Cambio Dasyatidae

Zootaxa 4139 (3): 345–368 ISSN 1175-5326 (print edition)
http://www.mapress.com/j/zt/
Copyright © 2016 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://doi.org/10.11646/zootaxa.4139.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:DCF4220B-4A73-407C-837C-54DEAE29F435
A revised classification of the family Dasyatidae (Chondrichthyes:

Myliobatiformes) based on new morphological and molecular insights
PETER R. LAST1,4, GAVIN J.P. NAYLOR2 & B. MABEL MANJAJI-MATSUMOTO3

1
CSIRO National Research Collections Australia, Australian National Fish Collection, Castray Esplanade, Hobart, TAS, 7001, AUS-
TRALIA. E-mail: peter.last@csiro.au
2
Department of Biology, College of Charleston, Charleston, SC 29401, USA. E-mail: gjpnaylor@gmail.com
3
Endangered Marine Species Research Unit, Borneo Marine Research Institute, Universiti Malaysia Sabah, 88400 Kota Kinabalu,
Sabah, MALAYSIA.. E-mail: mabel@ums.edu.my
4
Corresponding author
Abstract
The higher-level taxonomy of the stingrays (Dasyatidae) has never been comprehensively reviewed. Recent phylogenetic
studies, supported by morphological data, have provided evidence that the group is monophyletic and consists of four ma-
jor subgroups, the subfamilies Dasyatinae, Neotrygoninae, Urogymninae and Hypolophinae. A morphologically based re-
view of 89 currently recognised species, undertaken for a guide to the world’s rays, indicated that most of the currently
recognised dasyatid genera are not monophyletic groups. These findings were supported by molecular analyses using the
NADH2 gene for about 77 of these species, and this topology is supported by preliminary analyses base on whole mito-
chondrial genome comparisons. These molecular analyses, based on data generated from the Chondrichthyan Tree of Life
project, are the most taxon-rich data available for this family. Material from all of the presently recognised genera (Dasya-
tis, Pteroplatytrygon and Taeniurops [Dasyatinae]; Neotrygon and Taeniura [Neotrygoninae]; Himantura and Urogymnus
[Urogymninae]; and Makararaja and Pastinachus [Hypolophinae]), are included and their validity largely supported. Ur-
ogymnus and the two most species rich genera, Dasyatis and Himantura, are not considered to be monophyletic and were
redefined based on external morphology. Seven new genus-level taxa are erected (Megatrygon and Telatrygon [Dasyati-
nae]; Brevitrygon, Fluvitrygon, Fontitrygon, Maculabatis and Pateobatis [Urogymninae], and an additional three (Bathy-
toshia, Hemitrygon and Hypanus [Dasyatinae]) are resurrected from the synonymy of Dasyatis. The monotypic genus
Megatrygon clustered with ‘amphi-American Himantura’ outside the Dasyatidae, and instead as the sister group of the
Potamotrygonidae and Urotrygonidae. Megatrygon is provisionally retained in the Dasyatinae pending further investiga-
tion of its internal anatomy. The morphologically divergent groups, Bathytoshia and Pteroplatytrygon, possibly form a
single monophyletic group so further investigation is needed to confirm the validity of Pteroplatytrygon. A reclassification
of the family Dasyatidae is provided and the above taxa are defined based on new morphological data.
Key words: Dasyatidae, stingrays, classification, group review, subfamilies, new genera, Brevitrygon, Fluvitrygon, Fon-
titrygon, Maculabatis, Megatrygon, Pateobatis, Telatrygon, resurrected genera, Bathytoshia, Hemitrygon, Hypanus
Introduction
The phylogenetic relationships and higher-level structure of the batoids has been the subject of much recent debate
since the first modern, morphology-based classification by Compagno (1973). Compagno’s initial phenetic study
was followed by an expanded cladistic analysis (Compagno, 1977). Subsequent morphological studies (e.g.
Lovejoy, 1996; Carvalho, 1996; McEachran et al., 1996; McEachran & Aschliman, 2004) were followed by a
sequence of molecular studies (e.g. Dunn et al., 2003; Douady et al., 2003; Naylor et al., 2005; Aschliman et al.,
2012a, Naylor et al., 2012), and the historical significance of most of these have been succinctly summarised by
Aschliman et al. (2012b). The batoids are widely viewed as monophyletic within the chondrichthyan fishes, but
unresolved structural uncertainty still exists within some family-level taxa.
Non-monophyletic interrelationships within the family Dasyatidae have been suggested (e.g. Compagno &
Accepted by M.R. de Carvalho: 16 Jun. 2016; published: 21 Jul. 2016 345

