Alatina alata ( Reynaud, 1830 )

Lewis, Cheryl, Bentlage, Bastian, Yanagihara, Angel, Gillan, William, Blerk, Johan Van, Keil, Daniel P., Bely, Alexandra E. & Collins, Allen G., 2013, Redescription of Alatina alata (Reynaud, 1830) (Cnidaria: Cubozoa) from Bonaire, Dutch Caribbean, Zootaxa 3737 (4), pp. 473-487 : 475-482

publication ID

https://doi.org/ 10.11646/zootaxa.3737.4.8

publication LSID

lsid:zoobank.org:pub:B12F5D90-E13A-4ACA-8583-817DB0F6FD18

persistent identifier

https://treatment.plazi.org/id/03D287BF-FFE3-FFB2-FF1F-12D5FB567AA4

treatment provided by

Felipe

scientific name

Alatina alata ( Reynaud, 1830 )
status

 

Species Alatina alata ( Reynaud, 1830) View in CoL

Table 1, Figs 1–6 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6

Carybdea (medusa) alata

Reynaud, 1830 (in Lesson 1830, pl. 33, Fig 1a View FIGURE 1 )

Marsupialis alata

Lesson, 1837, p. 9, n. 26; Lesson, 1843, p. 278

Tamoya alata

Agassiz, 1862, p. 174

Charybdea alata

Haeckel 1880, p. 441; p. 42; 1940a, p. 5

Carybdea alata

Mayer 1910, p. 508–510; Bigelow, 1918, p. 400; 1938, pp. 144–151, Text–Figs 11–16; Kramp, 1961, p. 304; Arneson 1976, pp. 36, Figs 1 View FIGURE 1 , 2 View FIGURE 2 , Table 1,2, pl. I–V; Arneson and Cutress, 1976, pp. 227–236, Table 1, pl. I A–G; Cutress, 1971, p. 19, pl. 1; Larson, 1976, pp. 242; Larson et al. 1991, p. 313, Table 2; Humann & Deloach, 2002; Morandini, 2003, p. 15–17, Fig. 2 View FIGURE 2 ; Gershwin 2005 pp. 501–523; Calder 2009, pp. 12, 13, Fig. 1 View FIGURE 1 ; Bentlage 2010, p. 52; Bentlage et al. 2010, p. 498; Bentlage and Lewis, 2012, p. 2602

Carybdea alata var. grandis

Graham 1998, pp 28–30;

Material examined. Neotype: USNM 1195802 View Materials , 1 View Materials ind, female, BW 40 mm, BH 70 mm (live), BW 30 mm, BH 69 mm (preserved), 24 June 2011, Karel's Pier, Kralendijke, Bonaire, The Netherlands, 12 09' 06.37 N 6816' 40.84 W, depth = surface. GoogleMaps

