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.

SHARKS AND RAYS OF BORNEO

Sharks and Rays of Borneo

SHARKS and RAYS of

BORNEO

Peter R. Last 4 William T. White 4 Janine N. Caira 4 Dharmadi 4 Fahmi 4 Kirsten Jensen 4 Annie P. K. Lim 4 B. Mabel Manjaji-Matsumoto 4 Gavin J. P. Naylor 4 John J. Pogonoski 4 John D. Stevens 4 Gordon K. Yearsley 4

SHARKS and RAYS of

BORNEO Peter R. Last 4 William T. White 4 Janine N. Caira 4 Dharmadi 4 Fahmi 4 Kirsten Jensen 4 Annie P. K. Lim 4 B. Mabel Manjaji-Matsumoto 4 Gavin J. P. Naylor 4 John J. Pogonoski 4 John D. Stevens 4 Gordon K. Yearsley 4

© CSIRO 2010 All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact the author for all permission requests. The National Library of Australia Cataloguing-in-Publication entry

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

Sharks and rays of Borneo/ Peter R. Last, William T. White, Janine N. Caira, Dharmadi, Fahmi, Kirsten Jensen, Annie P.K. Lim, B. Mabel Manjaji-Matsumoto, Gavin J.P. Naylor, John J. Pogonoski, John D. Stevens, Gordon K.Yearsley.

9781921605598 (pbk.) 9781921605642 (pdf)

Includes index. Bibliography

Sharks – Borneo. Rays (Fishes) – Borneo.

597.3095983

Produced and distributed by CSIRO PUBLISHING 150 Oxford Street (PO Box 1139) Collingwood VIC 3066 Australia Telephone: +61 3 9662 7666 Local call: 1300 788 000 (Australia only) Fax: +61 3 9662 7555 Email: [email protected] Web site: www.publish.csiro.au 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)�� Rhinoidei Rhinidae (Shark Rays)�� Rhynchobatoidei Rhynchobatidae (Wedgefishes)�� Rhinobatoidei Rhinobatidae (Shovelnose Rays)�� Torpedinoidei Narcinidae (Numbfishes)�� Narkidae (Sleeper Rays)�� Rajoidei Rajidae (Skates)�� Anacanthobatidae (Leg Skates)�� Myliobatoidei Plesiobatidae (Giant Stingarees)�� Dasyatidae (Stingrays)�� Gymnuridae (Butterfly Rays)�� Myliobatidae (Eagle Rays)�� Rhinopteridae (Cownose Rays)�� Mobulidae (Devilrays)�� Chimaeriformes Chimaeridae (Shortnose Chimaeras)�� Acknowledgements�� Images sourced outside Borneo�� References�� Checklist of Borneo sharks and rays�� Scientific names index�� English common names index�� Indonesian common names index�� Malaysian common names index��

140 146 148 154 160 166 170 178 180 182 238 242 252 256 266 268 270 271 277 283 288 293 296

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 contributions to our knowledge of this fascinating fauna. Cartilaginous fishes are relatively diverse in both marine and freshwater environments of Borneo. The region is the type locality for Pristis zijsron, Carcharhinus borneensis, C. sealei, Lamiopsis tephrodes and Sinobatis borneensis, as well as 10 new species discovered recently, i.e. Orectolobus leptolineatus, Cephaloscyllium sarawakensis, Glyphis fowlerae, Rhynchobatus springeri, Okamejei cairae, Okamejei jensenae, Himantura lobistoma, Pastinachus gracilicaudus, Pastinachus solocirostris and Pastinachus stellurostris. Until recently, this fauna was poorly known and accounts by field scientists in the mid-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 SouthEast Asian countries, and over-harvesting of invertebrates such as triton, sea cucumber and giant clam, the area’s coral reefs and associated ecosystems are being increasingly threatened. While there are some Marine Parks in the region, the lack of enforcement reduces their effectiveness. However there are some recent encouraging initiatives. In March 2008, the Asia Development Bank provided a US$1.5 million grant for environmentalists and respective governments to protect the forests and seas of Brunei, Indonesia, Malaysia and the Philippines.

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 elasmobranchs as bycatch include those operating small-scale bottom trawls, trammel and gill nets, and longlines and droplines. In Malaysia, demersal trawling and drift nets account for the bulk of the catch (DOFM, 2006).

Darwin project In 1996, the first major investigation of the elasmobranchs of Sabah was initiated by an international research team led by Sarah Fowler and supported by funding from the UK Darwin Foundation. This project attempted to answer basic questions relating to the biodiversity, fisheries impact, biology, and conservation status of coastal marine and freshwater elasmobranchs of Sabah, and compile a preliminary checklist of chondrichthyans of greater Borneo. The survey work, which focused on both coastal marine and freshwater habitats, uncovered a relatively rich freshwater fauna. For example, Fowler & Payne (1995) recorded two species of rays, several species of rare sharks and a species of sawfish, from the Kinabatangan River in East Malaysia. A preliminary report on the project was published by Cook & Compagno (1996) providing data on 36 elasmobranch species in markets and freshwater habitats of Sabah. At the completion of the formal survey of elasmobranchs from fish markets, and river and coastal villages throughout Sabah, conducted by the IUCN Shark Specialist Group (see Fowler et al., 1999), 69 species of elasmobranchs (34 sharks and 35 rays) were recorded, 23 of which were new records for the state. Of particular note was the listing of new species of the river shark, genus Glyphis, and new species of Mustelus, Orectolobus and Rhynchobatus. A checklist of chondrichthyan fishes of all of Borneo was compiled at the same time, which included 82 species (40 sharks, 41 rays and 1 chimaera). Following on from this project (1999-2004), a primarily Malaysian led study of the chondrichthyans of Brunei and Malaysia, including Sabah and Sarawak, led to the


collection of 760 specimens of 79 species, comprised of 43 sharks, 35 rays and one chimaera (Yano et al., 2005), but some of these species appear to be invalid. A new and valid species of catshark, Cephaloscyllium sarawakensis, was also described from this research.

Figure 1 Map of Borneo showing the locations of landing sites and villages surveyed. The different landmass colours used correspond to the different countries making up Borneo: grey (Indonesia), green (Malaysia) and yellow (Brunei).

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 development of a resource management plan for elasmobranchs of the region. Several inshore species exhibit regional patterns in distribution that appear to be closely related to habitat availability. For example, several micro-endemic stingrays (genera Himantura and Pastinachus) occur in the environs of the large Kapuas River estuary in western Kalimantan, and the region has additional undocumented, extreme, intraspecific variability in some shark groups.

Characteristics of the fauna The chondrichthyan fauna of Borneo presently contains 118 validated species, including 52 sharks, 65 rays and 1 chimaera. The representation of the group at highest taxonomic levels (Table 1) is similar to regions adjacent (i.e. northwestern Australia, Philippines and Indonesia), but there are some differences in the relative importance of some subgroups. For example, ground sharks (Carcharhiniformes) and eagle rays and their allies (Myliobatoidei) are the dominant subgroups in each region comprising more than 20% of the species; the latter are more dominant in Borneo than any other region. Conversely, dogsharks (Squaliformes) and skates (Rajoidei) are relatively depauperate in Borneo and


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

Higher taxa

NW Australia

Philippines

Indonesia

Borneo

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

179

164

139

118

TOTAL

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

Triakidae

4

4

4

Orectolobidae

3

Centrophoridae

3

4

5

Etmopteridae

3

Squalidae

3

4

4

Urolophidae

3

Pristidae

3 4

4

4

Mobulidae

4

4

Rhinobatidae

4

3

Sphyrnidae

3

3

Hemigaleidae

3

3

Myliobatidae

4

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 IndoMalay Archipelago.


Table 3 Proportional species richness (%) for primary biomes across the region. Habitat type Freshwater/estuarine

NW Australia

Philippines

Indonesia

Borneo

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


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.

12


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).

14


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.

16


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 sawsharks. rostrum (adj. rostral) – a projecting snout. rounded – margin evenly convex. saddle – a blotch extending across dorsal surface from one side to another. salinity – the concentration of salt in water. school – a close aggregation of fish swimming in association with each other. scientific name – the formal binomial name of an organism consisting of the genus and species names; only one valid scientific name exists per species.

18


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

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.

19

20


1st dorsal fin

anterior

posterior

dorsal-fin spine eye

2nd dorsal fin

gill opening

spiracle

precaudal pit

snout tip caudal fin

nostril mouth pelvic fin

labial furrows

anal fin

clasper (males)

structural features of sharks

caudal keel

pectoral fin

anterior margin

apex

spine

posterior margin

origin lobe

free rear tip insertion

inner margin

shark dorsal fin barbel

circumnarial groove

nostril

circumnarial fold

oronasal groove

upper labial furrow

mouth symphysial groove

shark undersurface of head

lower labial furrow

21

Glossary

terminal lobe

posterior tip

upper lobe terminal margin

dorsal margin

subterminal margin

upper origin

subterminal notch upper postventral margin caudal fork lower origin

lower postventral margin

preventral margin

ventral tip lower lobe

shark caudal fin

ventral internasal flap

dorsal rostral cartilage

mouth eye

shoulder

gill opening

spiracle

pectoral fin

cloaca pectoral fin insertion thorn pelvic fin

caudal fin clasper (male)

lateral tail fold 2nd dorsal fin 1st dorsal fin

structural features of rays

stinging spine

22


nostril lobes

internasal flap fringe mouth

papillae

ray nostrils and mouth (oronasal) stinging spine

enlarged thorn

dorsal fold

thorn ventral fold

granulations

ray tail

spine

1st dorsal fin

head clasper (male) mucous canal

lateral line 2nd dorsal fin

eye

caudal-fin upper lobe

mouth gill opening

pre-pelvic clasper (male)

clasper (male) pelvic fin

pectoral fin

structural features of chimaerids

anal fin caudal filament

caudal-fin lower lobe

23

Running head

Key to families and genera 1. Single gill opening on each side of head (fig. 1) .................................................. ............ Family Chimaeridae, Chimaeras fig. 1; 266 5–7 gill openings on each side of head ........................................................... 2

fig. 1

2. Body flattened, ray-like; gill openings on undersurface of head (except in Angel Sharks) ..................................................... 3 Body more or less fusiform, shark-like; gill openings on sides of head ........... 17 3. Gill openings partly on sides of head (fig. 2); pectoral fins very large, clearly detached from head (fig. 2) ................. ......... Family Squatinidae, Angel Sharks fig. 3; 42

fig. 2

Gill openings entirely on undersurface of head; pectoral fins wholly or partly joined to head .................................... 4 4. Snout saw-like, flattened, armed with lateral teeth (fig. 4) ................................. ..................... Family Pristidae, Sawfishes

Rostral teeth absent from basal quarter of saw (fig. 4); lower lobe of caudal fin large, more than half length of upper lobe (fig. 4) .......................... Anoxypristis fig. 4; 140

fig. 3

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

24


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 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. 6

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 ....... .... Family Rhynchobatidae, Wedgefishes fig. 8; 148

fig. 8

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, 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. 11

fig. 13

fig. 12

25

Key to families and genera

9. 2 dorsal fins present (fig. 14); a deep groove around mouth; teeth extending onto outer surfaces of upper and lower jaws in most species ............................... ............. Family Narcinidae, Numbfishes fig. 14; 160

fig. 14

0–1 dorsal fins present (figs 15 and 16); a shallow groove around mouth; teeth not extending onto outer surfaces of upper and lower jaws in most species ............... Family Narkidae, Sleeper Rays

No dorsal fins present (fig. 15) ....... Temera fig. 15; 168

One dorsal fin present (fig. 16) ...... Narke fig. 16; 166

fig. 15

10. Pelvic fin divided into two distinct lobes (fig. 17); no enlarged stinging spine on tail ......................................................... 11

fig. 16

Pelvic fin with only one lobe (fig. 18); usually with 1 or more enlarged, serrated stinging spines (scar visible when spine absent) .............................................. 12 11. Entire dorsal surface smooth (except for alar thorns of male); preorbital snout more than 8 times eye diameter (fig. 19); tail very short and thin, without dorsal fins (fig. 19) .......................................... ..... Family Anacanthobatidae, Leg Skates fig. 19; 178 Thorns or fine denticles present on dorsal surface; preorbital snout much less than 8 times eye diameter (fig. 20); tail slender (but not filamentous), with 2 dorsal fins when undamaged (fig. 20) ..................... ........................... Family Rajidae, Skates fig. 20; 170 12. Anterior part of head not extending forward beyond disc (fig. 21); eyes located

fig. 17

fig. 19

fig. 20

fig. 18

26


dorsally and well inward from disc margin (fig. 21) ............................................ 13 Anterior part of head extending forward beyond disc (fig. 22); eyes located laterally on side of head (fig. 22) ...................... 15 13. Disc very broad, width more than 1.5 times length (fig. 23); tail very short and filamentous (fig. 23) .............................. ...... Family Gymnuridae, Butterfly Rays fig. 23; 238 Disc narrower, width less than 1.5 times length; tail much longer and thicker .... ........................................................ 14

fig. 21

fig. 22

fig. 23

14. Caudal fin present (fig. 24); no thorns or tubercles on disc or tail ......................... ... Family Plesiobatidae, Giant Stingarees fig. 24; 180 Caudal fin absent; central disc and dorsal surface of tail normally with some thorns or tubercles ................................. ................... Family Dasyatidae, Stingrays a. No skin folds on tail; base of tail narrow and typically rounded to slightly compressed in cross-section (fig. 25) ..... b

fig. 24

fig. 25

Skin folds present on undersurface of tail, and sometimes on dorsal surface after sting; base of tail relatively broad, distinctly depressed (fig. 26) .................. c

fig. 27

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

c. Disc oval or subcircular; ventral skin fold tall, extending to tail tip ..................... d

fig. 26

fig. 28

27


Disc more or less quadrangular; ventral skin fold terminating well before tail tip when undamaged ................................ e

d. Disc oval (fig. 29); dorsal surface with numerous blue spots over a brownish background (fig. 29); tail with a blue stripe along each side ................ Taeniura fig. 29; 232

Disc subcircular (fig. 30); dorsal surface with fine black and white mottling (fig. 30); no blue spots or stripes on body or tail ......................................... Taeniurops fig. 30; 234

fig. 29

fig. 30

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 less than half of disc width; ventral skin fold relatively low, its maximum height much less than spiracle length .............. g

fig. 32

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

fig. 34

28


Mouth terminal, located at end of snout tip ............................................... Manta fig. 35; 256

Mouth subterminal, located on undersurface of head .......................... Mobula fig. 36; 258

No lateral, paddle-like processes on head, instead with a pair of broadly rounded lobes (fig. 37) or a single, fleshy subrostral lobe (fig. 38); teeth large and plate-like, fewer than 10 rows in each jaw ......... 16

fig. 35

fig. 36

16. Margin of subrostral lobe with a deep central notch (fig. 37); floor of mouth without papillae .................................... ... Family Rhinopteridae, Cownose Rays fig. 37; 252

fig. 37

Margin of subrostral lobe rounded (fig. 38); floor of mouth with fleshy papillae .......... Family Myliobatidae, Eagle Rays

Posterior margin of internasal flap deeply notched (inverted V-shape) (fig. 39); teeth in a single row in each jaw ............................................... Aetobatus fig. 38; 242

Posterior margin of internasal flap almost straight or only undulating slightly (fig. 40); teeth in more than one row in each jaw ............................. Aetomylaeus fig. 41; 246

17. A single dorsal fin (fig. 42); 6 or 7 gill openings on each side of head (fig. 42) ............ Family Hexanchidae, Cowsharks fig. 42; 36

fig. 38

fig. 39

fig. 40

fig. 41

2 dorsal fins; 5 gill openings on each side of head ..................................... 18 18. Anal fin absent ................................ 19 Anal fin present, sometimes small or

fig. 42

29


obscure (situated close to caudal fin) ... 20 19. Teeth in both jaws similar in shape and size (fig. 44); no subterminal notch on caudal fin (fig. 43) ............................... ................... Family Squalidae, Dogfishes fig. 43; 38 Teeth in upper jaw differing in shape and/or size to those of lower jaw (fig. 45); strong subterminal notch present (fig. 46) ................................................. ... Family Centrophoridae, Gulper Sharks fig. 46; 40 20. Head very broad, hammer-shaped, with very widely separated eyes (fig. 47) ........ Family Sphyrnidae, Hammerhead Sharks