Roberts, 1982; Last, 1987; Manjaji, 2004). Maisey (1984) and McEachran (1990) also revealed that the lower
taxonomic level (genera and species) classification of dasyatids in use at the time had never been critically
reviewed. The extent of alpha-level, taxonomic confusion, particularly for stingrays of the Indo–Pacific, was
further highlighted by Last & Stevens (1994). This was further demonstrated by Manjaji (2004) in a detailed study
of whip-tail stingrays (Himantura). Observations of non-monophyly in several of the genera have been supported
by several morphology-based studies of batoid phylogeny (e.g. Lovejoy, 1996; McEachran et al., 1996; Lovejoy et
al., 1998; Rosenberger, 2001; Carvalho et al., 2004; González-Isáis & Domínguez, 2004).
Nishida (1990) produced arguably the most detailed, modern-day, morphological treatment of myliobatiform
fishes (as a suborder), and proposed a revised classification in which the family Dasyatidae was represented by four
genera, Potamotrygon, Taeniura, Dasyatis and Himantura. Compagno (1999) in his checklist of living
chondrichthyans, which was based on contributions from the broader ichthyological community, listed 68 valid
nominal species (two dubious and possibly 11 others undescribed) from seven genus-level taxa: Dasyatis,
Himantura, Pastinachus, Pteroplatytrygon, Taeniura, Urogymnus, and a probable unnamed taxon referred to as
Western Hemisphere Himantura. Recent changes to the classification include the resurrection of Neotrygon, for a
subgroup of Dasyatis (Last & White, 2008), and Taeniurops, a subgroup of Dasyatis and formerly of Taeniura
(Last & Stevens, 2009; Ebert et al., 2013). Several other recent papers have treated new taxa and contributed to the
discussion of stingray taxonomy.
An important contribution by Manjaji (2004) concluded that Himantura is a monophyletic group consisting of
at least three species complexes. Also, the amphi-American Himantura species were found to be non-monophyletic
with Indo–Pacific Himantura, as suggested by Lovejoy (1996) and McEachran & Aschliman (2004), and more
recent molecular analyses have demonstrated a close link between amphi-American Himantura and the
potamotrygonids (Aschliman et al., 2012a).
Changes to the classification have been proposed at family-level. Roberts (2007) erected a new genus
Makararaja which he assigned to the Pastinachinae, a new subfamily of the Dasyatidae. More recently, Lim et al.
(2015) identified four clades within the contemporary definition of the family Dasyatidae which they assigned to
family-level taxa: Dasyatidae (including genera Dasyatis and Taeniurops) and three new families, the
Neotrygonidae (Neotrygon and Taeniura), Himanturidae (Himantura) and Pastinachidae (Pastinachus). They used
a suite of nine ‘unreliable’ morphological characters to support their argument, but did not appear to be aware of
the establishment of Roberts’ subfamily Pastinachinae. They also failed to apply for obligatory Zoobank
registration of their names (as requested by PLosOne, an electronic publication) so their proposed new names are
not available (R. van der Laan, pers. comm.).
Morphological research by the authors for the compilation of a guide to rays of the world (Last et al., in press)
identified several inconsistencies in the dasyatid classification. Naylor et al. (2012), using NADH2 sequence data,
proposed a new hypothetical phylogeny of the Myliobatiformes (see their Figure 2.11), and also flagged major
problems in the existing classification. Several stingray genera were found to be non-monophyletic, and ‘amphi-
American Himantura’, H. schmardae (Werner) and H. pacifica (Beebe & Tee-Van), as well as the large
benthopelagic stingray, Dasyatis microps (Annandale), clustered with the outgroups outside the Dasyatidae. Their
analysis recovered the four higher level taxonomic groups discussed by Lim et al. (2015), supported the existence
of several genus-level taxa within Dasyatis, and the recognition of Taeniurops (but not Pteroplatytrygon) as a valid
genus. Himantura was found to be polyphyletic, being represented by several genus-level taxa, with four species
clustering with the monotypic Urogymnus rather than the other Himantura subgroups. The family-level group
containing the Indo–Pacific taxa Himantura and Urogymnus, also included West African coastal and freshwater
species, Dasyatis margarita (Günther) and D. margaritella Compagno & Roberts, which are unique within
whiprays in having a well-developed ventral tail fold (absent in Urogymnus and all but one Himantura). A family-
level group, recognised by Lim et al. (2015) as the Neotrygonidae, contained two monophyletic groups coinciding
with the genera Taeniura and Neotrygon. Pastinachus formed a natural group with two paraphyletic species pairs.
Inconsistencies in the dasyatid classification led to more detailed morphological investigation of the group
when drafting the stingray chapter for the Rays of the World book (Last et al., in press). Datasets were expanded
and reanalysed, and more marker-rich molecular studies using more taxa, have provided new insights into the
generic interrelationships of this group. The patterns observed were used to propose a new generic framework for
the group.
346 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Materials and methods
A checklist of the world’s dasyatids (89 species) was compiled as part of the Tree of Life for Chondrichthyan fishes
project (Appendix 1), using data extracted largely from long-term anatomical and morphological studies of the
group (by PL & MM) and enhanced by molecular data (GN). The following characters were collated for all
nominal taxa: maximum disc width (cm DW), disc shape, pectoral-fin apex angularity, body thickness (extent of
depression), snout shape and length (also expressed as a proportion of combined orbit and spiracle length), eye size
and level of protrusion, nasal curtain shape, mouth size, oral papillae number, tail base breadth and cross-sectional
shape, tail shape and length relative to disc width, pelvic-fin size and extent of exposure beyond the disc margin,
dorsal tail fold presence/absence and shape, ventral tail fold presence/absence and depth and base length, insertion
of caudal sting(s) on tail (expressed as distance from pectoral-fin insertions to the sting origin/interspiracular
width), skin roughness, presence/absence of a denticle band, development of thorn-like denticles on the nape,
shoulder and median thorn rows on the tail, distribution of other thorns and denticles on the tail, dorsal and ventral
coloration (plain or patterned), and primary habitat. Other data collected, including squamation, and tooth row and
meristic counts, follow standards used in Manjaji (2004) and Manjaji-Matsumoto & Last (2006). Species
demarcated with an asterisk in the checklist are provisional assignments as they are based on either incomplete
morphological or molecular support (or both) and need further investigation. Morphometric methods, including tail
fold measurements, follow Last & White (2008), with one notable modification: distance measurement from the
pectoral-fin insertions to the caudal sting was taken horizontally to minimise errors from distortions of the tail.
Authority/date for species and genera of the family Dasyatidae largely follow Eschmeyer et al. (2016) and
have not been referenced separately. Protocols for naming family-level taxa follow van der Laan et al. (2014).
Comparative material used to define genera were sourced from museums worldwide and tissues used for molecular
analyses were accessed from a large collection assembled for the Tree of Life Project. Extensive use was also made
of digital images and line illustrations, as not all species were observed directly. Unnamed taxa are known to exist
in some genera and several more detailed revisions are in progress by PL and MM.
The mitochondrial DNA sequence data were generated following methods described in Naylor et al. (2012).
Briefly, specimens were sampled for liver or muscle tissue by project team members. Samples were stored in 95%
alcohol or in dimethyl sulphoxide solution (20%) in the field. DNA was extracted using the E.Z.N.A Tissue DNA
Kit (Omega Bio-Tek, Inc Norcross, GA). Extracted total DNA was stored at -20 °C until used for amplification of
the NADH dehydrogenase subunit 2 (NADH2) region of the mitochondrial DNA via the Polymerase Chain
Reaction (PCR). A single set of universal primers (Naylor et al., 2005) designed to bind to the ASN and ILE tRNA
regions of the mitochondrial genome were used to amplify the target fragment. PCR reactions were carried out in
25 µl volume comprising 0.3 µM primers, 2.5 mM MgCl2, 200 µM each dNTP, 10X Ex Taq buffer (20 mM Tris-
HCl pH 8.0, 100 mM KCl, 0.1mM EDTA, 1mM DTT, 0.5% Tween20, 05% Nonidet P-40, 50% Glycerol), 0.25 U
TaKaRa Ex Taq (Takara, Mountain View, CA), and 50–100 ng template DNA. The reaction cocktail was denatured
at 94˚C for 3 minutes, after which it was subjected to 35 cycles of denaturation at 94° C for 30s, annealing at 48° C
for 30s and extension at 72° C for 90s. PCR products were purified with ExoSAP-IT (USB, Cleveland, Ohio), and
bi-directionally Sanger sequenced using BigDye®. Terminator chemistry on an ABI 3730xl genetic analyzer
(Applied Biosystems®, Life Technologies, Grand Island USA) at Retrogen Inc. Custom DNA Sequencing Facility
(San Diego USA). DNA sequences were edited using Geneious® Pro v. 6.1.7 (Biomatters Ltd Auckland, New
Zealand. Available at http://www.geneious.com).
Edited sequences were translated to amino acids and aligned with corresponding NADH2 sequences from
representatives of closely related species using the MAFFT module within the Geneious Package (Biomatters Ltd
Auckland, New Zealand). The aligned amino acid sequences were translated back, in frame, to their original
nucleotide sequences, to yield a nucleotide alignment. The full protein-coding alignment was 1044 nucleotides
long. The alignment was subjected to a constrained Maximum Likelihood analysis under the GTR +I + G model of
molecular evolution using the software package Paup*4.0 version a148. Estimates for the parameter values for the
model were estimated from the data. Topological constraints were derived from an independent ML analysis of
whole mitochondrial genomes for selected representative species of stingrays (unpublished). It should be noted that
the unconstrained ML analysis of the NADH2 sequences presented in the current study (Figs. 1–5) resulted in a
tree topology that was very similar to that seen for the analysis of the whole mitochondrial genomes. However,
internal branch lengths and associated branch support values differed considerably between the two analyses.
REVISED CLASSIFICATION OF THE STINGRAYS (DASYATIDAE) Zootaxa 4139 (3) © 2016 Magnolia Press · 347
Dasyatis longicauda Papua New Guinea GN 15791
Dasyatis longicauda Papua New Guinea GN 15792
Dasyatis longicauda Indonesia GN 11199
Dasyatis ﬂuviorum Australia GN 10855
Dasyatis bennetti Taiwan GN 13895 Hemitrygon
Dasyatis bennetti Taiwan GN 9933
Dasyatis bennetti Taiwan GN 9934
Dasyatis bennetti Indonesia GN 11193
Dasyatis bennetti Indonesia GN 11194
Dasyatis akajei Japan GN 12904
Dasyatis akajei Japan GN 12919
Dasyatis akajei Korea GN 13849
Dasyatis laevigata Japan GN 12917
Dasyatis izuensis Japan GN 12922
Dasyatis parvonigra Malaysia GN 10992
Dasyatis parvonigra Sulu Sea GN 16042
Dasyatis sp. Borneo GN 3424
Dasyatis sp. Borneo GN 4448 Telatrygon
Dasyatis zugei Vietnam GN 7021
Dasyatis zugei Vietnam GN 7048
Dasyatis zugei Taiwan GN 13896
Dasyatis zugei Taiwan GN 10079
Dasyatis zugei Japan GN 12928
Dasyatis acutirostra Japan GN 12930
Dasyatis acutirostra Japan GN 12931
Dasyatis centroura Gulf of Mexico GN 12771
Dasyatis centroura NW Atlantic GN 4639
Dasyatis centroura NW Atlantic GN 4638
Dasyatis centroura Caribbean Sea GN 14617
Dasyatis lata Japan GN 12921
Dasyatis lata Indonesia GN 11302
Dasyatis lata New Zealand GN 15449 Bathytoshia
Dasyatis lata Oman GN 9760
Dasyatis lata Hawaii GN 15451
Dasyatis brevicaudata Australia GN 10852
Dasyatis brevicaudata Japan GN 12924
Dasyatis brevicaudata Japan GN 15450
Pteroplatytrygon violacea GN 16154
Pteroplatytrygon violacea Pakistan GN 14947
Pteroplatytrygon violacea South Africa GN 10378
Pteroplatytrygon violacea USA GN 6498 Pteroplatytrygon
Dasyatis tortonesei E Atlantic GN 16082
Dasyatis hypostigma E Atlantic GN 16151
Dasyatis chrysonota South Africa GN 14824
Dasyatis marmorata Senegal GN 6009
Dasyatis pastinaca E Atlantic GN 16086 Dasyatis
Dasyatis pastinaca E Atlantic GN 16128
Taeniurops grabata Senegal GN 3294
Taeniurops grabata Cape Verde GN 16153
Taeniurops grabata Israel GN 16195
Taeniurops grabata Israel GN 16196
Taeniurops meyeni Japan GN 12932 Taeniurops
Taeniurops meyeni Australia GN 2109
Taeniurops meyeni Australia GN 11013
Taeniurops meyeni Borneo GN 3711
Taeniurops meyeni Taiwan GN 16461
Dasyatis say USA GN 13502
Dasyatis say GN 13504
Dasyatis say Gulf of Mexico GN 12770
Dasyatis dipterura Mexico GN 1598
Dasyatis sabina Gulf of Mexico GN 6229
Dasyatis sabina USA GN 13503
Dasyatis sabina USA GN 13516 Hypanus
Dasyatis americana Gulf of Mexico GN 6232
Dasyatis americana USA GN 13493
Dasyatis americana NW Atlantic GN 1594
Dasyatis longa Mexico GN 1599
Dasyatis guttata Belize GN 13939
Neotrygon orientale Borneo GN 3405
Neotrygon varidens Thailand GN 1588
Neotrygon australiae Australia GN 2077
Neotrygon trigonoides New Caledonia GN 16023
Neotrygon kuhlii Solomon Islands GN 16863
Neotrygon leylandi Australia GN 10860 Outgroup
Neotrygon picta Australia GN 2046
Neotrygon annotata Indonesia GN 11247
Neotrygon ningalooensis Australia GN 10859
Taeniura lymma Borneo GN 4816
0.01 substitutions/site
FIGURE 1. Dasyatidae (subfamily Dasyatinae). Topology and branch lengths resulting from a constrained Maximum
Likelihood analysis of aligned NADH2 sequence data using the GTR +I + G model. Topological constraints were derived from
an unpublished ML analysis of whole mitochondrial genomes for a representative subset of the taxa (unpublished). Members of
the Neotrygoninae were used as outgroups. Names shown at branch tips are those currently in use. Revised generic names
proposed are shown to the right.

Neotrygon orientale Philippines GN 2211
Neotrygon orientale Philippines GN 2216
Neotrygon varidens Borneo GN 3067
Neotrygon varidens Borneo GN 3621
Neotrygon varidens Vietnam GN 7105
Neotrygon australiae Papua New Guinea GN 15782
Neotrygon australiae Papua New Guinea GN 15783
Neotrygon picta Australia GN 2061 Neotrygon
Neotrygon leylandi Australia GN 10860
Neotrygon annotata Papua New Guinea GN 16552
Neotrygon annotata Papua New Guinea GN 16553
Taeniura lymma Australia GN 2027
Taeniura sp. Solomon Islands GN 14084 Taeniura
Taeniura sp. Solomon Islands GN 14100
Outgroup
FIGURE 2. Dasyatidae (subfamily Neotrygoninae). Topology and branch lengths resulting from a constrained Maximum
the Dasyatinae and Neotrygoninae were used as outgroups. Names shown at branch tips are those currently in use. Revised
generic names proposed are shown to the right.
Himantura sp. 2 Oman GN 9858
Himantura sp. 2 India GN 14700
Himantura sp. 2 Iran GN 14164
Himantura randalli Iran GN 6649
Himantura randalli Kuwait GN 7801
Himantura cf. pastinacoides Borneo GN 3383
Himantura cf. pastinacoides Borneo GN 3452
Himantura pastinacoides Borneo GN 3373
Himantura gerrardi Borneo GN 4828 Maculabatis
Himantura gerrardi Borneo GN 4795
Himantura gerrardi Borneo GN 4800
Himantura gerrardi Oman GN 9439
Himantura gerrardi Iran GN 6638
Himantura sp. 1 Tanzania GN 12175
Himantura sp. 1 Tanzania GN 12185
Himantura macrura Borneo GN 4824
Himantura macrura Thailand GN 1589
Himantura toshi Australia GN 2038
Himantura sp. 4 Australia GN 5082
Himantura sp. 4 Papua New Guinea GN 15784
Himantura undulata Borneo GN 4796
Himantura undulata Borneo GN 3524 Himantura
Himantura leoparda Oman GN 9444
Himantura leoparda Oman GN 9630
Himantura leoparda Bahrain GN 12144
Himantura leoparda Iran GN 14334
Himantura leoparda Iran GN 14480
Himantura uarnak Tanzania GN 12181
Himantura uarnak Borneo GN 4221
Himantura uarnacoides Borneo GN 4218
Himantura bleekeri India GN 13947
Himantura jenkinsii Australia GN 2084
Himantura jenkinsii Borneo GN 4479 Pateobatis
Himantura jenkinsii Tanzania GN 12180
Himantura fai Oman GN 9713
Himantura fai Oman GN 9632
Himantura fai Philippines GN 2231
Himantura fai Australia GN 21058
Himantura fai Papua New Guinea GN 15780
Himantura hortlei Indonesia GN 11211
Himantura hortlei Indonesia GN 11214
Himantura hortlei Australia GN 17241
Himantura hortlei Australia GN 17242
Himantura hortlei Papua New Guinea GN 15738
Himantura heterura Borneo GN 3415
Himantura imbricata India GN 13957
Himantura imbricata Myanmar GN 17062 Brevitrygon
Himantura walga Iran GN 6639
Himantura walga Qatar GN 6593
Himantura javaensis Indonesia GN 11216
Himantura signifer Borneo GN 4550
Himantura kittipongi Borneo GN 4840
Himantura oxyrhyncha Borneo GN 4409
Fluvitrygon
Himantura granulatus Borneo GN 4809
Himantura granulatus Australia GN 5569
Himantura granulatus Solomon Islands GN 14083
Himantura granulatus Solomon Islands GN 14091
Himantura granulatus Borneo GN 2929
Himantura polylepis Borneo GN 4823
Urogymnus
Himantura dalyensis Australia GN 17237
Urogymnus asperrimus Australia GN 5550
Urogymnus asperrimus Solomon Islands GN 14106
Urogymnus asperrimus Papua New Guinea GN 15786
Urogymnus asperrimus Oman GN 9833
Urogymnus asperrimus Philippines GN 4385
Himantura lobistomus Borneo GN 2964
Dasyatis margarita Senegal GN 3296
Dasyatis margaritella Senegal GN 6066
Fontitrygon
Dasyatis lata Japan GN 12921
Outgroup
Dasyatis americana NW Atlantic GN 1594
FIGURE 3. Dasyatidae (subfamily Urogymninae). Topology and branch lengths resulting from a constrained Maximum
the Dasyatinae and Neotrygoninae were used as outgroups. Names shown at branch tips are those currently in use. Revised

