Macroscapha, SP., 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 : 656-667

? MACROSCAPHA SP. ( FIG. 47O )

1989 in part Macrocyprina tensa, Hartmann, 1989b: 253 .

Material: 1 valve.

1 RV, was contained on a micropalaeontological slide, labelled ‘ K- 33970 . Macrocyprina cf. tensa ., 1 ♀, Antarktis, Winterreise, Stat . PS 09 /V-1: 115(4), l/d. Hartmann, 1986 ’. ZMH K-33970 . This specimen was studied by Hartmann (1989b). This specimen is herein illustrated in Figure 47O .

Remarks: This specimen is conspicuously more globulose than the species of the informal group Mh. tensaopaca ( Fig. 47O ).

INFORMAL GROUP MACROSCAPHA TURBIDA

( FIGS 59–65 )

Remarks: Following previous publications, the geographical distribution of Mh. turbida would be circum-Antarctic, including both Antarctic (Weddell, Scotia, Davis, and Ross Seas) and Subantarctic regions (Scotia Sea, southern South America, and Macquarie Island) ( Fig. 59 ) ( Müller, 1908; Hartmann, 1986, 1987, 1991, 1997; Maddocks, 1990). Additionally the bathymetric distribution is thought to be quite wide: from the upper littoral of King Georg Island (north-western Antarctic Peninsula) to 494 m in the Ross Sea ( Maddocks, 1990; Hartmann, 1991). The same authors have reported very wide morphologic variability in Macroscapha turbida :

1. Carapace outline rectilinear ( Müller, 1908: pl. 11.11, 11.12; Maddocks, 1990: figs 14.12, 15.13) or rounded ( Hartmann, 1986: fig 90, pl. 6.3-5; Maddocks, 1990: figs 14.11, 15.12);

2. Outline of terminal podomere of male appendage V rounded ( Maddocks, 1990: figs 27.6, 27.7) or rectilinear ( Müller, 1908: pl. 11.13, 11.14; Hartmann, 1986: figs 92, 93; Maddocks, 1990: figs 27.8, 27.9);

3. Furca more ( Maddocks, 1990: fig. 47.35) or less ( Müller, 1908: pl. 12.12) asymmetrical;

4. Hemipenis subhemispherical ( Müller, 1908: pl. 12.11; Maddocks, 1990: pl. 94.1. 94.2, 94.4, 94.6, 94.9) or elongated ( Hartmann, 1986: fig. 94; Maddocks, 1990: pl. 94.3, 94.5, 94.7, 94.8, 94.13);

5. Cylindrical tube of Zenker’s organ thick ( Maddocks, 1990: pl. 107.2, 107.3, 107.6-8, 107.10-14) or thin ( Maddocks, 1990. pl. 107.4, 107.5, 107.9);

6. Terminal bulb of Zenker’s organ small ( Maddocks, 1990. pl. 107.7, 107.8, 107.12) or medium-sized ( Maddocks, 1990: pl. 107.4, 107.5, 107.9).