Lateral blades of head very broad, narrow and wing-like; width of head close to half of total length ................... Eusphyra fig. 47; 134

Lateral blades of head much shorter and not wing-like; width of head less than 40% of total length .................... Sphyrna fig. 48; 136

fig. 43

fig. 44

fig. 45

fig. 46

fig. 47

Head much narrower, not hammershaped .............................................. 21 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

30


....... Family Heterodontidae, Hornsharks fig. 50; 44 Dorsal-fin spines absent; no crest over eye ................................................... 23

fig. 51

23. Whole mouth forward of front margin of eye (fig. 51) ................................... 24 Mouth partly beneath or behind front margin of eye (fig. 52) ....................... 28 24. Mouth very broad, terminal; caudal fin forked (fig. 53) ....................................... ... Family Rhincodontidae,Whale Sharks fig. 53; 60 Mouth smaller, subterminal; caudal fin not forked ...................................... 25

fig. 52

fig. 53

25. No fleshy lobe or groove on outer margin of nostril (fig. 54) .................. 26 Fleshy lobe and groove present on outer margin of nostril (fig. 55) ................ 27

fig. 54

26. Caudal fin very long, about equal to trunk length (fig. 56); ridges present along sides of body (fig. 56) ........................... ... Family Stegostomatidae, Zebra Sharks fig. 56; 58 Caudal fin shorter, less than half trunk length (fig. 57); no ridges on body (fig. 57) ..... Family Ginglymostomatidae, Nurse Sharks fig. 57; 56

fig. 56

fig. 57

27. Head and body strongly depressed (fig. 58); skin flaps present on sides of head (fig. 58) ......................................... ....... Family Orectolobidae,Wobbegongs fig. 58; 46 Head and body cylindrical (fig. 59); no skin flaps on sides of head (fig. 59) ........

fig. 55

fig. 58

fig. 59

31


.......................... Family Hemiscylliidae, Longtail Carpetsharks fig. 59; 48 28. Caudal fin almost symmetrical, lunate (fig. 60); a strong keel on either side of caudal peduncle (fig. 60) ....................... ........ Family Lamnidae, Mackerel Sharks fig. 60; 64

fig. 60

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 than 3 times spiracle diameter, mostly extending to level of front of eye (fig. 62) ........................................................... b

Upper labial furrows much shorter or rudimentary, when present much less than 3 times spiracle diameter, never extending to level of front of eye (fig. 63) ............ c

fig. 62

fig. 63

fig. 64

b. Anterior nasal flaps greatly expanded, reaching mouth (fig. 65); nasoral grooves present ............................... Atelomycterus fig. 64; 68

Anterior nasal flaps much shorter, smaller than nostrils, not reaching mouth (fig. 66); nasoral grooves absent ............. Apristurus fig. 67; 66

c. 2nd dorsal fin much smaller than 1st dorsal fin (fig. 68); origin of 2nd dorsal fin only slightly behind origin of anal fin (fig. 68); stomach inflatable .................... ....................................... Cephaloscyllium fig. 68; 70

Dorsal fins about equal in size (fig. 69); origin of 2nd dorsal fin behind midbase of anal fin (fig. 69); stomach not inflatable ................................ Halaelurus fig. 69; 72

fig. 65

fig. 67

fig. 68

fig. 69

fig. 66

32


1st dorsal-fin origin well in front of pelvic-fin base .................................. 30 30. Precaudal pits absent (fig. 71); anterior margin of caudal fin smooth (fig. 71) ............ Family Triakidae, Hound Sharks fig. 70; 74

fig. 70

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

b. Pelvic, dorsal and caudal fins not falcate (fig. 76); mouth long, narrowly arched; lower teeth near symphysis with erect cusps and slightly arched roots; cusplets present on lower teeth……….. Paragaleus fig. 76; 84

Pelvic, dorsal and caudal fins strongly falcate (fig. 77); mouth very short, broadly arched; lower teeth near symphysis with erect cusps and highly arched roots; no cusplets on lower teeth ........... Hemigaleus fig. 77; 80

c. Snout obtusely wedge-shaped in dorsoventral view; fins not falcate, posterior margins of pelvic and pectoral fins straight or slightly concave (fig. 78) ....... .......................................... Chaenogaleus fig. 78; 78

fig. 74

fig. 76

fig. 77

fig. 78

fig. 75

33


Snout bluntly rounded in dorsoventral view; fins strongly falcate, posterior margins of pelvic and pectoral fins deeply concave (fig. 79) ................... Hemipristis fig. 79; 82

Spiracles absent (except in Tiger Shark); posterior margin of 2nd dorsal fin not deeply concave; intestine with a scroll valve (fig. 80) ........................................ ... Family Carcharhinidae, Whaler Sharks a. Caudal peduncle with lateral keels (fig. 81); spiracles present; upper labial furrows, reaching forward to front of eyes; teeth cockscomb-shaped (fig. 82)........... .............................................. Galeocerdo fig. 83; 114

Caudal peduncle without lateral keels (weak ones present in Prionace); spiracles absent; upper labial furrows short, never reaching forward to in front of eyes; teeth not cockscomb-shaped .............. b

fig. 79

fig. 80

fig. 81

fig. 82

fig. 83

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

2nd dorsal fin less than half height of 1st dorsal fin ............................................ e

fig. 84

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

dorsal 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; 120

fig. 85

1st

fig. 87

2nd dorsal fin half to two thirds height of 1st dorsal fin (fig. 88) .................. Glyphis fig. 88; 116

e. Head greatly depressed and trowelshaped (fig. 89); pectoral fins triangular

fig. 88

fig. 86

34


(fig. 89); 1st dorsal-fin free rear tip almost over mid-base of pelvic fin (fig. 89) .................................. Scoliodon fig. 89; 130

Head varying from conical to slightly depressed; pectoral fins narrower; 1st dorsal-fin free rear tip over or slightly anterior to pelvic-fin origin ................. f

fig. 89

fig. 90

f. 2nd dorsal-fin origin well behind anal-fin origin (usually over or just anterior to analfin insertion) (fig. 90); posterior margin of anal fin nearly straight or shallowly concave (fig. 90); preanal ridges very long, subequal to anal-fin base length ... g

2nd dorsal-fin origin usually about opposite anal-fin origin (sometimes more posterior but always well anterior of analfin insertion) (fig. 91); posterior margin of anal fin deeply concave or deeply notched (fig. 91); preanal ridges short, about half anal-fin base or less .............. h

h. Notch present at posterior edge of eye (fig. 93); 1st dorsal-fin origin posterior to adpressed pectoral-fin free rear tips by a distance exceeding 4th gill opening 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 rear tips (fig. 95) …….… Rhizoprionodon fig. 95; 126

i. 1st dorsal-fin base much closer to pelvicfin than pectoral-fin bases (fig. 96); colour brilliant dark blue above;weak keels present on caudal peduncle ......... Prionace fig. 96; 124

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; 86

fig. 91

fig. 92

fig. 93 fig. 94

fig. 95

fig. 96

fig. 97

SPECIES INFORMATION

36


Bluntnose Sixgill Shark Hexanchus griseus (Bonnaterre, 1788) 3 2

4

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, elasmobranchs, cephalopods and crustaceans, and adults also feed on larger bony fishes (e.g. billfishes), cetaceans and seals. Utilisation: Caught rarely so of minimal commercial value. Conservation status: IUCN Red List: Near Threatened. Local synonyms: None. References: Ebert (2003); White & Dharmadi (2010).

37

Hexanchiformes

In Borneo: Probably widespread, but known only from near Layang-layang, off western Sabah.

Ventral head

Upper tooth

Lower tooth

6

Hexanchidae (Cowsharks)

5

38


Western Highfin Spurdog Squalus altipinnis Last, White & Stevens, 2007

4 5

4 3

1 7

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

39

Squaliformes

2

7 Late-term embryo (19 cm)

6

Flank denticle

Squalidae (Dogfishes)

Ventral head

In Borneo: Northern Borneo, from Sabah and northern Sarawak.

40


Smallfin Gulper Shark Centrophorus moluccensis Bleeker, 1860 3 4

3

1

Other names: Endeavour Dogfish (English), Hiu Botol Danten, Hiu Taji, Hiu Senget, Cucut Botol (Indonesian),Yu Minyak (Malaysian) Identifying features: 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). Conservation status: IUCN Red List: Data Deficient. Local synonyms: None. References: White & Dharmadi (2010).

41

Squaliformes

In Borneo: Sparse records from off Sarawak and eastern Sabah. Ventral head

Upper teeth

6 5 Lower teeth Patch of flank denticles

Centrophoridae (Gulper Sharks)

2

42


Ocellated Angelshark Squatina tergocellatoides Chen, 1963

1

6 3

5 8

8

8

7

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

43

Squatiniformes

Utilisation: Caught opportunistically rather than being targeted. Used for its meat. Fins of no commercial value. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None.

Ventral head and abdomen

In Borneo: Off western Sabah and

Sarawak.

2

4 Front view of head

Anterior nasal flap (from front)

Squatinidae (Angelsharks)

References: None.

44


Zebra Hornshark Heterodontus zebra (Gray, 1831)

1 1 2 3

Other names: Zebra Bullhead Shark, Zebra Port Jackson Shark, Bullhead Shark (English), Hiu (Indonesian),Yu Kebut (Malaysian) Identifying features: 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.

45

Heterodontiformes

Conservation status: IUCN Red List: Least Concern. Local synonyms: Cestracion amboinensis Regan, 1906. References: None.

4

Ventral head In Borneo: Northern Borneo, includ-

ing Sarawak and Sabah.

6

Anterior upper tooth

7

Posterior upper tooth

Heterodontidae (Hornsharks)

5

46


Indonesian Wobbegong Orectolobus leptolineatus Last, Pogonoski & White, 2010

1 4 5

2 3

Other names: Indo Wobbegong (English), Hiu Kodok, Hiu Lepang, Hiu Jenggot (Indonesian),Yu Misai (Malaysian) Identifying features: 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).

47

Orectolobiformes

2

Ventral head

Embryo (14 cm)

In Borneo: Known from a few specimens collected off southeastern Sabah (Semporna) and Sarawak.

Orectolobidae (Wobbegongs)

3

48


Indonesian Bambooshark Chiloscyllium hasseltii Bleeker, 1852

4

1 6

4 3 1

5

2

Other names: Hasselt’s Bambooshark (English), Hiu Tekek (Indonesian),Yu-cicak Lampai,Yu Bodoh (Malaysian) Identifying features: 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 bottomdwelling invertebrates, and possibly small fishes. Utilisation: Caught occasionally by gillnet, longline and trawl fisheries. Used for its meat. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Chiloscyllium obscurum Gray, 1851; Chiloscyllium dolganovi Kharin, 1987; Chiloscyllium indicum var. obscura Günther, 1870; Chiloscyllium hasselti Bleeker: Compagno (2001), Compagno et al. (2005a),Yano et al. (2005). References: Compagno (2001).

Ventral head

In Borneo: Central western Borneo, including northwestern Kalimantan and Sarawak.

6

Dorsal head and pectoral fins

7

Juvenile (34.5 cm)

Hemiscylliidae (Longtail Carpetsharks)

49

Orectolobiformes

50


Slender Bambooshark Chiloscyllium indicum (Gmelin, 1789) 4

1 6

4

3 1

5

2

Other names: Ridgeback Bambooshark, Ridgeback Shark (English), Hiu Bongo, Cucut Dolok, Hiu Tekek (Indonesian),Yu-cicak Tembaga,Yu Bodoh,Yu Tokeh,Yu Pasir (Malaysian) Identifying features: 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 4 posterior margins of dorsal fins straight to convex 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 Size: To at least 65 cm; males and females mature at about 39 and 43 cm respectively; 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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Chiloscyllium phymatodes Bleeker, 1852. References: Compagno (2001).

In Borneo: Patchy; recorded from southern and western Kalimantan and Sarawak.

Ventral head

3 6


7

Juvenile (19 cm)


51

Orectolobiformes

52


Whitespotted Bambooshark Chiloscyllium plagiosum (Bennett, 1830)

4

4

1 6

3 1

5

2

Other names: None (English), Hiu Bongo, Cucut Dolok, Hiu Tekek (Indonesian),Yucicak Bintik,Yu Bodoh (Malaysian) Identifying features: 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 4 posterior margins of dorsal fins straight to convex 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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Chiloscyllium plagiosum var. interruptum Bleeker, 1852; Chiloscyllium margaritiferum Bleeker, 1863. References: Compagno (2001); Chen et al. (2007).

Ventral head

In Borneo: South China Sea, from northwestern Kalimantan to northern Sabah.

6


6

Juvenile (13 cm)


53

Orectolobiformes

54


Brownbanded Bambooshark Chiloscyllium punctatum Müller & Henle, 1838

1

4 6

4

3 1

5

2

Other names: Grey Carpetshark, Brownbanded Catshark, Spotted Catshark (English), Hiu Bongo, Hiu Batu, Hiu Gedok, Hiu Tekek (Indonesian),Yu-cicak Insang Putih,Yu Bodoh,Yu Tokeh,Yu Punai (Malaysian) Identifying features: 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. Conservation status: IUCN Red List: Near Threatened.

55

Orectolobiformes

Local synonyms: None.

Ventral head

In Borneo: Southern and western Kalimantan, Sarawak and Sabah.


7 Sub-adult (51 cm)


References: Compagno (2001).

56


Tawny Nurse Shark Nebrius ferrugineus (Lesson, 1830)

4 3

4

1

5

Other names: Tawny Shark, Spitting Shark, Sleepy Shark, Madame X (English), Hiu Gedebong, Hiu Gedok, Hiu Bisu (Indonesian),Yu Semilang,Yu Bodoh (Malaysian) Identifying features: 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 Size: To at least 320 cm; males and females mature at about 225 and 230 cm respectively; 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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Ginglymostoma rueppellii Bleeker, 1852; Nebrius concolor Rüppell 1837. References: None.

57

Orectolobiformes

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 from dive operator in Kota Kinabalu.

Upper tooth

Lower tooth

4 4

Lateral precaudal tail region

Ginglymostomatidae (Nurse Sharks)

2

58


Zebra Shark Stegostoma fasciatum (Hermann, 1783) 3 5 1

4

Other names: Leopard Shark (English), Hiu Belimbing, Kluyu Blimbingan (Indonesian),Yu Rimau,Yu Kebut,Yu Cicak,Yu Tokeh (Malaysian) Identifying features: 1 colour pattern yellowish brown with numerous, dark brown spots in adults 2 dark brown with vertical white bars and spots in juveniles (smaller than 70 cm) 3 caudal fin very long, blade-like 4 prominent ridges on dorsal surface and sides 5 spiracle similar in size to eye Size: To at least 235 cm (reports of 354 cm need validation); males and females mature at about 145–185 and 170 cm respectively; hatch at about 20–36 cm. Distribution: Tropical Indo–West Pacific, from southeastern Africa to New Caledonia and Japan. Habitat and biology: A common inshore, demersal shark found over coral reefs and soft bottoms in shallow waters to at least 40 m. Oviparous; their large egg cases (13–17 cm long, 8 cm wide), which are dark brown to purplish black with longitudinal striations, are anchored to the substrate by lateral masses of hair-like fibres. Diet consists primarily of gastropods and bivalves, but also crustaceans and small fishes. Active mostly at night, resting on the bottom during day. Utilisation: Used primarily for its meat and skin (exported to Thailand for leather). Protected under the Malaysian Fisheries Act, so catches are opportunistic and landings not reported. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Squalus pantherinus Kuhl & van Hasselt in Bleeker, 1852 (name

59

Orectolobiformes

mentioned in synonmy with S. fasciatum); Scyllia quinquecornuatum van Hasselt, 1823; Squalus tigrinus Forster, 1781; Stegostoma varium Garman, 1913.

Ventral head

Upper tooth

In Borneo: Western and northern Borneo, including Sarawak, Sabah and north-west and north-east Kalimantan.

Lower tooth

2

Juvenile (49.5 cm)

Stegostomatidae (Zebra Sharks)

References: None.

60


Whale Shark Rhincodon typus Smith, 1828

1

6

5

4

3

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

61

Orectolobiformes

Local synonyms: Rhiniodon typus Smith, 1829. References: None.