Pastinachus gracilicaudus Borneo GN 4797
Pastinachus sephen Iran GN 15400
Pastinachus sephen Pakistan GN 14965
Pastinachus sephen Kuwait GN 7803
Pastinachus sephen Iran GN 15397
Pastinachus solocirostris Borneo GN 4217 Pastinachus
Pastinachus solocirostris Borneo GN 3049
Pastinachus stellurostris Borneo GN 4600
Pastinachus ater Borneo GN 4208
Pastinachus ater Papua New Guinea GN 16669
Pastinachus ater Australia GN 5502
Pastinachus ater Philippines GN 2226
Pastinachus ater Oman GN 9860
Himantura dalyensis Australia GN 17237 Outgroup
Urobatis halleri Mexico GN 1578
FIGURE 4. Dasyatidae (subfamily Hypolophinae). Topology and branch lengths resulting from a constrained Maximum
the Urogymninae and Urotrygonidae were used as outgroups. Names shown at branch tips are those currently in use. Revised
Molecular analysis
Topologies of estimated trees for subfamilies of the Dasyatidae (Figs. 1–4) and across the entire family (Fig. 5)
provide strong support for a revision of the existing dasyatid classification with the inclusion of several new as well
as resurrected genus-level taxa. Seven genus-level groups were identified in a cluster representing the subfamily
Dasyatinae (Fig. 1), corresponding to Dasyatis, Pteroplatytrygon and Taeniurops, and five new or resurrected
genera formerly placed in Dasyatis (Bathytoshia, Hemitrygon, Hypanus and Telatrygon). Of these families, as
defined, only Telatrygon and Bathytoshia are not inferred to be monophyletic. Telatrygon constitutes a paraphyletic
group made up of a monophyletic subdivision and a single species Dasyatis acutirostra Nishida & Nakaya, while
Bathytoshia is non-monophyletic when the monotypic genus Pteroplatytrygon is included. Taeniurops, which has
been considered a synonym of both Dasyatis and Taeniura, forms a strongly monophyletic subdivision within the
subfamily. Dasyatis, the type genus of the subfamily, is restricted to the eastern Atlantic (including the
Mediterranean Sea) and southwestern Indian Ocean. One of the largest subdivisions, the resurrected genus
Hemitrygon, is comprised solely of Western Pacific species. Hypanus, sister to the rest of the subfamily, is
represented by Atlantic and Eastern Pacific species. The very large benthopelagic stingray, Dasyatis microps
(Annandale), clustered outside the four dasyatid subfamilies in the molecular analyses (Fig. 5), instead, displaying
an apparent close relationship with the potamotrygonids. However, it has been provisionally retained in the
Dasyatinae as a new monotypic genus (Megatrygon) pending further analysis.
The Neotrygoninae is represented by two strongly monophyletic subdivisions (Fig. 2), corresponding to
presently recognised taxa Neotrygon and Taeniura. Further work is needed to resolve species-level structure in the
difficult Neotrygon kuhlii species complex.
Hemitrygon fluviorum Australia GN 10855
Hemitrygon longicauda Indonesia GN 11199
Hemitrygon bennetti NW Pacific GN 15971
Hemitrygon akajei Japan GN 12904
Hemitrygon izuensis Japan GN 12922
Hemitrygon laevigata Japan GN 12916
Hemitrygon parvonigra Malaysia GN 10992
Telatrygon sp. Borneo GN 2884
Telatrygon zugei Japan GN 12928
Telatrygon acutirostra Japan GN 12930
Bathytoshia brevicaudata Japan GN 12924
Pteroplatytrygon violacea California GN 6498
Bathytoshia centroura Gulf of Mexico GN 12771
Bathytoshia lata Japan GN 12921 Dasyatinae
Dasyatis hypostigma GN 16151
Dasyatis tortonesei GN 16004
Dasyatis pastinaca GN 16086
Taeniurops meyeni Japan GN 12932
Hypanus americanus NW Atlantic GN 1594
Hypanus longus Mexico GN 1599
Hypanus guttatus Belize GN 13939
Hypanus dipterurus Mexico GN 1598
Hypanus say Gulf of Mexico GN 12768
Hypanus sabinus Gulf of Mexico GN 6229
Neotrygon leylandi Western Australia GN 10860 Neotrygoninae
Maculabatis sp. 2 Oman GN 9858
Maculabatis randalli Oman GN 9714
Maculabatis sp. 3 India GN 13965
Maculabatis cf. pastinacoides Borneo GN 3383
Maculabatis pastinacoides Borneo GN 3373
Maculabatis sp. 1 Tanzania GN 12175
Maculabatis gerrardi Oman GN 9439
Maculabatis macrura Borneo GN 4824
Maculabatis toshi Australia GN 15665
Pateobatis bleekeri India GN 13947
Pateobatis uarnacoides Borneo GN 4209
Pateobatis jenkinsii Australia GN 2084
Pateobatis fai Oman GN 9632 Urogymninae
Pateobatis hortlei Indonesia GN 11211
Brevitrygon heterura Borneo GN 3415
Brevitrygon imbricata Myanmar GN 17062
Brevitrygon walga Qatar GN 6593
Brevitrygon javaensis Indonesia GN 6954
Fluvitrygon kittipongi Borneo GN 4840
Fluvitrygon signifer Borneo GN 4550
Fluvitrygon oxyrhyncha Borneo GN 4409
Urogymnus sp. 5 Australia GN 13667
Urogymnus sp. 5 Australia GN 17253
Urogymnus granulatus Borneo GN 4809
Urogymnus dalyensis Australia GN 17237
Urogymnus polylepis Borneo GN 4823
Urogymnus lobistomus Borneo GN 4206
Fontitrygon margarita Senegal GN 3296
Fontitrygon margaritella Senegal GN 6066
Pastinachus solocirostris Borneo GN 4217 Hypolophinae
Pastinachus stellurostris Borneo GN 4600
Pastinachus ater Borneo GN 4208
Rhinoptera steindachneri Mexico GN 5214
Rhinoptera brasiliensis Belize GN 13937
Rhinoptera marginata Egypt GN 16460
Rhinoptera jayakari Iran GN 14162
Rhinoptera neglecta Australia GN 1624
Rhinoptera javanica Vietnam GN 7092
Rhinoptera bonasus NW Atlantic GN 5465
Rhinoptera peli Senegal GN 6089
Manta alfredi Mozambique GN 16688
Manta birostris Indonesia GN 6791
Mobula tarapacana Japan GN 10564
Mobula thurstoni Oman GN 9728
Mobula kuhlii Oman GN 9426
Mobula mobular Spain GN 15654
Mobula hypostoma Gulf of Mexico GN 5814
Mobula munkiana Baja California GN 5251 Outgroups
Potamotrygon magdalenae Colombia GN 16889
Potamotrygon orbignyi Colombia GN 16902
Potamotrygon yepezi Colombia GN 16915
Plesiotrygon iwamae Brazil GN 13458
Potamotrygon hystrix Peru GN 2725
Paratrygon aiereba Peru GN 5874
Heliotrygon rosai Peru GN 13456
“Himantura” schmardae Belize GN 13936
Megatrygon microps Indonesia GN 11295
Urotrygon rogersi Mexico GN 5250
Ur otrygon simulatrix Mexico GN 1585
Urobatis jamaicensis Cayman Islands GN 5911
Urobatis maculatus Mexico GN 5374
Urobatis concentricus Mexico GN 2275
Urobatis halleri Mexico GN 1578
FIGURE 5. Phylogenetic relationships and nomenclatural revision for Dasyatidae based on representative samples used in this
study. Topology and branch lengths result from a constrained Maximum Likelihood analysis of aligned NADH2 sequence data
using the GTR +I + G model. Topological constraints were derived from an unpublished ML analysis of whole mitochondrial
genomes for a representative subset of the taxa (unpublished). Outgroups are represented by a selection of taxa from the
families Rhinopteridae, Mobulidae, Potamotrygonidae and Urotrygonidae.

