, 102
Cucut Tokek 68 Hiu Lepang 46
Hiu Lonjor 90, 98
H Hiu Lontar 148, 150, 152
Hiu 44, 92, 148 Hiu Macan 114
Hiu Aer 124 Hiu Mada 106
Hiu Air 74, 76 Hiu Mako 64
Hiu Anjing 64 Hiu Martil 134, 136, 138
Hiu Aron 104 Hiu Minsong 150, 152
Hiu Barong 146 Hiu Monas 82
Hiu Batu 54 Hiu Monyet 62
Hiu Belimbing 58 Hiu Omas 114
Hiu Bingkoh 134, 136, 138 Hiu Parang 136, 138, 140, 142, 144
Hiu Bisu 56 Hiu Pasir 84
Hiu Bokem 132 Hiu Paus 60
Hiu Bongo 52, 54 Hiu Pilus 78, 80, 126, 128
Hiu Botol Danten 40 Hiu Pisang 126, 128
Hiu Buas 100 Hiu Plen 86, 94, 126
Hiu Bujit 88, 90, 94, 96, 98, 100, Hiu Pogak 266
102, 106, 112, 120 Hiu Senget 40
Hiu Capil 134, 136, 138 Hiu Sonteng 86
Hiu Caping 134, 136, 138 Hiu Tahu Putih 36
Hiu Cermin 156 Hiu Taji 40
Hiu Coklat 132 Hiu Tekek 48, 50, 52, 54
Hiu Tenggiri 64 Pari Boro 186

Hiu Teteri 108 Pari Burung 242, 244, 246, 248, 250, 252
Hiu Tikus 62 Pari Duren 236
Hiu Tokek 68, 72 Pari Duri 196
Pari Elang 252, 254
I Pari Gitar 156
Ikan Pari 184, 232 Pari Gunung 226
Pari Jantung 186
K
Pari Kekeh 156
Kluyu Blimbingan 58
Pari Kelelawar 238, 240
Kluyu Karang 106
Pari Kembang 232
Kupu-kupu Brangkas 146
Pari Kerbua 256
L Pari Kikir 196, 218
Pari Kodok 220
Lampengan 260
Pari Kontak 160, 162, 164, 166, 168
Lanyam 112
Pari Lambingan 260, 262, 264
Liongbun 148, 150, 152, 154
Pari Lampengan 262
M Pari Lampin 230
Merak Bulu 90, 94, 102, 112 Pari Lemer 230
Minsong 148 Pari Lempengan 264
Mungsing 98, 112, 126, 130 Pari Listrik 160, 162, 164, 166, 168
Mungsing Jara 114 Pari Lowo 238, 240
Pari Lumpur 202, 212
P Pari Luncur 172, 174
Paitpait 156 Pari Macan 200, 214, 216
Pamprang 140 Pari Merica 214, 234
Pangrum 148, 150, 152 Pari Minyak 190
Parangpang 140 Pari Pasir 206
Paredung 148, 150, 152 Pari Payung 240
Pari 172, 180, 204, 208 Pari Sapi 182, 194
Pari Aer 196, 206 Pari Sungai 198, 210
Pari Ayam 244 Pari Super 192
Pari Babi 182, 234 Pari Tanjung 202, 212
Pari Bandrong 156 Pari Tembaga 190
Pari Batu 192 Pari Totol 232
Pari Bendera 222, 224, 226, 228 Payubek 230
Pari Biasa 188 Pe Lamping 248
Pari Bintang 192 Pe Pasung 212
Pari Blentik 220 Petong 148, 150, 152, 156
Indonesian common names index 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 Pari Bendera 222

Bengkong 134 Pari Bintik Putih 192
Beroi 140 Pari Bulat 206
Pari Daun 222
C Pari Dedap 236
Chimara Perak 266 Pari Duri 236
Pari Durian 236
I Pari Ekor Duri 186
Ikan Tikus 266 Pari Gergasi 182
Pari Helang 244
J
Pari Hitam 230
Jerung-mako Sirip 64
Pari Karan 164, 166
Jerung Biru 124
Pari Karan Belangkas 168
Jerung Gigi Cerakah 82
Pari Karang 232
Jerung Sapi 100
Pari Karan Tompok Besar 160
Jerung Tenggiri 114
Pari Karas 168
K Pari Kelabu 234
Kan Sua 140 Pari Kelawar 240, 256, 262
Kemejan 152 Pari Kelawar Bintik 238
Kiampau 220, 232 Pari Kemejan 150
Pari Ketuka 188, 218
P Pari Lalat 220
Pari 152, 184 Pari Lang 242, 244, 252, 254
Pari-karan Hitam 164 Pari Lang Bintik Putih 244
Pari-kelawar Jawa 252 Pari Lang Bunga 250
Pari-kemejan Kepala Keras 146 Pari Lembik Merah 180
Pari-kemejan Muncung Panjang 154 Pari Letrik 164
Pari-kemejan Muncung Pendek 156 Pari Macan 220
Pari-kemejan Tompok Putih 148 Pari Merah 180
Pari-lang Jalur 248 Pari Mulut Kasar 226
Pari-lang Tompok Putih 246 Pari Mulut Tajam 202
Pari-raja Tompok 172 Pari Naga 196
Pari Air Tawar 208, 210 Pari Nyiru 188
Pari Ayam 252 Pari Nyonya 222
Pari Bakau 194 Pari Pasir 196, 212
Pari Batik 192, 216 Pari Paus 256, 258, 260, 262, 264
Pari Belanda 170, 174, 176 Pari Pucat 190
Malaysian common names index 297
Pari Reben 232, 234 Yu Gigi Cangkuk 78