Based on over 300 specimens studied herein (from the Weddell, Scotia, Davis, and Ross Seas) and on published illustrations, Mh. turbida as defined by previous authors ( Müller, 1908; Hartmann, 1986, 1987, 1991, 1997; Maddocks, 1990) is herein subdivided into at least four species, each one with less morphological variability: (1) Mh. turbida occurring in the Davis and Ross Seas; (2) Mh. rehmi sp. nov. occurs in the Weddell Sea. Perhaps the specimens (previously assigned to Mh. turbida ) from the Scotia Sea and from off Macquarie Island ( Maddocks, 1990) should also be included in Mh. rehmi ; (3) Mh. scotia sp. nov., inhabiting the Scotia Sea, includes at least some of the specimens that Hartmann (1986, 1987, 1989b, 1990, 1991) recorded as Mh. turbida ; (4) Mh. subhemispherica sp. nov. includes the specimens recorded as Mh. turbida by Hartmann (1991) from the upper littoral of King Georg Island. Additional records of Mh. turbida from the Pacific Sector of the SO and from the northern Scotia Sea (including the southern tip of South America; Maddocks, 1990) are herein considered dubious, as they include mostly females and subfossils, and are based on a much wider morphological species concept, which obviously comprises species other than Mh. turbida . The presence of more than one species in the material identified by previous authors as Mh. turbida was anticipated because of the continuous distribution of the specimens on the length ¥ height scatter plot ( Maddocks, 1990: graph 52). Each of the four species – Mh. turbida , Mh. rehmi , Mh. scotia , and Mh. subhemispherica – presents a discrete distribution of size ( Fig. 59 ). Additionally, several specimens recorded previously from the Scotia Sea ( Hartmann, 1987, 1989b, 1993) are herein left in open nomenclature, because (1) they are females and obviously lack the hemipenis and Zenker’s organs, which are the diagnostic characters differentiating Mh. turbida , Mh. rehmi , and Mh. scotia ( Figs 60 , 61 ); (2) and/or the valves are entirely decalcified.

Macroscapha subhemispherica sp. nov. is easily differentiable from the other three species of the ‘ Mh. turbida group’ ( Mh. rehmi , Mh. scotia , and Mh. turbida ) in valve outline and size: the former species is considerably smaller and subhemispherical in outline, with a highly and continuously arched dorsal margin.

Like many freshwater ostracods ( Meisch, 2000; Petkovski, Scharf & Keyser, 2002), in spite of the fact that the three remaining species show different valve outlines, their correct identification depends on the study of the male soft parts. In Mh. turbida , Mh. rehmi , and Mh. scotia the hemipenis is subhemispherical, sometimes slightly elongate in lateral view and presents a tetrasegmented copulatory process ( Fig. 60 ): (1) the most proximal segment is elongated and terminates distally in a bulbous structure that resembles the condyles of a human femur. Distally this proximal rod is articulated with two processes (2) one of those is conical (with sinuous tip) and perforated by the vas deferens, and transfers the sperm to the female ( Fig. 60B, F, J ); (3) the other process is short and varies in form amongst the three species ( Fig. 60C, G, K ); (4) attached to this last segment (3) is a lamelliform structure that is larger and flexible in Mh. turbida and Mh. scotia sp. nov. ( Fig. 60D, H ), but smaller and more sclerotized in Mh. rehmi ( Fig. 60L ). Additionally, Mh. rehmi sp. nov. is differentiable from Mh. turbida and Mh. scotia sp. nov. because the first species presents a Zenker’s organ with a very small terminal bulb ( Fig. 41T–X), whereas the other two species present medium-sized terminal bulbs ( Fig. 41J–S). Moreover, podomere III of the male appendage V (= palp podomere II) is thinner and smoothly rounded in Mh. rehmi . Finally, Mh. turbida is larger, with a more sinuous ventral margin and rectilinear anterior margin than Mh. scotia sp. nov., which is conspicuously smaller, with a rounded, subtriangular outline.

MACROSCAPHA TURBIDA (G. W. MÜLLER, 1908)

( FIGS 39C, I, R–U , 41N–S, Y, 1 , 59 , 60A–D , 61A–D , 62A–E , 63A–E , 64B–D )

1908 Macrocypris turbida G. W. Müller, 1908: 94–95 , pl. 9, figs 11–14, pl. 12, figs 12–13.

1977 in part Macrocyprina turbida, Maddocks, 1977 .

1979 non Macrocyprina turbida, Maddocks, 1979

1986 non Macrocyprina turbida, Hartmann, 1986: 174 .

1987 non Macrocyprina turbida, Hartmann, 1987: 132 .

1989b non Macrocyprina turbida, Hartmann,1989b: 253 .

1990 non Macrocyprina turbida, Hartmann, 1990: 212 .

1990 in part Macroscapha turbida, Maddocks, 1990: 105–106 .

1991 non Macroscapha cf. turbida, Hartmann, 1991: 216 .