The subfamily Urogymninae is a diverse group of whiprays formerly consisting of Himantura species and the
monotypic genus Urogymnus, for U. asperrimus (Bloch & Schneider). Seven morphologically identifiable,
monophyletic subdivisions were recovered and several of these are named here (Fig. 3). A monophyletic group of
Atlantic stingrays (Fontitrygon), formerly assigned to Dasyatis based largely on tail fold morphology, appear to be
sister to the rest of the subfamily. A more species-rich Urogymnus, including several large whiprays formerly in
placed in Himantura, appears to be the immediate sister group to Fontitrygon. Urogymnus is sister to a subdivision
containing two new morphologically homogeneous groups, an Indo–West Pacific marine group (Brevitrygon) and a
small freshwater/estuarine group from the Indo–Malay Archipelago (Fluvitrygon). Pateobatis consists of a
somewhat morphologically heterogeneous group of whiprays whose assignment to a single genus is not well
supported by morphological data. Two very large whiprays, Pateobatis fai (Jordan & Seale) and P. jenkinsii
(Annandale), which are more widely distributed throughout the Indo–West Pacific, have a more angular rhombic
disc with a much shorter and broader snout than their congeners. The genus Himantura, herein confined to a small
group of very large whiprays with a strong dorsal colour pattern consisting of spots, ocelli and/or reticulations, is
sister to a larger group of smaller whiprays (Maculabatis) with weaker markings. Both genera occur throughout the
Indo–West Pacific. A Western Atlantic species once placed in Himantura, H. schmardae (Werner), clustered well
outside the Dasyatidae, together with the outgroups (i.e. Potamotrygonidae and Urotrygonidae) and Megatrygon. A
sister group relationship between ‘amphi-American Himantura’ and South American freshwater stingrays
(Potamotrygonidae), has been suggested by earlier morphological and molecular studies (e.g. Lovejoy, 1996;
Naylor et al., 2012)
Only one of the two genera of the subfamily Hypolophinae was included in the analysis (Fig. 4). The
monotypic genus Makararaja is known only from the holotype and no fresh tissue was available for molecular
analysis.
Family Dasyatidae Jordan and Gilbert, 1879
Dasybatidae Jordan & Gilbert, 1879:386 (family), corrected to Dasyatidae by Jordan, 1888: 22) [van der Lann et al., 2014];
type genus Dasyatis (as “Dasybatis Rafinesque”).
Definition. Small to very large myliobatiform fishes (adults from 22 cm to 260 cm DW) and distinguished by the
following combination of characters: body variably depressed with a well-formed oval, circular or rhombic disc
that fully incorporates head; snout angular to obtuse and sometimes very elongate; nasal curtain well developed,
skirt-shaped, rectangular or bilobed; five gill slits; oral papillae usually present on floor of mouth; tail moderately
stout to slender-based and more or less elongated (sometimes very elongate and whip-like); dorsal surface variably
covered with dermal denticles, thorns and/or tubercles, smooth to very spiny and often with a median thorn row
and/or a median denticle band; no dorsal or caudal fins; 1–4 prominent caudal stings, positioned on tail well
posterior to pelvic fins; skin folds variably developed on the ventral and sometimes dorsal midline of tail; dorsal
surface plain to strongly patterned, usually darker than ventral surface.
Subfamilies. Dasyatinae Jordan and Gilbert, 1879, Hypolophinae Stromer, 1910, Neotrygoninae subfam. nov.,
and Urogymninae Gray, 1851.
Remarks. Four subfamilies are recognised that are consistent with the family groups identified by Lim et al.
(2015). Although, as discussed already, their proposed names are not available (R. van der Laan, pers. comm.), we
maintained use of one of their groups by erecting a new taxon at the rank of subfamily: Neotrygoninae.
Hypolophinae was resurrected and the Urogymninae resurrected and elevated from the rank of tribe.
Subfamily Dasyatinae
Dasybatidae Jordan & Gilbert, 1879:386 (corrected to Dasyatidae by Jordan, 1888:22; proposed herein as a subgenus of
Dasyatis); type genus Dasyatis Rafinesque, 1810, by original designation.
Definition. Morphologically variable subfamily with no unique character states; disc circular, flattened cone-
shaped but mostly rhombic; snout short to long; tail variably depressed, short and firm to long and whip-like; oral
papillae 1–7 or absent; dorsal tail fold developed or reduced to a fleshy ridge or absent; denticle band absent
(typically weak if present), no tail thorns in most species, and usually plain dorsal surface coloration with dark
posterior tail. All members with a short to long-based ventral tail fold that can be poorly developed to deep, and
caudal sting relatively well forward on tail (distance from pectoral-fin insertion to caudal-sting base 2–3 times
interspiracular width).
Genera. Bathytoshia, Dasyatis, Hemitrygon, Hypanus, Megatrygon, Pteroplatytrygon, Taeniurops and
Telatrygon.
Remarks. The group contains eight genera of which three are resurrected (Bathytoshia, Hemitrygon and
Hypanus) and two are new (Megatrygon and Telatrygon). Several nomenclatural issues need to be discussed that
relate specifically to each of the genera. Species are treated in more detail in an identification guide to rays of the
world (Last et al. in press) and other review papers in preparation, but synonymies are discussed briefly in Remarks
sections for each genus.
Genus Bathytoshia Whitley, 1933
Bathytoshia Whitley, 1933:61. Type species Dasyatis thetidis Ogilby, 1899; by original designation.
Definition. Gigantic dasyatids (adults to 210–260 cm DW) characterised by the following: robust, rhombic disc
with pectoral-fin apex narrowly angular; snout obtuse and moderately elongate (1.5–2.5 times combined orbit and
spiracle length); eye small and protruding slightly; nasal curtain broadly skirt shaped; mouth rather broad, with 3–7
oral papillae; tail firm and short to moderately elongate (length <1–2 times DW), its base broad to very broad and
depressed; pelvic fins medium-sized, protruding slightly to greatly beyond disc; dorsal fold absent or forming a low
ridge; ventral fold low with a short or long base; caudal sting not posterior on tail (distance from pectoral-fin
insertion to caudal-sting base 2.2–3 times interspiracular width); skin smooth or rough and denticle band absent;
median thorns absent or in a row extending along disc and onto tail, scapular thorns small or absent; tail spiny with
bucklers and tubercles on dorsal midline of adults; dorsal colour largely plain; ventral surface white, disc margin
sometimes dark; tail plain, usually black distally; marine, Atlantic and Indo–West Pacific.
Species. B. brevicaudata (Hutton, 1875), B. centroura (Mitchill, 1815), and B. lata (Garman, 1880).
Remarks. Former junior synonym of Dasyatis (Kottelat, 2013) includes a small group of very large and
widely distributed stingrays. Molecular data (G. Naylor, unpubl.) provides evidence that species complexes (A)
Bathytoshia (as Dasyatis) brevicaudata and D. matsubarai Miyosi, 1939, and (B) Bathytoshia (as Dasyatis) lata,
D. thetidis Ogilby, 1899, D. ushiei (Jordan & Hubbs, 1925), and eastern Atlantic stingrays identified as D.
centroura (e.g. McEachran & Capapé, 1984), are populations of just two species (i.e. D. brevicaudata and D. lata).
A western Atlantic species, Bathytoshia centroura (Mitchill, 1815), is not conspecific with Bathytoshia of the
eastern Atlantic. The pelagic stingray Pteroplatytrygon violacea (Bonaparte, 1832), which clusters with these
species (Fig. 1), is morphologically distinct from all other members of the family.
Genus Dasyatis Rafinesque, 1810
Dasyatis Rafinesque, 1810:16. Type species Dasyatis ujo Rafinesque, 1810 (=Raja pastinaca Linnaeus, 1758); designated by
monotypy.
Definition. Small to medium-sized dasyatids (adults to 38–84 cm DW) characterised by the following: robust,
rhombic disc with pectoral-fin apex narrowly rounded to angular; snout broadly angular and short (1.1–1.5 times
combined orbit and spiracle length); eye medium-sized to large and protruding slightly; nasal curtain broadly skirt-
shaped; mouth medium-sized, with 3–7 oral papillae (3 centrally); tail firm, somewhat filamentous distally (length
1.2–2.3 times DW), its base broad to very broad and depressed; pelvic fins medium-sized to large, protruding well
beyond disc; dorsal fold variably developed, low or reduced to form a ridge; ventral fold low (better developed in
D. pastinaca) with a short to long base; caudal sting typically forward on tail (distance from pectoral-fin insertion
to caudal-sting base 1.8–3 times interspiracular width); no median thorns on disc and denticle band absent; skin
typically smooth, except in largest adults which may have sparse coverage of denticles and sometimes well-

developed thorns on anterior tail; scapular thorns absent; tail smooth or prickly; dorsal coloration plain or with
prominent pattern; ventral surface white, disc margin sometimes dark; tail plain, black distally; marine, Atlantic
and South-West Indian Ocean.
Species. D. chrysonota (Smith, 1828), D. hypostigma Santos & Carvalho, 2004, D. marmorata (Steindachner,
1892), D. pastinaca (Linnaeus, 1758), and D. tortonesei Capapé, 1977.
Remarks. The polyphyletic genus Dasyatis, once thought to consist of 36 species with widespread
distributions in tropical and temperate seas, now applies to a monophyletic group of five small to medium-sized
species found in the Atlantic (including the Mediterranean Sea) and SW Indian Ocean (see Fig. 1). Molecular data
supports the non-conspecificity of similar blue-marbled forms, D. chrysonota and D. marmorata.
Genus Hemitrygon Müller & Henle, 1838
Hemitrygon (subgenus of Trygon) Müller & Henle, 1838:90. Type species Trygon bennettii Müller & Henle, 1841; by
subsequent monotypy.
Definition. Small to medium-sized dasyatids (adults typically to 31–66 cm DW, 93 cm DW in H. fluviorum)

characterised by the following: depressed to robust, weakly rhombic to rhombic disc with pectoral-fin apex
narrowly angular to broadly rounded; snout obtuse to broadly angular and moderately elongate (1.7–2.5 times
combined orbit and spiracle length); eye small to very small and protruding slightly; nasal curtain typically broadly
skirt shaped; mouth narrow, with 1–7 oral papillae (typically with 3 centrally); tail short and filamentous to long
and whip-like (length 0.8–2.9 times DW), its base depressed oval and not especially broad; pelvic fins medium-
sized to large, protruding slightly beyond disc; dorsal fold variably developed, typically low or reduced to form a
ridge; ventral fold low with a short to long base; caudal sting close to tail base (distance from pectoral-fin insertion
to caudal-sting base 1.9–2.6 times interspiracular width); median thorns on disc in a row, often poorly defined or
absent; denticle band poorly defined, small, diffuse or absent; skin smooth or with a patchy coverage of denticles;
scapular thorns small or absent; tail sometimes with large median thorns, otherwise smooth or partly covered with
fine denticles; dorsal coloration plain; ventral surface white, disc margin sometimes yellow or orange; tail plain,
usually black distally; estuarine/marine, Western Pacific.
Species. H. akajei (Müller & Henle, 1841), H. bennetti (Müller & Henle, 1841), H. fluviorum (Ogilby, 1908),
H. izuensis (Nishida & Nakaya, 1988), H. laevigata (Chu, 1960), H. laosensis (Roberts & Karnasuta, 1987), H.
longicauda (Last & White, 2013), H. navarrae (Steindachner, 1892), H. parvonigra (Last & White, 2008), and H.
sinensis (Steindachner, 1892).
Remarks. Resurrected genus and formerly a junior synonym of Dasyatis (Kottelat, 2013), it consists of 10
morphologically similar species formerly placed in Dasyatis. The group is under review (PL); some of the nominal
species listed above may be synonyms but other un-named species appear to exist.
Genus Hypanus Rafinesque, 1818
Hypanus Rafinesque, 1818:272. Type species Raja say Lesueur, 1817 (Eschmeyer et al., 2016).
Definition. Mixed group of small to very large dasyatids (adults 31–198 cm DW) characterised by the following:
depressed to robust, weakly rhombic to rhombic disc with pectoral-fin apex narrowly angular to broadly rounded;
snout obtuse to pointed and short to elongate (1–3 times combined orbit and spiracle length); eye typically large
and protruding slightly (small and sunken in H. guttata); nasal curtain skirt shaped or bilobed; mouth narrow to
broad, with 3–5 oral papillae (typically with 3 centrally); tail short to very long and whip-like or filamentous
distally (length 1.3–3 times DW), its base typically depressed oval and narrow to broad; pelvic fins medium-sized,
protruding slightly to well beyond disc; dorsal fold variably developed, prominent, reduced to form a ridge, or
absent; ventral fold low to rather deep with a short to long base; caudal sting close to tail base (distance from
pectoral-fin insertion to caudal-sting base 2.1–2.6 times interspiracular width); median thorns on disc in a distinct
row; denticle band typically very small or absent (well-defined in H. guttata); skin smooth or with a patchy
coverage of denticles (rough in H. rudis); scapular thorns present, small, 1–2 or in a short row; median thorns on
tail weak or continuous with median thorn row on disc, sometimes absent; remaining tail smooth or covered with
fine denticles; dorsal coloration plain; ventral surface white, disc margin dark or reddish; tail plain coloured or
dorsal surface darker than ventral surface; marine, Atlantic and Eastern Pacific.
Species. H. americanus (Hildebrand & Schroeder, 1928), H. dipterurus (Jordan & Gilbert, 1880), H. guttatus
(Bloch & Schneider, 1801), H. longus (Garman, 1880), H. marianae (Gomes, Rosa & Gadig, 2000), H. rudis
(Günther, 1870), H. sabinus (Lesueur, 1824), and H. say (Lesueur, 1817).
Remarks. Resurrected genus, formerly considered a junior synonym of Dasyatis (Kottelat, 2013), now
provisionally consists of eight somewhat morphologically heterogeneous species. Six of these species cluster
together based on molecular data (see Fig. 1), but provisional placement of H. marianae and H. rudis still needs to
be confirmed when tissues become available.
Genus Megatrygon gen. nov.
Type species Trygon microps Annandale, 1908:393; newly proposed, monotypic.
Definition. Very large dasyatid (adults to 220 cm DW) characterised by the following: very robust, broad rhombic
disc with pectoral-fin apex angular; snout broadly angular (~3 times combined orbit and spiracle length); eye very
small and sunken; nasal curtain skirt shaped; mouth narrow, with 5 oral papillae; tail short (length subequal to
DW), very broad-based and depressed anteriorly, very strongly tapered at caudal sting then becoming filamentous;
pelvic fins large, protruding greatly beyond disc; dorsal fold forming a low ridge; ventral fold low with a very short
base; caudal sting posterior on tail (distance from pectoral-fin insertion to caudal-sting base more than 3 times
interspiracular width); skin densely covered with minute stellate denticles but denticle band absent; no median
rows of thorns and scapular thorns absent; tail base and sides covered with thorny denticles; dorsal colour plain;
ventral surface white, disc margin dark; tail plain, black distally; marine, Indo–West Pacific.
Etymology. Combination of the Greek mégas (great, large, mighty) and Greek trygon (stingray) with reference
to the massive bulk of this gigantic stingray.
Species. M. microps (Annandale, 1908).
Remarks. Newly erected, monotypic genus and formerly assigned to Dasyatis. The placement of Megatrygon
microps in the family Dasyatidae is provisional as molecular data (see also Naylor et al., in press) suggest that it,
along with the ‘amphi-American Himantura’, are more closely related to the freshwater Neotropical stingrays
(Potamotrygonidae) of South America. Further investigations are needed to determine the position of this species
in the order Myliobatiformes, but it may belong in its own family.
Genus Pteroplatytrygon Fowler, 1910
Pteroplatytrygon (subgenus of Dasyatis) Fowler, 1910:474. Type species Trygon violacea Bonaparte, 1832; by original
designation and also monotypic.
Definition. Medium-sized dasyatid (adults to 80 cm DW) characterised by the following: very robust, broadly
cone-shaped disc with pectoral-fin apex angular; snout short and rounded (1.1–1.3 times combined orbit and
spiracle length); eye very small and sunken; nasal curtain skirt shaped; mouth very narrow, with numerous oral
papillae; tail moderately elongate (length >2 times DW), rather broad-based and depressed anteriorly, very strongly
tapered at caudal sting then becoming filamentous; pelvic fins large, protruding well beyond disc, with long inner
margins; dorsal fold rudimentary or absent; ventral fold low with a long base; caudal sting moderately well back on
tail (distance from pectoral-fin insertion to caudal-sting base ~1.6 times interspiracular width); dense median row
of small thorns and small denticles extending from nape onto tail; skin otherwise smooth or with sparse denticles;
denticle band and scapular thorns absent; dorsal and ventral surfaces similarly plain and dark; marine,
cosmopolitan.
Species. P. violacea (Bonaparte, 1832).
Remarks. Frequently assigned to Dasyatis (e.g. Krefft & Stehmann, 1973; Rosenberger, 2001),
Pteroplatytrygon violacea (Bonaparte, 1832) is a pelagic stingray with a body shape and coloration that is unique