Other material. Collected and identified by Lewis et al., depth = surface: USNM 1205450 View Materials , 1 View Materials ind, female, BW 26 mm, BH 78 mm ; USNM 1205449 View Materials , 1 View Materials ind, female, BW 38 mm, BH 77 mm ; USNM 1205448 View Materials , 1 View Materials ind, male, BW 24 mm, BH 64 mm ; USNM 1205447 View Materials , 1 View Materials ind, female, BW 31 mm, BH 83 mm ; USNM 1195807 View Materials , 1 View Materials ind, female, BW 27 mm, BH 80 mm ; USNM 1195806 View Materials , 1 View Materials ind, male, BW 30 mm, BH 75 mm ; USNM 1195805 View Materials , 1 View Materials ind, male, BW 49 mm, BH 77 mm ; USNM 1195804 View Materials , 1 View Materials ind, male, BW 29 mm, BH 84 mm ; USNM 1195803 View Materials , 1 View Materials ind, female, BW 32 mm, BH 50 mm ; USNM 1195801 View Materials , 1 View Materials ind, female, BW 30 mm, BH 83 mm, 25 June 2011, Karel's Pier, Kralendijke, Bonaire , The Netherlands , 12 09' 06.37" N 68 16' 40.84" W GoogleMaps . Collected by Ross et al. and identified by Bentlage, B .: USNM 1131246 View Materials , 1 View Materials ind, gonads absent, BW 22 mm, BH 47 mm, 28 Aug 2007, depth = 98–133 m (bottom depth = 468–595 m), Lease Block VK 826, Gulf of Mexico , 29 09' 34.99" N 88 01' 19.99" W GoogleMaps . USNM 1131245 View Materials , 1 View Materials ind, gonads absent, BW 19 mm, BH 25 mm, 25 Aug 2007, depth = surface (bottom depth = 2206–2282 m), Lease Block AT 340, Gulf of Mexico , 27 38' 38.00" N 88 20' 59.99" W GoogleMaps . Identified by Burnett, J.W .: USNM 94780 View Materials , 1 View Materials ind, female, BW 24 mm, BH 55 mm, Feb 1992, Guantanamo Bay, Cuba, Caribbean Sea. Collected by U S Navy, identified by Larson , R. J.: USNM 58692 View Materials , 1 View Materials ind, gonads absent, BW 17 mm, BH 30 mm, 3 June 1970, depth = 55m, Ocean Acre Area, Off Bermuda, North Atlantic , 31 55' 59.99 N 64 25' 00.00 W GoogleMaps . USNM 58691 View Materials , 1 View Materials ind, gonads absent, BW 7 mm, BH 17 mm, 7 Sept 1968, depth = 327–335 m, Ocean Acre Area, Off Bermuda, North Atlantic , 31 52' 59.99 N 64 25' 00.00 W GoogleMaps . USNM 58655 View Materials , 1 View Materials ind, male, BW 20 mm, BH 49 mm, 28 Oct 1967, depth = 550–675 m, Ocean Acre Area, Off Bermuda, North Atlantic , 32 34' 59.99 N 63 58' 00.00 W GoogleMaps . USNM 58316 View Materials , 1 View Materials ind, gonads absent, BW 8 mm, BH 16 mm, 28 Oct 1967, Depth = 55m, Ocean Acre Area, Off Bermuda , North Atlantic , 31 55' 59.99" N 64 25' 00.00 W GoogleMaps . USNM 54367 View Materials , 1 View Materials ind, gonads absent, BW 8 mm, BH 22 mm, Apr 27 1969, depth = 0–300 m, Open Ocean Area, Off Bermuda, North Atlantic , 31 55' 00.00 N 67 57' 00 W GoogleMaps . USNM 54366 View Materials , 1 View Materials ind, gonads absent, BW 16 mm, BH 55 mm, 25 Apr 1968, depth = 350 m, Open Ocean Area, Off Bermuda, North Atlantic , 31 55' 59.99 N 63 46' 00.00 W GoogleMaps . USNM 53694 View Materials , Open Ocean Area, 1 ind, gonads absent, BW 5 mm, BH 9 mm, 6 Apr 1967, depth = 224–298 m, Off Cape Hatteras, North Atlantic , 35 02' 59.99 N 74 40' 59.99 W GoogleMaps . U SNM 53659, 4 View Materials ind(s) gonads absent, BW 26 mm, BH 51 mm, BW 10 mm, BH 22 mm, BW 10 mm, BH 24 mm, BW 10 mm, BH 22.5 mm, 1 ind sex undetermined, BW 19 mm, BH 47 mm, 28 Sept 1965, depth = surface, Caracas , Caribbean Sea, 10 54' 00 N 67 58' 00.00 W GoogleMaps . Identified by Larson : USNM 58211 View Materials , 1 View Materials ind, female, BW 22 mm, BH 75 mm, 3 Apr 1978, Carrie Bow Cay, Lagoon, Dock , Belize, Caribbean Sea . USNM 58210 View Materials , 1 View Materials ind, female, BW 25 mm, BH 52 mm, 25 Mar 1978, Carrie Bow Cay , Lagoon, Dock . USNM 54472 View Materials , 4 View Materials ind(s), female 1, BW 16 mm, BH 58 mm, female 2, BW 18 mm, BH 64 mm, gonads absent, BW 17 mm, BH 32 mm, sex undetermined, BW 26 mm, BH 43 mm, 13 Oct 1974, Mona Island , Caribbean Sea, 18 04' 0.00 N 67 52' 59.99 W GoogleMaps . USNM 54398 View Materials , 5 View Materials ind(s), female, BW 8 mm, BH 17 mm, gonads absent, BW 7 mm, BH 19 mm, sex undetermined 1, BW 6 mm, BH 20 mm, sex undetermined 2, BW 8 mm, BH 16 mm, sex undetermined 3, BW 7 mm, BH 18 mm, 13 Oct 1974, Mona Island , Caribbean Sea, 18 04' 00.00 N 67 52' 59.99 W GoogleMaps . Collected by Lea and identified by Larson , R.J.: USNM 56737 View Materials , 1 View Materials ind, gonads absent, BW 10 mm, BH 20 mm, 11 Sept 1977, depth = 0–90 m, Open Ocean Area, Off Delaware, North Atlantic , 37 20' 41.99 N 69 10' 23.99 W GoogleMaps . USNM 56736 View Materials , 1 View Materials ind, gonads absent, BW 10 mm, BH 17 mm, 17 Sept 1977, depth = 0–50 m, Open Ocean Area, Off Delaware, North Atlantic , 37 18' 24.00 N 66 51' 24.00 W GoogleMaps . USNM 56735 View Materials , 1 View Materials ind, gonads absent, BW 6 mm, BH 8 mm, 10 Sept 1977, depth = 0–150 m, Open Ocean Area, Off Delaware, North Atlantic , 37 49' 59.99 N 67 25' 23.99 W GoogleMaps . Collected by Chase and Nicholson , identified by Larson , R.J.: USNM 54385 View Materials , 2 View Materials ind(s), female 1, BW 30 mm, BH 80 mm, female 2, BW 34 mm, BH 100 mm, 4 Apr 1956, Freemans Bay, English Harbor, Antigua Island , Caribbean Sea. Collected and identified by Bigelow, H.B : USNM 42017 View Materials , 1 View Materials ind, female, BW 29 mm, BH 78 mm, 30 Jan 1914, depth = 0–100 m, Open Ocean Area , E of Cape Romain, North Atlantic, 32 32' 59.99 N 72 13' 59.99 W GoogleMaps . USNM 41921 View Materials , 1 View Materials ind, gonads absent, BW 17 mm, BH 37 mm, collected 21 Mar 1914, depth = surface, N of Little Bahama Bank, Bahamas, North Atlantic , 27 46' 00.00 N 78 46' 00.00 W GoogleMaps . USNM 41920 View Materials , 1 View Materials ind, gonads absent, BW 18 mm, BH 40 mm, 18 Mar 1914, depth = surface, N of Havana, Cuba, Caribbean Sea , 22 31' 59.99 N 81 47' 59.99 W GoogleMaps . USNM 41919 View Materials , 1 View Materials ind, gonads absent, BW 19 mm, BH 36 mm, 3 Mar 1914, depth = surface, W of Eleuthera Island, Bahamas, North Atlantic , 25 26' 59.99 N 77 16' 00.00 W GoogleMaps .