1993 non Macrocyprina turbida, Hartmann, 1993: 232 .

1997 in part Macrocyprina turbida, Hartmann, 1997: 251–253 .

Material: 209 live specimens plus 2 valves.

Previously the type series used by Müller to describe ‘ Macrocypris turbida ’ consisted of seven slides and glass containing alcohol and labelled ‘ Macrocypris turbida G. W. Müller , Gauss-Station’ (65°S, 90°E, 385 m depth), ‘Deutsche Südpolar Exp., 1902- 03, ZMB 13133’. These specimens are catalogued as type-series 13133 in the Crustacea Division of the Museum für Naturkunde, Humboldt University, Berlin. Each of the slides contains the carapace and usually the soft parts of one specimen imbedded in Canada balsam. All are labelled ‘Gauss-Exp.,’ and most are labelled ‘178.’

Lectotype: 1 A M on a glass slide labelled ‘2-4.12.1, 1364’ and bearing Maddocks’ (1990) specimen number 1943M ( Maddocks, 1990: 106).

Paralectotypes: 7 glass slides plus 34 *live specimens preserved in alcohol, 1 RLV on a micropalaeontological slide .

The glass slides bear Maddocks’ specimen numbers 1942, 1944–48 ( Maddocks, 1990). Another specimen, catalogued in type series 13131 and mislabelled ‘ Macrocypris tensa ,’ bearing Maddocks’ specimen number 1934M, also belongs to Mh. turbida and was designated paralectotype of the last species ( Maddocks, 1990: 106).

Inside this larger glass there were eight small vials labelled ‘178-6.02’, ‘178-7.02’, ‘178-8.02’, ‘178-10.02’, ‘178-11.02’, ‘178-12.02’, ‘178-1.03’, ‘178-3.2.03’ (see below). These labels most probably indicate the date of collection of each sample.

*Part of the material listed below as live A (A-1) (A-2) (A-3) (A-?4) might or might not contain soft parts; as all the valves from the Gausstation were opaque, it was not possible to be sure if some specimens were live or subfossil.

1. Vial labelled ‘178-6.02’ – live: 1 A F (=RV> LV);

2. Vial labelled ‘178-7.02’ – live: 1 (A-1) M (=LV> RV), 4 decalcified specimens;

3. Vial labelled ‘178-8.02’ – live: 1 (A-1) F (=RV> LV), 1 decalcified (A-?4);

4. Vial labelled ‘178-10.02’ – live: 1 A F (=RV> LV);

5. Vial labelled ‘178-11.02’ – live: 2 (A-1) F (=RV> LV), 1 decalcified (A-1), 1 (A-2) F (=RV> LV), 1 (A-2) M (=LV> RV);

6. Vial labelled ‘178-12.02’ – live: 2 A M (LV> RV), 1 (A-1) F (=RV> LV), 1 (A-1) M (LV> RV), 2 (A-3) F (=RV> LV); 1 decalcified juvenile; subfossil 1 RLV A.

7. Vial labelled ‘178-1.03’ – live: 1 A F (=RV> LV), 1 A M (LV> RV), 2 (A-2) M (LV> RV), 2 (A-2) F (RV> LV), 1 (A-2) M (LV> RV), 1 (A-3) M (LV> RV), 3 decalcified juveniles.

8. Vial labelled ‘178-3.2.03’ – live: 2 A M (LV> RV) (SNB 0747). The specimen SNB 0747 is herein illustrated in Figures 39T–U , 41N–O, Y, 61A–B , 62A , 63A .

Additional material: 64 A F (SNB 0345-DNA 82, SNB 0348-DNA 85, SNB 0353, 0356), 62 A M (SNB 0342- DNA 79, SNB 0343-DNA 80, SNB 0344-DNA 81, SNB 0346-DNA 83, SNB 0347-DNA 84, SNB 0349-DNA 86, SNB 0350-DNA 87, SNB 0351-DNA 88, SNB 0352, SNB 0355, SNB 0709), 20 (A-1) F, 15 (A-1) M, 1 (A-2) F, 1 (A-2) M, 12 live specimens (SNB 0354), H in 5, ZMH K-41476.