within the family. Molecular data suggest it is not monophyletic with newly defined Dasyatis, but might be
monophyletic when assigned to Bathytoshia (Fig. 1). To avoid creating further confusion, this monotypic genus is
retained provisionally pending more thorough morphological and molecular investigations.
Genus Taeniurops Garman, 1913
Taeniurops (subgenus of Taeniura) Garman, 1913:399. Type species Taeniura meyeni Müller & Henle, 1841; by subsequent
designation (Fowler, 1941).
Definition. Large to very large dasyatids (adults to 100–180 cm DW) characterised by the following: robust,
almost circular disc; snout obtuse to broadly angular (2–2.2 times combined orbit and spiracle length); eye small
and protruding slightly; nasal curtain broadly skirt-shaped; mouth medium-sized, with ~7 oral papillae (in T.
meyeni); tail firm and short (length slightly exceeding DW), its base rather broad and depressed; pelvic fins small,
subrectangular, not protruding, inner margin long; dorsal fold absent; ventral fold deep with a long base; caudal
sting positioned rather anteriorly on tail (distance from pectoral-fin insertion to caudal-sting base 2–2.3 times
interspiracular width); skin rough in adults and denticle band absent; continuous strip of small median thorns on
disc and tail in most adults, scapular thorns present or absent; tail prickly; strong dorsal colour plain or heavily
black and white blotched; ventral disc white with dark margins; tail uniformly black; Atlantic (including the
Mediterranean Sea) and Indo–West Pacific.
Species. T. grabata (Geoffroy Saint-Hilaire, 1817) and T. meyeni (Müller & Henle, 1841).
Remarks. Formerly placed within Taeniura (e.g. Last & Stevens, 1994; Last & Compagno, 1999) and later
resurrected by Last & Stevens (2009), T. meyeni forms a monophyletic group with its cognate T. grabata, outside
the subfamily Neotrygoninae and within the Dasyatinae, close to Dasyatis (see Fig. 1). The two Taeniurops species
are morphologically distinct from other Dasyatinae in disc shape (almost circular) and ventral tail fold morphology
(fold unusually deep and continuous to tail tip).
Genus Telatrygon gen. nov.
Type species Trygon zugei Müller & Henle, 1841; newly proposed.
Definition. Small to medium-sized dasyatids (adults typically to 29–73 cm DW) characterised by the following:
strongly depressed, weakly rhombic to rhombic disc with pectoral-fin apex narrowly angular to broadly rounded;
snout acutely angular and elongate (2.9–4+ times combined orbit and spiracle length); eye small to very small and
sunken; nasal curtain typically narrowly skirt-shaped; mouth narrow and lacking oral papillae; tail rather long and
filamentous distally (length 1.4–3 times DW), its base narrow to medium width; pelvic fins medium-sized,
protruding slightly beyond disc; dorsal fold variably developed, from prominent to absent; ventral fold low with a
short to long base; caudal sting close to tail base (distance from pectoral-fin insertion to caudal-sting base 2–2.5
times interspiracular width); skin smooth; no scapular thorns or denticle band; median thorns on nape in a row,
usually separated from larger thorn-like denticles on tail; no other thorns or rarely denticles on tail; dorsal
coloration plain; ventral surface white, disc margin usually dark edged; tail plain or darker dorsally than ventrally;
marine, Northern Indian Ocean and Western Pacific.
Etymology. Combination of the Latin telum (javelin, spear, arrow, dart) and Greek trygon (stingray) with
reference to the long, narrowly pointed snout possessed by all members of the genus.
Species. T. acutirostra (Nishida & Nakaya, 1988), T. crozieri (Blyth, 1860), T. zugei (Müller & Henle, 1841),
and an undescribed species.
Remarks. Newly erected, morphologically conservative genus whose members were formerly assigned to
Dasyatis. Telatrygon acutirostra resembles other species of Telatrygon in disc shape (apart from having a much
longer snout and tail, smaller eyes, and less well-developed skin folds on the tail). It is otherwise similar in the
states of most external characters used to define the dasyatid genera. However, based on phylogenetic analysis of
NADH2 sequences (and unpublished analyses of whole mitochondrial genomes), T. acutirostra does not cluster
with other species of Telatrygon (Fig. 1), so its placement in the group is provisional pending more detailed
anatomical studies. Telatrygon is represented by an undescribed species in the Western Pacific (Last, White &
Naylor, submitted), and a cognate occurring in the northern Indian Ocean (identified by molecular data, GN)
appears to be conspecific with Trygon crozieri Blyth, 1860; placed in synonymy with Dasyatis (=Telatrygon) zugei
by Nishida & Nakaya (1988, 1990). Although Blyth’s description is brief, his species is undoubtedly a Telatrygon,
based on disc shape, squamation, and tail morphology. Given that no other member of this group is known from the
region, we propose the resurrection of Blyth’s species as Telatrygon crozieri.
Subfamily Neotrygoninae subfam. nov.
Type genus Neotrygon Castelnau, 1873, newly proposed.
Definition. Dasyatid subfamily with a short snout, large protruding eyes, 2 large median oral papillae, row of
enlarged caniniform teeth on each side of upper jaw, tail rather short (<1.7 times DW) and firm or flexible, rather
long and pointed pelvic fin, skin smooth, no dorsal denticle band but sometimes with weak median thorns, caudal
sting inserted posteriorly to well posteriorly (distance from pectoral-fin insertion to caudal-sting base 2–5 times
interspiracular width), a well-developed dorsal tail fold, much longer low ventral tail fold, and dorsal coloration
patterned (usually with bright blue spots and/or a prominent dark mask-like marking around eyes).
Genera. Neotrygon and Taeniura.
Remarks. The two genera of the subfamily Neotrygoninae are morphologically heterogeneous but they are the
only dasyatids known to have a row of enlarged canine teeth on each side of the upper jaw. All Taeniura and most
Neotrygon have vivid blue spots on the disc, but the disc is rhombic in shape in Neotrygon (vs. oval in Taeniura),
the tail less robust and more flexible with a more anteriorly positioned caudal sting, dorsal tail fold prominent
(otherwise rudimentary or absent), ventral tail fold less well-developed, and the posterior tail is marked with a
distinctive pattern of black and white bands.
Genus Neotrygon Castelnau, 1873
Neotrygon Castelnau, 1873:122. Type species Raya trigonoides Castelnau, 1873; by monotypy.
Definition. Small dasyatids (adults to 27–47 cm DW) characterised by the following: moderately depressed to
robust, rhombic disc with pectoral-fin apex typically narrowly angular; snout obtuse and short (1.2–2.3 times
combined orbit and spiracle length); eye large and protruding (often greatly); nasal curtain narrow, typically
subrectangular; mouth narrow with 2 large median oral papillae; tail flexible and short (length 1.1–1.5 times DW),
its base medium-broad and depressed to flattened oval; pelvic fins long and pointed, protruding slightly to well-
beyond disc; dorsal fold well developed; ventral fold low with a long base; caudal sting positioned moderately well
back on tail (distance from pectoral-fin insertion to caudal-sting base 2.1–2.8 times interspiracular width); skin
smooth and denticle band absent; row of small median thorns in most adults, no other thorns on disc; tail smooth;
typically with strong dorsal colour pattern, usually with dark mask-like marking around eyes and blue or black
spots on disc; posterior tail with black and white bands; ventral surface white, disc margin typically darker; marine,
Indo–West Pacific.
Species. N. annotata (Last, 1987), N. australiae Last, White & Séret, 2016, N. caeruleopunctata Last, White &
Séret, 2016, N. kuhlii (Müller & Henle, 1841), N. leylandi (Last, 1987), N. ningalooensis Last, White & Puckridge,
2010, N. orientale Last, White & Séret, 2016, N. picta Last & White, 2008, N. trigonoides (Castelnau, 1873), and
N. varidens (Garman, 1885).
Remarks. The taxonomy of a subgroup of Neotrygon, the kuhlii-complex, is not fully resolved (Puckridge et
al., 2013). Three new members of the kuhlii-complex were described recently and a Western Pacific species (N.
varidens) was resurrected (Last et al., 2016). Members of this subgroup are very similar in morphology and an
Indo–Pacific revision is needed.

