Zootaxa 5047 (5): 557–566 ISSN 1175-5326 (print edition)

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Copyright © 2021 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5047.5.5
http://zoobank.org/urn:lsid:zoobank.org:pub:B7207292-F004-4393-B24A-2D3DDC6E8D3F

First records of two species of reef-associated mantis shrimps (Crustacea:

Stomatopoda) from India
VINAY P. PADATE1, SHANE T. AHYONG2, ALEESHA K. SHAJI3, SHERINE SONIA CUBELIO1,4 & NARAY-
ANANE SARAVANANE1
1
Centre for Marine Living Resources and Ecology, Ministry of Earth Sciences, Government of India, Atal Bhavan, LNG Terminus Road,
Ochanthuruth (PO), Puthuvype, Kochi 682 508, Kerala, India.
� [email protected]; https://orcid.org/0000-0002-2244-8338
� [email protected]; https://orcid.org/0000-0003-3405-4923
2
Australian Museum Research Institute, 1 William St., Sydney, NSW 2010, Australia, and School of Biological, Earth & Environmental
Sciences, University of New South Wales, Kensington, NSW 2052, Australia.
� [email protected]; https://orcid.org/0000-0002-2820-4158
3
Department of Marine Biology, School of Ocean Studies and Technology, Kerala University of Fisheries and Ocean Studies, Kochi,
682016, Kerala, India.
� [email protected]; https://orcid.org/0000-0001-6231-3326
4
Corresponding author: � [email protected]; https://orcid.org/0000-0002-2960-7055

Abstract

The coastal marine environments in the Indian Exclusive Economic Zone harbour a rich diversity of mantis shrimps.
However, several regions have not been adequately surveyed for their stomatopod faunal composition. In this paper,
we report the first records of two species, Gonodactylopsis drepanophora (de Man, 1902) and Cloridina malaccensis
(Manning, 1968) from Indian waters. Samples were collected using a chain dredge deployed by the Fishery Oceanographic
Research Vessel Sagar Sampada on the shallow reefs (53–56 m depths) of the Andaman Sea and Bay of Bengal during
August–November 2019. Gonodactylopsis drepanophora was previously known only from Indonesia, and Cloridina
malaccensis from Southeast Asia, New Caledonia and Madagascar.

Key words: Stomatopoda, reef-associated, taxonomy, new records, India

Introduction

The stomatopod fauna of India is comparatively well-documented, with 72 species reported so far (Trivedi et
al. 2020). Taxonomic studies during the colonial period contributed significantly to the documentation of these
organisms (Fabricius 1798; Heller 1865; Wood-Mason 1875, 1895; Wood-Mason & Alcock 1891; Henderson 1893;
Alcock 1894; Thurston 1895; Alcock & Anderson 1899; Lanchester 1903; Jurich 1904; Kemp 1911, 1915; Kemp
& Chopra 1921; Gravely 1927; Chopra 1934), particularly by the land-mark monograph and of Kemp (1913).
Post-independence, the introduction of mechanized fishing and expansion of the demersal fisheries to the offshore
regions have enabled several regional-level studies resulting in the reporting of several new and rare species (Tiwari
& Biswas 1952; Chhapgar & Sane 1967; Manning 1969a, 1975; Ghosh 1975; Ahyong & Kumar 2018; Niveditha et
al. 2019; Trivedi et al. 2020).
The Centre for Marine Living Resources and Ecology, Kochi (India) has conducted several fishing operations
in the Indian Exclusive Economic Zone (EEZ) to collect spatially and taxonomically resolved records of marine
crustaceans from the region, and maintains voucher specimens as part of natural history collections from the seas
surrounding India particularly the EEZ. The present study reports the first Indian records of two shallow-water
species collected from Andaman Sea and Bay of Bengal.

Accepted by J. Goy: 15 Sept. 2021; published: 4 Oct. 2021 557

Material and methods

Two cruises were conducted on-board the Fishery Oceanographic Research Vessel Sagar Sampada in the Andaman
Sea (cruise number 388) and Bay of Bengal (cruise number 392). Samples were collected using Chain Dredge
at 43–407 m in the Andaman Sea and 56–860 m in the Bay of Bengal. The specimens were photographed and
preserved in 70% ethanol. Specimen measurements were made using vernier calipers with an accuracy of 0.01 mm.
Material examined during this study was deposited as reference vouchers at the Referral Centre of the Centre for
Marine Living Resources and Ecology, Kochi (India).
Morphological terminology follows Ahyong (2001, 2012). Abbreviations used in the following text are listed
in alphabetical order: CL = Carapace length, measured along the midline, excluding the rostral plate; FORVSS =
Fishery Oceanography Research Vessel Sagar Sampada; IO/SS/STO = Indian Ocean/ Sagar Sampada/ Stomatopoda;
TL = Total length, measured along the dorsal midline, from the tip to the rostral plate to the apices of the submedian
teeth of the telson.