1

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

Rhincodontidae (Whale Sharks)

3

62


Pelagic Thresher Alopias pelagicus Nakamura, 1935

1 3

2 4 5

Other names: None (English), Hiu Monyet, Hiu Tikus, Cucut Pedang, Tikusan (Indonesian),Yu Ekor Panjang (Malaysian) Identifying features: 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.

63

Lamniformes

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None.

In Borneo: Uncommon, South China Sea off Sabah and also the Natuna Islands off Sarawak; possibly more widespread. Ventral head

Upper tooth

Lower tooth

Alopiidae (Thresher Sharks)

References: White (2007b).

64


Shortfin Mako Isurus oxyrinchus Rafinesque, 1810

3

1

2

Other names: Mako Shark, Blue Pointer, Mackerel Shark, Snapper Shark (English), Hiu Tenggiri, Hiu Anjing, Hiu Mako, Hiu Kakap (Indonesian), Jerung-mako Sirip Pendek,Yu Jerung (Malaysian) Identifying features: 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 Size: To at least 390 cm; males and females mature at about 185–195 and 250–280 cm 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.

65

Lamniformes

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Oxyrhina glauca Müller & Henle, 1839. References: White (2007b).

5

In Borneo: Only recorded off Sabah, but almost certainly more widespread. Ventral head

Upper tooth

5

Lower tooth

Lamnidae (Mackerel Sharks)

4

66


Bigfin Catshark Apristurus platyrhynchus (Tanaka, 1909)

3 6

4 5 2

1

Other names: Spatulasnout Catshark (English), none (Indonesian), none (Malaysian) Identifying features: 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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Pentanchus verweyi Fowler, 1934; Apristurus verweyi (Fowler). References: Kawauchi et al. (2008).

67

Carcharhiniformes

8

In Borneo: Known only from off Sipadan Island, in the Celebes Sea (eastern Sabah). Ventral head

Flank denticles

Scyliorhinidae (Catsharks)

7

68


Coral Catshark Atelomycterus marmoratus (Bennett, 1830) 1 3

1

2

Other names: None (English), Hiu Tokek, Cucut Tokek (Indonesian),Yu Cicak,Yu Tokeh,Yu Pendek (Malaysian) Identifying features: 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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Scyllium marmoratum Bennett, 1830. References: Bor et al. (2003); White (2007a).

69

Carcharhiniformes

5

Ventral head


In Borneo: Coastal waters of Sabah and Sarawak.


4

70


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) Identifying features: 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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Cephaloscyllium circulopullum Yano, Ahmad & Gambang, 2005; Cephaloscyllium parvum Inoue & Nakaya, 2006. References: Schaaf-Da Silva & Ebert (2008).

71

Carcharhiniformes

5

4

In Borneo: South China Sea, off Sarawak.

Egg case



Ventral head

72


Indonesian Speckled Catshark Halaelurus maculosus White, Last & Stevens, 2007

1 1 2 3

4

Other names: None (English), Hiu Tokek (Indonesian),Yu Bodoh (Malaysian) Identifying features: 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. Conservation status: IUCN Red List: Least Concern. 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).

Carcharhiniformes

73

6

5

In Borneo: Known only from off Sarawak.

Egg case

Dorsal head and anterior trunk


Ventral head

74


Starspotted Smoothhound Mustelus manazo Bleeker, 1854 1 4 5

3

2

Other names: Gummy Shark, Japanese Smoothhound (English), Hiu Kacang, Hiu Air, Cucut Londer (Indonesian),Yu-jaras Bintik,Yu Jaras (Malaysian) Identifying features: 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. Conservation status: IUCN Red List: Data Deficient.

75

Carcharhiniformes

Local synonyms: Mustelus mosis Hemprich & Ehrenberg: Yano et al. (2005). References: None.

8 7 In Borneo: Known from the South China Sea, off Sarawak and Sabah. Ventral head

Triakidae (Houndsharks)

6

76


Whitefin Smoothhound Mustelus widodoi White & Last, 2006 1 4 5

2

2

3

Other names: None (English), Hiu Kacang, Hiu Air, Cucut Londer (Indonesian),Yu Jaras (Malaysian) Identifying features: 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 midcontinental shelf to upper slope in depths greater than 60 m. Reproductive strategy unknown; either viviparous, with histotrophy or viviparous, with a yolk sac-placenta as in other members of the genus. Diet unknown, presumably consisting of small fishes and invertebrates as in other members of the genus. Utilisation: Marketed infrequently, probably more abundant in the deeper parts of the South China Sea. Used for its meat and fins, but of limited use due to its rarity. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Mustelus sp. 1: White et al. (2006b). References: White & Last (2006); White (2007a).

Carcharhiniformes

77

6

8

In Borneo: Known only from northwestern Sabah. Ventral head

Triakidae (Houndsharks)

7

78


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) Identifying features: 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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None. References: None.

79

Carcharhiniformes

4

In Borneo: Uncommon, recorded from Sarawak, Sabah and northeastern Kalimantan, but probably more widely distributed. Ventral head

Upper tooth

Lower tooth

6

Hemigaleidae (Weasel Sharks)

5

80


Sicklefin Weasel Shark Hemigaleus microstoma Bleeker, 1852

3 3 1 2 3

3

Other names: Weasel Shark (English), Hiu Kacang, Hiu Pilus (Indonesian),Yu Bintik Putih,Yu Pasir (Malaysian) Identifying features: 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) Size: To at least 114 cm; males and females mature at about 74 and 78 cm respectively; born at about 45 cm. Distribution: Tropical Indo–West Pacific, from the Red Sea to Indonesia and north to China, excluding Australia and New Guinea. Habitat and biology: Occurs on or near the bottom on the continental shelf. Viviparous, with a yolk-sac placenta; gives birth possibly twice a year to litters of 2–4 pups after a gestation period of probably less than 6 months. Diet largely unknown, but a few stomachs examined contained cephalopods. Utilisation: Often caught by inshore gillnet, bottom trawl and, to a lesser extent, longline fisheries. Used for its meat and fins, but of limited value due to its small size. Conservation status: IUCN Red List: Vulnerable.

81

Carcharhiniformes

Local synonyms: Hemigaleus machlani Herre, 1929; Carcharias (Hypoprion) notatus Bleeker, 1850.

4 5 In Borneo: Widespread. Ventral head

Upper tooth

6

7 Lower tooth Pectoral fin - ventral


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

82


Fossil Shark Hemipristis elongata (Klunzinger, 1871)

3 3 1

2 3 3

Other names: Snaggletooth Shark (English), Hiu Monas (Indonesian), Jerung Gigi Cerakah,Yu Putih (Malaysian) Identifying features: 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 cephalopods and fishes, including small elasmobranchs. Utilisation: Caught often by inshore gillnet, bottom trawl and longline fisheries. Used for its meat (good quality), fins and cartilage. Population declines have been reported in some regions. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Chaenogaleus macrostoma (Bleeker): White et al. (2006b); Hemipristis elongatus (Klunzinger): Yano et al. (2005).

83

Carcharhiniformes

References: White (2007a).

5

In Borneo: Widespread. Ventral head

Upper tooth

6 Lower tooth Pectoral fin - ventral

Juvenile (63 cm)


4

84


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) Identifying features: 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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Negogaleus longicaudatus Bessednov, 1966. References: None.

85

Carcharhiniformes

4

5

In Borneo: Off Sabah, Sarawak and eastern Kalimantan.

Ventral head

Upper tooth

7

8 Lower tooth


6

86


Silvertip Shark Carcharhinus albimarginatus (Rüppell, 1837) 3 1 1 2

1 1 1

Other names: None (English), Cucut Lanjaman, Hiu Lanyam, Hiu Plen, Hiu Sonteng (Indonesian), none (Malaysian) Identifying features: 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 Size: To at least 275 cm; males and females mature at about 170 and 195 cm respectively; 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. Conservation status: IUCN Red List: Near Threatened.

87

Carcharhiniformes

Local synonyms: None. References: None.

In Borneo: Few confirmed local records, but observed by divers at the Spratly Islands in the South China Sea; probably also common around coral reefs off eastern Sabah.

Ventral head

Upper tooth

Lower tooth

5

6

Carcharhinidae (Whaler Sharks)

4

88


Graceful Shark Carcharhinus amblyrhynchoides (Whitley, 1934) 1

2

1

1

1 1

Other names: Queensland Shark (English), Cucut Lanjaman, Hiu Bujit (Indonesian), Yu Jalur Putih (Malaysian) Identifying features: 1 all fin tips usually black or dusky (except anal fin) 2 interdorsal ridge absent 3 snout rather short, narrowly rounded to pointed (viewed ventrally), internarial space 1–1.2 times preoral snout length 4 upper and lower teeth with a slender, erect cusp, no basal cusplets 5 precaudal vertebrae usually less than 82 Size: To at least 178 cm; both sexes mature at about 110–115 cm; born at 50–60 cm. Distribution: Tropical Indo–West Pacific, from the Gulf of Aden to northern Australia and the Philippines. Habitat and biology: Primarily pelagic in midwater over insular shelves, from close inshore to bottom depths of at least 50 m.Viviparous, with a yolk-sac placenta; gives birth to litters of an average of 3 pups (maximum 8) after a 9–10 month gestation period. Diet consists primarily of fish, with smaller amounts of crustaceans and cephalopods. Not dangerous to humans. Utilisation: Caught mainly as bycatch by longlining. Used for its fins and meat, but typically only small in size. Conservation status: IUCN Red List: Near Threatened.

89

Carcharhiniformes

Local synonyms: None. References: White (2007c); Ahmad et al. (2008).

In Borneo: Widespread. Ventral head

Upper tooth

4

Lower tooth


3

90


Grey Reef Shark Carcharhinus amblyrhynchos (Bleeker, 1856)

3 2

1

Other names: Longnose Blacktail Shark (English), Cucut Lanjaman, Hiu Bujit, Merak Bulu, Hiu Lonjor (Indonesian),Yu Ekor Hitam (Malaysian) Identifying features: 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 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. Conservation status: IUCN Red List: Near Threatened.

111

Carcharhiniformes

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


4

112


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) Identifying features: 1 2nd dorsal, pectoral and ventral tip of caudal fin strikingly black-tipped 2 interdorsal ridge present 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 respectively; born at 50–55 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. Conservation status: IUCN Red List: Near Threatened.

113

Carcharhiniformes

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.

In Borneo: Widespread.

Ventral head

Upper tooth

5

Lower tooth

6


4

114


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) Identifying features: 1 dark, vertical bars on sides (less distinct in adults) 2 spiracle present, small and slit-like 3 caudal peduncle with a low, rounded, lateral keel 4 snout very short and bluntly rounded (viewed ventrally) 5 upper labial furrows very long, subequal to preoral length 6 teeth in both jaws heavily serrated, cockscomb-shaped, one edge deeply notched, the other convex Size: To at least 600 cm (one record of 740 cm); males and females mature at 300–305 and 250–350 cm respectively; born at about 51–76 cm. Distribution: Cosmopolitan in all tropical seas, making seasonal incursions into warm temperate areas. Habitat and biology: Occurs close inshore to the outer continental shelf, including the intertidal zone and surface, to at least 150 m depth. The only non-placental whaler shark; viviparous, with histotrophy; gives birth to litters of an average of 33 pups (maximum 82) after a 12–16 month gestation period. Males and females mature at 7–8 and 7–12 years respectively. Omnivorous, feeding on a broad variety of prey including fishes, elasmobranchs, dugongs, birds, turtles, snakes, dolphins, birds, crustaceans and cephalopods. Potentially very dangerous to humans given its large size and presence close inshore, but normally not aggressive. Probably belongs to its own family of sharks.

115

Carcharhiniformes

Utilisation: Caught occasionally by the shark longline, tangle net and bottom trawl fisheries.Valuable for its fins, meat, skin, jaws and cartilage. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Carcharias (Prionodon) fasciatus Bleeker, 1852; Galeocerdo fasciatus van Kampen, 1907.

4

5

In Borneo: Almost certainly widespread; current records are from off Sarawak, Sabah and eastern Kalimantan.

Ventral head

Upper tooth

6

Lower tooth


References: None.

116


Mukah River Shark Glyphis sp.

5 4

2

6 1

5 5

Other names: None (English), none (Indonesian),Yu Sungai (Malaysian) Identifying features: 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 viviparous, with yolk-sac placenta. Diet also unknown, presumably primarily bony fishes. Conservation status needs investigating, likely to be highly threatened. Utilisation: Caught rarely. Probably utilised for its meat and fins where caught. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: None.

117

Carcharhiniformes

References: Fahmi & Adrim (2009); Compagno et al. (2010).

3

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.


7

118


Borneo River Shark Glyphis fowlerae Compagno, White & Cavanagh, 2010

4 3

2

6 1

7 5

5

Other names: None (English), none (Indonesian),Yu Sungai (Malaysian) Identifying features: 1 precaudal pits longitudinal (not crescentic) 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

119

Carcharhiniformes

lines and gill nets in the Kinabatangan River. Utilised for its meat. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: Glyphis sp. B: Compagno & Niem (1998); Compagno et al. (2005); Glyphis sp.: Yano et al. (2005).

In Borneo: Recorded only from the Kinabatangan River, northeastern Sabah. Ventral head

3 7

Upper tooth

Lateral head

Lower tooth

8


References: Compagno et al. (2010).

120


Borneo Broadfin Shark Lamiopsis tephrodes (Fowler, 1905)

3

2 1

4

Other names: None (English), Hiu Bujit (Indonesian),Yu Sirip Lebar (Malaysian) Identifying features: 1 precaudal pits longitudinal (not crescentic) 2 2nd dorsal fin very tall, similar to height of 1st dorsal fin 3 interdorsal ridge absent 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 Size: To at least 157 cm; males and females mature at about 114 and 130 cm respectively; 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. Conservation status: IUCN Red List: Not Evaluated.

121

Carcharhiniformes

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


5

122


Sliteye Shark Loxodon macrorhinus Müller & Henle, 1839

3 1

2 5

Other names: Jordan’s Blue Dogshark, Slender Dogshark (English), Hiu Kejen (Indonesian),Yu Mata Lekuk (Malaysian) Identifying features: 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 Size: To at least 99 cm; males and females mature at about 80–83 and 80–90 cm respectively; born at 54–55 cm. Distribution: Tropical waters of the Indo–West Pacific, from southeastern Africa to Australia and southern Japan. Habitat and biology: A common species occurring near the bottom in shallow, clear, inshore waters to at least 100 m depth.Viviparous, with a yolk-sac placenta; gives birth each year to litters of 2 pups (maximum 4); no reproductive seasonality. Diet consists primarily of small fishes and crustaceans, as well as cephalopods. Harmless to humans. Utilisation: Caught frequently by inshore demersal gillnet fisheries. Used for its fins and meat, but of limited value due to its small size. Conservation status: IUCN Red List: Least Concern. Local synonyms: Carcharias (Scoliodon) dumerilii Bleeker, 1856. References: None.

123

Carcharhiniformes

4

In Borneo: Mainly northern Borneo; off Sarawak, Sabah, and eastern and western Kalimantan. Ventral head

Upper tooth

1

7 Eye Lower tooth


6

124


Blue Shark Prionace glauca (Linnaeus, 1758)

4

1

3

2

Other names: Blue Whaler, Great Blue Shark (English), Hiu Aer, Hiu Lalaek, Hiu Karet, Cucut Selendang (Indonesian), Jerung Biru (Malaysian) Identifying features: 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.

125

Carcharhiniformes

Conservation status: IUCN Red List: Near Threatened. Local synonyms: None. References: None.

In Borneo: Known from a single record off Sabah; probably more widespread.

Ventral head

Upper tooth

Lower tooth

6

7


5

126


Milk Shark Rhizoprionodon acutus (Rüppell, 1837)

1

2

Other names: Longman’s Dogshark, Fish Shark, White-eye Shark (English), Hiu Pilus, Hiu Plen, Mungsing, Hiu Pisang (Indonesian),Yu Muncung Susu (Malaysian) Identifying features: 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 and meat, but of limited value due to its small size.

127

Carcharhiniformes

Conservation status: IUCN Red List: Least Concern. Local synonyms: Carcharias (Scoliodon) walbeehmi Bleeker, 1856; Carcharias (Prionodon) sorrahkowah Bleeker, 1854; Carcharhinus borneensis (Bleeker):Yano et al. (2005). References: None.

4

In Borneo: Mainly northern Borneo; from Sabah, Sarawak, and eastern and western Kalimantan.