Genus Taeniura Müller & Henle, 1837
Taeniura Müller & Henle, 1837:117. Type species Trygon ornatus Gray, 1830 (=Raja lymma Forsskål, 1775); by monotypy.
Definition. Small dasyatids (adults to 22–37 cm DW) characterised by the following: moderately robust, oval disc
with pectoral-fin apex broadly rounded; snout obtuse and short (1.6–2.2 times combined orbit and spiracle length);
eye large and protruding greatly; nasal curtain medium-sized, bilobed; mouth not small, with 2 large median oral
papillae; tail firm and typically moderately short (length 1.5–1.7 times DW), its base broad and depressed; pelvic
fins long and pointed, protruding slightly; dorsal fold rudimentary or forming a ridge; ventral fold deep with a long
base; caudal sting positioned well posterior on tail (distance from pectoral-fin insertion to caudal-sting base 4.5–5
times interspiracular width); skin smooth and denticle band absent; 1–2 rows of very small median thorns in most
adults, no other thorns on disc; tail smooth; strong dorsal colour pattern of vivid blue spots on disc (no dark mask-
like marking around eyes or black and white bands on tail); ventral surface white, disc margin yellowish; marine,
Species. T. lymma (Forsskål, 1775) and an undescribed species.
Remarks. Distinctive and locally abundant rays on coral reefs of the tropical Indo–West Pacific. An
undescribed species occurs in Melanesia.
Subfamily Urogymninae
Urogymni Gray, 1851:114 (tribe); type genus Urogymnus Müller & Henle, 1837, by monotypy.
Himanturidae Lim et al., 2015:13 (family); type genus Himantura Müller & Henle, 1837 (not available).
Definition. Dasyatids with a long, narrow-based, whip-like tail (except in Brevitrygon) lacking tail folds (ventral
fold present in Fontitrygon and one Urogymnus), small to very small eyes, no prominent thorns on tail (except in
most Brevitrygon, and some Pateobatis and Urogymnus) and a variably developed median denticle band (usually
with sharply defined edges) is present in adults. Morphologically variable, mostly with a rounded disc (sometimes
weakly rhombic); snout short to very long; oral papillae 1–7 or absent; and caudal sting well forward to posterior
on tail (distance from pectoral-fin insertion to caudal-sting base 1–3 times interspiracular width); and dorsal
surface coloration plain to variably patterned (sometimes strongly so) with black and white bands on tail of several
species.
Genera. Brevitrygon, Fluvitrygon, Fontitrygon, Himantura, Maculabatis, Pateobatis, and Urogymnus.
Remarks. A large subgroup of the family Dasyatidae known as whiprays, with almost all species once
collectively grouped in the genus Himantura. The group now consists of seven genera with five of these new
(Brevitrygon, Fluvitrygon, Fontitrygon, Maculabatis and Pateobatis). Manjaji (2004) recognised three main
subgroups in Himantura: the uarnak-complex (almost entirely Himantura and Maculabatis), uarnacoides-complex
(mostly Urogymnus), and signifer-complex (containing representatives of new genera, Brevitrygon and
Fluvitrygon). Nomenclatural issues relating specifically to each genus are discussed below, but more detailed
species treatments are provided in an identification guide to rays of the world (Last et al., in press) and other
associated papers in preparation.
Genus Brevitrygon gen. nov.
Type species Dasyatis javaensis Last & White, 2013; newly proposed.
Definition. Small dasyatids (adults typically to 23–32 cm DW) characterised by the following: strongly depressed,
oval to suboval disc with pectoral-fin apex broadly rounded; snout acutely angular and elongate (1.9–3.6 times
combined orbit and spiracle length); eye small and protruding slightly; nasal curtain broadly skirt shaped; mouth
medium-sized with 2–4 oral papillae; tail rather short and semi-rigid or filamentous distally (length 1–2.2 times
DW), its base broad and strongly or weakly depressed; pelvic fins small to medium-sized, protruding slightly
beyond disc; dorsal fold and ventral folds absent (low ridges present in B. walga); caudal sting close to tail base
(distance from pectoral-fin insertion to caudal-sting base 1.4–2 times interspiracular width); denticle band well
developed with edge sharply defined, skin on rest of disc naked; median thorns on disc weak, those on tail larger
when present; no scapular thorns and rarely denticles on posterior tail; dorsal coloration plain; ventral surface
white, disc margin dark edged; posterior tail plain or with pale lateral stripe; marine, Indo–West Pacific.
Species. B. heterura (Bleeker, 1852), B. imbricata (Bloch & Schneider, 1801), B. javaensis (Last & White,
2013), and B. walga (Müller & Henle, 1841).
Etymology. Combination of the Latin brevi (short) and Greek trygon (stingray) reflects the short and semi-
rigid tail of all members of the genus.
Remarks. Newly erected, morphologically conservative genus with four small species previously placed in
Himantura. The taxonomy of this group is very confused and is the subject of a generic review in progress (PL,
unpublished). Brevitrygon heterura (Bleeker, 1852) is the correct name of a common Western Pacific species,
misidentified as Himantura (=Brevitrygon) walga (Müller & Henle, 1841). Brevitrygon walga and B. imbricata
(Bloch & Schneider, 1801) are both confined to the Indian Ocean.
Genus Fluvitrygon gen. nov.
Type species Himantura signifer Compagno & Roberts, 1982; newly proposed.
Definition. Small dasyatids (adults typically to ~37 cm DW) characterised by the following: depressed, oval to
almost circular disc with pectoral-fin apex broadly rounded; snout acutely angular to obtuse, short or long (2.2–3.7
times combined orbit and spiracle length); eye very small or small and protruding slightly; nasal curtain broadly
skirt shaped; mouth narrow to medium-sized with 2–8 oral papillae; tail rather short or long, whip-like (length 1.3–
3.8 times DW), its base narrow and oval to almost circular in cross section; pelvic fins medium-sized, almost
entirely concealed by disc; dorsal fold and ventral folds absent; caudal sting rather forward on tail (distance from
pectoral-fin insertion to caudal-sting base 2.3–3.2 times interspiracular width); 0–2 pearl-shaped mid-scapular
thorns, no other scapular thorns; denticle band well developed with edge diffuse to sharply defined, skin on rest of
disc typically naked; median thorns on tail weak or absent, small denticles usually on posterior tail in adults; dorsal
coloration plain or with strong pattern; ventral surface white, disc margin sometimes dark edged; posterior tail
plain or with pattern; estuarine/freshwater, Indo–Malay Archipelago.
Etymology. Combination of the Latin fluvius (river) and Greek trygon (stingray) reflects the riverine
distribution of all members of the genus.
Species. F. kittipongi (Vidthayanon & Roberts, 2006), F. oxyrhyncha (Sauvage, 1878), and F. signifer
(Compagno & Roberts, 1982).
Remarks. Newly erected, morphologically similar genus consisting of three small species previously placed in
Himantura. These freshwater/estuarine species are sister to Brevitrygon (Fig. 3), but differ in tail morphology from
members of that marine group (having a longer, narrower-based, whip-like tail).
Genus Fontitrygon gen. nov.
Type species Trygon margarita Günther, 1870; newly proposed.
Definition. Heterogeneous group of small to large dasyatids (adults 28–165 cm DW) characterised by the
following: variably depressed, oval to almost circular disc (weakly rhombic in F. colarensis) with pectoral-fin apex
broadly rounded; snout acutely angular to obtuse, long to very long (1.8–4 times combined orbit and spiracle
length); eye very small or small and sunken to protruding slightly; nasal curtain skirt-shaped to bilobed; mouth
narrow to very narrow with 1–5 oral papillae; tail rather short or long, whip-like (length 1.5–3.8 times DW), its
base narrow and oval to almost circular in cross section; pelvic fins small to medium-sized, almost entirely
concealed by disc (except long, pointed and well exposed in F. geijskesi); dorsal fold reduced to a ridge or absent;
ventral fold low or very low, short to long based; caudal sting not positioned well back on tail (distance from
pectoral-fin insertion to caudal-sting base 1.9–3 times interspiracular width); 1–4 mid-scapular thorns or thorns on
nape in a row; small scapular thorns present; denticle band well developed with edge diffuse to sharply defined,

skin on rest of disc prickly or with patchy denticles; median thorns on tail either continuous with those of mid-disc
or absent; rest of tail naked or spiny; dorsal coloration plain; ventral surface white, disc margin sometimes dark
edged; posterior tail typically plain; marine/estuarine/freshwater, western Africa and Atlantic Ocean.
Etymology. Combination of the Latin fontis (spring, fountain) and Greek trygon (stingray) reflects an ability
of these stingrays to live in estuaries and freshwater.
Species. F. colarensis (Santos, Gomes & Charvet-Almeida, 2004), F. garouaensis (Stauch & Blanc, 1962), F.
geijskesi (Boeseman, 1948), F. margarita (Günther, 1870), F. margaritella (Compagno & Roberts, 1984), and F.
ukpam (Smith, 1863).
Remarks. Newly erected, morphologically heterogeneous genus consisting of six small to large stingrays,
with two species confined to freshwater and the others to estuarine/marine habitats. The tail is long and whip-like
(typical of the Urogymninae), but has a well-developed ventral skin fold so they have been traditionally placed in
the genus Dasyatis. Fontitrygon margarita and F. margaritella are assigned in this group based on molecular data
(Fig. 3), but placement of the other three species is provisional in the absence of sequence data. The four species
from western Africa form a natural group based on morphology. However, the two western Atlantic species, F.
colarensis and F. geijskesi, differ significantly from the other taxa (a much more elongated, narrowly pointed snout,
smaller and somewhat sunken eyes, and pelvic fins longer and protruding further beyond the disc margin), and may
prove to be non-congeneric with the African species. More data is needed for this group.
Genus Himantura Müller & Henle, 1837
Himantura Müller & Henle, 1837:400. Type species Raja sephen uarnak Forsskål, 1775; by subsequent designation (Jordan &
Evermann 1896:82).
Definition. Large dasyatids (adults to 130–160 cm DW) characterised by the following: robust, suboval to rhombic
disc with pectoral-fin apex broadly rounded to narrowly angular; snout broadly angular, moderately elongate (1.7–
2.8 times combined orbit and spiracle length); eye small and protruding; nasal curtain broadly skirt shaped; mouth
narrow with 4–5 oral papillae (lateral papillae always present); tail very long, whip-like (length 2.5–3.7 times DW),
its base narrow and oval to almost circular in cross section; pelvic fins small, almost entirely concealed by disc;
dorsal fold and ventral folds absent; caudal sting close to tail base (distance from pectoral-fin insertion to caudal-
sting base 1.7–2.3 times interspiracular width); 1–3 mid-scapular thorns or in row on nape, no other scapular
thorns; denticle band well developed with diffuse edge, skin on rest of disc with patchy denticles in adults; no row
of enlarged median thorns on tail, small thorns and denticles posteriorly in adults; dorsal surface with strong colour
pattern (spots, ocelli and/or reticulations); ventral surface white; posterior tail typically banded in young; marine,
Species. H. leoparda Manjaji-Matsumoto & Last, 2008, H. uarnak (Gmelin, 1789), H. undulata (Bleeker,
1852), and an undescribed species.
Remarks. The genus Himantura, which was once represented widely in the Indo–West Pacific by ~20 species,
is now restricted to a group of four large Indo–Pacific species with strong colour patterns. The taxonomy of this
group is under review (PL & MM) and a new species exists in Australasian seas.
Genus Maculabatis gen. nov.
Type species Trygon gerrardi Gray, 1851:116; newly proposed.
Definition. Small to large dasyatids (adults to 42–116 cm DW) characterised by the following: firm, rather
depressed to robust, suboval to rhombic disc with pectoral-fin apex narrowly angular to rounded; snout broadly
angular, short to moderately elongate (1.7–2.7 times combined orbit and spiracle length); eye small and protruding
slightly; nasal curtain skirt-shaped; mouth narrow with 2–5 oral papillae (lateral papillae present in most species);
tail long, whip-like (length 1.8–3.4 times DW), its base typically narrow and oval to almost circular in cross
section; pelvic fins small, almost entirely concealed by disc; dorsal fold and ventral folds absent; caudal sting close
to tail base (distance from pectoral-fin insertion to caudal-sting base 1.9–2.5 times interspiracular width); 1–3
variably developed mid-scapular thorns or thorns in row on nape, no other scapular thorns; denticle band well
developed with edge sharply defined, skin on rest of disc naked or with patchy denticles; no row of enlarged
median thorns on tail, fine denticles posteriorly in most adults; dorsal surface plain or with spotted colour pattern;
ventral surface white, disc sometimes dark or yellow edged; posterior tail typically half or fully banded in young;
marine, Indo–West Pacific.
Etymology. Combination of the Latin macula (stain, mark, spot) and batis (skate, ray, flatfish) alludes to the
spotted coloration and/or black-and-white banded tail of most members of the genus.
Species. M. astra (Last, Manjaji-Matsumoto & Pogonoski, 2008), M. gerrardi (Gray, 1851), M. macrura
(Bleeker, 1852), M. pastinacoides (Bleeker, 1852), M. randalli (Last, Manjaji-Matsumoto & Moore, 2012), M.
toshi (Whitley, 1939) and three undescribed species.
Remarks. Newly erected, morphologically similar genus consisting of nine medium to large, marine whiprays
previously placed in Himantura. The group contains at least three new species that are presently being described by
the authors and their associates. Molecular analyses have divided the group into two subgroups; one containing
largely black-spotted and/or white-spotted whiprays (gerrardi-complex), and another of largely plain-coloured
whiprays (pastinacoides-complex) (see Fig. 3).
Genus Pateobatis gen. nov.
Type species Trygon uarnacoides Bleeker, 1852; newly proposed.
Definition. Medium-sized to large dasyatids (adults to 71–150 cm DW) characterised by the following: rather
depressed to robust, suboval to rhombic disc with pectoral-fin apex narrowly to broadly rounded; snout angular to
obtuse, rather short to elongate (1.7–5.5 times combined orbit and spiracle length); eye very small to small and
protruding slightly; nasal curtain skirt shaped; mouth very narrow to broad, with 2–4 oral papillae (absent in P.
hortlei); tail short to very long, whip-like (length 1.1–4.1 times DW), its base typically narrow and almost circular
in cross section; pelvic fins small, produced slightly or almost entirely concealed by disc; dorsal fold and ventral
folds absent; caudal sting close to tail base (distance from pectoral-fin insertion to caudal-sting base 1.6–2.2 times
interspiracular width); 1–3 variably developed, pearl-shaped mid-scapular thorns or thorns in row on nape; no
shoulder thorns; denticle band well developed with edge typically sharply defined, skin on rest of disc naked or
with patchy denticles; no row of enlarged median thorns on tail (except well developed in P. jenkinsii and
connected with row on disc); posterior tail often with fine denticles or prickly; dorsal surface plain; ventral surface
white, disc often dark edged; tail plain, not banded; marine and estuarine, Indo–West Pacific.
Etymology. Cryptic combination of the Latin pateo (lie open, be exposed) and batis (skate, ray, flatfish)
alluding the eclectic nature of members of this group.
Species. P. bleekeri (Blyth 1860), P. fai (Jordan & Seale, 1906), P. hortlei (Last, Manjaji-Matsumoto &
Kailola, 2006), P. jenkinsii (Annandale, 1909), and P. uarnacoides (Bleeker, 1852).
Remarks. Newly erected, morphologically heterogeneous genus consisting of five medium-size to very large,
marine whiprays previously placed in the genus Himantura. Two widely distributed and relatively abundant
species, Pateobatis fai and P. jenkinsii, have a broad rhombic disc with a short obtuse snout, whereas the other
three species have a characteristic subcircular disc with a long, pointed snout. These species cluster together using
mitochondrial data (Fig. 3), but analyses using nuclear genes may provide other insights.
Genus Urogymnus Müller & Henle, 1837
Urogymnus Müller & Henle, 1837:434. Type species Raja asperrima Bloch & Schneider, 1801; replacement name for
Gymnura Müller & Henle, 1837 (preoccupied by Gymnura van Hasselt, 1823).
Definition. Large to gigantic dasyatids (adults to 100–192 cm DW) characterised by the following: strongly
depressed to robust, suboval to almost circular disc with pectoral-fin apex broadly rounded; snout angular to
obtuse, rather short to very elongate (1.5–4.6 times combined orbit and spiracle length); eye very small to small
and protruding; nasal curtain skirt shaped; mouth very narrow to narrow, with 3–7 oral papillae (absent in U.