Pari Riman 220 Yu Gigi Lurus 84
Pari Rimau 200, 214, 216, 220 Yu Giling 114
Pari Serban 164 Yu Insang Enam 36
Pari Sungai 198, 208, 210 Yu Jalur Putih 88
Pari Susun 252 Yu Jaras 74, 76
Pari Tanjung 188, 218, 220, 222 Yu Jereh 98, 102, 104, 130
Pari Tembikar 238, 240 Yu Jerung 64, 100
Pendek 64 Yu Kebut 44, 58
Yu Kembong 70
S
Yu Kemejan 146, 154, 158
Sirip Lekuk 264
Yu Kepak Hitam 102, 106, 112
T Yu Kia-kia 148, 150, 152, 154
Tanpa Duri 262 Yu Kulit Tebal 108
Tuka-tuka 218 Yu Mata Jauh 134, 136
Yu Mata Lekuk 122
Y Yu Minyak 38, 40
Yu 134 Yu Misai 46
Yu-buntal Sarawak 70 Yu Muncung Keras 104
Yu-cicak Bintik 52 Yu Muncung Minyak 128
Yu-cicak Insang Puih 54 Yu Muncung Panjang 94
Yu-cicak Lampai 48 Yu Muncung Susu 126
Yu-cicak Tembaga 50 Yu Nipah 106
Yu-gergaji Seragam 142 Yu Padi 130
Yu-gergaji Silih 144 Yu Palang 134, 136
Yu-jaras Bintik 74 Yu Parang 136, 138, 140, 142, 144
Yu-tukul Palang 134 Yu Parangan 140, 142, 144
Yu-tukul Sanggul 136 Yu Pari 42, 156
Yu Bengkong 136 Yu Pasir 50, 80, 96, 98, 104, 110, 130
Yu Bintik Putih 80 Yu Paus 60
Yu Bodoh 48, 50, 52, 54, 56, 72 Yu Pendek 68
Yu Borneo 92 Yu Punai 54
Yu Buncit 70 Yu Putih 82
Yu Buntal 70 Yu Rimau 58
Yu Cicak 58, 68 Yu Sanggul 134, 136
Yu Ekor Hitam 90 Yu Semilang 56
Yu Ekor Panjang 62 Yu Sirip Hitam 106
Yu Gergaji 140, 142, 144 Yu Sirip Lebar 120
Yu Gergaji Jarang 140 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
SHARKS AND RAYS OF BORNEO

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.
Peter R. Last 4
William T. White 4
Janine N. Caira 4
Dharmadi 4
Fahmi 4
Kirsten Jensen 4
Annie P. K. Lim 4
John J. Pogonoski 4
John D. Stevens 4

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SHARKS and RAYS of

SHARKS AND RAYS OF BORNEO

The National Library of Australia Cataloguing-in-Publication entry

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

Produced and distributed by

Cover design by Louise Bell

Early work on Borneo fishes

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

National Science Foundation project

Characteristics of the fauna

Higher taxa NW Philippines Indonesia Borneo

Family NW Australia Philippines Indonesia Borneo

Aims of this book

Habitat type NW Philippines Indonesia Borneo

How to use this book

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

Images and illustrations

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

Common and scientific names

Habitat and biology

acute – sharp or pointed.

transferring sperm to female during mating.

dorsal – relating to upper part or surface of back.

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

mouth – opening through which food enters the alimentary canal.

plain – uniformly coloured, without a contrasting colour pattern.

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

1st dorsal fin

free rear tip

shark dorsal fin

barbel circumnarial groove

lower labial furrow

shark undersurface of head

upper origin subterminal margin

upper postventral margin

lower origin lower postventral margin

structural features of rays

ray nostrils and mouth (oronasal)

1st dorsal fin

eye 2nd dorsal fin

structural features of chimaerids

Key to families and genera

1. Single gill opening on each side of head

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

3. Gill openings partly on sides of head

4. Snout saw-like, flattened, armed with fig. 3

Rostral teeth extending onto basal quarter

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

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

6. Caudal fin with a well developed

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

8. Snout triangular, forming an acute angle

Anterior nasal aperture almost rectangular,

Snout broadly rounded (fig. 14); body

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

10. Pelvic fin divided into two distinct lobes

11. Entire dorsal surface smooth (except fig. 17 fig. 18

12. Anterior part of head not extending

dorsally and well inward from disc margin

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

b. No stinging spine; many long, sharp

c. Disc oval or subcircular; ventral skin fold

Disc more or less quadrangular; ventral

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