Catalogued multimedia material (no specimen collected). USNM 1195809 View Materials , filmed by Harbison , R & Widde, E et al., identified by Larson , R (in Larson et al. 1991), still frame from a video voucher taken from the JSL manned submersible, 11 Nov 1989, depth = 540 m (range = 457–610 m), 26 04' 00.00 N 77 32' 59.99 W, Off Gordon Cay, 96 NM Off Rock Point GoogleMaps . USNM 1005621 View Materials , photographed by Continental Shelf Associates for BLM / MMS and Texas A & M University taken from ROV SeaROVER, identified by Lewis, C., photo voucher, Aug 1998, depth = 96.5–108.7 m, 29 19' 39 N 87 46' 00.00 W, Mississippi GoogleMaps , MMS Lease Block Destin Dome 661 , MMS –MAPTEM/M3–4, Gulf of Mexico .

Neotype locality. Bonaire, Dutch Caribbean (Atlantic Ocean)

Diagnosis. Alatina with tall narrow bell, flared at base, tapering into truncated pyramid at apex; 4 crescentric gastric phacellae at interradial corners of stomach; 3 simple to palmate branching velarial canals per octant, each with a velarial lappet bearing a row of 3 to 4 nematocyst warts; 4 long wing–like (sensu Reynaud 1830) pedalia, each with a pink tentacle. Cnidome consisting of heterotrichous microbasic p–euryteles and small birhopaloids in tentacles, and large isorhizas in nematocysts warts.

Description ( Figs 1–6 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 , Table 1). Neotype Fig 2a–f View FIGURE 2 .