Systematics

Order Stomatopoda Latreille, 1817

Superfamily Gonodactyloidea Giesbrecht, 1910

Family Gonodactylidae Giesbrecht, 1910

Genus Gonodactylopsis Manning, 1969b

Gonodactylopsis drepanophora (de Man, 1902)

(Figs. 1A, 2A–H, 3A–H)

Gonodactylus drepanophorus De Man, 1902: 914, pl. 27, fig. 68, 68a [type locality: Ternate, Indonesia].—Kemp 1913: 11 (list),
148 (key), 173.—Hansen 1926: 30, pl. 2, fig. 1a.
Gonodactylopsis drepanophora.—Manning 1969b: 150 (key).—Moosa 1974: 73, figs. 1a–c. —Manning, 1995: 20 (list).—Erd-
mann & Manning 1998: 621 (remarks).
Mesacturus drepanophorus.—Manning 1967: 2.—Shanbhogue 1975: 524 (list), 539 (key).

Material examined. IO/SS/STO/00008, 1♀, TL 24.0 mm, Andaman Sea, east of Little Andaman Island, FORVSS
38806, 10.72°N, 92.71°E, 53 m, chain dredge, coll. Vinay P. Padate, 10 August 2019.
Description of Indian specimen. Body subcylindrical, convex. Eye large, reaching beyond distal end
of second antennular segment. Cornea subglobular, as wide as eyestalk. Ocular scales rounded, separate; anterior
margin of ophthalmic somite triangular, small distal point. Antennular somite dorsal processes low, margin irregular.
Antennular peduncle 0.52 CL. Antennal protopod with articulated plate dorsally, small distoventral spine; antennal
scale with setose margins, length 0.31 CL.
Rostral plate trispinous, median spine slender, not reaching corneal base, lateral spines wider, triangular, directed
anterolaterally. Carapace anterolateral and posterolateral angles rounded; dorsal surface smooth, carinae absent;
gastric grooves present, cervical groove absent.
Mandibular palp present, with 2 articles. Maxillipeds 1–5 each with epipod, first epipod largest; maxillipeds 1,
3–5 with ischium elongate, merus moderately long, carpus short, propodus ovate, dactylus elongated, apposed to
propodus. Maxilliped 5 merus and carpus slightly expanded ventrally.
Raptorial claw propodus occlusal margin sparsely pectinate proximally, bearing 1 movable spine proximally;
dactylus outer margin strongly inflated proximally, with basal notch; occlusal margin finely serrated distally.
Thoracic somites 5–8 smooth dorsally, lateral margins rounded.
Abdominal somites 1–5 smooth dorsally, bearing low marginal carina and adjacent lateral depression, unarmed;
somite 6 dorsal surface with submedian, intermediate and lateral bosses, each terminating in spine.
Telson wider than long, with 3 pairs of primary teeth: submedian teeth long, with movable apices, bearing 5–6

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blunt dorsal spines; intermediate teeth moderately long, bearing 1 pair of blunt dorsal spines; lateral teeth sharp,
shorter than half length of intermediate teeth; 13 submedian denticles. Dorsal surface with well-separated carinae.
Median carina weakly inflated, terminating posteriorly in blunt tooth, followed by blunt median spine; accessory
median carinae with 2 pairs of posterior directed short spines; anterior submedian carinae terminating posteriorly in
blunt spine, followed by 5 blunt spines, of which anterior 4 spines arranged in two pairs, fifth spine located at base of
submedian tooth; short spine on inner side of base of submedian tooth; anterior intermediate carina bearing 3 spines,
posteriormost spine located at base of intermediate tooth. Ventral surface smooth, submedian and intermediate teeth
with longitudinal carina.

FIGURE 1. Dorsal habitus: A, Gonodactylopsis drepanophora (de Man, 1902), ♀, TL 24.0 mm, IO/SS/STO/00008; B, Cloridi-
na malaccensis (Manning, 1968), ♂, TL 68.0 mm, IO/SS/STO/00016. Scale: 5 mm.