5

Ventral head

Upper tooth

6

Lower tooth


3

128


Grey Sharpnose Shark Rhizoprionodon oligolinx Springer, 1964

1

2

Other names: None (English), Hiu Pilus, Hiu Pisang (Indonesian),Yu Muncung Minyak (Malaysian) Identifying features: 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 respectively; born at 21–26 cm. Distribution: Tropical Indo–West Pacific, from the Red Sea to Indonesia and southern Japan (only a single validated record from the Gulf of Carpentaria in northern Australia). Habitat and biology: A small, common, inshore shark, schooling close inshore to at least 36 m depth.Viviparous, with a yolk-sac placenta; gives birth to litters of 3–5 pups after an unknown gestation period. Diet probably consists primarily of small fishes. Utilisation: Caught by inshore demersal gillnet fisheries. Used for its fins and meat, but of limited value due to its small size. Conservation status: IUCN Red List: Least Concern.

129

Carcharhiniformes

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

6 Lower tooth


3

130


Pacific Spadenose Shark Scoliodon macrorhynchos (Bleeker, 1852)

2 3

1

5 4

Other names: None (English), Mungsing, Hiu Kejen (Indonesian),Yu Padi,Yu Pasir, Yu Jereh (Malaysian) Identifying features: 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. Conservation status: IUCN Red List: Not Evaluated.

131

Carcharhiniformes

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

6 Lower tooth


1

132


Whitetip Reef Shark Triaenodon obesus (Rüppell, 1837) 1 3

1

2

Other names: Whitetip Shark, Blunthead Shark (English), Hiu Bokem, Hiu Karang Buas, Hiu Coklat (Indonesian),Yu Sirip Putih (Malaysian) Identifying features: 1 1st dorsal and upper lobe of caudal fin with distinctive white tips 2 interdorsal ridge absent 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 respectively; 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. Conservation status: IUCN Red List: Near Threatened.

133

Carcharhiniformes


Ventral head

In Borneo: Mainly northern Borneo; from Sabah, Sarawak, and northwestern and eastern Kalimantan.

Upper tooth

5

Lower tooth


4

134


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) Identifying features: 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 Size: To at least 186 cm; males and females mature at about 108 and 120 cm respectively; 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.

135

Carcharhiniformes

Utilisation: Used for its meat but less important than other hammerheads due to its small average size. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Zygaena latycephala van Hasselt, 1823.

1

4 In Borneo: A few records off Sabah, Sarawak and northwestern Kalimantan but probably widespread.

Ventral head

Upper tooth

6

Lower tooth

Sphyrnidae (Hammerhead Sharks)

References: Stevens & Lyle (1989).

136


Scalloped Hammerhead Sphyrna lewini (Griffith & Smith, 1834)

3 1 4 6 5

Other names: Kidney-headed Shark (English), Hiu Capil, Hiu Bingkoh, Hiu Caping, Hiu Martil, Hiu Parang (Indonesian),Yu-tukul Sanggul,Yu Tukul,Yu Palang,Yu Mata Jauh,Yu Bengkong,Yu Sanggul,Yu Parang (Malaysian) Identifying features: 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. Distribution: Cosmopolitan in all tropical and warm temperate seas. 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,

137

Carcharhiniformes

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. Conservation status: IUCN Red List: Endangered. Local synonyms: Zygaena indica van Hasselt, 1823.

1

2

In Borneo: Probably widely distributed, but few records from southern Kalimantan.

Ventral head

Upper tooth

7

Lower tooth


References: Stevens & Lyle (1989); White et al. (2008).

138


Great Hammerhead Sphyrna mokarran (Rüppell, 1837)

3 4 6 1

5

Other names: None (English), Hiu Capil, Hiu Bingkoh, Hiu Caping, Hiu Martil, Hiu Parang (Indonesian),Yu Tukul,Yu Parang (Malaysian) Identifying features: 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. Distribution: Cosmopolitan in all tropical and warm temperate seas. Habitat and biology: Continental and insular shelves and semi-oceanic, from the surface, and from very shallow water, to at least 80 m depth.Viviparous, with a yolksac placenta; gives birth to litters of 6–42 pups after a gestation period of 11 months. Diet consists primarily of bony fishes and elasmobranchs, but also cephalopods and crustaceans. Potentially dangerous to humans and known to intimidate spear fishers occasionally, but most often timid when observed. Utilisation: Retained for their fins, meat and cartilage. Large population declines

139

Carcharhiniformes

recorded in some areas. Conservation status: IUCN Red List: Endangered. Local synonyms: None.

1

2

In Borneo: Recorded from Sabah and Sarawak, but probably widely distributed.

Ventral head

Upper tooth

7

Lower tooth


References: Stevens & Lyle (1989).

140


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) Identifying features: 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.

141

Pristoidei

Conservation status: IUCN Red List: Critically Endangered; Appendix I of CITES. Local synonyms: None.

2 In Borneo: Single records from southern Kalimantan and near Sandakan, northeastern Sabah.

Caudal fin

6

5

4 Ventral head

Pristidae (Sawfishes)

References: Manjaji (2002a, b).

142


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) Identifying features: 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 4 17–23 pairs of rostral teeth 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

143

Pristoidei

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. Local synonyms: None.

2 Caudal fin

In Borneo: Occurs in the Kinabatangan River; possibly once more widespread in larger rivers and adjacent marine bays of the region.

6

4

5

Ventral head


References: Manjaji (2002a, b).

144


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) Identifying features: 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 4 24–29 pairs of rostral teeth 5 rostral teeth present near base of saw 6 rostral teeth much closer together near tip than at mid-length of saw Size: To at least 540 cm, possibly 730 cm; both sexes mature at about 300 cm; born at about 80 cm. Distribution: Tropical Indo–West Pacific, from southeastern Africa to New Guinea and Australia, north in the Pacific to Vietnam. Habitat and biology: Demersal in coastal bays and estuaries, also well offshore.Viviparous, with yolk-sac dependency; gives birth to litters of about 12 pups. Matures at about 9 years in Australian populations. Diet consists of small schooling fishes and demersal 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. References: Manjaji (2002a, b).

145

Pristoidei

2 In Borneo: Recent records from central Sarawak (near Bintulu), and northeastern (Labuk River) and eastern Sabah (Kinabatangan River); holotype collected in the 19th century from southern Kalimantan.

6

4

5 Ventral head


Caudal fin

146


Shark Ray Rhina ancylostoma Bloch & Schneider, 1801

3 5

1 2

Other names: Bowmouth Guitarfish, Mud Skate (English), Hiu Barong, Kupukupu Brangkas,Yunbun Karang (Indonesian), Pari-kemejan Kepala Keras,Yu Kemejan (Malaysian) Identifying features: 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 dependency; biology poorly known, one pregnant female contained 9 mid-term embryos (27–31 cm). Diet consists primarily of bottom-dwelling crustaceans and molluscs. Handling large adults can be hazardous. They create a nuisance to trawl fishers as they are difficult to remove and their roughness can damage smaller species in the catch.

147

Rhinoidei, Rhynchobatoidei & Rhinobatoidei

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None. References: Devadoss & Batcha (1995); Frimodt (1995).

4

Lateral head In Borneo: Known from coastal areas of Sabah, Sarawak, and western and southern Kalimantan.

3

Ventral head

Rhinidae (Shark Rays)

5

148


Whitespotted Wedgefish Rhynchobatus australiae Whitley, 1939

6

4

2 5

4

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) Identifying features: 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 Size: To at least 270 cm; males and females mature at about 130 and 155 cm respectively; 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


149

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. Local synonyms: None.

6

In Borneo: Mainly western and northern Borneo.

Ventral head

7 Dorsal view of eyes and spiracles

Rhynchobatidae (Wedgefishes)

References: None.

150


Smoothnose Wedgefish Rhynchobatus laevis (Bloch & Schneider, 1801)

4

2

6

4 1

5

8

3

Other names: Giant Guitarfish, Sandshark, Shovelnose Shark (English), Paredung, Pangrum, Hiu Lontar, Liongbun, Petong, Hiu Kemenyan, Hiu Minsong (Indonesian), Pari Kemejan,Yu Kia-kia (Malaysian) Identifying features: 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 7 spiracles with two skin folds on posterior margin 8 caudal fin with a well-developed lower lobe 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.

151


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. Local synonyms: None.

6

In Borneo: Recorded from off northeastern Sabah. Ventral head



References: None.

152


Broadnose Wedgefish Rhynchobatus springeri Compagno & Last, 2010

2

6

4 1

5

4

3

8

Other names: None (English), Paredung, Pangrum, Hiu Lontar, Liongbun, Petong, Hiu Kemenyan, Hiu Minsong (Indonesian), Pari Kemejan,Yu Kia-kia (Malaysian) Identifying features: 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 7 spiracles with two skin folds on posterior margin 8 caudal fin with a well-developed lower lobe 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.

153


Conservation status: IUCN Red List: Not Evaluated; of long-term conservation concern. Local synonyms: None.

6

Ventral head


In Borneo: A few records from brackish habitats of southern Sarawak.


References: Compagno & Last (2010).

154


Clubnose Guitarfish Glaucostegus thouin (Anonymous, 1798)

2 1 3

6

4

Other names: None (English), Cucut Mandrong, Liongbun (Indonesian), Parikemejan Muncung Panjang,Yu Kemejan,Yu Kia-kia (Malaysian) Identifying features: 1 snout tip enlarged to form an elongate knob 2 dorsal surface plain, without blotches or spots 3 denticles along midline of dorsal surface distinct 4 clear part of snout sharply demarcated from rest of head near eyes 5 nostrils broad and slit-like with narrow nasal flaps 6 lower lobe of caudal fin short Size: To about 300 cm; largest observed from the region 240 cm. Distribution: Widespread in the Indo–West Pacific from the Red Sea through Thailand, Vietnam, Malaysia, Indonesia, New Guinea and Japan; apparently not known from Peninsula Malaysia or nearby Philippines. Habitat and biology: Demersal on insular and continental shelves to about 100 m depth; local depth information not reliable, reported from inshore but probably more abundant offshore on the continental shelf. Biology poorly known, presumably viviparous, with yolk-sac dependency. Diet consists of benthic invertebrates and small fishes. Utilisation: Caught infrequently by local demersal tangle net and trawl fisheries. Used opportunistically for its meat and fins (both very high value), as well as skin and cartilage; an esteemed food fish in India. Likely to become a major conservation concern if fished at existing levels. Conservation status: IUCN Red List: Vulnerable.

155


Local synonyms: Rhinobatos thouin (Anonymous [Lacepède]): Yano et al. (2005), White et al. (2006b), also misspelled as Rhinobatos thouini.

5

In Borneo: Known from Sarawak, Sabah and southeastern and western Kalimantan. Ventral head

6 Lateral tail and dorsal fins

Rhinobatidae (Shovelnose Rays)

References: None.

156


Giant Guitarfish Glaucostegus typus (Bennett, 1830)

2

1 4

3 6

Other names: Common Shovelnose Ray, Giant Shovelnose Ray, Shovelnose Ray, Shovelnose Shark (English), Hiu Cermin, Paitpait, Pari Bandrong, Pari Gitar, Pari Kekeh, Petong (Indonesian), Pari-kemejan Muncung Pendek,Yu Pari (Malaysian) Identifying features: 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 6 lower lobe of caudal fin short 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.

157


Marketed for its meat and fins (both very high value), and skins and cartilage; sold fresh and possibly dried and salted. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Rhinobatos typus Bennett: Yano et al. (2005), White et al. (2006b).

5

Ventral head

In Borneo: Primarily from eastern Borneo, also known from Sarawak and western Kalimantan.

6 Lateral tail and dorsal fins


References: None.

158


Taiwan Guitarfish Rhinobatos formosensis Norman, 1926

2 1 3 4

6

Other names: None (English), none (Indonesian),Yu Kemejan (Malaysian) Identifying features: 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 6 lower lobe of caudal fin short 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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Rhinobatos cf. schlegelii Müller & Henle: Compagno et al. (2005b). References: Compagno et al. (2005b).

159


In Borneo: Off Sarawak and Sabah. Ventral head

7

Embryo (19 cm)

1st dorsal fin

2nd dorsal fin


5

160


Shortlip Numbfish Narcine brevilabiata Bessednov, 1966

1

3 2 4 5

Other names: Shortlip Electric Ray (English), Pari Listrik, Pari Kontak (Indonesian), Pari Karan Tompok Besar (Malaysian) Identifying features: 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.

161

To r p e d i n o i d e i

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None.

7

6

Mouth and nostrils In Borneo: Off Sarawak and eastern Sabah.

Juvenile (14 cm)

Narcinidae (Numbfishes)

References: Carvalho (1999).

162


Rough Numbfish Narcine lingula Richardson, 1846

1

5 2

3

4

Other names: Chinese Numbfish, Rough Electric Ray (English), Pari Listrik, Pari Kontak (Indonesian), none (Malaysian) Identifying features: 1 disc subcircular to oval, thick and flabby 2 eyes not embedded in skin 3 two similar-sized dorsal fins 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.

163


Like other members of the genus, the electric organs in the disc are capable of delivering a moderate shock on direct contact. Utilisation: Caught occasionally in the region by bottom trawl. Not considered for food and used for fish meal. Conservation status: IUCN Red List: Data Deficient. Local synonyms: None.

7 6

Mouth and nostrils In Borneo: Uncommon locally, known from southern Sarawak and eastern Sabah.


References: Carvalho (1999); Carvalho et al. (1999).

164


Darkspotted Numbfish Narcine maculata (Shaw, 1804)

1

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) Identifying features: 1 disc subcircular, thick and flabby 2 eyes not embedded in skin 3 two similar-sized dorsal fins 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 Size: To at least 40 cm; males mature at about 24 cm. 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

165


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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: None.

6 7

Mouth and nostrils

In Borneo: Recorded locally from southern Sarawak; another specimen recorded in literature from unspecified locality off Sabah.


References: Carvalho (1999); Carvalho et al. (1999).

166


Spot-tail Sleeper Ray Narke dipterygia (Bloch & Schneider, 1801)

3 4

1

5

2

Other names: Numbray, Spot-tail Electric Ray (English), Pari Listrik, Pari Kontak (Indonesian), Pari Karan (Malaysian) Identifying features: 1 one dorsal fin 2 eyes very small, partly embedded in skin 3 disc moderately enlarged, its length (measured to cloaca) slightly longer than tail 4 spiracle lateral to eye 5 caudal fin longer than deep 6 nostrils slit-like 7 mouth very small and not strongly arched 8 undersurfaces of disc and pelvic fin uniformly pale Size: To at least 18 cm; males mature by about 15 cm. Distribution: Tropical Indo–West Pacific, from Oman to Malaysia and north to Japan; may occur off Sumatra. Habitat and biology: Demersal on the continental shelf, both inshore and offshore; locally in 70–95 m depth. Biology largely unknown, presumably viviparous, with yolksac dependency. Diet also unknown but probably consists of small invertebrates, such as polychaetes that live in mud and sand.

167


Utilisation: Infrequent bycatch of both inshore and offshore bottom trawl fisheries operating over soft bottoms on the continental shelf. Not used for food. Conservation status: IUCN Red List: Data Deficient. Local synonyms: None. References: Compagno & Last (1999b).

7

Mouth and nostrils In Borneo: Off central Sarawak.

8 8

Ventral

Narkidae (Sleeper Rays)

6

168


Finless Sleeper Ray Temera hardwickii Gray, 1831

3 4 5 2

1

Other names: Finless Electric Ray (English), Pari Listrik, Pari Kontak (Indonesian), Pari Karan Belangkas, Pari Karas (Malaysian) Identifying features: 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

169


a moderate shock on direct contact. Utilisation: Caught rarely by bottom trawl on the continental shelf. Not used for food. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None. References: Compagno & Last (1999b).

7 Mouth and nostrils In Borneo: Single record from the South China Sea off southwestern Sarawak.

8 8

Ventral

Narkidae (Sleeper Rays)

6

170


Kwangtung Skate Dipturus kwangtungensis (Chu 1960)

3

5

1 6

2

7 4

Other names: None (English), none (Indonesian), Pari Belanda (Malaysian) Identifying features: 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

171

Rajoidei

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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Raja (Okamejei) hollandi Jordan & Richardson:Yano et al. (2005).