Distribution ( Fig. 59 ): Recent. Davis and Ross Seas, depth 196– 385 m.

Valve measurements ( Fig. 59 ): A F RV L 1.90– 2.02 mm, H 0.80–0.88 mm; A M LV L 1.86–1.96 mm, H 0.74–0.83 mm; (A-1) F RV L 1.48–1.60 mm, H 0.58–0.66 mm; (A-1) M LV L 1.47–1.56 mm, H 0.58–0.65 mm; (A-2) F RV L 1.18–1.26 mm, H 0.48– 0.50 mm; (A-2) M LV L 1.18–1.24 mm, H 0.48–0.50 mm; (A-3) F RV L 0.92–1.02 mm, H 0.38– 0.44 mm; (A-3) M LV L 1.00 mm, H 0.38 mm; (A-?4) F RV L 0.82 mm, H 0.38 mm.

Diagnosis: Carapace medium-sized (for the genus). Lateral outline of valves subhemispherical. LV larger than RV in males, but RV larger than LV in females. Podomere V of female appendage V with two mediumsized claws and one short, dorsal claw. Male appendage V very asymmetrical, but in both sides podomere III pointed at 90°. Hemipenis subhemispherical, copulatory process tetrasegmented, second segment elongate, strongly sclerotized, distally bifurcated; distal segment conical, sinuous perforated by the vas deferens. Zenker’s organ with robust chitinous central tube and medium-sized terminal bulb. Furca slightly asymmetrical.

Description: Carapace medium-sized (for the genus), in dorsal view elongated with pointed ends, maximum thickness at mid-length. Ventral margin of larger valve conspicuously sinuous in ventral view. Lateral outline of valves subhemispherical; ventral margin sinuous, with conspicuous indentation on mouth region; anterior margin protruded and angulated; posterior margin rounded at 90°; maximum height posterior to mid-length. LV larger than and overlapping RV in males, but RV larger than and overlapping LV in females.

Antennae I and II thin and slender, with relatively elongated podomeres and long setae. Palp of mandible very elongate, especially podomeres II and III. Palp of maxilla I with three articulated podomeres, podomere III with six distal setae; vibratory plate with two long and one short strahlen, plus around 18–19 feathered setae. Podomere II of female appendage V long and wide; podomere V with two medium-sized claws and one short, dorsal claw. Male appendage V asymmetrical, right one with very elongated, modified, ventral seta (= peg), and one shorter, but robust modified seta (= peg), plus short seta; left male appendage V with two short, robust modified seta (= peg), plus short seta; podomere III of both sides pointed at 90°, with one modified, thick distal seta. Podomere VI of appendage VI with one short, ventral seta, one medium-sized, dorsal claw, and one long, mediodistal claw. Appendage VII slender, with very long podomere III. Furca slightly asymmetrical, with minute tubercles on distal, thick setae. Hemipenis subhemispherical, copulatory process tetrasegmented, proximal segment longest and very strongly chitinized, second segment elongate, strongly sclerotized, distally bifurcated; lamellar segment thin and flexible; distal segment conical, sinuous perforated by the vas deferens. Zenker’s organ with robust chitinous central tube and medium-sized terminal bulb; vas deferens arranged in few loops as long as half of the Zenker’s organ’s central tube.

