Yemanja, Brandão, 2010

Brandão, Simone N., 2010, Macrocyprididae (Ostracoda) from the Southern Ocean: taxonomic revision, macroecological patterns, and biogeographical implications, Zoological Journal of the Linnean Society 159 (3), pp. 567-672 : 583-590

YEMANJA GEN. NOV. ( FIGS 5 , 6 )

1990 in part Macrocyprina Maddocks, 1990: 117–118 , graph 63, figs 16.11, 16.12, 17.11, 17.12, 21.15, 23.7, 28.14-16, 31.18, 35.10, 45.7, 47.20, 54.11-14, 55.11- 13, 69.8, 80.30; pls. 48.1-6, 49.1-6, 81.5, 97.15-18, 109.16,17.

2004 in part Macrocyprina Brandão, 2004: 166–167 , figs 8, 9.

2005 Macrocyprina Brandão, 2005: 219–243 , figs 1– 15.

Etymology: After Yemanjá , the goddess of the oceans and seas in the Afro-Latin American religion Candomblé.

Type species (original binomen): Macrocyprina coimbrai Brandão, 2005 .

Additional species (original binomen): Macrocyprina hawkae Maddocks, 1990 ; Macrocyprina rocas Brandão, 2005 ; Macrocyprina youngi Brandão, 2005 .

Distribution: Recent. Tropical Western Atlantic, live 1–100 m, subfossil 1– 183 m.

Measurements: Holotype of Yemanja coimbrai ( Brandão, 2005) , RV L 1.56 mm, H 0.68 mm; LV L 1.54 mm, H 0.64 mm.

Diagnosis: Carapace small, adults with one to more than ten circular to irregularly shaped, opaque, white patches, which occupy different positions on valve surface. Lateral outline elongate oblong, with slightly arched dorsal margin, and rounded posteroventral margin. Male appendage V extremely asymmetrical. Right appendage robust, ~1.5 times larger than left appendage; podomere II (= palp podomere I) with one medium-sized dorsodistal seta; robust, hook-shaped podomere III curved at 90 to 110°. Left male appendage V slightly sclerotized, highly modified; podomere II with convex ventral margin, and with short to medium-sized, distal seta at lateral surface or postero-dorsal angle; podomere III partially or completely fused to podomere II, surface covered with numerous short setulae. Female appendage V with fairly short, thick, distal claws, the mediodistal one longest. Furca large, thick, and symmetrical. Hemipenis composed of two lamellae, the proximal one subrectangular in outline, posterodistal lamella variable in shape. Zenker’s organ with long and thin chitinous tube; terminal bulb small; vas deferens arranged in few loose loops that are about as long as the chitinous tube.

Description: Carapace small, adults with one to more than ten circular to irregularly shaped, opaque, white patches, which occupy different positions at valve surface. RV always larger than and overlapping LV ventrally, antero-, and posterodorsally. Lateral outline elongate oblong; fairly inequilateral; dorsal margin slightly arched, posterodorsal margin convex, concave, or straight; posteroventral margin always rounded; ventral margin straight to fairly indented in mouth region; anterior margin narrowly rounded. Zone of concrescence medium-sized with mostly straight, but also few ramified radial pore canals; vestibules wide.

