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Field Guide to the

Identification of Mobulid
Rays (Mobulidae):
Indo-West Pacific

Guy Stevens - Director

[email protected] / www.mantatrust.org
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Overview
The guide is intended to be used as a resource for both scientists and members of the general public who
have a keen interest in identifying and learning more about the Mobulidae family, which contains the
manta and devil rays. At present the taxonomy of this family of batoid rays is poorly understood, with
many questions remaining over the true distinction between, and validity of, the 11 described species
currently recognized globally. Due to the scarcity of data available for the mobula species which are
restricted to the Atlantic and Eastern Pacific, this guide focuses only on the 7 mobulid species which occur
in the Indo-West Pacific. Close underwater encounters of the smaller and more elusive devil rays (mobula)
are rare, making identification during these brief
encounters difficult without prior knowledge of the key
identifiable features of each of these species. Scientists
working in the field examining dead specimens at fish
markets often find the current literature unsatisfactory
for accurate identification.

This guide brings together a comprehensive pool of

images, knowledge and data pieced together by
scientists working with the Manta Trust in the Indo-
West Pacific to create the first field guide to the
identification of all the species which occur in this A Scientist measuring a dead Oceanic Manta Ray
region. It is hoped that the creation of this guide will aid (Manta birostris) at a fish market in Sri Lanka.
future scientific research, while simultaneously
increasing the general public’s knowledge of these animals, which together, will lead to better
conservation and protection measures for these fascinating animals.
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Contents Page

• Introduction to Mobulid Rays (Page 4-6)

o Taxonomy and Family Characteristics
o Mobulid Species of the Indo-West Pacific
o Basic Mobulid Anatomy
o Sexual Dimorphism
o Closely Related Species

• Threats to Mobulid Rays (Page 7-8)

o Global Mobulid Fisheries
o Why So Vulnerable?
o The Gill Raker Trade

• Key: Mobulid Species of the Indo-West Pacific (Page 9-10)

• Mobulid Identification Guide (Page 11-18)

• Acknowledgements & Thanks (Page 19)

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Introduction to Mobulid Rays

Taxonomy & Family Characteristics
Manta and mobula rays belong to the Suborder Myliobatoidei, which contains all of the Eagle Rays
(Myliobatidae), Cownose Rays (Rhinopteridae) and the Mobulid Rays. In total this suborder contains about
40 species which are characterized by diamond shaped bodies and wing-like pectoral fins which they use to
propel themselves through open water. Eagle Rays and Cownose Rays all feed on the seabed, using their
mouths to dig amongst the substrate in search of buried Molluscs and crustaceans, while the mobulid rays
have truly reverted back to a completely pelagic way
of life.

Scientifically the mobulid rays belong to one family

called Mobulidae, which contains two Genera; the
Mobula Rays (9 species) and the Manta Rays (2
species). All are filter feeders, using their mouths and
modified gill rakers to strain plankton and small fishes
from the water. In general, mobula rays are much
smaller than the mantas and can be distinguished by
morphological differences in their mouths and
cephalic fins (‘head fins’). Mobula rays have a bottom
jaw which is undercut, so that when their mouths are Large cephalic fins of a Reef Manta Ray (Manta alfredi),
closed the edge of the lower jaw rests much further scoop up tiny planktonic prey into their forward facing,
terminal mouths.
back than the upper, whereas manta ray's jaws are
aligned evenly. The other differentiating anatomical feature is the shape of the cephalic fins, which when
rolled up look like horns projecting off their heads, hence the name “Devil Rays”. The primary function of
these fins is to help funnel planktonic food into the gaping mouths of the ray's when they are feeding.
Unfurled, the mobula ray's cephalic fins are just a flap, but in the mantas these fins unravel to form much
larger paddle-like structures which touch in the centre to form a complete funnel around the mantas
mouth.