Adult chaetotaxy: Antenna I 1, 2(0/.2), 3(.1/.1.), 4(.1/.1.), 5(.1/.1), 6(.2/.3), 7(0/0:6). Antenna II 1(0/0:1), 2(0/0:1), Exopodite (0/0:2,1r), 3(0/5:4), 4[female (.2r./.2r.1c,3)] [male (.2r./.2r.1c,2mod,1)], 5(0/.1c,1: 4c,1), 6(0/0:3c,?1). Mandible 1(0/ 6t,+5.1.), 2(0/.2:1), Exopodite (0/0:1r,6), 3(.0-2/.1-4:4), 4(.2.2/.4), 5(0/ 0:3c,3). Maxilla I vibratory plate (3re,18-19), palp 1 (.1/0), 2(.4/0), 3(0/0:6). Appendage V 1(0/.2.1.1), Exopodite (0/0:3.6-7), [female 2, 3, 4(0/.1) 5(.1./0:1c,1)]; [male 2(0/.2mod,1), 3(0/0:1mod)] Appendage VI 1(0/.1- 2), 2(.1/.2.1), 3(0/.1), 4(0/.1), 5(0/.1,1r), 6(0/0:2c,1). Appendage VII 1(.1/0), 2(.1.1/.1), 3(0/.1), 4(0/.1), 5(0/.2), 6(0/0:1.1re,1). Furca 1(0/0:1.4r).

Remarks: Contrary to previous assumptions ( Maddocks, 1990: 11), all males of Mh. turbida studied herein present the left valve larger than, and overlapping, the right valve anterodorsally, posterodorsally, and ventrally ( Fig. 64B, C ). However, in all females the right valve is larger than and overlaps the left valve ( Fig. 64D ).

Based on the published illustrations the specimens recorded as Mh. turbida from the Scotia Sea and off Macquarie Island ( Maddocks, 1990) are herein considered to be a different species. Some of them may be conspecific with Mh. rehmi sp. nov.

MACROSCAPHA REHMI SP. NOV. ( FIGS 39B, N–O , 41T–X, 41.4, 59 , 60E –H, 61E–I , 62G–I , 63G–I , 64K–L )

1979 Macrocyprina turbida, Maddocks, 1979 : pl. 1.13.

1990?in part Macroscapha turbida, Maddocks, 1990: 105–106 ,?pls. 41.6, 94.10, 107.13.

1997 in part Macrocyprina turbida, Hartmann, 1997: 251–253 .

Etymology: In honour of Dr Peter Rehm (Universität Hamburg, Germany), who collected the Ross Sea samples studied in the present publication.

Material: 25 live specimens.

Holotype: 1 A M ( SNB 0744 ) (soft parts in glass slide, valves in micropalaeontological slide), EASIZ II, # 171, ZMH K-41482 .

Paratypes: 1 A F ( SNB 0742 ), 1 A, 1 (A-1), EASIZ II , # 48-111, ZMH K-40835 ;

10 A F (SNB 0028, 0059), 12 A M (SNB 0025, 0026, 0058, 0742, 0743), 1 (A-1), 1 RV, EASIZ II, # 171, ZMH K-40836.

Distribution ( Fig. 59 ): Recent. Eastern Weddell Sea, 231– 397 m.

Valve measurements ( Fig. 59 ): Holotype, RV L 1.66 mm, H 0.68 mm; LV L 1.64 mm, H 0.64 mm. Paratypes, A L 1.54–1.69 mm, H 0.64–0.72 mm; (A-1) L 1.38–1.40 mm, H 0.54 mm.

Diagnosis: Carapace small (for the genus); lateral outline very elongated; anterior margin protruded, posterior margin rounded. Podomere V of female appendage V with two long claws and one mediumsized, dorsal claw, the latter less than half as long as the long claws. Podomere III (= palp podomere II) of left and right male appendage V smoothly curved and thin. Furca slightly asymmetrical; terminal setae with minute tubercles. Hemipenis elongated, with sinuous outline, maximum height proximally; copulatory process tetrasegmented, second segment curved and distally pointed. Zenker’s organ with thick central, sclerotized tube and very small terminal bulb.

Description: Carapace small (for the genus). Lateral outline of valves very elongated; ventral margin slightly sinuous; anterior margin protruded and nar- rowly rounded; posterior margin obtusely rounded; maximum height conspicuously posterior to midlength. RV always larger than and overlapping LV. Females higher in relation to length than males. Zone of concrescence very narrow, with short, straight radial pore canals; vestibules medium sized.