8

1

In Borneo: Off Sarawak.

Ventral disc (female)

7

Lateral posterior tail

Rajidae (Skates)

References: Ishihara (1987).

172


Borneo Sand Skate Okamejei cairae Last, Fahmi & Ishihara, 2010

4 1

3

7 2

6

5

Other names: None (English), Pari, Pari Luncur (Indonesian), Pari-raja Tompok (Malaysian) Identifying features: 1 pelvic fin divided into two distinct lobes 2 snout supported by a firm cartilage 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.

173

Rajoidei

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. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: ?Okamejei boesemani (Ishihara): Compagno et al. (2005b); Raja (Okamejei) boesemani Ishihara: Yano et al. (2005).

Rajidae (Skates)

References: Last et al. (2010a).

8

1 Ventral disc (female)

In Borneo: Off western Borneo, including Sabah, Sarawak and northwestern Kalimantan.

7 6 Lateral tail

174


Yellowspotted Skate Okamejei hollandi (Jordan & Richardson, 1909)

4 3

1

2 6

7

5

Other names: Holland Skate, Thornback Ray (English), Pari Luncur (Indonesian), Pari Belanda (Malaysian) Identifying features: 1 pelvic fin divided into two distinct lobes 2 snout supported by a firm cartilage 3 snout broadly triangular, quite long (preorbital length ~3 times orbit diameter) 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

175

Rajoidei

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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Figured (Plate 211) as Raja (Okamejei) boesemani Ishihara: Yano et al. (2005).

Rajidae (Skates)

References: None.

8

1 In Borneo: Off western Borneo, including Sarawak and Kalimantan.


6

7

Lateral tail

176


Philippine Ocellate Skate Okamejei jensenae Last & Lim, 2010

4 1

3

7 2 6

5

Other names: Yellow-spotted Skate (English), none (Indonesian), Pari Belanda (Malaysian) Identifying features: 1 pelvic fin divided into two distinct lobes 2 snout supported by a firm cartilage 3 snout broadly triangular, quite long (preorbital length ~4 times orbit diameter) 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.

177

Rajoidei

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. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: Okamejei sp. 1: Compagno et al. (2005b).

8

1 In Borneo: Off Lahad Datu, eastern Sabah.


7

Lateral posterior tail

Rajidae (Skates)

References: Compagno et al. (2005b); Last & Lim (2010).

178


Borneo Leg Skate Sinobatis borneensis (Chan, 1965)

3 2 4 5

Other names: None (English), none (Indonesian), none (Malaysian) Identifying features: 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 Size: To at least 32 cm; males mature at about 26 cm. 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.

179

Rajoidei

Conservation status: IUCN Red List: Least Concern. Local synonyms: Previously as Anacanthobatis borneensis Chan.

Mouth and nostrils In Borneo: Known from the holotype collected in the South China Sea, off Sarawak.

1

Ventral

Anacanthobatidae (Leg Skates)

References: Last & Compagno (1999).

180


Giant Stingaree Plesiobatis daviesi (Wallace, 1967)

1

2

4 3

5

Other names: Deepwater Stingray (English), Pari (Indonesian), Pari Lembik Merah, Pari Merah (Malaysian) Identifying features: 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 Size: To at least 270 cm; males and females mature at about 130–172 and 189–200 cm 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

181

Myliobatoidei

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. Conservation status: IUCN Red List: Least Concern. Local synonyms: None.

Mouth and nostrils In Borneo: Known only from off Sarawak.

1 Lateral posterior tail with caudal fin

Plesiobatidae (Giant Stingarees)

References: None.

182


Smalleye Stingray Dasyatis microps (Annandale, 1908) 2

1

4

6 5

3

Other names: Thickspine Giant Stingray (English), Pari Sapi, Pari Babi (Indonesian), Pari Gergasi (Malaysian) Identifying features: 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

183

Myliobatoidei

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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: None.

Dasyatidae (Stingrays)

References: Pierce et al. (2008).

In Borneo: Northeastern Sabah.

184


Dwarf Black Stingray Dasyatis parvonigra Last & White, 2008

2 5

1

6

Other names: None (English), Ikan Pari (Indonesian), Pari (Malaysian) Identifying features: 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;

185

Myliobatoidei

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. Conservation status: IUCN Red List: Not Evaluated. 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 the eastern border with Kalimantan.

Ventral disc

1

Nuchal area

3 4

Lateral tail

186


Cow Stingray Dasyatis ushiei (Jordan & Hubbs, 1925) 2

5

6 1

Other names: None (English), Pari Boro, Pari Jantung (Indonesian), Pari Ekor Duri (Malaysian) Identifying features: 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.

187

Myliobatoidei

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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Dasyatis cf. ushiei (Jordan & Hubbs): White et al. (2006b), White & Dharmadi (2007); Dasyatis cf thetidis Ogilby: Yano et al. (2005).

7

In Borneo: Known from off Sarawak and Sabah.

Ventral disc

1

Nuchal area

6

3

Lateral tail

4


References: None.

188


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) Identifying features: 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.

189

Myliobatoidei

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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: None.


References: None.

9 In Borneo: Probably widespread, although currently few records from eastern Kalimantan.

Ventral disc

1

Nuchal area

5

Lateral tail

6

190


Pink Whipray Himantura fai Jordan & Seale, 1906 5

4

3 6

2 1

7

Other names: None (English), Pari Minyak, Pari Tembaga (Indonesian), Pari Pucat (Malaysian) Identifying features: 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.

191

Myliobatoidei

Habitat and biology: Lives over soft substrates near the coast and on the inner continental shelf, from the intertidal zone to about 70 m depth, occasionally deeper to 200 m; schools near coral reefs and sand cays in some areas. Biology poorly known; viviparous, with histotrophy, and diet probably consists of small fishes and bottomdwelling invertebrates. Utilisation: Occasional catch of the demersal tangle net and bottom trawl fisheries. Used mainly for its meat. Conservation status: IUCN Red List: Least Concern. Local synonyms: Image of MSR659 in Yano et al. (2005) is of H. pastinacoides.


References: None.

Ventral disc

Nuchal area


192


Whitespotted Whipray Himantura gerrardi (Gray, 1851) 5 4

2

3 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) Identifying features: 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) 4 no skin folds on tail 5 tail long, whip-like, with alternating light and dark bands (rarely faint) 6 sting situated anteriorly on tail 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

193

Myliobatoidei

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.

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). Conservation status: IUCN Red List: Vulnerable. Local synonyms: Trygon macrurus Bleeker, 1852. References: None.


Ventral disc

1 7 Nuchal area


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.

194


Mangrove Whipray Himantura granulata (Macleay, 1883) 6 5 2 7 4

3

9

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) Identifying features: 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) 5 no skin folds on tail 6 tail moderately long, whip-like, whitish posterior to sting 7 sting situated anteriorly on tail 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.

195

Myliobatoidei

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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: None.


References: None.

In Borneo: Known from off Sabah and northeastern Kalimantan. Ventral disc

1

8 Nuchal area

196


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) Identifying features: 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) 4 no skin folds on tail 5 tail moderately long, whip-like, uniformly dark (not banded) 6 sting situated anteriorly on tail 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.

197

Myliobatoidei

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. Conservation status: IUCN Red List: Least Concern. Local synonyms: Himantura draco Compagno & Heemstra, 1984.

Ventral disc

In Borneo: Mainly known from Sabah and Sarawak; rarely caught off eastern, northwestern and southern Kalimantan.

7

Dorsal tail thorns

1 7

Nuchal area


References: None.

198


Roughback Whipray Himantura kittipongi Vidthayanon & Roberts, 2005

6 5

3

9

1 8 7

2

Other names: None (English), Pari Sungai (Indonesian), Pari Sungai (Malaysian) Identifying features: 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 5 no skin folds on tail 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.

199

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered. Local synonyms: None.


References: None.

4 In Borneo: Known locally from the Pawan and Kapuas rivers, western Kalimantan. Ventral disc

7 Nuchal area

200


Leopard Whipray Himantura leoparda Manjaji-Matsumoto & Last, 2008 5 4

3 6 1

8

2

Other names: Undulate Whipray (English), Pari Macan (Indonesian), Pari Rimau (Malaysian) Identifying features: 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) 4 no skin folds on tail 5 tail long, whip-like, variably banded (rarely faint) 6 sting situated anteriorly on tail 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.

201

Myliobatoidei

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. Conservation status: IUCN Red List: Not Evaluated. 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 and northeastern Kalimantan.

Ventral disc

1 7 Nuchal area

3 5

Tail section - dorsal

Tail section - ventral

Juvenile (200 mm DW)

202


Tubemouth Whipray Himantura lobistoma Manjaji-Matsumoto & Last, 2006

6

5 2 3

1

9 8

Other names: None (English), Pari Lumpur, Pari Tanjung (Indonesian), Pari Mulut Tajam (Malaysian) Identifying features: 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 5 no skin folds on tail 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.

203

Myliobatoidei

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None.



Ventral disc

In Borneo: Known mainly from western Borneo; occasionally off southern and eastern Kalimantan.

7 Nuchal area

4

Protrusible mouth

Late-term embryo (18 cm DW)

204


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) Identifying features: 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 5 no skin folds on tail 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.

205

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered. Local synonyms: Dasybatus krempfi Chabanaud, 1923.

4

In Borneo: Known from north-west

Kalimantan in the environs of the Kapuas River. Ventral disc

7 Nuchal area


References: None.

206


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) Identifying features: 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 5 no skin folds on tail 6 tail long, whip-like and dark posteriorly (not banded) 7 sting situated anteriorly on tail 8 usually one large, greenish pearl thorn on central disc 9 no other enlarged thorns on disc or tail 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.

207

Myliobatoidei

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Trygon pareh Bleeker, 1852.


References: None.

4

Ventral disc

8 Nuchal area

In Borneo: Widespread, concentrated near large river estuaries.

208


Giant Freshwater Whipray Himantura polylepis (Bleeker, 1852) 6

5 2 3 7

1 9

Other names: Freshwater Whipray, Giant Freshwater Stingray (English), Pari (Indonesian), Pari Air Tawar, Pari Sungai (Malaysian) Identifying features: 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 5 no skin folds on tail 6 tail long, thin-based, whip-like and blackish (not banded) 7 sting situated anteriorly on tail 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.

209

Myliobatoidei

Distribution: Widespread but patchy in the Indo–West Pacific, from India to eastern Indonesia; a similar form off Australia, and possibly New Guinea, is a separate species (H. dalyensis Last & Manjaji-Matsumoto, 2008). Habitat and biology: A very large coastal ray living primarily in rivers and estuaries, often well upstream. Probably common in the Kinabatangan River but rarely caught; sometimes in coastal marine and brackish habitats in other parts of Indonesia.Viviparous, with histotrophy; biology not well known. Feeds mainly on small fishes and crustaceans. Utilisation: Caught accidentally by local fishers, mainly by line and occasionally in gillnets and traps. Used for food and possibly its cartilage. Conservation status: IUCN Red List:Vulnerable; Critically Endangered in Thailand (as Himantura chaophraya Monkolprasit & Roberts, 1990).


Local synonyms: Himantura chaophraya Monkolprasit & Roberts, 1990. References: Last & Manjaji-Matsumoto (2008).

4 In Borneo: Known from Sabah and eastern Kalimantan. Ventral disc

8 Nuchal area

210


White-edge Freshwater Whipray Himantura signifer Compagno & Roberts, 1982 6

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) Identifying features: 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 5 no skin folds on tail 6 tail long, broad-based, whip-like and whitish (not banded) 7 sting situated anteriorly on tail 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

211

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered.



4

In Borneo: Kapuas, Sampi and Rungan rivers, western and southern Kalimantan. Ventral disc

1

Nuchal area

212


Bleeker’s Whipray Himantura uarnacoides (Bleeker, 1852)

5 6

3

1 9 7 2

Other names: Whitenose Whipray (English), Pari Tanjung, Pe Pasung, Pari Lumpur (Indonesian), Pari Pasir (Malaysian) Identifying features: 1 denticle band very broad, margin rounded on snout 2 profile of disc weakly rhomboidal, snout narrowly triangular 3 greyish or brownish above (without a pattern) 4 whitish below, sometimes with a narrow dark posterior margin 5 no skin folds on tail 6 tail long, whip-like and pale (not banded) 7 sting situated anteriorly on tail 8 usually 1–3 large pearl thorns on central disc 9 no other enlarged thorns on disc or tail 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.

213

Myliobatoidei

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). Conservation status: IUCN Red List: Vulnerable. Local synonyms: Trygon bleekeri Blyth, 1860.

4

In Borneo: Widespread, less common off western Sabah. Ventral disc

8 Nuchal area

Juvenile (277 mm TL)


References: None.

214


Reticulate Whipray Himantura uarnak (Forsskål, 1775) 5

3

1

7

9

6 2

Other names: Reticulate Whipray, Coachwhip Ray, Longtail Ray, Leopard Stingray (English), Pari Macan, Pari Merica (Indonesian), Pari Rimau (Malaysian) Identifying features: 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 with fine dark spots and reticulations 4 in juveniles upper surface with relatively small dark spots 5 no skin folds on tail 6 tail long, whip-like, variably banded (often faint in adults) 7 sting situated anteriorly on tail 8 central disc usually with 2 heart-shaped thorns (not preceded by row of slightly smaller thorns) 9 midline of tail before sting without thorns Size: To at least 160 cm DW (450 cm TL); males mature at 82–84 cm DW; born at 21–28 cm DW.

215

Myliobatoidei

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Trygon punctata Günther, 1870; Himantura toshi Whitley: Yano et al. (2005).


References: None.

In Borneo: Widespread, but patchy. Ventral disc

4 8 Nuchal area Juvenile (262 mm DW)

216


Honeycomb Whipray Himantura undulata (Bleeker, 1852) 6

5

3

7

1 9

2

Other names: Bleeker’s Variegate Ray, Ocellate Whipray (English), Pari Macan (Indonesian), Pari Batik, Pari Rimau (Malaysian) Identifying features: 1 broad band of flat denticles on central disc (sparse or absent in small juveniles) 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 5 no skin folds on tail 6 tail long, whip-like, with dark pattern or bands 7 sting situated anteriorly on tail 8 central disc usually with dominant pearl-shaped thorn followed by 2–3 slightly smaller thorns 9 midline of tail before sting without 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.

217

Myliobatoidei

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: Trygon favus Annandale, 1909; Himantura fava Annandale: White et al. (2006b); Himantura sp. B: Yano et al. (2005).

Borneo: Sarawak, northeastern Sabah, and northwestern and eastern Kalimantan; probably more widespread.

In

Ventral disc

8

4

Nuchal area

Juvenile (350 mm DW)



218


Dwarf Whipray Himantura walga (Müller & Henle, 1841)

2 3 6 1 5 7

Other names: None (English), Pari Kikir, Toka Toka (Indonesian), Pari Ketuka, Tukatuka, Pari Tanjung (Malaysian) Identifying features: 1 narrow band of flat denticles on central disc in adults 2 profile of disc almost oval, snout broadly triangular 3 greyish or brownish above (without a pattern) 4 whitish below, sometimes with a yellowish grey posterior margin 5 no skin folds on tail 6 tail short, not whip-like (end bulbous in adult females) 7 sting situated anteriorly on tail 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

219

Myliobatoidei

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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: Trygon heterurus Bleeker, 1852; images of Himantura imbricata (Bloch & Schneider) in Yano et al. (2005) appear to be this species.

4

In Borneo: Widespread, but not recorded from eastern Kalimantan. Ventral disc

8

Nuchal area

9

Anterior tail - dorsal view

6 Dorsal post-sting tail - mature female


References: None.

220


Bluespotted Maskray Neotrygon kuhlii (Müller & Henle, 1841)

4

6 5

1

7

Other names: Bluespotted Stingaree, Bluespotted Stingray, Kuhl’s Stingray (English), Pari Blentik, Pari Kodok (Indonesian), Pari Lalat, Pari Macan, Pari Riman, Pari Rimau, Pari Tanjung, Kiampau (Malaysian) Identifying features: 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.

221

Myliobatoidei

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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Dasyatis kuhlii (Müller & Henle): White et al. (2006b), White & Dharmadi (2007).


References: None.