lobistoma); tail typically whip-like (but relatively short and stiff in U. asperrimus), short to long (length 1–3.1
times DW), its base typically narrow and oval to almost circular in cross section; pelvic fins very small to small,
almost entirely or fully concealed by disc; dorsal fold and ventral folds absent; caudal sting absent, or positioned
normally or well back on tail (distance from pectoral-fin insertion to caudal-sting base 1.8–3.3 times interspiracular
width); 1–3 mid-scapular thorns or thorns absent; no distinct shoulder thorn patch or median row on tail; denticle
band well developed, its edge typically diffuse; skin elsewhere, often very rough or prickly (extremely so in U.
asperrimus); posterior tail with fine denticles or very prickly; dorsal surface plain or almost plain; ventral surface
white, disc often dark edged; tail plain blackish or white, not banded; marine/estuarine/freshwater, Indo–West
Pacific.
Species. U. asperrimus (Bloch & Schneider, 1801), U. dalyensis (Last & Manjaji-Matsumoto, 2008), U.
granulatus (Macleay, 1883), U. lobistomus (Manjaji-Matsumoto & Last, 2006), U. polylepis (Bleeker, 1852), and
an undescribed species.
Remarks. Formerly monotypic, containing only the very spiny porcupine ray, Urogymnus asperrimus (Bloch
& Schneider, 1801). Molecular analysis confirms its placement in the Urogymninae (Fig. 5). Five additional, large
to very large species, including a new whipray from Australasia, were added to the group based on molecular data
(Fig. 3). These species all have a flexible, oval or subcircular disc, small or very small eyes, and relatively small
pelvic fins. Some live in freshwater. An undescribed species occurs in inshore habitats of Australasia.
Subfamily Hypolophinae
Hypolophinae Stromer, 1910:493 (subfamily); type genus Hypolophus Müller & Henle, 1837:117 (junior synonym of
Pastinachus Rüppell, 1829).
Pastinachinae Roberts, 2007:279 (subfamily); type genus Pastinachus Rüppell, 1829, by original designation.
Pastinachidae Lim, 2014:13 (family); type genus Pastinachus Rüppell, 1829 (not available).
Definition. Unique within the Dasyatidae in possessing a strongly arcuate upper jaw, quincuncial teeth in the lower
jaw in a long and broad plate, tail relatively broad based and depressed with the caudal sting inserted well
posteriorly (distance from pectoral-fin insertion to caudal-sting base exceeding 3.5 times interspiracular width), no
dorsal tail fold, a well-developed ventral tail fold that falls well short of the tail tip, and plain dorsal coloration
(usually greyish or black).
Genera. Makararaja and Pastinachus.
Remarks. The two hypolophin genera are morphologically distinct from each other, and from other genera of
the family Dasyatidae. Roberts (2007) characterised his new genus Makararaja by its very fine dermal denticles,
no denticle band on the median disc, a well-developed ventral tail fold, and a very posteriorly positioned caudal
sting. In Pastinachus, the disc is distinctly rhombic (rather than subcircular) with a well-developed denticle band
(usually with pearl-shaped midscapular thorns), the ventral tail fold is deep (shorter-based and less well-developed
in Makararaja) and terminates well before the tail tip, and the tail base of Pastinachus is relatively much broader.
Genus Makararaja Roberts, 2007
Makararaja Roberts, 2007:286. Type species Makararaja chindwinensis Roberts 2007; by original designation and monotypic.
Definition. Small dasyatids (to at least 39 cm DW) characterised by the following: relatively well-depressed, oval
to almost circular disc with pectoral-fin apex broadly rounded; snout obtuse and medium length (~1.7 times
combined orbit and spiracle length); eye small and protruding slightly; nasal curtain distinctly bilobed; mouth
narrow with 4 oral papillae; tail flexible and short (length ~1.5 times DW), its base moderately broad and
depressed; pelvic fins large, protruding slightly beyond disc; dorsal fold absent; ventral fold moderately deep with
a short base; caudal sting positioned very posteriorly on tail (distance from pectoral-fin insertion to caudal-sting
base ~5.3 times interspiracular width); skin on dorsal surface uniformly velvety, densely covered with minute
denticles and lacking a denticle band; no enlarged thorns on disc or tail (but with a short nuchal row of rudimentary
thorns); plain coloured dorsally; freshwater, South-East Asia.
Species. M. chindwinensis Roberts, 2007
Remarks. Poorly known (based on the holotype); more specimens and tissues needed.
Genus Pastinachus Rüppell, 1829
Pastinachus (subgenus of Trigon) Rüppell, 1829:51. Type species Raja sephen Forsskål, 1775; by subsequent designation.
Definition. Small to very large dasyatids (adults to 43–200 cm DW or more) characterised by the following:
depressed to robust, rhombic disc with pectoral-fin apex angular to narrowly rounded; snout obtuse or broadly
angular, and short to medium length (1.6–2.4 times combined orbit and spiracle length); eye small and protruding
slightly; nasal curtain distinctly bilobed; mouth narrow to medium-width with 5 oral papillae; tail firm and long to
very long (length 1.8–3.2 times DW), its base broad to very broad and noticeably depressed; pelvic fins large,
protruding slightly beyond disc; dorsal fold absent; ventral fold deep to very deep with a long base; caudal sting
positioned posteriorly on tail (distance from pectoral-fin insertion to caudal-sting base 3.5–4.6 times interspiracular
width); denticles on dorsal surface of disc patchy, denticle band well developed, but with diffuse edges; 1–3
variably developed, pearl or heart-shaped mid-scapular thorns, no other thorns on disc; tail prickly but lacking
enlarged thorns (except in Pastinachus stellurostris); plain coloured dorsally, ventrally white, disc margin often
dark; marine/estuarine, Indo–West Pacific.
Species. P. ater (Macleay, 1883), P. gracilicaudus Last & Manjaji-Matsumoto, 2010, P. sephen (Forsskål,
1775), P. solocirostris Last, Manjaji & Yearsley, 2005, and P. stellurostris Last, Fahmi & Naylor, 2010.
Remarks. Until recently, thought to consist of a single, widespread Indo–Pacific species, Pastinachus sephen
(Last & Stevens, 1994). Recent research (confirmed by molecular data) has demonstrated that the group is not
monotypic, and P. sephen is restricted to the northwestern Indian Ocean and another large stingray (P. ater) is more
widespread.
Acknowledgements
This project was supported by a National Science Foundation (NSF grant DEB-01132229) and the CSIRO Wealth
from Oceans Flagship. Many scientists and laypersons across the globe assisted this project in various ways over
the past few decades, by providing images, tissues and specimen data, or granting loans and permitting access to
invaluable research material at international museum collections. We thank all of you for your support in
completing this review. We particularly acknowledge John Pogonoski (CSIRO) for obtaining radiographs and
meristic data and providing valuable comments on the draft manuscript, Alastair Graham (CSIRO) for facilitating
loans, and William White (CSIRO) who modified and amended text in the molecular trees (all figures). The authors
received major input from Elisabeth Rochel, Lei Yang and Shannon Corrigan (College of Charleston, USA) who
provided input into molecular work to support the findings of this revision. Richard van der Laan and Bill
Eschmeyer provided advice on nomenclatural decisions and the Catalog of Fishes and associated papers were used
extensively during this project.
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533–561.
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with comments on the status of Dasyatis microphthalmus. Ichthyological Research, 53 (3), 290–297.
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Mitochondrial Estimate Based on 595 Species. In: Carrier, J.C., Musick, J.A. & Heithaus, M.R. (Eds.), Biology of Sharks
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W.T., Carvalho, M.R. de, Séret, B., Stehmann, M.F.W. & Naylor G.J.P. (Eds.), Rays of the World. CSIRO Publishing,
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Biology, 204, 379–394.
APPENDIX 1. Checklist of the family Dasyatidae.
Modified checklist (May 2016) Former identifications

