Revision of the Indo-West Pacific Coral Reef Mantis Shrimp Genus, Gonodactylopsis Manning, 1969 (Crustacea: Stomatopoda: Gonodactylidae)

. The Indo-West Pacific coral reef mantis shrimp genus genus, Gonodactylopsis Manning, 1969, was established for two Indo-West Pacific species: Gonodactylus herdmani Tattersall, 1906 (type species) described from the Gulf of Mannar, India; and Gonodactylus drepanophorus de Man, 1902, from Indonesia. A third species, Gonodactylopsis komodoensis Erdmann & Manning, 1998, was described from Komodo, Indonesia. Gonodactylopsis is revised herein based on type and other material of the three known species, and two new species are described based on recent collections from coral reefs: one from Macclesfield Bank, South China Sea, and the other from Fiji and Papua New Guinea. Reconsideration of all species of Gonodactylopsis revealed a probable synapomorphy of the genus that is unique in the Stomatopoda: the presence of membranous, partially de-chitinized posterior surfaces of the uropodal endopod and exopod distal article. All species of Gonodactylopsis are described and figured, and an identification key is provided.


Introduction
proposed the genus Gonodactylopsis to accommodate two species of mantis shrimp formerly placed in Gonodactylus Group II of Kemp (1913) (recognized as Mesacturus Miers, 1880by Holthuis, 1967, characterized by the combination of a trispinous rostral plate, anteriorly produced lateral plates of the carapace, presence of a mandibular palp, subterminal articulation of the uropodal exopod segments, and weakly curved, movable distal outer spines on the proximal article of the uropodal exopod. More recently, it was recognized that the upraised conical spine or boss at the base of the submedian and intermediate teeth of the telson links Gonodactylopis with Hoplosquilla Holthuis, 1964 and a group of species within Gonodactylellus Manning, 1995(Barber & Erdmann, 2000Ahyong, 2001;Ahyong & Erdmann, 2007).
Prior to the present study, three species of Gonodactylopsis, all from Indo-West Pacific coral reefs were recognized: G. herdmani (Tattersall, 1906) (type species, Gulf of Mannar), G. drepanophora (de Man, 1902) (type locality: Ternate, Indonesia) and G. komodoensis Erdmann & Manning, 1998 (type locality: Komodo, Indonesia). The few published reports of Gonodactylopsis are based on only a few specimens in collections worldwide. In the present study, the known species of Gonodactylopsis are redescribed based on type and other material, and two new species are described based on expeditionary materials from various Indo-West Pacific localities collected since the 1980s.

Materials and methods
Morphological terminology follows Ahyong (2001) and . Total length (TL) is measured along the dorsal midline from the tip of the rostral plate to the apices of the submedian teeth of the telson. Carapace length (CL) is measured along the dorsal midline of the carapace and excludes the rostral plate. The abdominal-width carapace-length index (AWCLI) is given as 1000 × (width of abdominal somite 5)/CL. Specimens are deposited in the collections of: the Australian Museum, Sydney (AM); Muséum national d'Histoire naturelle, Paris (MNHN); Museum Zoologicum Bogoriense, Cibinong, Indonesia (MZB); Senckenberg Museum Forschunginstitut, Frankfurt, Germany (SMF); National Museum of Natural History, Smithsonian Institution, Washington D.C. (USNM); National Taiwan Ocean University, Keelung (NTOU); the Natural History Museum, London (NHM); and the Western Australian Museum, Perth (WAM). The distribution map ( Fig. 1) was prepared using QGIS 3.4.
Diagnosis. Eye subcylindrical, cornea subglobular. Ocular scales low, separate, not wider than basal width of median spine of rostral plate. Rostral plate trispinous, with slender median spine and shorter, wider lateral spines. Carapace anterolateral margins convex, extending anteriorly beyond base of rostral plate. Mandibular palp present. Raptorial claw propodus with proximal movable spine. Telson with submedian, intermediate and lateral primary teeth; mid-dorsal surface with median, accessory median, anterior submedian and anterior intermediate carinae, each posteriorly armed; submedian and intermediate primary teeth with 1 or more upright basal spines or bosses; carina of intermediate tooth without accessory carina on mesial margin; anus located ventrally. Uropodal protopod without lobes between terminal spines; exopod inner margin glabrous, distal article outer margin setose; endopod inner margin glabrous, outer margin distal half setose; distal exopod article and endopod with soft, membranous, glabrous mesial surface.
