Integrative taxonomy reveals the presence of a new species of Cyanea (Scyphozoa: Discomedusae: Semaeostomeae: Cyaneidae) from the West coast of Africa Samsodien, Yusra Zootaxa 2024 2024-09-13 5507 3 401 426 Samsodien 2024 [151,379,572,599] Scyphozoa Cyaneidae Cyanea Animalia Semaeostomeae 6 407 Cnidaria species altafissura sp. nov.   Material examined.   Holotype: Ghana: Gulfof Guinea ( 29.5cmin diameter,  15 August 2019, in 7% formaldehyde in ambient seawater, area ( 4°N, 2°W), SAMC-A096867).   Paratypes: Ghana: Gulfof  Guinea( 23.3 cmin diameter,  12 August 2019, preserved in 7% formaldehyde in ambient seawater, area ( 5°N, 0°W), SAMC-A096868); Côte d'Ivoire: Gulf of  Guinea( 22.2 cmin diameter,  4 August 2019, preserved in 7% formaldehyde in ambient seawater, area ( 4°N, 5°W), SAMC-A096869).   One specimencollected by the RV  Dr. Fridtjof Nansenoff Côte d'Ivoireon the 6 thof  August 2019by bottom trawl at a sampling depth of  23 m( 5.12°N, 4.75°W);  Four specimenscollected by the RV  Dr. Fridtjof Nansenoff Ghanaon the on the 12 thof  August 2019by bottom trawl at a depth of  28 m( 5.05°N, 0.94°W);  Three specimenscollected by the RV  Dr. Fridtjof Nansenoff Accraoff Ghanaon the 13 thof  August 2019by bottom trawl at a depth of  26 m( 5.39°N, 0.36°W);  Two specimenscollected by the RV  Dr. Fridtjof Nansenoff Axim Ghanaon the 9 thof  August 2019by bottom trawl at a depth of  41 m( 4.80°N, 2.30°W);  Four specimenscollected by the RV  Dr. Fridtjof Nansenoff Akwidaa, Ghana, on the 15 thof  August 2019by bottom trawl at a depth of  26 m( 4.72°N, 2.03°W);  One specimenscollected by the RV  Dr. Fridtjof Nansenoff Assouindé, Côte d'Ivoireon the 8 thof  August 2019by bottom trawl at a depth of  25 m( 5.13°N, 3.44°W);  Four specimenscollected by the RV  Dr. Fridtjof Nansenoff Takoradi, Ghanaon the 10 thof  August 2019by bottom trawl at a depth of  26 m( 4.82°N, 1.78°W);  One specimencollected by the RV  Dr. Fridtjof Nansenoff Akrou, Côte d'Ivoireon the 7 thof  August 2019by bottom trawl at a depth of  25 m( 5.19°N, 4.34°W);  Four specimenscollected by the RV  Dr. Fridtjof Nansenoff Gazéko, Côte d'Ivoireon the 5 thof  August 2019by bottom trawl at a depth of  23 m( 5°N, 5.73°W);  Two specimenscollected by the RV  Dr. Fridtjof Nansenoff New Town, Ghanaon the 8 thof August by a bottom trawl at a depth of  26 m( 5.05°N, 3.08° W);  One specimencollected by the RV  Dr. Fridtjof Nansenoff Akwidaa, Ghanaon the 8 thof  August 2019by bottom trawl at a depth of  25 m( 4.59°N, 2.03° W);  One specimencollected by the RV  Dr. Fridtjof Nansenoff Kanga, Ghanaon the 28 thof  September 2019by pelagic trawl at a depth of  33 m( 4.94°N, 2.68° W);  One specimenwas collected by the RV  Dr. Fridtjof Nansenoff Nyanga, Gabon, on the 26 thof  September 2017by bottom trawl at a depth of  41 m( 3.06°S, 10.04°E);  One specimenwas collected by the RV  Dr. Fridtjof Nansenoff Gabonon the 25 thof  September 2017by trawl at a depth of  46 m( 2.48°S, 9.39°E);  One specimenwas collected by the RV  Dr. Fridtjof Nansenoff Gabon, on the 24 thof  September 2017by bottom trawl at a depth of  118 m( 2.57°S, 9.06°E);  One specimenwas collected by the RV  Dr. Fridtjof Nansenoff Nyanga, Gabon, on the 26 thof  September 2017by bottom trawl at a depth of  41 m( 3.06°S, 10.04°E);  Four specimenswere collected by the RV  Dr. Fridtjof Nansenoff the coast of Angola, on the 16 thof  October 2017by pelagic trawl at a depth of  20 m( 10.11°S, 13.34°E)  Type locality.Akwidaa, Ahanta West Municipal District, Ghana   Distribution.Range extends around the Gulf of Guinea, West Africa, from Côte d’Ivoireto Gabonand southwards to central Angola.   