Uropodal protopod bearing 2 gently curved primary spines, outer spine longer, wider, inner margin smooth,
overreaching posterior margin of exopod; slender spine above articulation of exopod; endopod shorter than exopod,
crescent-shaped, distal tip curved inwards, outer margin setose, inner margin without setae; exopod proximal article
with 12 movable spines on outer margin, posterior 8 spines distinctly recurved, distalmost spine not reaching end of
distal exopod article; distal article length about twice width, outer margin setose, inner margin glabrous.
Colour (preserved in ethanol). Body pale brown, carapace with scattered dark pigmentation anteriorly
between gastric groove and lateral plate, elongated dark spot on inner side of posterior half of gastric groove, dark
pigmentation on cervical region comprising one median and pair of submedian dark spots in addition to scattered
dark pigmentation. Thoracic sternite 6 with dark pigmentation including median spot. Posterior half of abdominal
somite 1 with dark median spot surrounded laterally and posteriorly by dark mottled patch; somites 3–5 with
small dark median spot flanked by 1 pair of submedian spots, somite 6 with minute dark spot anteriorly between
submedian and intermediate carinae. Telson with minute dark spot anterior to submedian carinae. Cornea with
dark pigmentation. Raptorial claw, pereiopods pale grey; telson, uropodal protopod and proximal segment brown;
uropodal endopod and distal segment of exopod white.
Remarks. Gonodactylopsis drepanophora was originally described in the genus Gonodactylus from a female
specimen (TL 18.5 mm) from Ternate, Indonesia (De Man 1902: pl. 27, fig. 68, 68a). Kemp (1913) provided a short
diagnosis of this species based on De Man’s account, and included it in the identification key to species. Hansen

FIRST RECORDS OF STOMATOPODS Zootaxa 5047 (5) © 2021 Magnolia Press · 559
(1926: pl. 2, fig. 1a) reported a small specimen (11.8 mm, sex indeterminate) from Timor, Indonesia, which differed
from the holotype in having: (1) the outer spine of the uropodal protopod posteriorly extending beyond exopod;
(2) uropodal exopod with the proximal segment more posteriorly prolonged, and bearing 9–10 recurved spines on
outer margin; (3) uropodal exopod with slightly oval-shaped distal segment with length lesser than two times width,
setose in distal half; (4) uropodal endopod with inner margin thickened and strongly curved distally, distal half of
outer margin setose. These observations of Hansen (1926) reflect the juvenile condition of his specimen. Manning
(1967) tentatively assigned G. drepanophora to Mesacturus Miers, 1880, and then Manning (1969b) established
Gonodactylopsis Manning, 1969b (for G. herdmani Tattersall, 1906 and G. drepanophora). Manning (1969b)
characterized Gonodactylopsis by the combination of a trispinous rostral plate, subglobular cornea, carapace lateral
plates with rounded anterior margins, presence of a mandibular palp, and subterminal articulation of the uropodal
exopod segments, with the proximal segment extending well beyond articulation with the distal segment. Moosa
(1974: fig. 1a–c) reported a TL 14 mm female specimen from Ambon Bay, Indonesia, and provided a detailed
morphological description for the same.

Figure 2. Gonodactylopsis drepanophora (de Man, 1902), ♀, TL 24.0 mm, IO/SS/STO/00008 (colour photographs). A, dor-
sal view of carapace; B, dorsal view of ocular and rostral plates; C, ischio-meral articulation of raptorial claw; D, external view
of raptorial claw dactylus and propodus; E, propodi of maxillipeds 3–4; F, thoracic somites 5–8; G, dorsal view of abdomen,
telson and uropods; H, ventral view of telson and uropods. Scale: A, G, H, 2 mm; C–F, 1 mm; B, 500 μm.

Since then, only two species of Gonodactylopsis were known until Erdmann & Manning (1998) described a
new species Gonodactylopsis komodoensis Erdmann & Manning, 1998, from Indonesia. The three known species
of Gonodactylopsis are distinguished in the key below.
The present Andaman Sea specimen agrees with the original description and illustrations of holotype of G.
drepanophora, but differs from the other previously reported Indonesian specimens in possessing greater number of
blunt spines on the dorsal surface of submedian and intermediate marginal teeth. These differences are attributed to
allometry, as the present specimen is the largest known of G. drepanophora.
Geographical distribution and habitat. Indonesia: Ternate, Halmahera (De Man 1902), Timor (Hansen 1926),
Ambon Bay, Maluku (Moosa 1974); margins delimiting mud and coral, among Lithothamnium (Hansen 1926), also

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in mud and rubble, 34–80 m depth (Hansen 1926; Moosa 1974). The present specimen was collected from coral reef
at 53 m depth and represents the first record of the species from Indian Andaman Sea waters.