Present reconsideration of all species of Gonodactylopsis revealed a likely synapomorphy supporting monophyly of the genus: all species have membranous, partially de-chitinized posterior surfaces of the uropodal endopod and exopod distal article, these features being best developed in adults, although still evident in juveniles (Figs 2H,J,3E,H,I,4F,H,5H,J,6H,J,9H,K). In all other stomatopods, the articles of the uropod are fully chitinized. Species of Gonodactylopsis usually also have more elaborately developed, sharply cristate dorsal uropodal carinae (except G. komodoensis), which instead are less pronounced in other gonodactylids. Preliminary phylogenetic analysis (Barber & Erdmann, 2000) suggests that Gonodactylopsis is most closely related to Hoplosquilla but the two genera are readily separated by the presence of the absence of the mandibular palp in the latter (in addition to the aforementioned uropodal differences). Gonodactylopsis is readily distinguished from the Gonodactyllelus molyneux group in the trispinous rostral plate, which, in species of Gonodactyllelus, has rounded or angular, rather than spinous anterolateral margins of the basal portion. The speciose genus Gonodactylellus is not monophyletic and the G. molyneux group is currently under evaluation to determine whether it warrants separate generic status or inclusion within a redefined Gonodactylopsis and Hoplosquilla. (Tattersall, 1906) Figs 1, 2
Abdominal Uropodal protopod terminal spines with outer slightly longer than inner, each with prominent ventral carina, inner with dorsal carina; upper proximal surface smooth behind dorsal carina, without obtuse swelling; stout spine above exopod articulation. Exopod proximal article dorsal surface with sharp, sinuous longitudinal carina and short carina mesioproximally; outer margin with 10 or 11 movable spines, distalmost reaching distal one-third of distal article; inner margin smooth, glabrous; distal margin with small ventral spine. Exopod distal article with outer distal margin setose, inner margin smooth, glabrous; dorsal and ventral surfaces with inner one-third membranous, soft, wrinkled, clearly demarcated from chitinized outer portion; outer chitinized portion with sharp, curved carina dorsally, ventrally with interrupted longitudinal carina adjacent to margin of membranous portion. Endopod length 2.8-3.2× width, crescentic, articulation with protopod posterior to anterior end; outer margin strongly convex, setose along distal half, inner margin weakly, irregularly concave, glabrous; dorsally with sharp, sinuous carina adjacent to outer margin; dorsal and ventral surfaces with inner half membranous, soft, wrinkled, clearly demarcated from chitinized outer half.
Colouration. According to Kemp (1913: 172): "Living specimens are quite pale in colour with dull yellow marbling, darkest at the antero-lateral corners of the abdominal somites and tending to a more reddish tone on the sixth somite and telson. The ridges and tubercles of these last two segments are pure white. The propodus of the raptorial claw has a red-brown patch at the extreme distal end, and near the apex of each of the rostral spines, there is a transverse red band".
Remarks. Gonodactylopsis herdmani is unique in the genus for lacking submedian denticles on the telson, having short, stout intermediate telson teeth, and possessing low, convex lateral telson teeth that do not project posteriorly. The species remains known only from the type material and specimens reported by Kemp (1913), collected from the pearl banks in the Gulf of Mannar, Sri Lanka.
The two specimens of G. herdmani examined agree well in most respects (Fig. 2), differing in minor aspects of telson tubercle arrangement, the number of spines on the accessory median carina (1 or 2), the more prominent lateral telson tooth in the TL 26 mm specimen, and in the segmentation of the mandibular palp. Three mandibular palp articles are clearly demarcated in the TL 28 mm specimen whereas the two distal articles (articles 2 and 3) are indistinctly differentiated in the TL 26 mm specimen. Kemp (1913: 171) also reported a "rather indistinct" division between mandibular palp articles in his material of G. herdmani, but between the two proximal articles rather than the distal two as observed here; Kemp's (1913) reference to the proximal palp articles is probably a lapsus given that the proximal article typically differentiates in advance of the distal articles in stomatopods. The TL 26 mm female (NHM 1906.10.27.1), being in more complete condition than the TL 28 mm specimen, is herein selected as the lectotype to fix the identity of the species.
