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Rays of the World

Book · December 2016

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Rays of the World
Peter Last, William White, Marcelo de Carvalho, Bernard Séret,
Matthias Stehmann, Gavin Naylor (Eds)

The first illustrated guide to the over 630 known species of rays
found on the planet.
Rays are among the largest fishes and evolved from shark-like
ancestors nearly 200 million years ago. Rays of the World is the
first complete pictorial atlas of the world’s ray fauna and includes
information on many species only recently discovered by scientists
while undertaking research for the book. It includes all 26 families and
633 valid named species of rays, but additional undescribed species
exist for many groups. December 2016
Rays of the World features a unique collection of paintings of all 800 pages, Hardback
living species by Australian natural history artist Lindsay Marshall. ISBN: 9780643109131
This comprehensive overview of the world’s ray fauna summarises $220.00
information such as general identifying features and distributional
information about these iconic but surprisingly poorly known fishes. It
Available as an eBook
will enable readers to gain a better understanding of the rich diversity
of rays and promote wider public interest in the group.
Rays of the World is an ideal reference for a wide range of readers, Sales in Australia and
New Zealand only
including conservationists, fishery managers, scientists, fishers, divers, For rest of world sales contact:
students and book collectors. Cornell University Press
www.cornellpress.cornell.edu

k e y t o r A j I d g e n e r A 205 k e y t o r A j I d g e n e r A 207 210 R A Y S O F T H E W O R L D

19 KE Y TO R AJID GENER A 8. Snout long and narrowly pointed (fig. 15); end of
greatly enlarged clasper flattened and spatula-like (fig. SOUTHERN THORNY SKATE 19.1
The hardnose skates are very difficult to characterise based 18); temperate South Pacific and South-West Atlantic

S K AT E S
solely on external features because skeletal structures, ........................ Zearaja (4 species; fig. 15, pp. 360–363)
particularly the claspers, are more important in distin- Amblyraja doellojuradoi (Pozzi, 1935)
guishing groups. The following key is incomplete; clearly Snout short to moderately elongate; end of moderately
separating some genera but failing to distinguish others. large clasper narrowly rounded (fig. 19) or pointed (fig.
20) ....................................................................................... 9
Family Rajidae
For example, genera in couplet 14 could not be distinguished
so readers need to use distributional ranges (which largely 9. Long interdorsal and postdorsal tail sections (fig. 16);
do not overlap) to help identify species. Where a specimen end of clasper in all species, except O. ornata, with
P.R. Last, B. Séret, M.F.W. Stehmann & S. Weigmann fails to key out correctly, we suggest proceeding again using small soft spiny structure near its tip (component
the alterative option in couplets. The key below has incorpo- funnel) (fig. 21); Indo–West Pacific ..................................
rated some technical characters in order to distinguish .... Okamejei (in part, 12 species; fig. 16, pp. 302–313)
between genera (see the Glossary for further explanation). fig. 1 fig. 15
Short interdorsal and postdorsal tail sections (fig. 17);
Members of the family Rajidae, known as skates or hardnose skates, are small to very large rays (adults 33 cm to 1. Anterior margin of snout very obtuse (fig. 1); anterolat- clasper without component funnel; Australia ...............
eral margin of ventral disc with band of claw-like ...... Dentiraja (in part, 8 species; fig. 17, pp. 231–240)
more than 2 m TL) with a depressed body, almost circular to rhombic disc, and pectoral-fin apices broadly rounded
denticles (fig. 2); Western Atlantic ...................................
to angular. A firm, slender tail is well demarcated from the disc. The snout is more or less elongated and pointed, ............... Dactylobatus (2 species; fig. 1, pp. 229–230) 10. Median thorns on disc very large and with stellate
supported by a stiff rostral cartilage, and often has a short lobe at its tip. The anterior nasal flaps are expanded to bases (fig. 22); end of massive, broad clasper club-
Anterior margin of snout somewhat obtuse (fig. 7) to shaped and widened (fig. 24); cosmopolitan ..................
form an incomplete nasal curtain. These flaps usually reach the mouth but their posterior margins are not joined like acute (fig. 10); no enlarged claw-like denticles on ante- .................. Amblyraja (8 species; fig. 25, pp. 210–217)
some other ray groups, such as the stingrays. Pelvic fins are notched with distinct anterior and usually larger posterior rolateral margin of ventral disc .................................... 2
fig. 2 Median thorns on disc (if present) small to moderate in