Antenna I with seven slender, elongate podomeres, and long, thin, flexible setae; podomeres II and III fused with incomplete suture; podomere VII with four setae. Antenna II with six robust podomeres; podomere II long; podomere IV two times as long as podomere III. Mandible with a broad masticatory jaw armed with one dorsal, conical tooth followed by six tricuspidate teeth and several setae; exopodite with one reduced plus six or seven distal setae. Vibratory plate of maxilla I with two long strahlen (anteriorly directed setae) plus 13–19 feathered setae; palp podomeres I and II flexibly articulated; ventral endite with one or two basal setae; other two endites without basal seta. Exopodite of appendage V with 10–12 setae. Female appendage V symmetrical, slender; podomere II (= palp podomere I) two times as long as podomere IV, podomeres IV and V with fairly short, thick, distal claws, the mediodistal claw of podomere V longest. Male appendage V very asymmetrical. Right appendage robust, larger than left appendage; podomere II (= palp podomere I) subrectangular, ventral margin straight to slightly sinuous, with one medium-sized seta present on dorsodistal margin, plus one reduced seta and two or three modified setae (= peg) on ventrodistal margin; robust, hookshaped podomere III curved at 90 to 110°, one modified seta (= peg) present distally. Left male appendage V slightly sclerotized, highly modified; podomere II with very sinuous ventral margin, distal seta short to medium-sized at lateral surface or at dorsal margin, ventrodistal angle with two modified setae (= pegs) and one reduced, thin seta; podomere III partially or completely fused to podomere II, surface covered with numerous short setulae, one short, modified seta (= peg) present on dorsodistal margin. Ventral margin of podomeres III and IV of appendage VI finely setulate; podomere VI with one short seta and two long claws, dorsodistal claw slightly shorter than mediodistal one. Furca large, thick, and symmetrical, terminal setae robust, separated from furcal rods by a conspicuous suture; distal two thirds of posterior margin of each ramus with thin, short barbs. Hemipenis composed of two lamellae, the proximal one suboval to subrectangular in outline, posterodistal lamella very variable in shape amongst species (subtriangular, subrectangular, or bilobated). Zenker’s organ with long and thin chitinous, central tube and small terminal bulb; vas deferens arranged in few loose loops that are about as long as the central tube.

Adult chaetotaxy: Antenna I 1, 2(0/.2), 3(.1/.1.), 4(.1/.1.), 5(.1/.1), 6(.2/.2-3), 7(0/0:3-4). Antenna II 1(0/0:1-2), 2(0/0:1), Exopodite (0/0:2,1r), 3(0/6-7.1c,3), 4[female (.1./.2.1c,2)] [male (.1./.2.1c,2mod,1)], 5(0/.1c, 1:3c,2), 6(0/0:2c,1-2). Mandible 1(0/ 6t,+5.1.), 2(0/.2:1), Exopodite (0/0:1r,5-7), 3(0/.4-5:1-2), 4(.3./.4:2), 5(0/ 0:2c,0-3). Maxilla I vibratory plate (2re,13-19), palp 1(.1), 2(.3-4/0), 3(0/0:4-5). ApV 1(0/1:1,0-2), Exopodite (0/0.1.1.0-2.6-7), [female 2, 3, 4(0/.1) 5(.1./0:2)] [male 2(.0-1/2-3mod,0-1r:0-1), right 3(0/.0:1mod), left 3(.1mod/.0)]. Appendage VI 1(0/.1-2), +2(.1/.1.1.1), 3(0/.1), 4(0/.1), 5(0/.1), 6(0/0:2c,1). Appendage VII 1(.1/ 0), +2(.1/.1.1.1), 3(0/.1), 4(0/.1), 5(0/.2), 6(0/0:1.1r,1). Furca 1(0/0:1.0-2r).

Remarks: The genus Yemanja gen. nov. is closely related to the genus Macrocyprina ; both genera share small, oblong valves with white, opaque spots; antenna I with partially fused podomeres II and III (= palp podomeres I and II); podomeres I and II of maxilla I palp united by a complete suture; and the symmetrical furcae. The modified male left appendage V is diagnostic of Yemanja . As all previously described genera of family Macrocyprididae , and also most other Cypridocopina , have the left male appendage V similar or equal to the right appendage V, it is more parsimonious to hypothesize that the modified state is an apomorphy of the new genus Yemanja . Moreover, the largest diversity and wider geographical distribution of Macrocyprina indicates that this genus is geologically older and may be ancestral (and then polyphyletic) to Yemanja .

The reduced seta (= dorsoproximal peg from Brandão, 2005) on the medial surface of the exopodite of the mandible had been suggested to be an important taxonomic character for the three Brazilian species of Yemanja ( Brandão, 2005) because it had never been described nor figured in any other species of Macrocyprididae . However, all the macrocypridid species studied herein do present this character.

Additionally, because of its geographical distribution (Caribbean, Rocas Atoll, north-eastern and south-eastern Brazil), I suggest that the genus Yemanja evolved off the north-eastern coast of Brazil (at or near the Rocas Atoll) and migrated northwards and southwards according to the superficial Equatorial Current and Brazilian Current.