Very little is known about the mobula rays which, unlike the
mantas, are generally very shy in the presence of divers making
it hard for scientists to observe behaviour in the wild. Like the
mantas they are found throughout tropical and sub-tropical
oceans, sometimes aggregating in vast shoals of many hundreds
which probably come together to seek safety in numbers. These
aggregations may also occur more frequently at breeding times
and it is not uncommon to encounter these large groups
attracting attention from those at the surface as the rays leap
The mouths of all Devil Rays, like this Bent-fin several metres out of the water.
Devil Ray (Mobula thurstoni), are all under
slung, unlike the mantas, which face forward.
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Mobulid Species of the Indo-West Pacific

Basic Mobulid Anatomy

Ventral Surface
1 2

5
6 7

4
3

Mobulid Anatomy
11 12
8 10 1. Disc Width (DW)
2. Tail
3. Mouth
st
4. 1 Gill Slit
5. Spiracle
6. Cephalic Fin
7. Eye
8. Dorsal Fin
9. Pelvic Fin
10. Tail Spine
11. Dorsal Ridge 13
12. Neck Dorsal Surface
9 2 13. Pectoral Fin
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Sexual Dimorphism

1 1 2

Female Mature Male Immature Male

4
Sexual Dimorphism

1. Pelvic fin.
2. Enlarged claspers in mature male.
3. Undeveloped claspers in juvenile male.
4. Pregnancy bulge in mature female.

Closely Related Species

Spotted Eagle Ray – Aetobatus narinari Javanese Cow-nose Ray – Rhinoptera javanica
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Threats to Mobulid Rays

Global Mobulid Fisheries
One of the first country’s to commercially fish
their Mobulid populations was Mexico, when in
the early 1980’s, fishermen in the Sea of Cortez
switched from subsistence and bycatch fishing
of the locally abundant oceanic manta and
mobula ray species to directed target fisheries.
Using harpoons to impale the surface feeding
animals, and gill nets to entangle and drown
them, the rays were easy targets, and their
numbers soon began to plummet. The giant
carcases were towed back to the beaches
where only the choicest flesh was sold for A pile of Spine-tail Devil Ray (Mobula japanica) in Sri Lanka,
consumption, while the remainder was often fished for their gill rakers which are sent to Asia to be used in
Chinese Medicines.
used as bait in lobster pots, or simply discarded.
Within just a decade populations of the large mobulid ray species within the Sea of Cortez were virtually
wiped out, and the fishery collapsed. It was not until 2005 that the Mexican Government finally passed
legislation protecting the oceanic manta rays in Mexican waters, but by then the damage had already been
done. Even today, after nearly two decades of protection, very few mantas are recorded in this area and
those that are still fall victim to illegal fishing or bycatch.

Many other countries have also targeted their mobulid populations with similar results, switching from
local artisanal fisheries to commercial export fisheries wherever a market for their products can be found.
The Philippines, Indonesia, Mozambique, Madagascar, India, Pakistan, Sri Lanka, Brazil and Tanzania have
all chosen the same path as Mexico, with similar trends of population declines reported in many of these
countries. Yet out of all these countries, only in the Philippines are there now official laws in place to
protect these vulnerable species.

Globally Threatened Status of Mobulid Species Found in the Indo-West Pacific

Common Name Scientific Name IUCN “Red List” Status
Oceanic Manta Ray Manta birostris Near Threatened (NT)
Reef Manta Ray Manta alfredi -
Spine-tail Devil Ray Mobula japanica Near Threatened (NT)
Sickle-fin Devil Ray Mobula tarapacana Data Deficient (DD)
Bent-fin Devil Ray Mobula thurstoni Near Threatened (NT)
Short-fin Pygmy Devil Ray Mobula kuhlii Data Deficient (DD)
Long-horned Pygmy Devil Ray Mobula eregoodootenkee Near Threatened (NT)
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Why So Vulnerable?
The reason for these rapid declines is simple; Mobulid
rays live for a long time and reproduce infrequently.
They are large animals with few natural predators that
have long gestation periods which result in the birth of
just a single pup (most of the time), which themselves
are likely to take over a decades to reach sexual
maturity. As a result of these life history strategies, and
like many other large marine animals, manta ray
populations simply cannot survive or sustain any
commercial fisheries for an extended period of time.
Drying manta and devil ray gill rakers in Sri Lanka, Any target fishery which annually removes even a
destined for China.
relatively small percentage of the breeding adults results
in a rapid decline in the overall population within just a few years, as the remaining mature individuals
simply cannot breed fast enough to replace the loses. This is why, even with complete protection from
anthropogenic threats, an overfished population of manta rays will take decades to recover to its natural
state. A situation which, in the realities of today’s global fisheries management and protective
enforcement (or lack thereof), is never likely to happen to these populations which have already been
overfished.