Antennae I and II thin and slender, with relatively elongated podomeres and long setae. Mandible with one conical plus five tricuspidate teeth on masticatory process, exopodite with one reduced plus seven medium-sized setae. Palp of maxilla I with three articulated podomeres, podomere III with six setae; vibratory plate with two long strahlen, plus 19 feathered setae. Podomere II (= palp podomere I) of female appendage V longer than two times the length of podomere III; podomere IV with one medium-sized, ventrodistal claw; podomere V with two long claws and one medium-sized, dorsal claw, the latter less than half as long as the long claws. Male appendage V asymmetrical, right one with very elongated, modified, ventral seta (= peg), and one shorter, but robust, modified seta (= peg), plus one short seta; left male appendage V with two short, robust modified seta (= peg), plus one short seta; podomere III of both sides thin and smoothly rounded, with one modified, thick distal seta. Podomere VI of appendage VI with one short, ventral seta, one medium-sized, dorsal claw, and one long, medial claw. Appendage VII slender, with very long podomere III. Furca slightly asymmetrical, with tubercles on distal, thick setae; slight suture between rods and terminal setae. Hemipenis elongated, with sinuous outline, maximum height proximally; copulatory process tetrasegmented; proximal segment longest, and very strongly sclerotized; second segment curved, strongly sclerotized, and distally pointed; lamellar segment large, thin, and flexible; distal segment conical, sinuous perforated by the vas deferens. Zenker’s organ with thick, strongly sclerotized, central tube and very small terminal bulb; vas deferens arranged in concentric loops half as long as central tube.

Adult chaetotaxy: Antenna I 1, 2(0/.2), 3(.1/.1.), 4(.1/.1.), 5(.1/.1), 6(.2/.3), 7(0/0:6). Antenna II 1(0/0:1), 2(0/ 0:1), Exopodite (0/0:2,1r), 3(0/6.2,2r), 4[female (.1r./.1r.1c,2)] [male (.1r./.1r.1c,2mod,1)], 5(0/.1c,1:4c,1), 6(0/0:3c,1?-?2). Mandible 1(0/ 6t,+5.1.), 2(0/.2:1), Exopodite (0/0:1r,7), 3(.0/.4:4), 4(.3.2/.4), 5(0/0:3c,2). Maxilla I vibratory plate (2re,19), palp 1 (.1/0), 2(.4/0), 3(0/0:6). Appendage V 1(0/.1-2.1), Exopodite (0/0:3.6- 7), [female 2, 3, 4(0/.1) 5(.1./0:1c,1)]; [male 2(0/.2mod,1:1r), 3(0/0:1mod)] Appendage VI 1(0/.2), 2(.1/.2.1), 3(0/.1), 4(0/.1), 5(0/.1,1r), 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/ 0:1.1re,1). Furca 1(0/0:1.3r).

Remarks: The two specimens [1 A M, 1 F (A-1)] recorded as Mh. turbida from the Scotia Sea and off Macquarie Island ( Maddocks, 1990) may belong to Mh. rehmi sp. nov.

The valve lateral outline of Mh. rehmi sp. nov. is conspicuously more elongated (less high in relation to length) than most of the previously described Macroscapha species (i.e. Mh. atlantica Mh. heroica , Mh. inaequalis , Mh. inaequata , Mh. jiangi , Mh. marchilensis , Mh. opaca , Mh. sinuata , Mh. tensa , Mh. turbida ). Macroscapha gyrae is more oval shaped and equilateral in lateral view, with a more acute posterior angle than Mh. rehmi . Furthermore the lateral outline of the hemipenis of all Macroscapha species (with male soft parts illustrated) is conspicuously different from Mh. rehmi . The Zenker’s organ of Mh. rehmi bears a very small terminal bulb (similar to some species of the genus Macrocyprina ) that is considerably smaller than in other Macroscapha species.