Mature female specimen (BW 40 mm, BH 70 mm live; BW 30 mm, BH 69 mm preserved), with tall narrow bell flared at base ( Fig 2a View FIGURE 2 ), tapering into truncated pyramid at apex. Each interradial corner bearing a pedalium: 3 of the 4 pedalia long and broad (approximately 15 mm wide) and wing–like, each bearing a pink tentacle (2a, c) about 2 mm in diameter, with bands of nematocysts along the entire length ( Fig 2c, e View FIGURE 2 ). Fourth pedalium bearing a remnant of a tentacle (only several millimeters in height and width), much smaller and thinner than the other three (possibly damaged while being photographed together with other live specimens in an aquarium). Bell transparent and colorless in life ( Fig 2a View FIGURE 2 ), translucent in fixed specimens ( Fig 2e View FIGURE 2 ); exumbrella speckled with nematocyst warts. Stomach shallow ( Fig 2b View FIGURE 2 ), lacking mesenteries. Manubrium short (2–3 mm long), wide and flat, with 4 mouth lips curled at the tips. Each of the four corners of the stomach housing a crescentric gastric phacella bearing about 20 gastric cirri ( Fig 2a, b View FIGURE 2 ). A pair of leaf–like gonads, flanking each interradial septa and extending into the gastro– vascular cavity, filled with developing oocytes ( Fig 2a, b View FIGURE 2 ). Adjacent gonads (i.e., ovaries) overlapping in the gastrovacular cavity, disassociated into large pieces following rupture due to internal fertilization event prior to preservation in formalin. Clumps of eggs found in various parts of the gastrovascular system (e.g., the velarial and pedalial canals, and the gastric pockets). Velarium wide, suspended by 4 perradial muscular brackets (frenulae) bracing the subumbrellar wall. Each octant bearing 3 simple to palmate branching velarial canals (variable with each octant), with the pair flanking the peradial frenulum simple to bifurcating, those flanking the pedalia at the interadius bifurcating as two main branches each with 3 to 4 distal branches (up to 7 branches in total), with the velarial canal in between bearing three distal branches ( Fig 2d View FIGURE 2 ). Velarial canals in the two octants flanking the diminutive pedalium are not organized in the regular orderly fashion seen in the 6 other octants. Additionally, the velarium is torn near the base of the smaller pedalium, making it difficult to count the number of velarial canal branches there (N.B. the damaged pedalium is not readily apparent in the live photographs of the neotype specimen, and its small size may be exaggerated due to shrinkage following fixation in formalin). A new cubozoan character was discovered that appears to have been overlooked by previous workers. The structure which we call velarial lappets is found in sets of three in each octant overlaying the proximal two thirds of each velarial canal, each bearing a row of 3 to 4 nematocyst warts ( Fig 2d View FIGURE 2 ). Four club–shaped rhopalia ( Fig 2a View FIGURE 2 ), each situated just above the point where the frenulum connects to the subumbrella bearing 2 median lensed eyes, 2 lateral slit eyes, 2 lateral pit eyes, and 1 statocyst ( Fig 2f View FIGURE 2 ). Rhopaliar niche opening T–shaped (sensu Gershwin 2005), with a single upper scale and 2 lower scales ( Fig 2f View FIGURE 2 ). Nematocyst warts are scattered over the entire exumbrella and bell apex; occurring in rows varying in number along the pedalial keel and the rhopalial niche scales ( Fig 2f View FIGURE 2 ). Cnidome characterized below.

Other material. Medusae measurements are given as min– mean –max values, and were made on preserved specimens unless otherwise stated.