Diagnosis.Medium-sized  Cyanea; exumbrella surface slightly papillose throughout; 16 rectangular marginal lobes; rhopalial clefts deeper than velar clefts; radial and coronal muscle fields prominent, without gastrovascular intrusions, with 5 and 7 muscle folds respectively. With up to 200 strap-like, hollow tentacles arranged in eight Ushaped fields or clusters. Rhopalium small; statocyst on short stalk lacking bulbs/warts located at base of long, tubular rhopalial pit. Mouth arms greatly thickened basally; gastrovascular sinus in 16 lobes separated by interrupted septa of the  nozakii type, penetrating marginal lappets on a broad front as a fine network of branching and anastomosing canals.  Biological data:Found at depths between 20–118 m.   Etymology.  altafissura, Latin, refers to the comparatively deep depth of the rhopalial cleft.    Holotypedescription.Umbrella a flattened hemisphere with a diameter of 295 mm; thickest centrally and thinning to margin. Exumbrella light brown in colour, with nematocyst warts that impart a rough surface texture throughout (papillose). Umbrella margin divided into 16, broad and approximately quadratic lappets, 44.8 mmin width; two lappets per octant; (secondary) clefts between rhopalial lappets deeper than those between (primary) velar lappets. Coronal muscle in 16 fields, rhopalial fields ( 9.8 mm) narrower than velar ( 18.4 mm) fields; each field separated from neighbour by prominent thickened septal ridge; fields with 7–11 coronal muscle folds. Radial muscles in 16 fields extending distally from coronal muscles to bell margin, not into lappets, each field with 3–6 muscle folds. Coronal and radial muscles without gastrovascular pits. With eight rhopalia, each situated at base of long pit, protected by margins of neighbouring lappets; statocyst at end of short, smooth rhopalial stalk, which lacks a basal bulb. Eight, deep adradial U-shaped tentacle bundles situated between radial muscle fields opposite velar lappets; tentacle bundles comprised of three rows of tentacles proximally but a single row distally; tentacle bundles span the entire length of the radial muscle fields; tentacles hollow, between 160–190 innumber, largest proximally and smallest distally. Mouth broadly quadratic with four oral arms, thickened at base; thin, curtain-like tissue extending between oral arms; shorter than the bell diameter. Central stomach without septa or gastric cirri; extending to bell margin as 16 gastric pouches, separated by interrupted septa of the  nozakiitype, penetrating lappets on a broad front as dendritic canals, with obvious anastomoses. Gonads complexly folded, cream in colour, pendulous; attached to the subumbrella.   Description of other specimens and additional data  Medusae:The bell colour was light brown or beige, although one specimenhad purple oral arms and gonads. Bell a flattened saucer; thicker at the centre and thin at the periphery; maximum bell thickness increasing with increasing bell diameter ( Table 3). Exumbrella surface texture slightly rough throughout ( Figure 4). Bell margin divided into 16 broad, rectangular lappets of approximately similar size and shape ( Figure 5); lappet width increases isometrically with bell diameter; each approximately 16% bell diameter in width; the depth of the (primary) velar and (secondary) rhopalial marginal clefts increases with increasing bell diameter ( Table 3), isometric; standardised depth of rhopalial clefts (mean = 0.09, SE = 0.006, n = 22) significantly deeper than velar clefts (mean = 0.06, SE = 0.005; n = 22) ( F= 17.012, p= 0.0005). Only four of the thirty-eight analysed individuals (~10%) had tertiary clefts, dividing the bell margin into 32 marginal lobes.   