Odontozona lopheliae Goy & Cardoso 2014

(Figs. 4 – 8)

Description. (holotype male, MNRJ 22552). Rostrum (Fig. 4 A) long, almost reaching distal end of antennular peduncle, straight, upturned at tip; dorsal margin bearing 10 teeth, posterior 3 situated posterior to hind margin of orbit, ventral margin with two teeth on distal fourth; no lateral teeth. Carapace (Fig. 4 A) robust, with small hepatic and strong antennal spines, without inferior orbital and branchiostegal spines; 7 pterygostomial spines. Cervical groove deep, with 34 spines along posterior margin; smooth behind cervical groove; with 4 spines at supraorbital region. Ventrolateral carapace angle and branchiostegite slightly rounded. Abdomen (Fig. 4 A) smooth, dorsally unarmed, without median carina and without transverse grooves; margins of pleura 1 – 4 and 6 rounded, spineless; that of pleomere 5 triangular, spineless. Telson (Fig. 4 C) lance-shaped; deeply sulcate mesially at middle length; with 4 rows of dorsolateral spines, inner pair of rows with 3 spines, outer pair with 7 spines; each lateral margin bearing distinct tooth at midlength, provided with numerous long plumose setae along posterior three-fifths; distal margin blunt, fringed with setae, with a pair of lateral spines. Eyes (Fig. 4 B) well-developed with unpigmented cornea twice length of peduncle. Dorsal surface of peduncle with 3 spines, anterior margin with spines. Antennular peduncle (Fig. 4 E) short, reaching third of scaphocerite; proximal segment longest with short, subquadrate stylocerite proximally, bearing 3 slender spines on distal margin, one strong distal tooth on outer margin, one strong mesial spine on inner margin; middle segment 0.6 length of proximal segment with bifid strong distal spine at outer margin, 2 strong spines on inner margin; distal segment half of middle segment length, unarmed. Upper and lower flagella short reaching to middle of second pleomere. Antenna (Figs. 4 A, D) with large basicerite, with 4 ventral teeth, one dorsomesial tooth; scaphocerite twice as long as broad, lateral margin slightly convex, with 5 teeth including terminal tooth. Dorsal surface of scaphocerite with distinct longitudinal carina, no spinules, ventral surface unarmed. Flagellum well developed extending beyond tip of telson. Mandible (Fig. 5 A) robust, with short, fused molar and incisor processes. Molar surface nearly smooth, incisor with 2 large distal teeth, 4 smaller proximal teeth. Palp well developed, 3 - segmented, segments of subequal lengths; proximal segment glabrous, middle and distal segments setose. Maxillule (Fig. 5 B) with slender, undivided endopod bearing 7 simple setae distally. Proximal endite moderately broad, truncate distally, with numerous distal compound spinose and simple setae. Distal endite of subequal size, rounded distally, with numerous plumose setae. Maxilla (Fig. 5 C) with numerous plumose setae on both lobes of coxal and basal endites. Endopod long, slender, exceeding anterior margin of scaphognathite with distal and outer marginal simple setae. Scaphognathite long, narrow, fringed with numerous plumose setae, rounded terminal margin with 5 long plumose setae. First maxilliped (Fig. 5 D) with 3 - segmented endopod; proximal segment longer than wide, with 11 simple setae laterally along outer margin, middle segment about 0.25 length of proximal with 1 simple seta on outer margin; distal segment short, slender. Basipodite large, rounded anterior, nearly straight middle, rounded posterior, bearing dense fringe of plumose setae; coxopodite with numerous plumose setae. Exopod well developed, twosegmented, basis with plumose setae on outer margin; flagellum twice as long as endopod with 32 long plumose distolateral setae. Large epipod with slender proximal and distal lobes of equal length. Second maxilliped (Fig. 5 E) with 4 - segmented endopod; dactylus suboval with dense fringe of long setae along distodorsal margin; propodus equal dactylar length, densely setose on dorsal margin; carpus short, 0.5 propodal length, with 3 long distal plumose setae; merus 3 times dactylar length, ventral margin with fringe of numerous long simple setae, dorsal margin with distal simple seta; ischium fused to basis, both lobate with dense fringe of short setae; coxa with 8 short plumose setae. Exopod long, slender, undivided with distal third bearing numerous plumose setae; tiny epipod present; arthrobranch and podobranch also present. Third maxilliped (Fig. 6 A) with 7 - segmented endopod, dactyus, propodus and