Distribution. Central Indian Ocean, presently known only from the Gulf of Mannar, Sri Lanka.
Telson wider than long; 10-14 submedian denticles arising from inner margin of submedian teeth; 2 intermediate and 1 lateral denticles usually distinct; submedian and intermediate teeth slender, sharp; submedian teeth subparallel to slightly divergent; intermediate teeth length at least twice width, extending posteriorly to slightly beyond midlength of submedian teeth; lateral teeth stout, triangular, apex acute, directed posteriorly. Median carina inflated, more so in males than females, with posterior spine; accessory median with 2 (rarely 3) spines; anterior submedian carina composed of 2-4 spines in longitudinal row or cluster; submedian tooth with anterior cluster and irregular row of 1-7 dorsal spines ( Uropodal protopod terminal spines with outer slightly longer than inner, each with ventral carina, inner with dorsal carina; upper proximal surface with obtuse swelling behind dorsal carina in specimens > TL 14 mm; slender spine above exopod articulation. Exopod proximal article dorsal surface with sharp, straight longitudinal carina and two short subequal carinae mesial to main carina; outer margin with 10-12 movable spines, distalmost reaching distal one-third of distal article; inner margin smooth, glabrous; distal margin with small ventral spine. Exopod distal article with outer distal margin setose, inner margin glabrous; dorsal and ventral surfaces with inner quarter to half membranous, soft, wrinkled, clearly demarcated from chitinized outer portion; outer chitinized portion with prominent, curved carina dorsally, ventrally smooth. Endopod length 2.1-3.1× width, crescentic, articulation with protopod posterior to anterior end; outer margin strongly convex, setose along distal half, inner margin strongly, irregularly concave, glabrous; dorsally with sharp, curved carina adjacent to outer margin; dorsal and ventral surfaces with inner half membranous, soft, wrinkled, clearly demarcated from chitinized outer half.

Colouration in preservative.
Largely faded to pale yellowbrown. Carapace with scattered chromatophores, mottling and pair of small spots across cervical region. Thoracic somite 6 and abdominal somite 1 with rectangular field of small irregular chromatophores and larger median black spot. Abdominal somites 2-5 with transverse row of 2 or 3 small rounded spots.
Remarks. Gonodactylopsis drepanophora is distinguished by the combination of a crescentic uropodal endopod and a telson having slender, pointed primary teeth and a multispinose dorsal surface. Of the known species of the genus, G. drepanophora most closely resembles G. maqqaba from the South China Sea, differing chiefly in the crescentic rather than linear uropodal endopod; other distinguishing features are discussed under the account of the latter. Gonodactylopsis drepanophora was described from Ternate, Indonesia, and subsequently recorded from Timor Leste (Hansen, 1926), Ambon, Indonesia (Moosa, 1974) and the Andaman Sea (Padate et al., 2021). The present records of G. drepanophora from Japan and northwestern Australia are the first for these localities.
The specimens of G. drepanophora accord well, particularly those from Japan, Indonesia, and northwestern Australia. The holotype, however, exhibits an abnormality in the absence of the left lateral primary tooth of the telson (Fig. 3E), possibly the result of injury during moulting. The record from the Andaman Sea represents the largest known specimen of the species (TL 24 mm; Padate et al., 2021: figs 1A, 2, 3), and differs subtly from the smaller specimens in having a 2-rather than 3-articled mandibular palp, slightly more divergent submedian telson teeth, and a cluster of 3 or 4 spines, rather than a single spine, at the base of the intermediate telson teeth. These differences, for which the significance is not clear, are presently regarded as intraspecific variation.