The Gill Raker Trade

While the meat from the fished rays is often consumed locally,
the driving factor behind the dramatic increases in fishing
pressure for mobulid rays seen throughout South-east Asia
and Eastern Africa in recent decades is the growing demand
for the mantas branchial filaments, or gill rakers, which are
dried and exported for the Asian medicinal market. The dried Gill rakers being chopped out of a large
rakers are crushed into a powder which is added to a soup or Oceanic Manta Ray (Manta birostris).
broth, along with crushed pipefishes, ginseng and other ingredients, to be used as a treatment for a variety
of ailments. The gill rakers are marketed as being; anti-inflammatory, clearing away heat and toxic
material, and eliminating stasis to activate blood circulation. There is
absolutely no scientific proof to back up these claims, and there are
not even any records in the traditional Chinese medicinal texts which
list manta or mobula rays gills as being used in this way. So it appears
as though this is a fairly new product, clever marketing of a readily
available and cheap bycatch source from the fisheries. Marketed as a
“medicine” on the basis that because mobulid rays are capable of
filtering the water to catch their food, their gill rakers when consumed
Gill Raker Soup – Peng Yu Sai. can also filter and remove toxins from our human bodies.
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Key to the Identification of Mobulid Rays

(Mobulidae) in the Indo-West Pacific
IS THE MOUTH TERMINAL?

YES NO

MANTA:
Terminal Mouth
MOBULA:
Ventral Mouth

MANTA RAYS

KNOB-LIKE BULGE AT THE BASE OF THE TAIL JUST POSTERIOR TO THE DORSAL FIN?
VENTRAL SPOT PATTERN CLUSTERED AROUND THE LOWER ABDOMINAL REGION?

YES NO

REEF MANTA RAY:

Manta alfredi
(DW up to 4.5m/14.8ft)

OCEANIC MANTA RAY:

Manta birostris
(DW large up to 6.8m/22.3ft)

GO TO DEVIL RAYS
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DEVIL RAYS

IS THE TAIL LONGER THAN THE DISC WIDTH?

YES
IS THERE A SPINE AT THE BASE OF THE TAIL?

NO

IS THE SPIRACLE ABOVE THE MARGIN OF THE PECTORAL FIN

SPINE-TAIL DEVIL RAY:
SITUATED UNDER A DISTINCT RIDGE?
Mobula japanica
AND SITUATED POSTERIORALLY TO THE INTERSECTION OF THE (DW up to 3.1m/10.2ft)
PECTORAL FIN WITH THE HEAD?
NO
IS THE DORSAL FIN PLAIN IN COLOUR?

YES DOES THE PECTORAL FIN'S ANTERIOR MARGIN

HAVE A DISTINCTIVE DOUBLE CURVATURE?

YES
SICKLE-FIN DEVIL RAY:
Mobula tarapacana
(DW up to 3m/9.8ft)

NO

DOES THE ANIMAL HAVE A LONG-NECKED APPEARANCE?

BENT-FIN DEVIL RAY:
VERY LONG CEPHALIC FINS? Mobula thurstoni
(Length, from the tip of each fin to the corner of the mouth, is more (DW up to 2.2m/7.2ft)
than 16% of the total disc width)

NO YES

SHORT-FIN PYGMY DEVIL RAY:

Mobula kuhlii
(DW up to 1m/3.3ft)

LONG-HORNED PYGMY DEVIL RAY:

Mobula eregoodootenkee
(DW up to 1m/3.3ft)
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Mobulid Identification Guide

Manta Rays

• Oceanic Manta Ray – Manta birostris (Page 12)

• Reef Manta Ray – Manta alfredi (Page 13)

Mobula Rays

• Spine-tail Devil Ray – Mobula japanica (Page 14)

• Sickle-fin Devil Ray – Mobula tarapacana (Page 15)