FIGURE 3. Gonodactylopsis drepanophora (de Man, 1902), ♀, TL 24.0 mm, IO/SS/STO/00008 (line illustrations). A, dorsal
view of carapace; B, dorsal view of ocular and rostral plates; C, ischio-meral articulation of raptorial claw; D, external view of
raptorial claw dactylus and propodus; E, propodi of maxillipeds 3–4; F, thoracic somites 5–8; G, dorsal view of abdomen, telson
and uropods; H, ventral view of telson and uropods. Scale: A, G, H, 2 mm; C–F, 1 mm; B, 500 μm.

Key to species of Gonodactylopsis

1. Telson lateral teeth indicated as rounded lobe projecting from telson margin, separated from margin of intermediate teeth by
V-shaped incision; submedian denticles absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. herdmani
– Telson lateral teeth separated from margin of intermediate teeth by deep V-shaped incision; submedian denticles present . . . 2
2. Uropodal protopod inner primary spine longer than outer spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. komodoensis
– Uropodal protopod inner primary spine shorter than outer spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. drepanophora

Superfamily Squilloidea Latreille, 1802

Family Squillidae Latreille, 1802

Genus Cloridina Manning, 1995

Cloridina malaccensis (Manning, 1968)

(Figs. 1B, 4A–H, 5A–H)

Clorida malaccensis Manning, 1968: 244, fig. 2a–e.—Manning 1978: 26.—Naiyanetr 1980a: 43 (list).—Naiyanetr 1980b:
54.—Moosa 1986: 399.—Moosa 1991: 202.

FIRST RECORDS OF STOMATOPODS Zootaxa 5047 (5) © 2021 Magnolia Press · 561
Cloridina malaccensis.—Manning 1995: 24 (list), 180 (remarks), 192 (key).—Moosa 2000: 410 (list), 439.—Ahyong 2001:
230 (list), 231 (key).—Ahyong & Naiyanetr 2002: 306 (list).—Ahyong 2007: 335.—Naiyanetr 2007: 142 (list).—Poupin
2010: 39 (list).

Material examined. IO/SS/STO/00016, 2♂ (TL 62.0–68.0 mm), Bay of Bengal, India, off Nagapattinam, Tamil
Nadu, 10.79°N, 80.11°E, 56 m, FORVSS 39214, chain dredge, coll. Vinay P. Padate, 30 November 2019.
Description of Indian specimens. Body depressed. Eye small, reaching distal end antennular article 1. Eye-
stalk bulging, widest in proximal half. Cornea bilobed, dark, as wide as eyestalk. Ocular scales fused into single
plate, rounded distally; anterior margin of ophthalmic somite rounded. Antennular somite dorsal processes triangu-
loid, with sharp apices directed anterolaterally. Antennular peduncle 0.69–0.71 CL. Antennal protopod unarmed;
antennal scale margins fully setose, length 0.46 CL.

FIGURE 4. Cloridina malaccensis (Manning, 1968), ♂, TL 68.0 mm, IO/SS/STO/00016 (colour photographs). A, dorsal view
of carapace; B, dorsal view of ocular and rostral plates; C, ischiomeral articulation of raptorial claw; D, external view of rapto-
rial claw dactylus and propodus; E, propodi of maxillipeds 3–4; F, thoracic somites 5–8; G, dorsal view of abdomen, telson and
uropods; H, ventral view of telson and uropods. Scale: A, C–H, 10 mm; B, 5 mm.

Rostral plate subtriangular, longer than wide. Carapace with anterolateral spine, posterolateral margin rounded.
Dorsal surface with only marginal carina on posterolateral margin; gastric and cervical groove present.
Mandibular palp present, with 3 articles. Maxillipeds 1–4 each with epipod, epipod 1 largest; maxillipeds 1, 3–5
ischium elongate, merus moderately long, carpus short, propodus ovate, dactylus elongated, apposed to propodus.
Raptorial claw ischiomeral articulation terminal. Carpus bearing sharp dorsal ridge. Propodus occlusal margin
closely pectinate in sinuous row, bearing 3 movable spines proximally. Dactylus outer margin with proximal notch
and large triangular lobe; occlusal margin bearing 5 teeth, basal tooth minute.
Thoracic somites 5–8 smooth, without carinae; somite 5 lateral process short slender spine, directed anterolater-
ally, with short ventral spine. Somites 6–7 lateral process rounded, undivided.