Allometric variation in G. drepanophora follows a similar trajectory to that observed for G. lata in the increasing slenderness and elongation of the primary telson teeth, increasing density of dorsal telson spination, and more pronounced curvature of the uropodal endopod with increasing body size (Hansen, 1926: pl. 2 fig. 1a; Figs 3E,H, 4F,K). The dorsal tubercle on the uropodal protopod is present in all except the smallest specimen (juvenile male, TL 12 mm, WAM C54274; Fig. 3I). As with G. lata, the lateral lobe on the posterior "endite" of pleopod 1 in the juvenile male is yet to be fully developed (Fig. 3J). Notably, the proportionally shorter primary telson teeth of the juvenile male of G. drepanophora (Fig. 3I) closely resemble those of adults of members of the G. molyneux group, such as G. barberi Erdmann, 2007 andG. snidvongsi (Naiyanetr, 1987) (Ahyong & Erdmann, 2007: fig. 4;Ahyong, 2008: fig. 2), suggesting these and allied species could be paedomorphic.

Gonodactylopsis komodoensis
Rostral plate slightly wider than long; median spine about twice length of basal portion (medially), laterally compressed, with obtusely angular ventral keel; lateral spines divergent with weakly arcuate margins.
Telson wider than long to as long as wide; 6-14 (usually 10 or 11) submedian denticles arising from inner margin of submedian teeth; Uropodal protopod terminal spines with outer slightly longer than inner, smooth ventrally, neither with longitudinal carina, inner with low dorsal carina; upper proximal surface without obtuse swelling behind dorsal carina; slender spine above exopod articulation. Exopod proximal article dorsal surface with 2 longitudinal carinae, one submedially and one adjacent to inner margin; outer margin with 9-11 movable spines, distalmost reaching almost to end of distal article; inner margin smooth, glabrous; distal margin with small ventral spine. Exopod distal article with outer distal margin setose, inner margin glabrous; dorsal and ventral surfaces with inner one-fourth to one-fifth thickened, smooth, membranous but firm, diffusely demarcated from chitinized outer portion; outer chitinized portion dorsally and ventrally unadorned. Endopod length 2.7-3.3× width, spatulate, linear, spatulate, articulation with protopod at anterior end; outer margin gently convex, setose along distal half, inner margin almost straight glabrous; dorsally with curved carina adjacent to outer margin; dorsal inner quarter and ventral inner half or slightly less membranous but firm, diffusely demarcated from chitinized outer portion.

Colouration in preservative.
Completely faded to pale yellow-brown.
Remarks. Gonodactylopsis komodoensis is the least ornamented species of the genus, lacking the numerous tubercles or spines on the dorsal telson carinae that are present in other congeners, and in fewer and lower uropodal carinae (rather than sharply cristate) as in other gonodactylids. Instead, the dorsal telson carinae of G. komodoensis are unarmed apart from the tiny posterior spinule on the carinae and basal tubercles of the submedian and intermediate teeth. Similarly, the primary spines of the uropodal protopod of G. komodoensis lack the ventral longitudinal carina that is present in congeners. Additionally, the de-chitinized membranous posterior surfaces of the uropodal endopod and exopod distal article are comparatively less pronounced in G. komodoensis than in congeners, lacking wrinkling, and with a diffuse rather than distinct line of demarcation from the chitinized cuticle. In other species of Gonodactylopsis, the soft, membranous portion of the uropodal cuticle is irregularly wrinkled and sharply demarcated from the adjacent chitinized surface but in G. komodoensis, the membranous surface is smooth, and although swollen, it is diffusely demarcated from the adjacent chitinized cuticle. Additionally, the membranous portion in G. komodoensis is more extensive ventrally than dorsally, rather than similar on both sides in congeners. As such, G. komodoensis presents uropod features that could be transitional between other species of Gonodactylopsis and members of the Gonodactylellus molyneux group and Hoplosquilla, which have fully chitinized uropod articles.