Ventral disc

1

Nuchal area

2

3 Lateral tail

222


Eastern Cowtail Stingray Pastinachus atrus (Macleay, 1883) 4 5 2 7

3

1

6

8

Other names: Banana-tail Ray, Bull Ray, Fantail Ray, Feathertail Ray, Feathertail Stingray (English), Pari Bendera (Indonesian), Pari Bendera, Pari Daun, Pari Nyonya, Pari Tanjung (Malaysian) Identifying features: 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.

223

Myliobatoidei

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. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: Pastinachus sephen (Forsskål):Yano et al. (2005), White et al. (2006b).

Ventral disc


References: None.


5 Nuchal area Lateral tail - at midlength of ventral fold

224


Narrowtail Stingray Pastinachus gracilicaudus Last & Manjaji-Matsumoto, 2010 4

5

7

3

1

6

8

2

Other names: None (English), Pari Bendera (Indonesian), none (Malaysian) Identifying features: 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 typically pale, relatively slender (maximum depth usually < 3 times height of tail above) 6 tail relatively broad based 7 sting situated posteriorly on tail (its distance from cloaca more than half disc width) 8 no thorn row along middle of tail Size: To almost 83 cm DW (>200 cm TL); males mature by 67 cm DW; free-swimming by 30 cm DW. Distribution: Not well defined; known from the Indo–Malay Archipelago.

225

Myliobatoidei

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. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: Confused with Pastinachus sephen (Forsskål).

Ventral disc

In Borneo: Patchy, in western and northern Borneo; not observed from most of southern and eastern Kalimantan.

5 Nuchal area

Lateral tail - at midlength of ventral fold


References: Last & Manjaji-Matsumoto (2010).

226


Roughnose Stingray Pastinachus solocirostris Last, Manjaji & Yearsley, 2005 4 5 2

1

7

3 6 8

Other names: None (English), Pari Gunung, Pari Bendera (Indonesian), Pari Mulut Kasar (Malaysian) Identifying features: 1 broad band of flat denticles on central disc 2 disc profile strongly quadrangular with an acute, triangular snout 3 snout apex covered with enlarged, spear-shaped or fork-like denticles 4 ventral skin fold terminating well before tail tip (when undamaged) 5 fold relatively slender (maximum depth usually 150 cm TL); males mature at about 36–39 cm DW; born at about 20 cm DW. Distribution: Confined to Indonesia and Malaysia (Borneo and Sumatra). Habitat and biology: Habitat and biology largely unknown; demersal inshore on

227

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered. Local synonyms: Pastinachus cf. sephen (Forsskål): Yano et al. (2005).

In Borneo: Widespread off western Borneo; isolated records from southern and southeastern Kalimantan.

Ventral disc

3

Nuchal area

5 Lateral tail - at midlength of ventral fold

Denticles at apex of snout


References: Last et al. (2005).

228


Starynose Stingray Pastinachus stellurostris Last, Fahmi & Naylor, 2010 4

5

7

1 3

6

8

2

Other names: None (English), Pari Bendera (Indonesian), none (Malaysian) Identifying features: 1 broad band of flat denticles on central disc 2 disc profile strongly quadrangular with an acute, triangular snout 3 snout apex covered with enlarged, star-shaped denticles 4 ventral skin fold terminating well before tail tip (when undamaged) 5 fold relatively slender (maximum depth 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

243

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered. Local synonyms: None.

3

Ventral head In Borneo: Uncommon, known from

western Kalimantan and northeastern Sabah. Upper tooth plate

4

Lower tooth plate

Myliobatidae (Eagle Rays)

References: None.

244


Whitespotted Eagle Ray Aetobatus ocellatus (Kuhl, 1823)

2

5

1

Other names: Bonnet Skate, Duckbill Ray, Spotted Eagle Ray (English), Pari Burung, Pari Ayam (Indonesian), Pari Lang Bintik Putih, Pari Lang, Pari Helang (Malaysian) Identifying features: 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.

245

Myliobatoidei

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. Conservation status: IUCN Red List: Not Evaluated.


Local synonyms: Aetobatus narinari (Euphrasen, 1790): Yano et al. (2005), White et al. (2006b); Aetobatus cf. narinari (Euphrasen, 1790): Compagno et al. (2005b); Aetobatus guttatus (Shaw, 1804): Compagno & Last (1999a); Raia quinqueaculeata Quoy & Gaimard, 1824. References: White et al. (2010d).

3

Ventral head In Borneo: Widespread.

Upper tooth plate

4

Juvenile - dorsal disc (465 mm DW)

Lower tooth plate

246


Mottled Eagle Ray Aetomylaeus maculatus (Gray, 1834)

1 6

4

3

2 Other names: Ocellate Eagle Ray, Ornate Eagle Ray (English), Pari Burung (Indonesian), Pari-lang Tompok Putih (Malaysian) Identifying features: 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.

247

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered. Local synonyms: None. Authority previously considered to be Gray, 1832 but publication date listed on Eschmeyer’s Catalog of Fishes as 1 October 1834.

5


Upper tooth plate

4

Juvenile - dorsal disc (~250 mm DW)

Lower tooth plate


References: None.

248


Banded Eagle Ray Aetomylaeus nichofii (Bloch & Schneider, 1801)

1 6

4

3

2

Other names: Barbless Eagle Ray, Nieuhof ’s Eagle Ray (English), Pari Burung, Pe Lamping (Indonesian), Pari-lang Jalur (Malaysian) Identifying features: 1 fleshy ridge on side of head not connected to edge of disc 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) 5 edge of nasal curtain near mouth almost straight 6 spiracles lateral on head 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.

249

Myliobatoidei

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. Conservation status: IUCN Red List: Vulnerable. Local synonyms: None.


References: None.

5


Upper tooth plate

Lower tooth plate

250


Ornate Eagle Ray Aetomylaeus vespertilio (Bleeker, 1852)

1 6

4

3 2

Other names: Reticulate Eagle Ray (English), Pari Burung (Indonesian), Pari Lang Bunga (Malaysian) Identifying features: 1 fleshy ridge on side of head not connected to edge of disc 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 5 edge of nasal curtain near mouth almost straight 6 spiracles lateral on head 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-

251

Myliobatoidei

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. Conservation status: IUCN Red List: Endangered. Local synonyms: None.

5

Ventral head In Borneo: Known locally from a few individuals collected off Sabah and Kalimantan.

Upper tooth plate

Lower tooth plate


References: None.

252


Javanese Cownose Ray Rhinoptera javanica Müller & Henle, 1841 1 2

5 7 4

Other names: Cow Ray, Flapnose Ray (English), Pari Burung, Pari Elang (Indonesian), Pari-kelawar Jawa, Pari Ayam, Pari Lang, Pari Susun (Malaysian) Identifying features: 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.

253

Myliobatoidei

Distribution: Central western Pacific, from Thailand to Australia and north to the East China Sea; poorly defined due to confusion with other species. Habitat and biology: Preferred habitat and biology poorly known; patchy, but most common inshore in bays, estuaries and near coral reefs.Viviparous, with histotrophy; one large pregnant female in Indonesia (~130 cm DW) contained a single, late-term embryo of about 60 cm DW. Diet presumably consists mainly of bivalves and other bottomdwelling invertebrates. Utilisation: Caught occasionally by bottom trawl, demersal inshore gill and tangle net fisheries. Sold mainly for its meat, but not of high value. Conservation status: IUCN Red List: Vulnerable.

Rhinopteridae (Cownose Rays)

Local synonyms: Rhinoptera adspersa Müller & Henle, 1841. References: None.

1

3

Ventral head

In Borneo: Apparently uncommon, known only from eastern Sabah and western Kalimantan.

5

6 7

3 Lateral dorsal fin and anterior tail Lateral head

254


Short-tail Cownose Ray Rhinoptera jayakari Boulenger, 1895 1 2

5

7 4

Other names: None (English), Pari Elang (Indonesian), Pari Lang (Malaysian) Identifying features: 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.

255

Myliobatoidei

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). References: None.

1

3

In Borneo: Known off eastern Sabah, southern Sarawak and northwestern Kalimantan.

Ventral head

5

6 7

3 Lateral head

Lateral dorsal fin and anterior tail

Rhinopteridae (Cownose Rays)

Conservation status: IUCN Red List: Not Evaluated.

256


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) Identifying features: 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. Distribution: Circumtropical, occasionally making seasonal incursions into warm temperate waters. Habitat and biology: Wide-ranging, pelagic species found near the surface in tropical

257

Myliobatoidei

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. Conservation status: IUCN Red List: Near Threatened.

Mobulidae (Devilrays)

Local synonyms: None. References: White et al. (2006a); Marshall et al. (2009).

3

5 Ventral head In Borneo: Single record from a fish market in northwestern Kalimantan presumably this species; observed regularly by divers at Derawan and Sipadan islands off eastern Borneo.

4 Embryo

Lateral dorsal fin and anterior tail

258


Borneo Devilray Mobula sp. 1 2

4 5

Other names: None (English), none (Indonesian), Pari Paus (Malaysian) Identifying features: 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

259

Myliobatoidei

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. Conservation status: IUCN Red List: Not Evaluated. Local synonyms: None. References: None.

2 3 7

In Borneo: Known from off Kuching, Sarawak. Ventral head

6

Dorsolateral head


1

260


Japanese Devilray Mobula japanica (Müller & Henle, 1841)

1

Other names: Spinetail Mobula, Spinetail Devilray (English), Cawang Genul, Lampengan, Pari Lambingan, Plampangan (Indonesian), Pari Paus (Malaysian) Identifying features: 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 5 mouth subterminal, located on underside of head Size: To at least 310 cm DW (usually less than 250 cm); males mature at 198–205 cm; late-term embryos observed up to 113 cm DW. Distribution: Circumtropical, occasionally making seasonal incursions into warm temperate waters. Habitat and biology: A large pelagic inshore and offshore (possibly oceanic) devilray. Viviparous, with histotrophy; gives birth to a single pup after an unknown gestation

261

Myliobatoidei

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. Conservation status: IUCN Red List: Near Threatened. 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

4 5

Dorsolateral head

Ventral head


References: White et al. (2006a).

262


Lesser Devilray Mobula kuhlii (Müller & Henle, 1841) 1 2

5

Other names: Shortfin Devilray, Shorthorn Devilray (English), Pari Lambingan, Pari Lampengan (Indonesian), Pari Kelawar, Pari Paus, Tanpa Duri (Malaysian) Identifying features: 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 7 mouth subterminal, located on underside of head 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.

263

Myliobatoidei

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. Conservation status: IUCN Red List: Data Deficient. Local synonyms: Dicerobatis draco Günther, 1872.


References: White et al. (2006a).

1

2 3 7

Ventral head

4

In Borneo: Northeastern Borneo, off Sabah.

6

5 Lateral dorsal fin and anterior tail Dorsolateral head

264


Bentfin Devilray Mobula thurstoni (Lloyd, 1908) 1 2

5

6

Other names: Smoothtail Mobula (English), Pari Lempengan, Pari Lambingan, Plampangan (Indonesian), Pari Paus, Sirip Lekuk (Malaysian) Identifying features: 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 8 mouth subterminal, located on underside of head 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

265

Myliobatoidei

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. Conservation status: IUCN Red List: Near Threatened. Local synonyms: None. References: White et al. (2006a).

2 3 8

Ventral head

In Borneo: Single record from the Sandakan fish market, northeastern Sabah.

4

7 6 Lateral dorsal fin and anterior tail Dorsolateral head


1

266


Silver Chimaera Chimaera phantasma Jordan & Snyder, 1900

4

6

5

2

7 3 1

Other names: Ghost Shark (English), Hiu Hantu, Hiu Pogak (Indonesian), Chimara Perak, Ikan Tikus (Malaysian) Identifying features: 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.

267

Chimaeriformes

Conservation status: IUCN Red List: Data Deficient. Local synonyms: Chimaera sp.: Yano et al. (2005). References: Masuda et al. (1975); Dagit (2006).

8

In Borneo: Known only from off Sarawak.

Lateral head

7 1

Lateral tail

2

Trunk lateral line canal

Chimaeridae (Shortnose Chimaeras)

6

268


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).

270


Images sourced outside Borneo Hexanchus griseus: lateral (Brazil). Centrophorus moluccensis: lateral (Lombok, Indonesia). Orectolobus leptolineatus: ventral head, embryo (Lombok, Indonesia). Chiloscyllium plagiosum: juvenile (Hong Kong). Nebrius ferrugineus: lateral, ventral head (Western Australia); dorsal fins (Singapore). Stegostoma fasciatum: lateral, ventral head (Queensland, Australia). Rhincodon typus: lateral (Japan). Alopias pelagicus: lateral, ventral head (Bali, Indonesia). Isurus oxyrinchus: lateral, ventral head (Lombok, Indonesia). Apristurus platyrhynchus: lateral, ventral head (New South Wales, Australia); flank denticles (Western Australia). Halaelurus maculosus: lateral, ventral head, dorsal head and anterior trunk (Bali, Indonesia); egg case (Lombok, Indonesia). Chaenogaleus macrostoma: lateral (Qatar); ventral head (West Java, Indonesia). Carcharhinus albimarginatus: lateral, ventral head (Western Australia). Carcharhinus amblyrhynchos: embryo (Lombok, Indonesia). Carcharhinus dussumieri: ventral head (West Java, Indonesia). Carcharhinus falciformis: lateral, ventral head (Lombok, Indonesia). Carcharhinus plumbeus: lateral (Bali, Indonesia). Galeocerdo cuvier: lateral (Bali, Indonesia); ventral head (Queensland, Australia). Prionace glauca: lateral (Tasmania, Australia); ventral head (Queensland, Australia).Triaenodon obesus: lateral (Bali, Indonesia). Eusphyra blochii: lateral (West Java, Indonesia); ventral head (northern Australia). Anoxypristis cuspidata: dorsal, ventral head, caudal fin (Queensland, Australia). Pristis microdon: dorsal, ventral head, caudal fin (Queensland, Australia). Pristis zijsron: dorsal, ventral head, caudal fin (Queensland, Australia). Rhina ancylostoma: dorsal (Bali, Indonesia); lateral head (East Java, Indonesia). Glaucostegus typus: ventral head (Queensland, Australia); lateral tail (West Papua, Indonesia). Plesiobatis daviesi: dorsal (West Java, Indonesia); oronasal (New South Wales, Australia). Dasyatis microps: dorsal (Gulf of Carpentaria, Australia). Dasyatis ushiei: dorsal, ventral disc, nuchal area (Lombok, Indonesia). Himantura fai: nuchal area (Queensland, Australia). Himantura granulata: dorsal (East Java, Indonesia); nuchal area (Queensland, Australia). Himantura jenkinsii: dorsal tail thorns, nuchal area (West Java, Indonesia). Himantura leoparda: ventral disc, nuchal area (Queensland, Australia), tail section - dorsal and ventral (Manila, Philippines). Himantura uarnacoides: nuchal area (West Java, Indonesia). Pteroplatytrygon violacea: dorsal (Lombok, Indonesia); ventral disc (Bali, Indonesia). Taeniurops meyeni: dorsal, ventral disc (Bali, Indonesia). Urogymnus asperrimus: dorsal (Marshall Islands). Gymnura zonura: lateral anterior tail (Bali, Indonesia). Aetobatus flagellum: dorsal, ventral head (West Java, Indonesia). Aetomylaeus vespertilio: dorsal, ventral head (Queensland, Australia). Rhinoptera javanica: ventral head, lateral head, lateral dorsal fin and anterior tail (Queensland, Australia). Rhinoptera jayakari: lateral head, lateral dorsal fin and anterior tail (Lombok, Indonesia). Manta birostris: dorsal, lateral dorsal fin and anterior tail (Lombok, Indonesia); embryo (West Java, Indonesia). Mobula japanica: dorsal, dorsolateral head (Lombok, Indonesia); lateral dorsal fin and sting (East Java, Indonesia); ventral head (Bali, Indonesia). Mobula thurstoni: dorsal, ventral head, dorsolateral head, lateral dorsal fin and anterior tail (Lombok, Indonesia).