• Bent-fin Devil Ray – Mobula thurstoni (Page 16)

• Short-fin Pygmy Devil Ray – Mobula kuhlii (Page 17)

• Long-horned Pygmy Devil Ray – Mobula eregoodootenkee (Page 18)

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Oceanic Manta Ray - Manta birostris (Walbaum, 1792)

6 7

3 4
8 5

Species Characteristics
Size: Disc width large, up to 6.8m/22.3ft
Range: Circumtropical; Often in association with offshore oceanic islands
Distinctive Features:
1. Terminal mouth.
2. Knob-like bulge at base of tail.
3. Cephalic fins large and unfurl to meet together in centre of the mouth.
4. Ventral spot pattern clustered around lower abdominal region.
5. Trailing underside edge of pectoral fins usually shaded black.
6. Gill covers often with black shading/flaring.
1
7. Inside of mouth and cephalic fins often coloured black.
8. White shoulder markings form two mirror image right angled triangles
which create the letter ‘T’ in black across the top of the head.

Black Colour
Morph

2
Gil Rakers
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Reef Manta Ray - Manta alfredi (Krefft, 1868)

7

5 8
1&6

Species Characteristics
Size: Disc width up to 4.5m/14.8ft
Range: Circumtropical; Often in association with warm water coral reefs
Distinctive Features:
1. Terminal mouth.
2. Slight depression at base of tail; no bulge.
3. Cephalic fins large and unfurl to meet together in centre of the mouth.
4. Often ventral spots between the branchial gill slits.
5. Ventral spots spread across trailing edge of pectoral fins and abdominal region. 2
6. Inside of mouth and cephalic fins usually pale (excluding black morph).
7. Dorsal markings more varied than M. birostris, with some individuals almost completely
white across the whole dorsal surface, while others are completely black.
8. Transition between the white and black markings on the dorsal surface is blurred along
the colour boundary (unlike M.birostris which has a more defined boundary), forming
more of a ‘Y’ shaped pattern across the animals head and down the centre of its back.

Black Colour
Morph

7
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Spine-tail Devil Ray - Mobula japanica (Müller & Henle, 1841)

6
2

4 3
7

Species Characteristics
9 Size: Disc width up to 3.1m/10.2ft
Range: Probably circumglobal in temperate and tropical waters
Distinctive Features:

1. Ventral mouth.
2. White-tipped dorsal fin.
3. Spine at the base of the tail.
4. Very long tail, much longer than disc width (whip-like).
5. Coloration: general deep blue-mauve and purple on dorsal
surface, bright white across entire ventral surface.
6. Top of the head has a thick dark black band which stretches
from eye-eye (fades when animal is dead).
7. White ventral markings wrap up behind, and above, the eyes to
10
meet the black dorsal head band.
8. Tail is ventrally flattened (ovoid) near the base of the dorsal fin.
9. Tail is covered in distinctive rows of tiny white bumps
(tubricles) along either side.
10. The spiracle is above the margin of the pectoral fin near where
the fin meets the body; and under a distinct ridge.

Gil Rakers

1
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Sickle-fin Devil Ray - Mobula tarapacana (Philippi, 1892)

8 13
Species Characteristics
Size: Disc width up to 3m/9.8ft
Range: Circumtropical; Oceanic
Distinctive Features:
1. Ventral mouth.
3
2. Long-necked appearance.
3. Long cephalic fins.
4. Plain colored dorsal fin.
5. No spine.
6. Tail is shorter than disc width.
7. Distinctive bicoloured gill rakers.
8. Dorsal surface entirely olive green-brown in colour.
9. Spiracle is above and behind the margin of the pectoral fin where it
joins the body; and under a distinct ridge.
10. The trailing edge of the Pectoral fins are distinctly falcate, or crescent 10
moon-shaped, hence the name ‘Sickle-fin Mobula Ray’.
11. Trailing underside region of pectoral fins shaded grey, while anterior
region is white; with a zigzagged messy margin between the two areas.
12. Gill covers often with grey shading/flaring.
13. Distinctive pronounced ridge along dorsal midline.
1