As observed by Erdmann & Manning (1998), allometric changes are evident in the proportional elongation of the primary telson teeth (particularly the submedians) and reduction in the number of submedian denticles. The primary telson teeth are stoutest and proportionally shortest in the smallest specimens in which the submedian teeth are distinctly shorter than the length of the remaining telson (Fig.  5M), are almost as long as the remaining telson by TL 14-20 mm (Fig. 5H,K), and longer than the remaining telson by TL 22 mm (Fig. 5N). The submedian denticles being most numerous in the smallest specimens (10-14 at TL 10-11 mm), occupy most of the mesial margin of the submedian teeth (Fig. 5M). Specimens of TL 13-18 mm have 9-12 submedian denticles distributed along approximately the anterior half or more of the mesial margin of the submedian teeth (Fig. 5H,K); by TL 20 mm, 8 or 9 submedian denticles are present and restricted to the proximal half of the submedian teeth margin, and by TL 22 mm, 7 or 8 denticles are present and restricted to the anterior one-quarter of the mesial margin of the submedian teeth (Fig. 5N). Other sizerelated changes include a proportionally smaller posterior spinule on the telson carinae in larger specimens, and a tendency to lose the posterior spinule on the submedian bosses of abdominal somite 6 with increasing body size. The somite 6 submedian bosses are posteriorly armed in specimens to TL 15 mm, usually unarmed in specimens TL 16-18 mm, and always unarmed above this size. A minute intermediate telson denticle is usually present in specimens to TL 17 mm, but is lost in larger specimens.
The male pleopod 1 endopod is fully modified and penes well developed in the smallest males examined (TL 11 mm). The mandibular palp is clearly 2-articled except in one female (TL 14 mm, Siladen Island) in which the palp articles on one side appear to be undifferentiated. Barber et al. (2012) and Huffard et al. (2012) listed three unidentified species of Gonodactylopsis from Indonesia and New Guinea, referred to as Gonodactylopsis sp. A-C. Gonodactylopsis sp. A, from central-east Indonesia and New Guinea, exhibited a strong genetic break between more westerly populations (Sulawesi to Bali area) and those further east (New Guinea) (Barber et al., 2012). Gonodactylopsis sp. A, at least the Sulawesi-Bali population (material of which is represented in the present study), corresponds to G. komodoensis but further taxonomic scrutiny of New Guinean populations is warranted. The identities of Gonodactylopsis sp. B (western Sumatra, southern Java) and sp. C (Halmahera, western New Guinea) remain to be determined when specimens become available for study; they could be undescribed or belong to named species, such as G. drepanophora and G. lata, which also occur in the region.
The spectral properties of the compound eyes of G. komodoensis have been analysed in detail under the names G. sp. A. by Cronin et al. (2000) and G. spongicola by , , Marshall et al. (2003), Marshall et al. (2007) and Cronin et al. (2014). The name G. spongicola has never been formally proposed nor accompanied by a description, and is therefore a nomen nudum.
Habitat. Exposed vertical rock faces and reefs subject to strong tidal flow and currents; in cavities or burrows in coral, coralline algae, or barrel and petrosid sponges. Description. Eyes elongate; cornea subconical, reaching anteriorly almost to end of antennular article 3. Ocular scales low, subtruncate.
Rostral plate slightly wider than long; median spine twice length or less of basal portion (medially), laterally compressed, with obtusely angular ventral keel; lateral spines divergent with arcuate to almost straight margins.
Thoracic somites 6 lateral process slightly wider than that of somite 7, both with lower margins subtruncate. Thoracic somite 8 anterolateral margin rounded; sternal keel obsolete.
Uropodal protopod terminal spines with outer slightly longer than inner, each with ventral carina, inner with dorsal carina; upper proximal surface with obtuse swelling behind  cristate dorsal carina in specimens > TL 20 mm; slender spine above exopod articulation. Exopod proximal article dorsal surface with sharp, curved carina, distally flanked on either side two short subequal carinae, proximally flanked mesially with 0-2 small spines; outer margin with 9-12 movable spines, distalmost reaching distal one-third of distal article; inner margin smooth, glabrous; distal margin with small ventral spine. Exopod distal article with outer distal margin setose, inner margin glabrous; dorsal and ventral surfaces with inner half dorsal membranous, soft, wrinkled, clearly demarcated from chitinized outer portion; outer chitinized portion with prominent, curved carina dorsally, ventrally with longitudinal carina. Endopod length 2.5-3.2× width, crescentic, articulation with protopod posterior to anterior end; outer margin strongly convex, setose along distal half, inner margin strongly, irregularly concave, glabrous; dorsally with sharp, cristate, curved carina adjacent to outer margin; dorsal and ventral surfaces with inner dorsal half membranous, soft, wrinkled, clearly demarcated from chitinized outer half.