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FIGURE 5. Cloridina malaccensis (Manning, 1968), ♂, TL 68.0 mm, IO/SS/STO/00016 (line illustrations). A, dorsal view of
carapace; B, dorsal view of ocular and rostral plates; C, ischiomeral articulation of raptorial claw; D, external view of raptorial
claw dactylus, propodus, carpus and distal merus; E, maxilliped 3 propodus; F, thoracic somites 5–8; G, dorsal view of abdomen,
telson and uropods; H, ventral view of telson and uropods. Scale: A–E, 2 mm; F–G, 10 mm, H, 5 mm.

Abdominal somites 1–5 with intermediate, lateral and marginal carinae; somite 6 with submedian, intermediate
and lateral carinae. Abdominal carinae spined as follows: submedian 6, intermediate 5–6, lateral 5–6, marginal 5.
Telson trianguloid-subquadrate, inflated, dorsal surface with rows of tubercles, median carina sharp, elevated,
with small posterior spine; submedian, intermediate and lateral carinae slender; apices of submedian carinae mov-
able; prelateral lobe slightly shorter than margin of lateral tooth; denticles: submedian 3, intermediate 8–9, lateral 1.
Ventral surface with tuberculate postanal-carina.
Uropodal protopod bearing 2 curved primary spines, inner spine longer; lobe on outer margin of inner spine
wider than adjacent spine; protopod inner margin with row of 7 spines; endopod shorter than exopod; exopod distal
article longer than proximal; proximal article with 6 blunt movable spines on outer margin, distalmost not reaching
midlength of distal exopod article.
Colour (in life). Body (including carapace, thoracic sternites, abdomen, telson, and raptorial claw merus) light
brown with brownish mottling; cornea with dark pigmentation; raptorial claw carpus and propodus, as well as
pereiopods pale grey; raptorial claw dactylus greyish with basal half reddish; uropodal endopod with distal two-
thirds black, exopod proximal article with distal half of inner side black, distal segment with inner half black.
Remarks. Cloridina malaccensis was originally described (as Clorida malaccensis) from a single female (TL
55.8 mm) from Malacca Strait, Indonesia (Manning 1968: fig. 2a–e) and subsequently from Madagascar (Manning
1978), Thailand (Naiyanetr 1980a, 1980b), the Philippines (Moosa 1986), and New Caledonia (Moosa 1991; Ahy-
ong 2007). Manning (1978) synonymised Clorida malaccensis var. moluccensis Moosa, 1973 with C. malaccensis
but Ahyong (2001) showed the two species to be distinct and provided a key to the species of the genus. The present

FIRST RECORDS OF STOMATOPODS Zootaxa 5047 (5) © 2021 Magnolia Press · 563
specimens agree well with the original description of C. malaccensis except that the postanal carina is tuberculate
rather than smooth. We regard this variation as part of normal intraspecific variation that is often present in members
of the Clorida-group of genera (Ahyong 2005; Van Der Wal et al. 2019) including species of Clorida Eydoux &
Souleyet, 1842, Cloridina Manning, 1995, Fallosquilla Manning, 1995 and Levisquilla Manning, 1995 (see Ahyong
2001).
Geographical distribution and habitat. Malacca Straits (Manning 1968), Madagascar (Manning 1978),
Andaman Sea (Naiyanetr 1980a, 1980b), Philippines (Moosa 1986), New Caledonia (Moosa 1991; Ahyong 2007),
Gulf of Thailand (Naiyanetr 2007); 42 m in muddy sand (Manning 1978), 36–37 m (Moosa 1986), 29–80 m in mud,
fine sandy mud with bryozoans to coarse sand and blocks (Moosa 1991). The present specimens were collected
from sandy silt substratum at 56 m depth. The present observation is the first record from the Indian waters.

Acknowledgements

The authors are grateful to the Director, CMLRE, Kochi, India, for providing the opportunity to work on the
taxonomy of deep-water crustaceans of the Indian EEZ. We thank the chief scientists, fishing hands and participants
of the FORV “Sagar Sampada” cruise numbers 388 and 392 for meticulously collecting the specimens. The authors
wish to express their gratitude to the Associate Editor, Dr. Joseph Goy and the two anonymous reviewers for the
critical review that improved the quality of the manuscript.

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