MACROSCAPHA SCOTIA SP. NOV. ( FIGS 37D–G , 38D–G , 39A, L–M , 41J–M, 41.3, 60I–L , 61J–M , 64E–F, M–N , 65 )

1986 Macrocyprina turbida, Hartmann, 1986: 174 , figs 90–94, pl. 6, figs 3–7.

1987 Macrocyprina turbida, Hartmann, 1987: 132 .

1989b Macrocyprina turbida, Hartmann, 1989b: 253 .

1990 Macrocyprina turbida, Hartmann, 1990: 212 .

1990 in part Macrocyprina tensa, Hartmann, 1990: 212 .

1991? Macroscapha cf. turbida, Hartmann, 1991: 216 .

1993 Macrocyprina turbida, Hartmann, 1993: 232 .

1997 in part Macrocyprina turbida, Hartmann, 1997: 251-253 .

Etymology: For its occurrence in the Scotia Sea, name used in apposition.

Material: 99 live specimens plus 14 valves.

Holotype: 1 A M ( SNB 0739 ) (soft parts in glass slide, valves in micropalaeontological slide), EASIZ II, # 310, ZMH K-40820 .

Paratypes: 18 A F ( SNB 0024 , 0056 , 0740 , 0741 ), 28 A M ( SNB 0022 , 0023 , 0027 , 0738 , 0793 , 0794 ), 8 A (0785-0792), 4 (A-1), 3 (A-2), 7 damaged live specimens, EASIZ II , # 310, ZMH K-40837 .

1 live juvenile, 3 live specimens, plus partially dissected specimens (at least 4 A F, 2 A M) were contained in a glass, labelled ‘ K- 33192 , 12 ♂♂, ♀♀ , Macrocyprina turbida G. W. Müller, 1908 , Antarktis PS 158 , leg. Mühlenhardt-Siegel, det. Hartmann 1985’. This specimen was studied by Hartmann (1986). ZMH K-33192 .

1 RLV, contained in micropalaeontological slide, labelled ‘K- 33192a. Macrocyprina turbida G. W. Müller, 1908 , Antarktis PS 158, Joinville, leg. Mühlenhardt-Siegel, det. Hartmann, 1986 ’. This specimen was studied by Hartmann (1986). ZMH K-33192.

1 LV, 3 RLV ( SNB 0870 ), were contained on a micropalaeontological slide, labelled ‘ K- 33305 b. Macrocyprina tensa . Antarktis PS 06 /158, Joinville , leg. Müh. – Sie., 26.11.1984, det. Hartmann’. These specimens were studied by Hartmann (1987) . ZMH K-33192 . 1 RLV is herein illustrated in Figure 64F .

1 RV, 1 LV (plus broken valves) were contained on a micropalaeontological slide, labelled ‘ K- 33305 c. Macrocyprina tensa . Antarktis PS 06 / 158, Joinville , leg. Müh. – Sie., 26.11.1984, det. Hartmann’. These specimens were studied by Hartmann (1987) . ZMH K-33192 .

2 A M ( SNB 0850 , 0851 ), plus 10 live specimens, were contained in a glass with glycerine, labelled ‘ K- 33237 . Macrocyprina turbida Müller, 1908 , Antarktis PS 207 , Bransfield Str., leg. Mühlenhardt-Siegel, det. Hartmann 1985, 12 males and females’. This specimens were studied by Hartmann (1986). ZMH K-33237 .

1 dissected specimen was contained in a glass with glycerine labelled ‘K- 33300, Macrocyprina cf. turbida, Antarktis, Sta. WH 142, lg. Mühlenhardt- Siegel, 4.12.1984, det. Hartmann, 1986 ‘ This specimen was studied by Hartmann (1987). ZMH K-33300.

1 A M, 1 A F, 3 live juveniles, contained in a glass with glycerine labelled ‘K- 33301, Macrocyprina turbida, Antarktis, Sta. WH 266, lg. Mühlenhardt- Siegel, 4.12.1984, det. Hartmann, 1986 ‘ These specimens were studied by Hartmann (1987). ZMH K-33301.