In addition to the neotype, forty five A. alata specimens were examined (BH = 8– 48 – 100 mm, BW = 5– 20 – 49 mm). Bells tall, narrow truncated pyramids like the neotype ( Fig 1a View FIGURE 1 ; 4g View FIGURE 4 , 6a–c View FIGURE 6 ). Four long wing–like pedalia ( Fig 1b View FIGURE 1 ; 3c View FIGURE 3 ; 6a–c View FIGURE 6 ), each with a single highly contractile tentacle, round in cross–section, ‘fluorescent’ pink in both live and preserved specimens, with numerous bands of nematocysts along the length ( Fig 5a View FIGURE 5 ). Wide velarium ( Fig 1c, d View FIGURE 1 ; 3a–d View FIGURE 3 ), suspended by 4 perradial muscular brackets (frenulae) bracing the subumbrellar wall ( Fig 3a–d View FIGURE 3 ). Twenty four simple to branching velarial canals, 3 per octant, extending from the gastro–vascular space of the bell into the velarium ( Fig 1c, d View FIGURE 1 ; 3a–c & e View FIGURE 3 ). Velarial canals flanking perradial frenulae simple, bifurcating distally, or giving rise to side branches; those flanking interradial pedalia splitting into two distinct sub–branches in first half to two– thirds of main branch; subsequent branching increasing in complexity (on average 4 sub–branches per primary branch; maximum 10), making it difficult to trace velarial canal tips back to the original branch, and often giving the appearance of greater than 3 primary velarial canals per octant ( Fig 1c, d View FIGURE 1 ; 3a, b View FIGURE 3 ). Bifurcating velarial canals in individuals as small as BH = 8 mm; some canals with three branches in individuals of BH = 16 mm. Larger medusae generally displaying more highly branched velarial canals; wide variation exists between each octant in a single individual (1 to 5 branches seen per canal). Velarial lappets are thick gelatinous pouch–like structures comprising the proximal 50–75% of each velarial canal (24 total; 3 per octant), each bearing a row of 3–4 nematocyst warts ( Fig 3a, b, e View FIGURE 3 ), function unknown. Bell transparent and colorless in life ( Fig 4g View FIGURE 4 , 6a–c View FIGURE 6 ), translucent in fixed specimens, speckled with nematocyst warts (nematocyst warts in older material were often lacking, and may have rubbed off in collection gear or after many years in preservation fluids, despite velarial lappet warts being still recognizable in most specimens).

Rhopaliar niche opening T–shaped (sensu Gershwin 2005), with a single upper scale and 2 lower scales enclosing the rhopalial niche ( Fig 1e View FIGURE 1 ). Four club–shaped rhopalia, each with 2 median lensed eyes, 2 lateral slit eyes, 2 lateral pit eyes, and 1 statocyst ( Fig 3f, g View FIGURE 3 ). All eyes are conspicuous in newly collected material from Bonaire, but in older preserved museum specimens the lens eyes are discolored (having a brown tinge), and bilateral paired pigmented pit eyes and slit eyes are faded to absent, leaving only the complex lens eyes visible. This may be an artifact of prolonged exposure to fixatives.

Stomach shallow, lacking mesenteries; manubrium short (2–3 mm long), wide and flat, 4 mouth lips curled at the tips ( Fig 4a, f View FIGURE 4 ). Four crescentric gastric phacellae, 1 in each corner of the shallow stomach, each with 6–24 basal trunks branching several times at the base, giving rise to up to 100 terminal filaments (gastric cirri) in larger specimens ( Fig 1f, g View FIGURE 1 ; 4a, b, d–g View FIGURE 4 ). Smaller individuals (BH<20 mm) possessing long filaments (6–8 basal trunks) extending from each corner into the stomach, giving appearance of a central mass ( Fig 4c View FIGURE 4 ). Eight narrow leaf–like gonads extend in pairs within the gastric pockets, along either side of the interradial septa; filled with developing sperm or eggs in mature males and females ( Fig 4d, g, h View FIGURE 4 ). We examined preserved individuals with gonads (BH = 16– 61 –100, BW = 6– 23 –34, n = 23), as well as preserved medusae without gonads (BH = 8– 32 –88, BW = 5– 15 –32, n = 22). All medusae <BH = 16 mm lacked gonads, as did many larger medusae (n = 18). Gonads presumably are shed in mature medusae following a spawning event (see Arneson 1976). Gonads, translucent in live medusae ( Fig 4g View FIGURE 4 ; 6a, b View FIGURE 6 ), turning cream to pale amber in spawning medusae; and opaque in fixed material. Ripe gonads overlap along the perradial plane, becoming increasingly pleated ( Fig 4g View FIGURE 4 ).

Cnidome Table 1; Fig 5a–h View FIGURE 5 . Nematocyst measurements are provided as min– mean –max; L = length of capsule in µm, W = width of capsule at widest point in µm, SD = standard deviation, n = number of nematocysts measured. Nematocyst identification follows Mariscal (1974), Östman (2000), and Collins et al. (2011).

R

Departamento de Geologia, Universidad de Chile

SNM

Slovak National Museum

MMS

Montshire Museum of Science

Kingdom

Animalia

Phylum

Cnidaria

Class

Cubozoa

Order

Carybdeida

Family

Alatinidae

Genus

Alatina

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