Colour in life. (Fig. 8) Overall translucent with light green and white mottling or speckling and scattered black-brown and orange chromatophores; with 2 narrow, irregular, with white transverse bands, one across carapace at position of cervical groove, and a second across posterior quarter of abdominal somite 6, continuing onto uropodal protopod and anterior half of telson. Rostral plate with white spines, median with red apex. Carapace with white transverse band followed by irregular green patch with pair of small iridescent blue spots and scattered black chromatophores. Iridescent blue spot medially on thoracic somite 6 and abdominal somites 1, 3, 4, somite 1 also with rectangular green and black speckled patch; 3 blue iridescent spots in transverse row on abdominal somite 5. Pair of dark spots across anterior margin of telson. Antennae and antennules transparent with few scattered white and dark spots. Raptorial claw dactylus white, propodus translucent with small distal orange-brown spot at dactyl articulation; carpus and merus translucent with scattered white speckling or mottling; meral spot colourless. Pereopods 1-3 transparent with pale articulations. Uropodal protopod white proximally; primary spines transparent proximally, white with few scattered orange-brown spots distally; exopod proximal article transparent with pale orange-brown highlights; exopod distal article diffuse white; endopod white with irregular patch clear at outer midlength, marginal setae clear. Measurements. Male (n = 13) TL 10-20 mm; female (n = 12) TL 9-22 mm. Other measurements of holotype: CL 5.3 mm, antennular peduncle length 2.5 mm, antennal scale length 1.9 mm, abdominal somite 5 width 4.0 mm.
Etymology. Named lata, Latin for wide, alluding to the comparatively broad submedian and intermediate teeth of the telson.
Remarks. Gonodactylopsis lata sp. nov. is unique in the genus for the broad, lamellate, distally rounded intermediate and lateral teeth and dorsally placed submedian denticles of the telson in adults. In other species of the genus, the intermediate and lateral telson teeth are distally pointed or angular rather than rounded and marginally lamellate. The submedian denticles of the telson line the mesial margins of the submedian telson teeth in G. drepanophora, G. maqqaba and G. komodoensis (Figs 3E,G,4F,J,K,5H,K,M,N,9H,L) or are absent, as in G. herdmani (Fig. 2H,K). In G. lata, the submedian denticles are present along the mesial margins of the submedian telson teeth in juveniles but migrate mesiodorsally with increasing body size (Figs 6H,K,7,8).
Gonodactylopsis lata most closely resembles G. drepanophora and G. maqqaba, sharing similar mid-dorsal telson ornamentation composed of multiple prominent conical spines. Apart from the lamellate primary telson teeth and dorsally placed submedian denticles, G. lata is further distinguished from G. drepanophora and G. maqqaba in lacking the longitudinal carina on the ventral surface of the submedian and intermediate telson teeth. Gonodactylopsis lata and G. drepanophora further differ from G. maqqaba in the strongly crescentic rather than linear shape of the uropodal endopod.
The primary variation observed in G. lata is allometric change in increasing prominence of the telson primary teeth and dorsal ornamentation, namely the increasing density and prominence of dorsal spination, the increasingly lamellate primary teeth, migration of the submedian denticles from marginal to dorsal position (Figs 6H,K,7,8), as well as more pronounced curvature of the uropodal endopod. The main structures of the petasma are developed in all of the males examined, although the lateral lobe of the posterior "endite" is rudimentary in the smallest male examined (TL 10 mm, MNHN IU-2014-966), but fully developed by TL 13 mm (MNHN IU-2014-967). The mandibular palp is 2or 3-articled; two articles are evident in smaller specimens, with the third article typically becoming differentiated by TL 15-21 mm. The dorsal tubercle on the uropodal protopod becomes evident in specimens above about TL 20 mm.
Small specimens of G. lata in which the submedian telson denticles are yet to reach dorsal position may superficially resemble G. drepanophora and G. maqqaba. Nevertheless, at any given size, the intermediate and lateral telson teeth of G. lata (Figs 6H,K,7,8) are proportionally wider and blunter than in G. drepanophora (Figs 3E,G,I, 4F,J,K) and G. maqqaba (Fig. 9H,L) Description. Eyes elongate; cornea subconical, reaching anteriorly almost to end of antennular article 3. Ocular scales low, subtruncate.