GENUS MACROMCKENZIEA MADDOCKS, 1990

( FIGS 7–13 )

Type species (original binomen): Macrocypris siliquosa (Brady, 1887) (original designation).

Additional species (listed by original binomen): Macrocypris siliquosa Brady, 1887 ; Macrocypris australiana Neale, 1975 ; Macromckenziea glacierae Maddocks, 1990 ; Macromckenziea gregalis Maddocks, 1990 ; Macrocypris ligustica Bonaduce, Masoli & Pugliese, 1977 ; Macrocypris porcelanica Whatley & Downing, 1983 ; Macromckenziea swansoni Maddocks, 1990 .

MACROMCKENZIEA GLACIERAE MADDOCKS, 1990

( FIGS 7–10 )

1976 Macrocypris sp. nov. Maddocks, 1976: 42.

1987 ‘aff. Macrocypris spec. (= glacierae Maddocks , unpublished)’ of Hartmann, 1987: 132–133, figs 78– 90.

1988 ‘aff. Macrocypris spec. (= glacierae Maddocks , unpublished)’ of Hartmann, 1988: 149

1989a ‘aff. Macrocypris spec. (= glacierae Maddocks , unpublished)’ of Hartmann, 1989a: 219, fig.

1989b ‘aff. Macrocypris spec. (= glacierae Maddocks , unpublished)’ of Hartmann, 1989b: 253-

1990 in part Macromckenziea glacierae Maddocks, 1990: 51–52 .

1990 aff. Macromckenziea glacierae, Hartmann, 1990: 212 .

1992 Macromckenziea glacierae, Hartmann, 1992: 418 , tabs.

1997 Macromckenziea glacierae, Hartmann, 1997: 242–243 , fig. 101.

Material: 85 specimens plus 10 valves.

10 A F ( SNB 0014, 0016), 3 A M ( SNB 0020), 16 (A-1), 5 (A-2), 4 (A-3), 4 damaged specimens, 1 RV, 2 LV, EASIZ II, # 107, ZMH K- 40811; 1 (A-2), 1 damaged specimen, EASIZ II, # 111, ZMH K- 40812; 10 A F, 4 A M ( SNB 0060), 11 (A-1), 8 (A-2), 3 damaged J, 2 RLV, 1 LV, EASIZ II, # 310, ZMH K- 40813; 1 A F, EASIZ II, # 316, ZMH K- 40814; 2 A F, 1 A M, 1 (A-1), 1 RLV, EASIZ II, # 320, ZMH K- 40815; 2 A F, 4 A M ( SNB 0120-DNA 23, SNB 0130- DNA 31, SNB 0443- DNA 150, SNB 0444-DNA 151), 4 A, 9 (A-1) F, +11 (A-1) M, 1 (A-1), 8 (A-2), 5 (A-3), ANDEEP III, # 74-6-E, ZMH K- 41365; 9 A F ( SNB 0375-DNA 108, SNB 0376-DNA 109, SNB 0377-DNA 110), 3 A M ( SNB 0410-DNA 117, SNB 0411-DNA 118), 5 (A-1) ( SNB 0412-DNA 119), 3 (A-2), 3 (A-3), ANDEEP III, # 74-6-S, ZMH K- 41366; 4 A F, 3 (A-1) F, ANDEEP III, # 151-1. ZMH K- 41367; 4 A F ( SNB 0424-DNA 131, SNB 0425-DNA 132, SNB 0426-DNA 133), 4 A M ( SNB 0373-DNA 106, SNB 0374-DNA 107), 4 (A-1) F ( SNA 0427-0430-DNA 134-137), ANDEEP III, # 151- 7-E, ZMH K- 41368; 3 (A-1) F ( SNB 0431-DNA 138), ANDEEP III, # 151-7-S, ZMH K- 41369.

Distribution ( Fig. 7 ): Recent. Atlantic sector of the SO (Subantarctic and Antarctic regions), 90–1187 m (the only previous abyssal record is herein considered a misidentification, see Remarks section below).

Right valve measurements ( Fig. 7 ): A F L 1.72– 1.91 mm, H 0.77–0.86 mm; A M L 1.63–1.83 mm, H 0.65–0.68 mm; (A-1) L 1.44–1.60 mm, H 0.64– 0.71 mm; (A-2) L 1.18–1.32 mm, H 0.54–0.60 mm; (A-3) L 1.03–1.08 mm, H 0.48–0.50 mm.