Editors: Peter R. Last, William T. White, Marcelo R. de Carvalho,

lobes. Two small dorsal fins are located near the end of the tail and their bases are often joined. The caudal fin is 2. Anteriormost extension of pectoral-fin skeleton abut- size and without stellate bases (fig. 23); end of clasper
greatly reduced in size with the lower lobe (when present) smaller than the upper lobe. The skin is sometimes naked, ting or nearly abutting tip of snout, not separated from narrowly rounded (fig. 19) or pointed (fig. 20) ........... 11
pectoral fin ray
snout by largely translucent area (fig. 3); tip of snout
but the disc and tail are usually partly covered with dermal denticle patches, particularly along the anterior disc pliable ................................................................................ 3 narrowly rounded
margins. Thorns, usually present on the upper disc of juveniles (and most adults), are variably located on the orbital spatula-like LC
Anteriormost extension of pectoral-fin skeleton fig. 16
rims (around upper half of eye), nuchal (nape) and scapular (shoulder) regions, and along the median disc and tail distinctly separated from tip of snout by semi-translu- IDENTIFICATION. Meduin-sized skate with a weak
(additional lateral thorn rows on the tail of several species). Adult males have a well-defined, longitudinally arranged cent area (fig. 4); tip of snout rather firm and not pliable fig. 19 rhombic disc (width ~1.3 times length), snout short and stiff
............................................................................................. 4 fig. 18 with firm rostral cartilage, medium-sized eyes, prominent
patch of retractable alar thorns on the mid-outer regions of each pectoral fin, and some species have additional club-shaped
3. Triangular patch of thorns present on nuchal and pointed thorns on disc, tail short (~72% in precloacal length in
anteriorly positioned thorn patches, known as malar thorns. The family includes at least 154 valid described species

Bernard Séret, Matthias F. W. Stehmann and Gavin J. P. Naylor

shoulder region (fig. 7); length of anterior pelvic-fin adults), and upper disc much darker than undersurface. Disc
in 17 genera: Amblyraja, Beringraja, Breviraja, Dactylobatus, Dentiraja, Dipturus, Hongeo, Leucoraja, Malacoraja, fig. 3 fig. 4 rather thick, somewhat flabby, anterior margin weakly
lobe 65–80% of posterior lobe (fig. 5); tail width at base
Neoraja, Okamejei, Orbiraja, Raja, Rajella, Rostroraja, Spiniraja and Zearaja. However, several unnamed genera fig. 20 fig. 24 undulate (strongly so in adult males); apex abruptly angular.
slightly greater than orbit diameter; Western Atlantic skeleton  of head
....................... Breviraja (5 species; fig. 7, pp. 224–228) shape of right claspers Head broad; snout length ~2.3 times and interorbital space
and species are known to exist, and some species are likely to be re-assigned to other genera. Specialised skeletal
subequal to orbit length respectively; snout tip projecting
features, particularly those of the clasper cartilages, are important in distinguishing genera and species. The rostral No triangular patch of thorns on nuchal and shoulder forward slightly as broad, bluntly rounded lobe. Mouth
cartilage varies from being moderately slender to stiff and stout from its base to the snout tip (except in Breviraja), region (nuchal thorns and 1 or 2 scapular thorns are
wide; tooth rows in upper jaw ~30–38. Skin of upper surface
sometimes present, but these do not form a distinct
and its base extends anteriorly to, or noticeably forward of, the leading edges of the nasal capsules. The anteriormost rough, covered with coarse denticles; ventral surface
triangular patch) (fig. 8); length of anterior pelvic-fin
smooth. Thorns on upper disc massive, more enlarged in
part of pectoral-fin skeleton is well separated from the tip of snout (by a semi-translucent area, except in Breviraja), lobe 80–100% of posterior lobe (fig. 6); tail width at base fig. 17 SIZE. Attains ~69 cm TL. Hatches from egg case at ~9 cm
young than adults; usually 3 pairs near eyes, 1–2 on nape,
and the ventral terminal cartilage of the clasper has a characteristic sharp, lateral edge forming an external less than orbit diameter; Eastern and North-West TL.
fig. 5 fig. 6 2–3 on each shoulder; median row of 12–15 from shoulder to
Atlantic ........... Neoraja (5 species; fig. 8, pp. 297–301)
component, the shield (see Glossary). Most skates are demersal in cold water on continental slopes and abyssal first dorsal fin; malar thorns well developed; none HABITAT AND BIOLOGY. South-West Atlantic and
interdorsally; scattered thornlets on snout and pectoral fins. South-East Pacific; Uruguay to southern Chile, including
plains to depths of more than 4000 m, but some temperate and polar species occur inshore on continental shelves,
Tail rather robust, tapering from base to tip; lateral folds Falkland Islands. Primarily on mid- and outer continental
and 1 lives primarily in brackish/freshwater habitats. The group is cosmopolitan with representatives in all oceans. clasper
extend along most of tail. Pelvic-fin anterior lobe much shelf and upper slope at 50–640  m depths, occasionally
Some larger, more mobile benthopelagic species are wide-ranging, but most are primarily benthic with restricted shorter than posterior lobe. Dorsal fins small, separated caught deeper (to 1000  m). Feeds on benthic invertebrates
distributions, often within small geographical confines. Skates are mainly carnivorous, feeding on benthic invertebrates slightly; caudal fin very small. Clasper of adult male very and small bony fishes.
fig. 21
broad but clearly not reaching first dorsal fin.
(i.e. crustaceans, polychaetes, bivalve molluscs, and cephalopods) and fishes living on or in sediments of the seafloor. SIMILAR SPECIES. Other species of Amblyraja in the
All species are oviparous, with females laying their eggs in cases (also known as mermaid’s purses). Egg cases are COLOUR. Medium to dark brown above, with indistinct region are either almost entirely dark on both disc surfaces
pattern of scattered darker spots and blotches; usually with or have dark blotches on the undersurface. Validity of this
rectangular and flattened in shape with a tough protective outer coating of keratin, and often a horny structure at
pair of large pale blotches near front of eyes and at pectoral species has been confirmed by molecular data, but the
each corner. Embryos develop slowly, taking up to 2 years to hatch. Skates are caught worldwide, mainly as bycatch triangular
axils. Undersurface predominantly whitish, often with some taxonomy of some other species in the genus needs further
thorn patch side view of
of trawl fisheries. The flesh is edible and some species are retained as food for humans. fig. 7 fig. 8 fig. 22 median thorns fig. 23 fig. 25 grey specks and spots on disc and tail. investigation.