carpus subequal in length, with numerous long plumose setae; propodus with setiferous organ at distomesial angle; merus and ischium equal length, merus with 6 outer marginal spines, inner margin fringed with simple setae and with 8 small distal spines, ischium with 5 large outermarginal spines, inner margin fringed with simple setae; basis and coxa fused, unarmed; 2 - segmented exopod long, with distal half bearing numerous plumose setae. First pereiopod missing. Second pereiopod (Fig. 6 B) slender and elongate, glabrous; carpus longest segment, twice ischial length; cutting edge with indistinct ridge; tips of fingers with small tufts of long setae (Fig. 6 C). Third pereiopod (Fig. 6 D, E) largest and strongest, longer than entire length of body; propodus, carpus, merus subequal in length. Palm with dorsal row of 19 – 22 spinules, ventral margin with short simple setae. Fingers elongate with hooked, crossing tips; dactylus glabrous, cutting edge with straight, indistinct ridge. Carpus with 11 or 12 dorsal marginal spines, 1 or 2 ventral marginal spinules. Merus with 5 or 6 dorsal and ventral marginal spines. Ischium unarmed Fourth pereiopod (Fig, 6 F) long and slender. Dactylus biunguiculate with long, slender unguis separated from dactylar corpus. Propodus subdivided into 4 segments, bearing 19 movable spines. Carpus subdivided in 5 segments, no ventral movable spines; merus and ischium not subdivided. Fifth pereiopod is missing. Pleopods 1 – 5 missing at right side, 1, 2, 5 missing at left side. Pleopods 3, 4 at left side leaf like, biramous, lacking appendices (Fig. 7 A), ventral margin of basipodite with spinule; endopods, exopods with plumose seta on inner surface and posterior margin. Uropods (Fig. 4 C) shorter than telson; protopodite stout ending with 4 or 5 distal teeth. Exopod wider than endopod, with 8 – 10 teeth on outer margins; dorsal surface with one longitudinal carina. Endopod without outer marginal teeth, longer than exopod. Branchial formula same as O. edwardsi (Tab. 1.). Variations. The species shows some variation in spination of appendages. Female specimens have the abdominal pleura more rounded than males; there is a ventral medial spinule on each male pleomere; male thoracic sternites 4 – 6 bear a few blunt spinules, female sternites are broader, glabrous; first pleopods are uniramous and glabrous in both sexes, basipodite of male pleopods 2 – 5 bear a ventral spinule. Rostrum bears 6 – 12 dorsal, 1 – 4 ventral spines; cervical groove cincture with 26 – 36 spines. Epistome (Fig. 7 B) triangular anteriorly with 2 submedian spines followed by 4 spines. Labrum normally developed. Paragnath rounded with median fissure. Third maxilliped (Fig 7 C) with ischium bearing 1 – 8 outer marginal spines, 0 – 6 inner marginal spinules; merus with 1 – 6 strong outer marginal spines, 0 – 8 inner marginal spinules. First pereiopod (Fig. 7 D, E) slender, elongate, well-developed carpo-propodal setiferous organ, cutting edge with indistinct ridge, fingers of chela bearing long setae at tips. Fourth and fifth pereiopods equal in length, carpus subdivided into 4 or 5 segments, propodus subdivided into 3 – 6 segments. Measurements. (mm) holotype: carapace length, 5.5; carapace and rostrum length, 6.9; total length, from telson tip to scaphocerite tip, approximately 19.8; third pereiopod length, approximately 20.4. Additional examined specimens: carapace length, 3.6 – 5.9; carapace and rostrum length, 4.3 – 6.8; total length, 12.3 – 20.7; third pereiopod length, 10.2 – 20.5. Color pattern. (Fig. 8) An ovigerous female was photographed off Sapelo Island, Georgia (SERTC 1008) taken from a depth of 665 m. Carapace, abdominal pleomeres, telson and uropods have numerous red spots along edges; spines on uropodal exopodite margin and dorsal longitudinal carinae of telson white. Dorsal region of carapace in front of cervical groove bright red, coloration extends along rostrum, over ocular peduncle, antennular peduncle and antennal scaphocerite; cornea of eyes white. Edges of third maxillipeds, first and second pereiopods red. Third pereiopods merus, carpus red; ischium, proximal propodus edged in red; distal propodus, dactylus white. Fourth, fifth pereiopods mostly whitish to transparent with few red spots along edges. Ovary and eggs blue. Development. Ovigerous females ranged in size from 5.2 – 5.4 mm postorbital carapace length, 18.1 – 18.4 mm total length and carried 10 – 11 eggs. Eggs at blastula stage with undifferentiated yolk cells were 1.20 × 1.62 mm in size.