Adult chaetotaxy: Antenna I 1, 2(0/.2), 3(.1/.1.), 4 (.1/.1.), 5(.1/.1), 6(.1-2/.3), 7(0/0:4). Antenna II 1(0/0:1), 2(0/0:1), Exopodite (0/0:2,1r), 3(0/6.1c,5), 4[female (.2./.2.1c,3)] [male (.2./.2.1c,2mod,1)], 5(0/.1c,1:4c,1), 6(0/ 0:2c,3s). Mandible 1(0/ 5t,+5.1.), 2(0/.2:1), Exopodite (0/0:1r,6-7), 3(0-1/2-5:1-3), 4(3-7/2-4:0-5), 5(.2c./0:1c,3- 4). Maxilla I vibratory plate (2-3re,~19), palp 1+2(4/0), 3(0/0:5). Appendage V 1(0/0.2-3), Exopodite (0/0:6-8), [female 2, 3, 4(0/.1) 5(.1./0:2)]; [male 2(0/2mod,1:1r), 3(0/.1r:1mod)]. Appendage VI 1(.1/0:1.1.1), 2(.1.1.1/1), 3(0/.1), 4(.1/0), 5(1,1r/0) 6(0/0:1,2c). Appendage VII 1(.1/0), 2(0/.1) 3(.1/0), 4(.1/0), 5(.2/0), 6(0/0:2,1re). Furca 1(0/0:3r.1).

Remarks: Macromckenziea glacierae is an Antarctic and Subantarctic shelf species, which also occurs in the shallow continental slope. The only female recorded from abyssal depths ( Maddocks, 1990, Drake Passage, station GL-2-0022, Weddell Sea, 3111 m) is most probably a misidentification, as its size is much smaller [~ 1.4 mm, similar to other (A-2)] than the adult male holotype (~ 1.82 mm). Furthermore, the extreme size variability of other adult paratypes (~ 1.60–2.10 mm), and the continuous distribution of

590 S. N. BRANDÃO

adult and (A-1) specimens in the length ¥ height scatter plot ( Maddocks, 1990: graph 8) show that more than one species are most probably present on the ‘ type material’ of Mk. glacierae .

Genetic data (Brandão et al., in press) strongly indicate that the morphospecies Mk. glacierae is composed of at least two genetically distinct groups. Based on mitochondrial [Cytochrome Oxidase I ( COI)] and on nuclear [Internal Transcribed Spacer ( ITS)] markers, the two populations of Mk. glacierae – one in the eastern Weddell Sea (no. 74) and the other in the western Weddell Sea (no. 151) – are genetically discrete. However, no difference could be noted between these populations in the chaetotaxy of the appendages (antenna I, antenna II, mandible, maxilla I, appendage V, appendage VI, and appendage VII) and furca, or on the outline of the hemipenises. It is also not possible to separate the two populations (i.e. Eastern and Western) in the length ¥ height scatter plot ( Fig. 7 ), as eastern and western specimens plot mixed within other specimens of the same sex (when adult) or instar (when juvenile). Other cases of genetically distinct species that are morphologically indistinguishable are known in marine shallow and deep-waters for several taxonomic groups, such as for example, the Foraminifera ( Pawlowski et al., 2002), nematodes ( Derycke et al., 2008), gastropods ( Etter et al., 1999), and asselotan isopods ( Brökenland & Raupach, 2008).

Figures

Figure 5. Valves of Yemanja gen. nov. Yemanja coimbrai (Brandão, 2005) gen. nov.: A, B, holotype adult male (MNRJ 15706). C, F, (A-1) (MNRJ 18608). D, paratype (A-1) (MNRJ 15715). E, (A-2) (MNRJ 18610). G, (A-1) (MNRJ 18611). Yemanja youngi (Brandão, 2005) gen. nov.: H, I, paratype adult male (spm 8), J, K, paratype adult female (spm 2) (MNRJ 15701). Yemanja rocas (Brandão, 2005) gen. nov.: L, M, paratypes adult male (spm 7, MNRJ 15703). A, C, E, G, H, L, right valve (RV) external view; B, D, F, I, M, left valve (LV) external view; J, RLV ventral view; K, RLV dorsal view. Scale bar = 500 Mm. Dotted lines representing opaque patches. The arrows point towards the anterior of the animal. Modified after Brandão (2004, 2005).