011601 Rays 3pp.indd 204 22/08/2016 09:09:18.893 011601 Rays 3pp.indd 205 22/08/2016 09:09:20.377 011601 Rays 3pp.indd 207 22/08/2016 09:09:24.311 011601 Rays 3pp.indd 210 22/08/2016 09:09:30.606
 CONTENTS FEATURES
Foreword • Colour paintings by renowned natural history
Acknowledgements artist Lindsay Marshall
1. Introduction • Covers all 26 families and 633 valid named species
2. Phylogeny and classification of rays • Describes characteristics of each ray family, as well
3. Human interactions as providing identification details and distribution
maps for each individual species
4. Ray conservation
5. About this book
6. Glossary EDITOR AFFILIATIONS
7. Key to families of living rays
Peter Last: CSIRO National Research Collections,
8. PRISTIDAE (Sawfishes) Australian National Fish Collection (former)
9. RHINIDAE (Wedgefishes) William White: Australian National Fish Collection
10. RHINOBATIDAE (Guitarfishes) Marcelo de Carvalho: University of São Paulo,
11. GLAUCOSTEGIDAE (Giant guitarfishes) American Museum of Natural History
12. TRYGONORRHINIDAE (Banjo rays) Bernard Séret: National Museum of Natural History,
13. PLATYRHINIDAE (Fanrays) Paris (former)
14. ZANOBATIDAE (Panrays) Matthias Stehmann: Zoological Museum of Hamburg
15. NARCINIDAE (Numbfishes) (retired)
16. NARKIDAE (Sleeper rays) Gavin Naylor: College of Charleston
17. HYPNIDAE (Coffin rays)
18. TORPEDINIDAE (Torpedo rays)
ALSO AVAILABLE
19. RAJIDAE (Skates)
20. ARHYNCHOBATIDAE (Softnose skates)
Full table of contents available: www.publish.csiro.au/book/7053

31. AETOBATIDAE (Pelagic eagle rays)

32. RHINOPTERIDAE (Cownose rays)
33. MOBULIDAE (Devilrays)
Suggested reading
Checklist of the world’s rays
Scientific names index
Common names index

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