Diagnosis. Small, robust shrimp with laterally compressed body, with few spinous processes; carapace with cincture of spines marking cervical groove; smooth behind cervical groove; small hepatic spine; antennal spine strong; few spines at supraorbital region; antennal spine strong; few spines between antennal spine and pterygostomian border. Abdomen glabrous. Eyes without pigment. Second maxilliped without ventral tooth on propodus. Third maxilliped with spinous ischial and meral borders. Merus, carpus and propodus of third pereiopod with marginal spinules. Carpus and propodus of fourth and fifth pereiopod with 5 and 6 segments respectively.

Remarks. Both Odontozona edwardsi and O. lopheliae sp. nov. closely follow the definition of the genus Odontozona given by Holthuis (1946) and are most closely similar to each other as well as O. spongicola (Alcock & Anderson 1899) from deep water in the Pacific. The genus Odontozona includes 14 species (De Grave & Fransen 2011), O. lopheliae sp. nov. differs from 12 species of the genus in having the posterior half of carapace behind cervical groove cincture smooth. Odontozona edwardsi and Odontozona spongicola are close related with O. lopheliae sp. nov. sharing the feature mentioned above. Odontozona edwardsi is easily separated from O. lopheliae and O. spongicola by the transverse grooves on the first three abdominal pleomeres, dorso-tranverse carina on the sixth pleomere, and the posterior pleural teeth versus the smooth pleon of these two species. Odontozona spongicola lacks the spinous propodal margins of the third perieopods (Goy 1992) seen in O. edwardsi and O. lopheliae. The other deep water Pacific species, O. foresti Hendrickx, 2002, lacks the spination of the third pereiopods and telson seen on O. edwardsi and O. lopheliae. As mentioned above, the remaining Atlantic species of Odontozona are much more spinous on the carapace, abdomen, and appendages of both O. edwardsi and O. lopheliae. Atlantic Odontozona are distributed as follows: Odontozona striata Goy, 1981 and O. meloi Anker & Tavares, 2013 are known only from their respective holotypes, the first sampled in deep waters off Cape San Antonio, Cuba and the latter sampled on Brazilian continental shelf (off Espírito Santo state) at a depth of 81.6 m. Odontozona minoica Dounas & Koukouras, 1989 is a deep-water species known from 125 m and 300 m in Aegean Sea off Crete (Koukouras & Dounas 2000). Besides that, two of the Atlantic species are restricted to shallow waters: Odontozona libertae Gore, 1981 presently known from 56 m off Key Largo, Florida and from 23 m in Granate Bay, Columbia (Criales 1997) and Odontozona anaphorae Manning & Chace, 1990, known only from the holotype from 10 m off Ascension Island in the South Atlantic Ocean. Odonotozona addaia Pretus, 1990 is presently known from a marine cave in the western Mediterreanean on the coast of Menorca at a depth of 5 m. The Atlantic species of Odontozona can be separated base on the key below.

Etymology. The species is named for its association with the deep-sea coral Lophelia pertusa.

Material examined. (1) male holotype, cl 5.5, Ecoprof 5, TOISA Voyager, 22 ° 24 ' 12 " S 46 ° 06 ' 18 " W, 621 m, among deep sea corals Lophelia pertusa, MNRJ 22552. (2) 2 female paratypes, cl 3.8, 3.9, off Sapelo Island, Georgia, U. S. A., 31 ° 23.160 ' N 78 ° 36.198 ' W, 665 m, 24 Aug 2004, SERTC – S 1008. (3) 2 female paratypes, cls 4.2, 5.0, R / V Seward Johnson II, Green Canyon 234 a, Gulf of Mexico, JSL 4714, 27 ° 35.89 ' N 91 ° 49.60 ' W, 524 m, on Lophelia pertusa, June 2004, USNM 1175271. (4) 1 male paratype, 4 female paratypes (2 ov. females), cls 3.6, 5.9, 5.7, 5.4, 5.2, R / V Seward Johnson II, Green Canyon 234 a, Gulf of Mexico, JSL 4740, 27 ° 44.81 ' N 91 ° 13.44 ' W, 509 m, on Lophelia pertusa, June 2004, USNM 1222546.

Type locality. 22 ° 24 ' 12 " S 46 ° 06 ' 18 " W at Campos Basin, Rio de Janeiro, Brazil. Distribution: 31 ° 386 ' N 78 ° 6033 ' W off Sapelo Island, Georgia, USA; 27 ° 35.89 ' N 91 ° 49.60 ' W and 27 ° 44.81 ' N 91 ° 13.44 ' W Green Canyon, Gulf of Mexico. Habitat. The holotype was sampled at a deep sea coral reef at a depth of 621 m by means of a ROV (remotely operated vehicle). The species’ deepest record is from off Sapelo Island at 665 m. Six cruises from 2002 to 2005 recorded 84 specimens of the species by means of the deep-sea submersible Johnson-Sea-Link at primarily two locations, Green Canyon and Viosca Knoll in the Gulf of Mexico at depths from 459 to 525 m with an average depth of 489 m. The use of deep-sea human operated orb remotely operated vehicles allowed the observation of the species’ association with live or dead coral branches of Lophelia pertusa, a foundation species that is found throughout the world’s oceans on topographic features (Rogers 1999).