TABLE 3.Raw and standardised morphometric measures of  Cyanea altafissura  sp. nov.in the Gulf of Guinea. Bold values indicate significant ( p<0.01) correlations between measure and bell diameter. See Figure 2.     Abbreviation  Morphological characteristic description  RawMean Standard deviation n Max Min  StandardisedStandard Mean deviation n Max Min   bd Bell diameter (mm) 206,42 70,42 38 382 89   bt Maximum bell thickness (mm)  16,55 7,92 38 36,53 1,67 0,078 0,030 38 0,155 0,018   cmwr Rhopalial coronal muscle width (mm) 9,49 3,40 7 13,88 4,88   cmwv Velar coronal muscle width (mm) 18,04 3,95 7 23,06 11,13   cmwv:cmwr Velar: Rhopalial coronal muscle width 2,00 0,38 7 2,68 1,66   dcmf Depth of coronal muscle folds (mm)  5,30 3,17 38 13,87 1 0,024 0,012 38 0,058 0,009   dpmc Depth of primary marginal clefts (mm)  12,85 7,28 32 37,8 3,13 0,060 0,021 32 0,101 0,013   dsmc Depth of secondary marginal clefts  (mm)  19,08 9,29 32 43 2,44 0,089 0,024 32 0,137 0,010   loa Length oral arms (mm) 85,83 44,70 28 220,00 35,50 0,375 0,200 28 0,861 0,160   mlw Marginal lappet width (mm) 36,79 6,71 6 44,79 28,93 0,151 0,009 6 0,165 0,138  ncmf Number of coronal muscle folds per  field 7,38 1,04 38 9,69 5,4 0,041 0,017 38 0,079 0,019   nrmf Number of radial muscle folds per field  4,38 0,73 38 6,13 3 0,024 0,008 38 0,043 0,013   ntr Number of tentacle rows per cluster 2,14 0,42 37 3 1   woa Width oral arms (mm) 28,28 7,91 28 50,25 14,50 0,125 0,048 28 0,246 0,083   FIGURE 5.  Cyanea altafissura  sp. nov.View of subumbrella showing rectangular shape of marginal lappets, the comparative depths of the primary/velar ( pmc) and secondary/rhopalial ( smc) clefts, and the dendritic and anastomosing nature of marginal canals. Rhopalium relatively small; statocyst on a short, smooth stalk lacking basal bulbs, papillae or other protuberances ( Figure 6); located at the base of long tube-like pit, bounded on the exumbrella surface by a hood formed from the fusion of neighbouring lappet edges, and on the subumbrella surface by overlapping lappet margins ( Figure 6); ocelli not observed.   FIGURE 6.  Cyanea altafissura  sp. nov.A) Subumbrella view of the bell margin, showing a secondary marginal cleft ( smc), the position of radial muscle fields ( rmf), the rhopalium ( r) and the overlapping edges of the marginal lappets that in effect form a rhopalial tube (see schematic inset). B) subumbrella view of bell margin showing a secondary marginal cleft ( smc) and the rhopalial hood ( rh). C) detail of rhopalium at base of rhopalial “tube”, note the absence of bulbs/papillae on stalk. Sixteen coronal muscle fields, each field separated from neighbour by prominent thickened septal ridge ( Figure 7); rhopalial fields (mean = 4% bell diameter, SE = 0.2%, n = 68) significantly narrower than velar (mean = 7.2% bell diameter, SE = 0.2%, n = 67) ( F= 182.86, p<0.0005). Number of muscle folds per field varying from between 4–14 (mode = 7) in a size independent way, as too does the depth of individual muscle folds (mean = 6.5 mm, SE = 0.1 mm). Radial muscles in 16 fields, extending from a position between rhopaliar and velar coronal muscles towards base of lappets at bell margin ( Figure 7); with between 3–8 muscle folds (mode = 5) proximally, fanning out to a greater number distally. Gastrovascular pits in the coronal and radial muscle folds absent.   FIGURE 7.  