271

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White, W.T., Bartron, C. & Potter, I.C. (2008) Catch composition and reproductive biology of Sphyrna lewini (Carcharhiniformes, Sphyrnidae) in Indonesian waters. Journal of Fish Biology 72: 1675–1689. White, W.T., Last, P.R. & Stevens, J.D. (2007) Halaelurus maculosus n. sp. and H. sellus n. sp., two new species of catshark (Carcharhiniformes: Scyliorhinidae) from the IndoWest Pacific. Zootaxa 1639: 1–21. White, W.T., Giles, J., Dharmadi & Potter, I.C. (2006a) Data on the bycatch fishery and reproductive biology of mobulid rays (Myliobatiformes) in Indonesia. Fisheries Research 82: 65–73. White, W.T., Last, P.R. & Lim, A.P.K. (2010a) Rediscovery of the rare and endangered Borneo Shark Carcharhinus borneensis (Bleeker, 1858) (Carcharhiniformes: Carcharhinidae). In: P.R. Last, W.T. White & J.J. Pogonoski (eds). Descriptions of New Sharks and Rays from Borneo. CSIRO Marine and Atmospheric Research Paper 032. White, W.T., Last, P.R. & Naylor, G.J.P. (2010b) Scoliodon macrorhynchos (Bleeker, 1852), a second species of spadenose shark from the Western Pacific (Carcharhiniformes: Carcharhinidae). In: P.R. Last, W.T. White & J.J. Pogonoski (eds). Descriptions of New Sharks and Rays from Borneo. CSIRO Marine and Atmospheric Research Paper 032. White, W.T., Last, P.R., Naylor, G.J.P. & Harris, M. (2010c) Resurrection and redescription of the Borneo Broadfin Shark Lamiopsis tephrodes (Fowler, 1905) (Carcharhiniformes: Carcharhinidae). In: P.R. Last, W.T. White & J.J. Pogonoski (eds). Descriptions of New Sharks and Rays from Borneo. CSIRO Marine and Atmospheric Research Paper 032. White, W.T., Last, P.R., Naylor, G.J.P., Caira, J.N. & Jensen, K. (2010d) Clarification of Aetobatus ocellatus (Kuhl, 1823) as a valid species, and a comparison with Aetobatus narinari (Euphrasen, 1790) (Rajiformes: Myliobatidae). In: P.R. Last, W.T. White & J.J. Pogonoski (eds). Descriptions of New Sharks and Rays from Borneo. CSIRO Marine and Atmospheric Research Paper 032. White, W.T., Last, P.R., Stevens, J.D.,Yearsley, G.K., Fahmi & Dharmadi (2006b) Economically important sharks and rays of Indonesia. ACIAR Monograph Series, No 124, ACIAR Publishing, Canberra, 329 pp. Yano, K., Ahmad, A., Gambang, A.C., Idris, A.H., Solahuddin, A.R. & Aznan, Z. (2005) Sharks and rays of Malaysia and Brunei Darussalam. SEAFDEC, MFRDMD, 557 pp.

277

Checklist of Borneo sharks and rays Order Hexanchiformes: Cow and Frilled Sharks

Family Hexanchidae: Sixgill and Sevengill Sharks

 Hexanchus griseus (Bonnaterre, 1788)

Bluntnose Sixgill Shark

Order Squaliformes: Dogfish Sharks

Family Squalidae: Dogfishes

 Squalus altipinnis Last et al., 2007

Family Centrophoridae: Gulper Sharks

 Centrophorus moluccensis Bleeker, 1860

Western Highfin Spurdog 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

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)

Family Stegostomatidae: Zebra Sharks

 Stegostoma fasciatum (Hermann, 1783)

Indonesian Wobbegong

Tawny Nurse Shark Zebra Shark

278


Family Rhincodontidae: Whale Sharks

 Rhincodon typus Smith, 1828

Whale Shark

Order Lamniformes: Mackerel Sharks

Family Alopiidae: Thresher Sharks

 Alopias pelagicus Nakamura, 1935

Family Lamnidae: Mackerel Sharks

 Isurus oxyrinchus Rafinesque, 1810

Pelagic Thresher

Shortfin Mako

Order Carcharhiniformes: Ground Sharks

Family Scyliorhinidae: Catsharks

 Apristurus platyrhynchus (Tanaka, 1909)

Bigfin Catshark

 Atelomycterus marmoratus (Bennett, 1830)

Coral Catshark

 Cephaloscyllium sarawakensis Yano et al., 2005 Sarawak Swellshark

 Halaelurus maculosus White et al., 2007

Family Triakidae: Houndsharks

 Mustelus manazo Bleeker, 1854

Starspotted Smoothhound

 Mustelus widodoi White & Last, 2006

Whitefin Smoothhound

Family Hemigaleidae: Weasel Sharks

 Chaenogaleus macrostoma (Bleeker, 1852)

Indonesian Speckled Catshark

Hooktooth Shark

 Hemigaleus microstoma Bleeker, 1852

Sicklefin Weasel Shark

 Hemipristis elongata (Klunzinger, 1871)

Fossil Shark

 Paragaleus tengi (Chen, 1963)

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)

Straight-tooth Weasel Shark

Silky Shark

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Checklist of species

 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

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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



 Okamejei cairae Last et al., 2010a

Borneo Sand Skate

 Okamejei hollandi (Jordan & Rich., 1909)

Yellowspotted Skate

 Okamejei jensenae Last & Lim, 2010

Philippine Ocellate Skate

Dipturus kwangtungensis (Chu 1960)

Kwangtung Skate

281

Checklist of species

Family Anacanthobatidae: Leg Skates

 Sinobatis borneensis (Chan, 1965)

Borneo Leg Skate

Suborder Myliobatoidei: Stingrays

Family Plesiobatidae: Giant Stingarees

 Plesiobatis daviesi (Wallace, 1967)

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

Giant Stingaree

 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

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 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 Aetobatus flagellum 242 244 Aetobatus ocellatus Aetomylaeus maculatus 246 Aetomylaeus nichofii 248 Aetomylaeus vespertilio 250 albimarginatus, Carcharhinus 86 Alopias pelagicus 62 Alopiidae 29, 62–63 altipinnis, Squalus 38 amblyrhynchoides, Carcharhinus 88 amblyrhynchos, Carcharhinus 90 Anacanthobatidae 25, 178–179 146 ancylostoma, Rhina 140 Anoxypristis cuspidata 66 Apristurus platyrhynchus 236 asperrimus, Urogymnus 68 Atelomycterus marmoratus 222 atrus, Pastinachus australiae, Rhynchobatus 148

B birostris, Manta blochii, Eusphyra borneensis, Carcharhinus borneensis, Sinobatis brevilabiata, Narcine brevipinna, Carcharhinus

256 134 92 178 160 94

C cairae, Okamejei 172 Carcharhinidae 33, 86–133 66–139 Carcharhiniformes Carcharhinus albimarginatus 86 Carcharhinus amblyrhynchoides 88 Carcharhinus amblyrhynchos 90 Carcharhinus borneensis 92

Carcharhinus brevipinna Carcharhinus dussumieri Carcharhinus falciformis Carcharhinus leucas Carcharhinus limbatus Carcharhinus macloti Carcharhinus melanopterus Carcharhinus plumbeus Carcharhinus sealei Carcharhinus sorrah Centrophoridae Centrophorus moluccensis Cephaloscyllium sarawakensis Chaenogaleus macrostoma Chiloscyllium hasseltii Chiloscyllium indicum Chiloscyllium plagiosum Chiloscyllium punctatum Chimaera phantasma Chimaeridae Chimaeriformes cuspidata, Anoxypristis cuvier, Galeocerdo

94 96 98 100 102 104 106 108 110 112 29, 40–41 40 70 78 48 50 52 54 266 23, 266–267 266–267 140 114

D Dasyatidae Dasyatis microps Dasyatis parvonigra Dasyatis ushiei Dasyatis zugei daviesi, Plesiobatis dipterygia, Narke Dipturus kwangtungensis dussumieri, Carcharhinus

26, 182–237 182 184 186 188 180 166 170 96

E elongata, Hemipristis

82

284

Eusphyra blochii


134

F fai, Himantura falciformis, Carcharhinus fasciatum, Stegostoma ferrugineus, Nebrius flagellum, Aetobatus formosensis, Rhinobatos fowlerae, Glyphis

190 98 58 56 242 158 118

G Galeocerdo cuvier gerrardi, Himantura Ginglymostomatidae glauca, Prionace Glaucostegus thouin Glaucostegus typus Glyphis fowlerae Glyphis sp. gracilicaudus, Pastinachus granulata, Himantura griseus, Hexanchus Gymnura poecilura Gymnura zonura Gymnuridae

114 192 30, 56–57 124 154 156 118 116 224 194 36 238 240 26, 238–241

H Halaelurus maculosus hardwickii,Temera hasseltii, Chiloscyllium Hemigaleidae Hemigaleus microstoma Hemipristis elongata Hemiscylliidae Heterodontidae Heterodontiformes Heterodontus zebra Hexanchidae Hexanchiformes

72 168 48 32, 78–85 80 82 31, 48-55 30, 44–45 44–45 44 28, 36–37 36–37

Hexanchus griseus Himantura fai Himantura gerrardi Himantura granulata Himantura jenkinsii Himantura kittipongi Himantura leoparda Himantura lobistoma Himantura oxyrhyncha Himantura pastinacoides Himantura polylepis Himantura signifer Himantura uarnacoides Himantura uarnak Himantura undulata Himantura walga hollandi, Okamejei

36 190 192 194 196 198 200 202 204 206 208 210 212 214 216 218 174

I indicum, Chiloscyllium Isurus oxyrinchus

50 64

J japanica, Mobula javanica, Rhinoptera jayakari, Rhinoptera jenkinsii, Himantura jensenae, Okamejei

260 252 254 196 176

K kittipongi, Himantura kuhlii, Mobula kuhlii, Neotrygon kwangtungensis, Dipturus

198 262 220 170

L laevis, Rhynchobatus Lamiopsis tephrodes Lamnidae Lamniformes

150 120 31, 64–65 62–63

285

SR cuinennitni gf i ch enaadm e s i n d e x

leoparda, Himantura leptolineatus, Orectolobus leucas, Carcharhinus lewini, Sphyrna limbatus, Carcharhinus lingula, Narcine lobistoma, Himantura Loxodon macrorhinus lymma,Taeniura

200 46 100 136 102 162 202 122 232

M macloti, Carcharhinus macrorhinus, Loxodon macrorhynchos, Scoliodon macrostoma, Chaenogaleus maculata, Narcine maculatus, Aetomylaeus maculosus, Halaelurus manazo, Mustelus Manta birostris marmoratus, Atelomycterus melanopterus, Carcharhinus meyeni,Taeniurops microps, Dasyatis microstoma, Hemigaleus Mobula japanica Mobula kuhlii Mobula sp. Mobula thurstoni Mobulidae mokarran, Sphyrna moluccensis, Centrophorus Mustelus manazo Mustelus widodoi Myliobatidae Myliobatoidei

104 122 130 78 164 246 72 74 256 68 106 234 182 80 260 262 258 264 27, 256–265 138 40 74 76 28, 242–251 180–265

N Narcine brevilabiata Narcine lingula

160 162

Narcine maculata Narcinidae Narke dipterygia Narkidae Nebrius ferrugineus Neotrygon kuhlii nichofii, Aetomylaeus

164 25, 160–165 166 25, 166–169 56 220 248

O obesus,Triaenodon ocellatus, Aetobatus Okamejei cairae Okamejei hollandi Okamejei jensenae oligolinx, Rhizoprionodon Orectolobidae Orectolobiformes Orectolobus leptolineatus oxyrhyncha, Himantura oxyrinchus, Isurus

132 244 172 174 176 128 30, 46–47 46–61 46 204 64

P Paragaleus tengi 84 parvonigra, Dasyatis 184 Pastinachus atrus 222 Pastinachus gracilicaudus 224 Pastinachus solocirostris 226 Pastinachus stellurostris 228 pastinacoides, Himantura 206 62 pelagicus, Alopias phantasma, Chimaera 266 plagiosum, Chiloscyllium 52 platyrhynchus, Apristurus 66 Plesiobatidae 26, 180–181 Plesiobatis daviesi 180 plumbeus, Carcharhinus 108 poecilura, Gymnura 238 polylepis, Himantura 208 Prionace glauca 124 Pristidae 23, 140–145

286

Pristis microdon Pristis zijsron Pristoidei Pteroplatytrygon violacea punctatum, Chiloscyllium


142 144 140–145 230 54

R Rajidae Rajoidei Rhina ancylostoma Rhincodontidae Rhincodon typus Rhinidae Rhinobatidae Rhinobatoidei Rhinobatos formosensis Rhinoidei Rhinoptera javanica Rhinoptera jayakari Rhinopteridae Rhizoprionodon acutus Rhizoprionodon oligolinx Rhynchobatidae Rhynchobatoidei Rhynchobatus australiae Rhynchobatus laevis Rhynchobatus springeri

25, 172–179 172–179 146 30, 60–61 60 24, 146–147 24, 154–159 154–159 158 146–147 252 254 28, 252–255 126 128 24, 148–153 147 148 150 152

138 29, 134–139 152 29, 38–39 38–41 38 42 23, 42–43 42–43 58 30, 58–59 228

T Taeniura lymma Taeniurops meyeni Temera hardwickii tengi, Paragaleus tephrodes, Lamiopsis tergocellatoides, Squatina thouin, Glaucostegus thurstoni, Mobula Torpedinoidei Triaenodon obesus Triakidae typus, Glaucostegus typus, Rhincodon

232 234 168 84 120 42 154 264 160–169 132 32, 74–77 156 60

U

S sarawakensis, Cephaloscyllium Scoliodon macrorhynchos Scyliorhinidae sealei, Carcharhinus signifer, Himantura Sinobatis borneensis solocirostris, Pastinachus sorrah, Carcharhinus sp., Glyphis sp., Mobula Sphyrna lewini

Sphyrna mokarran Sphyrnidae springeri, Rhynchobatus Squalidae Squaliformes Squalus altipinnis Squatina tergocellatoides Squatinidae Squatiniformes Stegostoma fasciatum Stegostomatidae stellurostris, Pastinachus

70 130 31, 66–73 110 210 178 226 112 116 258 136

uarnacoides, Himantura uarnak, Himantura undulata, Himantura Urogymnus asperrimus ushiei, Dasyatis

212 214 216 236 186

V vespertilio, Aetomylaeus violacea, Pteroplatytrygon

250 230

W walga, Himantura

218

287

Scientific names index

widodoi, Mustelus

76

Z zebra, Heterodontus zonura, Gymnura zugei, Dasyatis

44 240 188

288


English common names index A Angelsharks

42–43

B Banana-tail Ray Banded Eagle Ray Banded Whiptail Ray Barbless Eagle Ray Bentfin Devilray Bigfin Catshark Blackblotched Stingray Blackspot Shark Blackspotted Stingray Blacktip Reef Shark Blacktip Shark Blacktip Whaler Bleeker’s Butterfly Ray Bleeker’s Variegate Ray Bleeker’s Whipray Blotched Fantail Stingray Blue-spotted Lagoon Ray Blue-spotted Ribbontail Ray Blue Pointer Blue Shark Bluespotted Fantail Ray Bluespotted Maskray Bluespotted Stingaree Bluespotted Stingray Blue Stingray Blue Whaler Blunthead Shark Bluntnose Sixgill Shark Bluntnose Whiptail Ray Bonnet Skate Borneo Broadfin Shark Borneo Devilray

222 248 192 248 264 66 234 110 234 106 106 102 240 216 212 234 232 232 64 124 232 220 220 220 230 124 132 36 192 244 120 258

Borneo Leg Skate 178 Borneo River Shark 118 Borneo Sand Skate 172 Borneo Shark 92 146 Bowmouth Guitarfish Broadnose Wedgefish 152 Brownbanded Bambooshark 54 54 Brownbanded Catshark 108 Brown Shark 44 Bullhead Shark 222, 234 Bull Ray Bull Shark 36, 100 238–241 Butterfly Rays

C Catsharks 66–73 Chinese Numbfish 162 Clubnose Guitarfish 154 194, 214 Coachwhip Ray Common Blacktip Shark 102 156 Common Shovelnose Ray Coral Catshark 68 252–255 Cownose Rays 252 Cow Ray 36 Cow Shark 36–37 Cowsharks Cow Stingray 186

D Darkfin Numbfish Darkspotted Electric Ray Darkspotted Numbfish Deepwater Stingray Devilfish Devilrays Dogfishes Duckbill Ray