Rostral plate slightly wider than long; median spine about twice length of basal portion (medially), laterally compressed, with obtusely angular ventral keel; lateral spines divergent with almost straight margins.
Telson wider than long; 12 or 13 submedian denticles arising from inner margin of submedian teeth; 2 intermediate and 1 lateral denticles; submedian and intermediate teeth slender, sharp; submedian teeth slightly divergent; intermediate teeth length at least twice width, extending posteriorly to midlength of submedian teeth; lateral teeth stout, triangular, apex acute, directed posteriorly. Median carina weakly inflated, with 1 large posterior spine and 2 smaller dorsally; accessory median carina with 2 spines; anterior submedian carina composed of 5 or 6 spines in longitudinal cluster; submedian tooth with 6 dorsal spines distributed from base almost to end of tooth; anterior intermediate carina composed of 2 spines; intermediate tooth with 1 dorsal spine; knob absent; submedian and intermediate teeth with distinct ventral carina.
Uropodal protopod terminal spines with outer slightly longer than inner, each with ventral carina, inner with dorsal carina; upper proximal surface with obtuse swelling behind dorsal carina; slender spine above exopod articulation. Exopod proximal article dorsal surface with sharp, curved longitudinal carina and two short carinae mesial to main carina, distal longest; outer margin with 10 or 11 movable spines, distalmost reaching distal one-fifth of distal article; inner margin smooth, glabrous; distal margin with small ventral spine. Exopod distal article with outer distal margin setose, inner margin glabrous; dorsal and ventral surfaces with inner half membranous, soft, wrinkled, clearly demarcated from chitinized outer half; outer chitinized portion with prominent, curved carina dorsally, ventrally smooth. Endopod length 2.4× width, linear, spatulate, articulation with protopod at anterior end; outer margin gently convex, setose along distal half, inner margin almost straight glabrous, with small proximal and distal point; dorsally with sharp, curved carina adjacent to outer margin; dorsal and ventral surfaces with inner half membranous, soft, wrinkled, clearly demarcated from chitinized outer half.

Colouration in preservative.
Largely faded to pale yellowbrown. Carapace with scattered chromatophores and pair of small spots across cervical region. Thoracic somite 6 and abdominal somite 1 with rectangular field of small irregular chromatophores and larger median black spot. Abdominal somites 2-5 with transverse row of 2 or 3 small rounded spots.
Etymology. The name is derived from maqqaba, Aramaic for hammer, alluding to the "smashing" mode of raptorial hunting employed by the species. Used as a noun in apposition.
Remarks. Gonodactylopsis maqqaba sp. nov. closely resembles G. drepanophora in similar telson ornamentation, in which the mid-dorsal carinae are composed of multiple prominent conical spines, the intermediate primary teeth are slender with a pointed apex, and the lateral primary tooth is prominently triangular with a pointed apex (Figs 3E,I, 4F, 9H). The two species differ chiefly in the shape of the uropodal endopod, being spatulate with a more-or-less straight mesial margin in G. maqqaba (versus lunate with a distinctly concave mesial margin in G. drepanophora) and in the position of the articulation with the protopod, being at the proximal end in G. maqqaba (rather than distal to the anterior end in G. drepanophora). In addition, in size-matched specimens the submedian teeth of the telson are dorsally spinose only on the anterior half in G. drepanophora, rather than along almost the full length in G. maqqaba. The telson armature observed in G. drepanophora (as with other spinose gonodactylids; Ahyong, 2001;Ahyong & Erdmann, 2007;Ahyong, 2008) becomes more extensive and prominent with increasing body size. At TL 16 mm, the submedian telson teeth of the holotype of G. maqqaba are spinose for almost their full length. The submedian telson teeth in similarlysized G. drepanophora, however, reach a similar degree of spination to that of the holotype of G. maqqaba only by TL 24 mm; by TL 16 mm, spination is yet to reach the midlength of the submedian teeth. Thus, the degree of spination of the submedian telson teeth can be useful in separating G. maqqaba from G. drepanophora provided body size is taken into account.
Distribution. South China Sea, from Macclesfield Bank; 30 m.