Figure 6. Soft part morphology of Yemanja gen. nov. Yemanja coimbrai (Brandão, 2005) gen. nov.: A, paratype adult female (spm 7, MNRJ 15714). B, E, K–M, holotype adult male (MNRJ 15706). Yemanja rocas (Brandão, 2005) gen. nov. C, F, J, paratype adult male (spm 7, MNRJ 15703). Yemanja youngi (Brandão, 2005) gen. nov.: D, G–I, paratype adult male (spm 8, MNRJ 15701). A, female appendage V; B–D, right male appendage V; E–G, left male appendage V; H, detail of podomere III of G; I, vibratory plate of maxilla I; J, maxilla I; K, appendage VI; L, appendage VII; M, furca. Scale bars: A–G, I–M = 100 Mm; H = 10 Mm. Modified after Brandão (2004, 2005).

Figure 7. Geographical distribution and height ¥ length scatter plot of valves of Macromckenziea glacierae Maddocks, 1990, Macromckenziea giambonini sp. nov., and Macromckenziea sp. cf. Mk. giambonini. In the scatter plot of Mk. glacierae: F+J, females and juveniles; M, males; EW, eastern Weddell Sea; WW, western Weddell Sea; Mk. sp. Maddocks, Macromckenziea sp. 3 from Maddocks (1990).

Figure 8. Valves of Macromckenziea glacierae Maddocks, 1990. A, B, I, J, adult female (SNB 0016); C, D, adult female (SNB 0014); E, F, K, L, adult male (SNB 0020) (ZMH K-40811); G, H, adult male (SNB 0060, ZMH K-40813). A, C, E, G, right valve (RV) external view; B, D, F, H, left valve (LV) external view; I, K, RV internal view; J, L, LV internal view. Scale bar = 500 Mm.

Figure 9. Appendages of Macromckenziea glacierae Maddocks, 1990. A, C, D, adult male (SNB 0060, ZMH K-40813); B, adult female (SNB 0016, ZMH K-40811). A, appendage VI, B–D, appendage V. Scale bars: A = 100 Mm; B–D = 50 Mm.

Figure 10. Hemipenis, Zenker’s organ, and adductor muscle scars of Macromckenziea glacierae Maddocks, 1990. A, F, H, adult male (SNB 0060, ZMH K-40813). B, E, adult male (SNB 0373-DNA 106, ZMH K-41368). C, adult male (SNB 0020, ZMH K-40811). D, G, adult male (SNB 0410-DNA 118, ZMH K-41366). A–D, hemipenis; E–G, Zenker’s organ; H, adductor muscle scars. Scale bars: A–G = 200 Mm; H = 100 Mm.

Figure 11. Valves and appendages of Macromckenziea giambonini sp. nov. and Macropyxis ghartmanni sp. nov. Macromckenziea giambonini sp. nov.: A–E, paratype adult female (SNB 0031, ZMH K-40810). Macropyxis ghartmanni sp. nov.: F, G, holotype adult female (SNB 0145, ZMH K-41360). A, right valve (RV) external view; B, left valve (LV) external view; C, RV internal view; D, LV internal view; E, adductor muscle scars; F, female appendage V; G, furca. Scale bars: A–D = 500 Mm; E–G = 100 Mm.

Figure 12. Valves and appendages of Macromckenziea giambonini sp. nov. A–H, holotype adult female (SNB 0030, ZMH K- K-40809). A, left valve external view; B, right valve external view; C, antenna I; D, antenna II; E, mandible; F, maxilla I; G, female appendage V; H, appendage VI. Scale bars: A, B = 500 Mm; C–H = 100 Mm.

Figure 13. Appendages of Macromckenziea giambonini sp. nov. A, paratype adult female (SNB 0031), D, paratype adult female (SNB 0032), (ZMH K- K-40810). B, C, holotype adult female (SNB 0030, ZMH K- K-40809). A, B, appendage VII; C, D, furca. Scale bars: A, B = 50 Mm; C, D = 100 Mm.