164 164 164 180 256 256–265 38–39 244

289

English common names index

Dwarf Black Stingray Dwarf Whipray

184 218

E Eagle Rays 242–251 Eastern Cowtail Stingray 222 40 Endeavour Dogfish

F Fantail Ray Fantail Stingray Feathertail Ray Feathertail Stingray Finless Electric Ray Finless Sleeper Ray Fish Shark Flapnose Ray Fossil Shark Freshwater Sawfish Freshwater Stingray Freshwater Whaler Freshwater Whipray

222 234 222 222 168 168 126 252 82 142 210 100 208

G Ghost Shark 266 Giant Freshwater Stingray 208 Giant Freshwater Whipray 208 Giant Guitarfish 150, 156 256 Giant Manta 234 Giant Reef Ray 156 Giant Shovelnose Ray Giant Stingaree 180 180–181 Giant Stingarees Graceful Shark 88 142 Great-tooth Sawfish 124 Great Blue Shark Great Hammerhead 138 Green Sawfish 144 54 Grey Carpetshark Grey Reef Shark 90

128 106 40–41 74

Grey Sharpnose Shark Guliman Gulper Sharks Gummy Shark

H Hammerhead Sharks Hardnose Shark Hasselt’s Bambooshark Holland Skate Honeycomb Whipray Hooktooth Shark Hornsharks Houndsharks

134–139 104 48 174 216 78 44–45 74–77

I Indonesian Bambooshark Indonesian Speckled Catshark Indonesian Wobbegong Indo Wobbegong Inkytail Shark

48 72 46 46 94

J Japanese Devilray Japanese Smoothhound Javanese Cownose Ray Jenkins’ Whipray Jordan’s Blue Dogshark

260 74 252 196 122

K Kidney-headed Shark Kuhl’s Stingray Kwangtung Skate

136 220 170

L Lagoon Ray Leg Skates Leopard Shark Leopard Stingray Leopard Whipray Lesser Devilray

232 178–179 58 214 200 262

290


Lesser Fantail Ray 232 Long-tailed Butterfly Ray 238 Longhead Eagle Ray 242 126 Longman’s Dogshark 90 Longnose Blacktail Shark 94 Longnose Grey Whaler Longnose Marbled Whipray 204 Longtail Butterfly Ray 238 48–55 Longtail Carpetsharks 214 Longtail Ray

M Mackerel Shark 64 Mackerel Sharks 64–65 194 Macleay’s Coachwhip Ray 56 Madame X 64 Mako Shark 194 Mangrove Ray 194 Mangrove Stingray Mangrove Whipray 194 256 Manta Manta Ray 256 204 Marbled Freshwater Stingray 204 Marbled Whipray 234 Marble Ribbontail Ray Milk Shark 126 Mottled Eagle Ray 246 36 Mud Shark 146 Mud Skate Mukah River Shark 116

N Narrow Sawfish Narrowsnout Sawfish Narrowtail Stingray Nieuhof ’s Eagle Ray Numbfishes Numbray Nurse Sharks

140 144 224 248 160–165 166 56–57

O Ocellated Angelshark Ocellate Eagle Ray Ocellate Whipray Ornate Eagle Ray

42 246 216 246, 250

P Pacific Spadenose Shark Pale-edged Stingray Pale Whipray Pelagic Stingray Pelagic Thresher Philippine Ocellate Skate Pink Whipray Pointed Sawfish Porcupine Ray

130 188 210 230 62 176 190 140 236

Q Queensland Shark

88

R Reef Ray Reticulate Eagle Ray Reticulate Whipray Ribbontail Stingray Ridgeback Bambooshark Ridgeback Shark River Whaler Rough-back Stingray Roughback Whipray Rough Electric Ray Roughnose Stingray Rough Numbfish Roughskin Stingaree Round Ribbontail Ray Round Whipray

232 250 214 232 50 50 100 196 198 162 226 162 236 234 206

S Sandbar Shark Sandshark

108 150

291

English common names index

Sarawak Pygmy Swell Shark 70 Sarawak Pygmy Swellshark 70 Sarawak Swellshark 70 144 Sawfish 140–145 Sawfishes Scalloped Hammerhead 136 112 School Shark Shark Ray 146 146–147 Shark Rays Sharpnose Stingray 188, 192 Short-tail Cownose Ray 254 262 Shortfin Devilray 64 Shortfin Mako 262 Shorthorn Devilray 160 Shortlip Electric Ray Shortlip Numbfish 160 266–267 Shortnose Chimaeras 156 Shovelnose Ray 154–159 Shovelnose Rays 150, 152, 156 Shovelnose Shark Sicklefin Weasel Shark 80 Silky Shark 98 Silver Chimaera 266 Silvertip Shark 86 36 Sixgill Shark 170–177 Skates 166–169 Sleeper Rays 56 Sleepy Shark Slender Bambooshark 50 122 Slender Dogshark 134 Slender Hammerhead Sliteye Shark 122 Smalleye Stingray 182 Smallfin Gulper Shark 40 94 Smoothfang Shark Smoothnose Wedgefish 150 264 Smoothtail Mobula 82 Snaggletooth Shark 64 Snapper Shark

236 Solander’s Ray Sorrah Shark 112 66 Spatulasnout Catshark 234 Speckle Stingray 260 Spinetail Devilray 260 Spinetail Mobula Spinner Shark 94 56 Spitting Shark 166 Spot-tail Electric Ray Spot-tail Shark 112 Spot-tail Sleeper Ray 166 54 Spotted Catshark 244 Spotted Eagle Ray Starspotted Smoothhound 74 Starynose Stingray 228 182–237 Stingrays Straight-tooth Weasel Shark 84

T Taiwan Guitarfish Tawny Nurse Shark Tawny Shark Thickskin Shark Thickspine Giant Stingray Thornback Ray Thorny Ray Thresher Sharks Tiger Shark Tubemouth Whipray

158 56 56 108 182 174 236 62–63 114 202

U Undulate Whipray

200

V Violet Stingray

230

W Weasel Shark Weasel Sharks Wedgefishes

80 78–85 148–153

292


Western Highfin Spurdog 38 Whaler Sharks 88–133 Whale Shark 60 60–61 Whale Sharks White-edge Freshwater Whipray 210 126 White-eye Shark 210 White-rim Stingray Whitecheek Shark 96 Whitefin Smoothhound 76 212 Whitenose Whipray 148 Whitespot Ray 148 Whitespot Shovelnose Ray Whitespotted Bambooshark 52 Whitespotted Eagle Ray 244 148 Whitespotted Guitarfish Whitespotted Wedgefish 148 Whitespotted Whipray 192 194 Whitetail Stingray 194 Whitetail Whipray Whitetip Reef Shark 132 132 Whitetip Shark 96 Widemouth Blackspot Shark 142 Wide Sawfish Winghead Shark 134 46–47 Wobbegongs

Y Yellow-spotted Skate Yellowspotted Skate

176 174

Z Zebra Bullhead Shark Zebra Hornshark Zebra Port Jackson Shark Zebra Shark Zebra Sharks Zonetail Butterfly Ray

44 44 44 58 58–59 240

293

Indonesian common names index C Cawang Genul 260 Cawang Kalung 256 100 Cucut Bekeman 40 Cucut Botol 50, 52 Cucut Dolok 140 Cucut Gergaji 86, 88, 90, 94, 96, 98, Cucut Lanjaman 102, 108, 110, 112 74, 76 Cucut Londer 154 Cucut Mandrong 36 Cucut Meong 62 Cucut Pedang 124 Cucut Selendang 68 Cucut Tokek

H Hiu 44, 92, 148 Hiu Aer 124 74, 76 Hiu Air 64 Hiu Anjing 104 Hiu Aron 146 Hiu Barong 54 Hiu Batu 58 Hiu Belimbing 134, 136, 138 Hiu Bingkoh 56 Hiu Bisu 132 Hiu Bokem 52, 54 Hiu Bongo 40 Hiu Botol Danten 100 Hiu Buas 88, 90, 94, 96, 98, 100, Hiu Bujit 102, 106, 112, 120 Hiu Capil 134, 136, 138 134, 136, 138 Hiu Caping 156 Hiu Cermin 132 Hiu Coklat

56 Hiu Gedebong Hiu Gedok 54, 56 142, 144 Hiu Gergaji 266 Hiu Hantu 46 Hiu Jenggot 74, 76, 78, 80, 84 Hiu Kacang 64 Hiu Kakap 132 Hiu Karang Buas 124 Hiu Karet 102, 122, 130 Hiu Kejen 148, 150, 152 Hiu Kemenyan 42, 46 Hiu Kodok 124 Hiu Lalaek 86, 98, 102 Hiu Lanyam 46 Hiu Lepang 90, 98 Hiu Lonjor 148, 150, 152 Hiu Lontar 114 Hiu Macan 106 Hiu Mada 64 Hiu Mako 134, 136, 138 Hiu Martil 150, 152 Hiu Minsong 82 Hiu Monas 62 Hiu Monyet 114 Hiu Omas Hiu Parang 136, 138, 140, 142, 144 84 Hiu Pasir 60 Hiu Paus 78, 80, 126, 128 Hiu Pilus 126, 128 Hiu Pisang 86, 94, 126 Hiu Plen 266 Hiu Pogak 40 Hiu Senget 86 Hiu Sonteng 36 Hiu Tahu Putih 40 Hiu Taji 48, 50, 52, 54 Hiu Tekek

294


Hiu Tenggiri Hiu Teteri Hiu Tikus Hiu Tokek

64 108 62 68, 72

I Ikan Pari

184, 232

K Kluyu Blimbingan Kluyu Karang Kupu-kupu Brangkas

58 106 146

L Lampengan Lanyam Liongbun

260 112 148, 150, 152, 154

M Merak Bulu Minsong Mungsing Mungsing Jara

90, 94, 102, 112 148 98, 112, 126, 130 114

P Paitpait Pamprang Pangrum Parangpang Paredung Pari Pari Aer Pari Ayam Pari Babi Pari Bandrong Pari Batu Pari Bendera Pari Biasa Pari Bintang Pari Blentik

156 140 148, 150, 152 140 148, 150, 152 172, 180, 204, 208 196, 206 244 182, 234 156 192 222, 224, 226, 228 188 192 220

Pari Boro 186 Pari Burung 242, 244, 246, 248, 250, 252 236 Pari Duren 196 Pari Duri 252, 254 Pari Elang 156 Pari Gitar 226 Pari Gunung 186 Pari Jantung 156 Pari Kekeh 238, 240 Pari Kelelawar 232 Pari Kembang 256 Pari Kerbua 196, 218 Pari Kikir 220 Pari Kodok Pari Kontak 160, 162, 164, 166, 168 260, 262, 264 Pari Lambingan 262 Pari Lampengan 230 Pari Lampin 230 Pari Lemer 264 Pari Lempengan Pari Listrik 160, 162, 164, 166, 168 238, 240 Pari Lowo 202, 212 Pari Lumpur 172, 174 Pari Luncur 200, 214, 216 Pari Macan 214, 234 Pari Merica 190 Pari Minyak 206 Pari Pasir 240 Pari Payung 182, 194 Pari Sapi 198, 210 Pari Sungai 192 Pari Super 202, 212 Pari Tanjung 190 Pari Tembaga 232 Pari Totol 230 Payubek 248 Pe Lamping 212 Pe Pasung 148, 150, 152, 156 Petong

295

Indonesian common names index

Plampangan

256, 260, 264

T Tikusan Toka-toka Toka Toka Tuka-tuka

62 188 218 188

Y Yunbun Karang

146

296


Malaysian common names index B Bengkong Beroi

134 140

C Chimara Perak

266

I Ikan Tikus

266

J Jerung-mako Sirip Jerung Biru Jerung Gigi Cerakah Jerung Sapi Jerung Tenggiri

64 124 82 100 114

K Kan Sua Kemejan Kiampau

140 152 220, 232

P Pari 152, 184 Pari-karan Hitam 164 252 Pari-kelawar Jawa 146 Pari-kemejan Kepala Keras 154 Pari-kemejan Muncung Panjang 156 Pari-kemejan Muncung Pendek 148 Pari-kemejan Tompok Putih 248 Pari-lang Jalur 246 Pari-lang Tompok Putih 172 Pari-raja Tompok 208, 210 Pari Air Tawar 252 Pari Ayam 194 Pari Bakau 192, 216 Pari Batik 170, 174, 176 Pari Belanda

Pari Bendera 222 Pari Bintik Putih 192 206 Pari Bulat 222 Pari Daun 236 Pari Dedap 236 Pari Duri 236 Pari Durian 186 Pari Ekor Duri 182 Pari Gergasi 244 Pari Helang 230 Pari Hitam 164, 166 Pari Karan 168 Pari Karan Belangkas 232 Pari Karang 160 Pari Karan Tompok Besar 168 Pari Karas 234 Pari Kelabu 240, 256, 262 Pari Kelawar 238 Pari Kelawar Bintik 150 Pari Kemejan 188, 218 Pari Ketuka 220 Pari Lalat 242, 244, 252, 254 Pari Lang 244 Pari Lang Bintik Putih 250 Pari Lang Bunga 180 Pari Lembik Merah 164 Pari Letrik 220 Pari Macan 180 Pari Merah 226 Pari Mulut Kasar 202 Pari Mulut Tajam 196 Pari Naga 188 Pari Nyiru 222 Pari Nyonya 196, 212 Pari Pasir Pari Paus 256, 258, 260, 262, 264 190 Pari Pucat

297

Malaysian common names index

Pari Reben Pari Riman Pari Rimau Pari Serban Pari Sungai Pari Susun Pari Tanjung Pari Tembikar Pendek

232, 234 220 200, 214, 216, 220 164 198, 208, 210 252 188, 218, 220, 222 238, 240 64

S Sirip Lekuk

264

T Tanpa Duri Tuka-tuka

262 218

Y Yu 134 Yu-buntal Sarawak 70 52 Yu-cicak Bintik 54 Yu-cicak Insang Puih 48 Yu-cicak Lampai 50 Yu-cicak Tembaga 142 Yu-gergaji Seragam 144 Yu-gergaji Silih 74 Yu-jaras Bintik 134 Yu-tukul Palang 136 Yu-tukul Sanggul 136 Yu Bengkong 80 Yu Bintik Putih 48, 50, 52, 54, 56, 72 Yu Bodoh 92 Yu Borneo 70 Yu Buncit 70 Yu Buntal 58, 68 Yu Cicak 90 Yu Ekor Hitam 62 Yu Ekor Panjang 140, 142, 144 Yu Gergaji 140 Yu Gergaji Jarang

78 Yu Gigi Cangkuk Yu Gigi Lurus 84 114 Yu Giling 36 Yu Insang Enam 88 Yu Jalur Putih 74, 76 Yu Jaras 98, 102, 104, 130 Yu Jereh 64, 100 Yu Jerung 44, 58 Yu Kebut 70 Yu Kembong 146, 154, 158 Yu Kemejan 102, 106, 112 Yu Kepak Hitam 148, 150, 152, 154 Yu Kia-kia 108 Yu Kulit Tebal 134, 136 Yu Mata Jauh 122 Yu Mata Lekuk 38, 40 Yu Minyak 46 Yu Misai 104 Yu Muncung Keras 128 Yu Muncung Minyak 94 Yu Muncung Panjang 126 Yu Muncung Susu 106 Yu Nipah 130 Yu Padi 134, 136 Yu Palang Yu Parang 136, 138, 140, 142, 144 140, 142, 144 Yu Parangan 42, 156 Yu Pari 50, 80, 96, 98, 104, 110, 130 Yu Pasir 60 Yu Paus 68 Yu Pendek 54 Yu Punai 82 Yu Putih 58 Yu Rimau 134, 136 Yu Sanggul 56 Yu Semilang 106 Yu Sirip Hitam 120 Yu Sirip Lebar 132 Yu Sirip Putih

298

Yu Sungai Yu Tanduk Yu Tenggiri Yu Todak Yu Tokeh Yu Trihitam Yu Tukul


116, 118 134 114 140, 142, 144 50, 54, 58, 68 102 136, 138

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.

SHARKS AND RAYS OF BORNEO


SHARKS and RAYS of

BORNEO

Peter R. Last 4 William T. White 4 Janine N. Caira 4 Dharmadi 4 Fahmi 4 Kirsten Jensen 4 Annie P. K. Lim 4 B. Mabel Manjaji-Matsumoto 4 Gavin J. P. Naylor 4 John J. Pogonoski 4 John D. Stevens 4 Gordon K. Yearsley 4

borneo

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