Cyanea altafissura  sp. nov.Subumbrella view of U-shaped tentacular bundle ( tb), hollow tentacle ( t), velar ( vcmf) and rhopalial ( rcmf) coronal muscle fields, radial muscle fields ( rmf), primary marginal cleft ( pmc) and rhopalium ( r). Tentacles arranged in eight, deeply rectangular, U-shaped bundles that extend from distal edge of coronal muscles towards base of lappets ( Figure 7); tentacle bases very close to each other, and distinct rows hard to observe clearly, especially proximally. As a rule, tentacles in a single row towards bell margin and clustered in up to four rows adjacent to coronal muscle fields. Tentacles hollow, strap-like ( Figure 7) and with a clear endodermal core. Proximal tentacles longer and thicker than distal ones. Up to 200 tentacles per field, size dependant.   FIGURE 8.  Cyanea altafissura  sp. nov.Subumbrella view showing one of the four gonads ( g) emerging from between adjacent oral pillars ( op) at base of oral arms ( oa). Velar ( vcmf) and rhopalial ( rcmf) coronal muscle fields, radial muscle fields ( rmf) and tentacle bundles ( tb) also shown. The mesoglea at the base of the manubrium a strongly thickened ring ( Table 3) supporting four oral arms on short pillars ( Figure 8); oral arms greatly expanded laterally, membranous and curtain-like. Unfortunately, it was not possible to accurately measure oral arm length, but generally shorter than diameter of bell. Mouth approximately quadratic in shape; stomach without septa; small specimens with numerous short gastric cirri arranged in four inter-radial groups attached to subumbrella ( Figure 9). Stomach extending to bell margin as 16 gastric pouches that penetrate lappets on a broad front as a network of fine dendritic canals, with obvious anastomoses ( Figure 5); pouches separated by indistinct septa distal of coronal muscle fields that are interrupted and of the nozakii- type( Figure 10) with anastomosing connections between the tentacular and rhopalial gastrovascular pouches. Four, complex, folded gonads originate from stomach wall, and emerge from between the thickened bases of adjacent oral pillars ( Figure 8) to hang freely below subumbrella, decumbent. 2019-08-15 Ghana 4.0 Gulf 43160 -2.0 6 407 SAMC-A096867 1 holotype [800,1103,716,743] Ghana Gulf 6 407 1 paratype 2019-08-12 Guinea-Bissau 5.0 Guinea 43062 -0.0 6 407 SAMC-A096868 1 paratype 2019-08-04 Guinea-Bissau 4.0 Guinea 43160 -5.0 6 407 SAMC-A096869 1 paratype 2019-08 RV Dr. Fridtjof Nansen Ivory Coast 23 5.12 Cote d'Ivoire 783 -4.75 6 407 1 2019-08 RV Dr. Fridtjof Nansen Ghana 28 5.05 Ghana 783 -0.94 6 407 4 2019-08 RV Dr. Fridtjof Nansen & Accra Ghana 26 5.39 Ghana 783 -0.36 6 407 3 2019-08 RV Dr. Fridtjof Nansen & Axim Ghana 41 4.8 Ghana 783 -2.3 6 407 2 2019-08 RV Dr. Fridtjof Nansen & Akwidaa Ghana 26 4.72 Ghana 783 -2.03 6 407 4 2019-08 RV Dr. Fridtjof Nansen & Assouinde Ivory Coast 25 5.13 Cote d'Ivoire 783 -3.44 6 407 1 2019-08 RV Dr. Fridtjof Nansen & Takoradi Ghana 26 4.82 Ghana 783 -1.78 6 407 4 2019-08 RV Dr. Fridtjof Nansen & Akrou Ivory Coast 25 5.19 Cote d'Ivoire 783 -4.34 6 407 1 2019-08 RV Dr. Fridtjof Nansen & Gazeko Ivory Coast 23 5.0 Cote d'Ivoire 0 -5.73 6 407 4 RV Dr. Fridtjof Nansen Ghana 26 5.05 New Town 783 -3.08 6 407 2 2019-08 RV Dr. Fridtjof Nansen & Akwidaa Ghana 25 4.59 Ghana 783 -2.03 6 407 1 2019-09 RV Dr. Fridtjof Nansen & Kanga Ghana 33 4.94 Ghana 783 -2.68 6 407 1 2017-09 RV Dr. Fridtjof Nansen Gabon 41 -3.06 Nyanga 784 10.04 6 407 1 Nyanga 2017-09 RV Dr. Fridtjof Nansen Gabon 46 -2.48 Gabon 784 9.39 6 407 1 2017-09 RV Dr. Fridtjof Nansen Gabon 118 -2.57 Gabon 784 9.06 6 407 1 2017-09 RV Dr. Fridtjof Nansen Gabon 41 -3.06 Nyanga 784 10.04 6 407 1 Nyanga 2017-10 RV Dr. Fridtjof Nansen Angola 20 -10.11 Angola 779 13.34 6 407 4