Bathytoshia brevicaudata (Hutton, 1875) Dasyatis brevicaudata (Hutton, 1875)
Bathytoshia centroura (Mitchill, 1815) Dasyatis centroura (Mitchill, 1815)
Bathytoshia lata (Garman, 1880) Dasyatis lata (Garman, 1880)
Brevitrygon heterura (Bleeker, 1852) Himantura heterura (Bleeker, 1852)
Brevitrygon imbricata (Bloch & Schneider, 1801) Himantura imbricata (Bloch & Schneider, 1801)
Brevitrygon javaensis (Last & White, 2013) Himantura javaensis Last & White, 2013
Brevitrygon walga (Müller & Henle, 1841) Himantura walga (Müller & Henle, 1841)
Dasyatis chrysonota (Smith, 1828) Dasyatis chrysonota (Smith, 1828)
Dasyatis hypostigma Santos & Carvalho, 2004 Dasyatis hypostigma Santos & Carvalho, 2004
Dasyatis marmorata (Steindachner, 1892) Dasyatis marmorata Steindachner, 1892
Dasyatis pastinaca (Linnaeus, 1758) Dasyatis pastinaca (Linnaeus, 1758)
Dasyatis tortonesei Capapé, 1977 Dasyatis tortonesei Capape, 1977
Fluvitrygon kittipongi (Vidthayanon & Roberts, 2006) Himantura kittipongi Vidthayanon & Roberts, 2005
......continued on the next page

APPENDIX 1. (Continued)
Fluvitrygon oxyrhyncha (Sauvage, 1878) Himantura oxyrhyncha (Sauvage, 1878)
Fluvitrygon signifer (Compagno & Roberts, 1982) Himantura signifer Compagno & Roberts, 1982
*Fontitrygon colarensis (Santos, Gomes & Charvet-Almeida, Dasyatis colarensis Santos, Gomes & Charvet-Almeida,
2004) 2004
Fontitrygon garouaensis (Stauch & Blanc, 1962) Dasyatis garouaensis (Stauch & Blanc, 1962)
*Fontitrygon geijskesi (Boeseman, 1948) Dasyatis geijskesi Boeseman, 1948
Fontitrygon margarita (Günther, 1870) Dasyatis margarita (Günther, 1870)
Fontitrygon margaritella (Compagno & Roberts, 1984) Dasyatis margaritella Compagno & Roberts, 1984
Fontitrygon ukpam (Smith, 1863) Urogymnus ukpam (Smith, 1863)
Hemitrygon akajei (Müller & Henle, 1841) Dasyatis akajei (Müller & Henle, 1841)
Hemitrygon bennetti (Müller & Henle, 1841) Dasyatis bennetti (Müller & Henle, 1841)
Hemitrygon fluviorum (Ogilby, 1908) Dasyatis fluviorum Ogilby, 1908
Hemitrygon izuensis (Nishida & Nakaya, 1988) Dasyatis izuensis Nishida & Nakaya, 1988
Hemitrygon laevigata (Chu, 1960) Dasyatis laevigata Chu, 1960
Hemitrygon laosensis (Roberts & Karnasuta, 1987) Dasyatis laosensis Roberts & Karnasuta, 1987
Hemitrygon longicauda (Last & White, 2013) Dasyatis longicauda Last & White, 2013
Hemitrygon navarrae (Steindachner, 1892) Dasyatis navarrae (Steindachner, 1892)
Hemitrygon parvonigra (Last & White, 2008) Dasyatis parvonigra Last & White, 2008
Hemitrygon sinensis (Steindachner, 1892) Dasyatis sinensis (Steindachner, 1892)
Himantura leoparda Manjaji-Matsumoto & Last, 2008 Himantura leoparda Manjaji-Matsumoto & Last, 2008
Himantura uarnak (Gmelin, 1789) Himantura uarnak (Gmelin, 1789)
Himantura undulata (Bleeker, 1852) Himantura undulata (Bleeker, 1852)
Himantura sp. nov. Himantura sp. 4
Hypanus americanus (Hildebrand & Schroeder, 1928) Dasyatis americana Hildebrand & Schroeder, 1928
Hypanus dipterurus (Jordan & Gilbert, 1880) Dasyatis dipterura (Jordan & Gilbert, 1880)
Hypanus guttatus (Bloch & Schneider, 1801) Dasyatis guttata (Bloch & Schneider, 1801)
Hypanus longus (Garman, 1880) Dasyatis longa (Garman, 1880)
*Hypanus marianae (Gomes, Rosa & Gadig, 2000) Dasyatis marianae Gomes, Rosa & Gadig, 2000
Hypanus rudis (Günther, 1870) Dasyatis rudis (Günther, 1870)
Hypanus sabinus (Lesueur, 1824) Dasyatis sabina (Lesueur, 1824)
Hypanus say (Lesueur, 1817) Dasyatis say (Lesueur, 1817)
Maculabatis astra (Last, Manjaji-Matsumoto & Pogonoski, Himantura astra Last, Manjaji-Matsumoto & Pogonoski,
2008) 2008
Maculabatis gerrardi (Gray, 1851) Himantura gerrardi (Gray, 1851)
Maculabatis macrura (Bleeker, 1852) Himantura macrura (Bleeker, 1852)
Maculabatis pastinacoides (Bleeker, 1852) Himantura pastinacoides (Bleeker, 1852)
Maculabatis randalli (Last, Manjaji-Matsumoto & Moore, Himantura randalli Last, Manjaji-Matsumoto & Moore,
2012) 2012
Maculabatis toshi (Whitley, 1939) Himantura toshi Whitley, 1939
Maculabatis sp. nov. 1 Himantura sp. 1
Makararaja chindwinensis Roberts, 2007 Makararaja chindwinensis Roberts, 2007
......continued on the next page
APPENDIX 1. (Continued)
Megatrygon microps (Annandale, 1908) Dasyatis microps (Annandale, 1908)
Neotrygon annotata (Last, 1987) Neotrygon annotata (Last, 1987)
Neotrygon australiae Last, White & Seret, 2016 Neotrygon australiae Last, White & Seret, 2016
Neotrygon caeruleopunctata Last, White & Seret, 2016 Neotrygon caeruleopunctata Last, White & Seret, 2016
Neotrygon kuhlii (Müller & Henle, 1841) Neotrygon kuhlii (Müller & Henle, 1841)
Neotrygon leylandi (Last, 1987) Neotrygon leylandi (Last, 1987)
Neotrygon ningalooensis Last, White & Puckridge, 2010 Neotrygon ningalooensis Last, White & Puckridge, 2010
Neotrygon orientale Last, White & Seret, 2016 Neotrygon orientale Last, White & Seret, 2016
Neotrygon picta Last & White, 2008 Neotrygon picta Last & White, 2008
Neotrygon trigonoides (Castelnau, 1873) Neotrygon trigonoides (Castelnau, 1873)
Neotrygon varidens (Garman, 1885) Neotrygon varidens (Garman, 1885)
Pastinachus ater (Macleay, 1883) Pastinachus atrus (Macleay, 1883)
Pastinachus gracilicaudus Last & Manjaji-Matsumoto, 2010 Pastinachus gracilicaudus Last & Manjaji-Matsumoto, 2010
Pastinachus sephen (Forsskål, 1775) Pastinachus sephen (Forsskål, 1775)
Pastinachus solocirostris Last, Manjaji & Yearsley, 2005 Pastinachus solocirostris Last, Manjaji-Matsumoto &
Yearsley, 2005
Pastinachus stellurostris Last, Fahmi & Naylor, 2010 Pastinachus stellurostris Last, Fahmi & Naylor, 2010
Pateobatis bleekeri (Blyth 1860) Himantura bleekeri (Blyth 1860)
*Pateobatis fai (Jordan & Seale, 1906) Himantura fai Jordan & Seale, 1906
Pateobatis hortlei (Last, Manjaji-Matsumoto & Kailola, 2006) Himantura hortlei Last, Manjaji-Matsumoto & Kailola,
2006
*Pateobatis jenkinsii (Annandale, 1909) Himantura jenkinsii (Annandale, 1909)
Pateobatis uarnacoides (Bleeker, 1852) Himantura uarnacoides (Bleeker, 1852)
Pteroplatytrygon violacea (Bonaparte, 1832) Pteroplatytrygon violacea (Bonaparte, 1832)
Taeniura lymma (Forsskål, 1775) Taeniura lymma (Forsskål, 1775)
Taeniura sp. nov. Taeniura sp.
Taeniurops grabata (Geoffroy Saint-Hilaire, 1817) Taeniurops grabata (Geoffroy Saint-Hilaire, 1817)
Taeniurops meyeni (Müller & Henle, 1841) Taeniurops meyeni (Müller & Henle, 1841)
*Telatrygon acutirostra (Nishida & Nakaya, 1988) Dasyatis acutirostra Nishida & Nakaya, 1988
Telatrygon crozieri (Blyth, 1860) Dasyatis sp. 2
Telatrygon zugei (Müller & Henle, 1841) Dasyatis zugei (Müller & Henle, 1841)
Telatrygon sp. nov. Dasyatis sp. 1
Urogymnus asperrimus (Bloch & Schneider, 1801) Urogymnus asperrimus (Bloch & Schneider, 1801)
Urogymnus dalyensis (Last & Manjaji-Matsumoto, 2008) Himantura dalyensis Last & Manjaji-Matsumoto, 2008
Urogymnus granulatus (Macleay, 1883) Himantura granulata (Macleay, 1883)
Urogymnus lobistomus (Manjaji-Matsumoto & Last, 2006) Himantura lobistoma Manjaji-Matsumoto & Last, 2006
Urogymnus polylepis (Bleeker, 1852) Himantura polylepis (Bleeker, 1852)
Urogymnus sp. nov. Himantura sp. 5
*species with provisional placement requiring further investigation

Cambio Dasyatidae

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Zootaxa 4139 (3): 345–368 ISSN 1175-5326 (print edition)

A revised classification of the family Dasyatidae (Chondrichthyes:

PETER R. LAST1,4, GAVIN J.P. NAYLOR2 & B. MABEL MANJAJI-MATSUMOTO3

Accepted by M.R. de Carvalho: 16 Jun. 2016; published: 21 Jul. 2016 345

346 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

348 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

350 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

352 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Family Dasyatidae Jordan and Gilbert, 1879

Genus Bathytoshia Whitley, 1933

Genus Dasyatis Rafinesque, 1810

354 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Genus Hemitrygon Müller & Henle, 1838

Definition. Small to medium-sized dasyatids (adults typically to 31–66 cm DW, 93 cm DW in H. fluviorum)

Genus Hypanus Rafinesque, 1818

Genus Megatrygon gen. nov.

Type species Trygon microps Annandale, 1908:393; newly proposed, monotypic.

Genus Pteroplatytrygon Fowler, 1910

356 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Genus Taeniurops Garman, 1913

Genus Telatrygon gen. nov.

Subfamily Neotrygoninae subfam. nov.

Type genus Neotrygon Castelnau, 1873, newly proposed.

Genus Neotrygon Castelnau, 1873

358 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Genus Brevitrygon gen. nov.

Genus Fluvitrygon gen. nov.

Genus Fontitrygon gen. nov.

Type species Trygon margarita Günther, 1870; newly proposed.

360 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Genus Himantura Müller & Henle, 1837

Genus Maculabatis gen. nov.

Type species Trygon gerrardi Gray, 1851:116; newly proposed.

Genus Pateobatis gen. nov.

Type species Trygon uarnacoides Bleeker, 1852; newly proposed.

Genus Urogymnus Müller & Henle, 1837

362 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

Genus Makararaja Roberts, 2007

Genus Pastinachus Rüppell, 1829

364 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

APPENDIX 1. Checklist of the family Dasyatidae.

Modified checklist (May 2016) Former identifications

366 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

368 · Zootaxa 4139 (3) © 2016 Magnolia Press LAST ET AL.

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