11
5

Gil Rakers

12

7
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Bent-fin Devil Ray - Mobula thurstoni (Lloyd, 1908)

9 11

Species Characteristics
Size: Disc width up to 2.2m/7.2ft
Range: Probably circumglobal in tropical and subtropical waters
Distinctive Features: Gil Rakers
1. Ventral mouth.
2. White-tipped dorsal fin.
3. No spine at the base of the tail.
4. Tail long; equals disc width in length when fully intact.
5. Short necked appearance.
6. Short cephalic fins: length, from the tip of each fin to the corner of
the mouth, is less than 16% of the total disc width.
7. White ventral markings do not extend above the eyes.
8. Tail base dorso-ventrally compressed close to the dorsal fin.
9. Pectoral fins anterior margin have a distinctive double curvature.
10. Spiracle small in size, sub-circular and below the margin of the 7 10
pectoral fin where it meets the body.
11. Coloration: general deep blue-black on dorsal surface, white ventral
surface anteriorally, while the distal half has a silver-brown sheen.
12. The top of the head has a dark band which stretches across the
head behind the eyes (fades when animal is dead).

5
3&8

1 11 2
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Short-fin Pygmy Devil Ray - Mobula kuhlii (Müller & Henle, 1841)

3&5

8
Species Characteristics
Size: Disc width up to 1.2m/3.9ft
9
Range: Distributed throughout the coastal continental waters of
the tropical Indian Ocean and Western Central Pacific
Distinctive Features:

1. Ventral mouth.
2. White-tipped dorsal fin (not all specimens, or regions).
3. No spine at the base of the tail.
4. Tail shorter than disc width and counter-shaded.
5. Base of tail quadrangular (square) in section.
11 6. Short necked appearance.
7. Short cephalic fins: length, from the tip of each fin to the
corner of the mouth, is less than 16% of the total disc width.
8. White ventral markings do not extend above the eyes.
9. Spiracle very small in size, sub-circular and below the margin
of the pectoral fin where it meets the body.
10. Coloration: mauve to chocolate brown and grey dorsally,
ventral surface usually plain white, although distal halve of
pectoral fins sometimes shaded dark brown-black.
11. Top of the head has a narrow brown-black band which
6 stretches across the head behind the eyes (fades when
animal is dead).

7
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Long-horned Pygmy Devil Ray - Mobula

eregoodootenkee (Bleeker, 1859)
4
1 11

2
9

8
Species Characteristics
3&5
Size: Disc width up to 1m/3.3ft
Range: Widely distributed throughout the coastal 9
continental waters of the tropical Indo-West Pacific
Distinctive Features:

1. Ventral mouth.
2. White-tip to dorsal fin in most specimens, with
variations possibly occurring between regions.
3. No spine at the base of the tail.
4. Tail shorter than disc width.
5. Base of tail quadrangular (square) in section.
6. Long-necked appearance.
7. Long cephalic fins: length, from the tip of each fin
to the corner of the mouth, is more than 16% of
7 6
the total disc width.
8. Spiracle very small, sub-circular and below the
margin of the pectoral fin where it meets the body. 9 10
9. Coloration: Chocolate brown dorsally, white
ventrally, with a patch of the dorsal coloration
extending down onto the ventral area along the
leading edge of both pectoral fins at the mid-point.
10. Pectoral fins have a dark brown strip which runs
along the leading edge of each fin.
11. The angle of the leading edge of the pectoral fins
sweeps back from the head more acutely than in
the other small mobula species.
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Acknowledgements & Thanks

The Manta Trust would like to thank all those people who contributed their knowledge, time and images to
the creation of this guide. Without their help and assistance this guide would not have been possible. I
would like to say a special thanks to the Save Our Seas Foundation for supporting my research, enabling
me to focus my time and effort on this guide's completion. I would also like to thank the Manta Ray of
Hope project for supplying market research data and numerous images. Thanks also to the IUCN of the
species distribution maps.

All images are copyright to: Guy Stevens, Thomas Peschak, Daniel Fernando, Marloes Poortvliet, Paul Hilton, Owen
Bruce, Alec Moore, AIMS, Carlos Minguell, Marc Sentis, Andy Green & Mark Fuller.