1 live specimen was contained in a glass with glycerine, labelled ‘K- 33302, Macrocyprina cf. turbida, Antarktis, Sta. PS 06/158, lg. Mühlenhardt- Siegel, 4.12.84, det. Hartmann, 1986 ‘ This specimen was studied by Hartmann (1987). ZMH K-33302.

A few dissected specimens – at least 1 A M ( SNB 0864 ), 1 A F, and 2 juveniles – were contained in a glass with glycerine, labelled ‘ ZMH K-34502 , Macrocyprina cf. tensa, Antarktis, Stat. PS Ant VI-2 (118) , Tiefe 207– 188 m, rp. Hartmann et al. 5.11.87, 2 ♀, 2 subad., L., d. Hartmann (1989)’. These specimens were studied by Hartmann (1990). ZMH K-34502 .

1 A F, 5 live juveniles, labelled ‘ K- 34506 , Macrocyprina turbida, Antarktis, Stat. PS Ant VI-2 (90/5) , Tiefe 197–205 m, rp. Hartmann et al. 2.11.1987, 1 female, 5 larvae, det. Hartm. (1989)’. These specimens were studied by Hartmann (1990). ZMH K-34506 .

1 A M, 2 RV, 1 LV, contained in a glass and one micropalaeontological slide, labelled ‘ K- 34507 , Macrocyprina turbida, Antarktis, Stat. PS Ant VI-2 (207) , Tiefe 146 m, rp. Hartmann et al. 7.12.1987, 1 male, 1 Schale, det. Hartmann (1989)’. These specimens were studied by Hartmann (1990) . ZMH K-34507 .

A few dissected specimens – at least 3 A M (SNB 0868, 0869) – were contained in a glass with glycerine, labelled ‘ZMH K-35501, Macroscapha turbida (G. W. Müller, 1908) , Antarktis, zw. Joinville und d’Urville (Beyerschlitten), PS Ant X 1b, 27.12.91, l/d. Hartmann’. These specimens were studied by Hartmann (1993). ZMH K-35501.

4 RV, 3 LV, were contained on a micropalaeontological slide, labelled ‘ZMH K-35501, Macroscapha turbida (G. W. Müller, 1908) , Antarktis, Ant. X 1b, zwischen Joinvil und d’Urville, 27.12.91, l/d. Hartmann’. These specimens were studied by Hartmann (1993). ZMH K-35501. These specimens are herein illustrated in Figures 37C and 38C .

Distribution ( Fig. 59 ): Recent. Scotia Sea, 68– 290 m.

Figures

Figure 1. Localities for samples studied herein. A, localities from the cruise EASIZ II samples, plus localities from specimens re-studied herein (several cruises from the R. V. Polarstern and R. V. Walter Herwig). B, samples from ANDEEP I, II, and III cruises. Arabic numerals are the station numbers as in the cruise reports. For repeated station numbers: WH, samples collected on board of the R. V. Walter Herwig; PS, samples collected on board of the R. V. Polarstern (cruises other than EASIZ II, PS 48 / ANTXV-3); E, samples collected on board of the R. V. Polarstern during the cruise of EASIZ II (PS 48 / ANTXV-3). Some station names are abbreviated: 1nO, ‘1. Station nach O’Higgins’; Fa, ‘Faraday’; zJU, ‘Zwischen Joinville und d’Urville’. For further details see Table 1. The faunistic analyses discussed below were based on the EBS samples marked ‘*’.

Figure 37. External view of right valves of Macroscapha inaequata Maddocks, 1990, Macroscapha scotia sp. nov., and Macroscapha walterae sp. nov. Macroscapha inaequata Maddocks, 1990: A, adult female (SNB 0752, ZMH K-40821); B, adult male (SNB 0578-DNA 285. ZMH K-41487); H, (SNB 0052); I, (SNB 0053); J, adult female (SNB 0054) (ZMH 40819). Macroscapha scotia sp. nov.: C, paratype A M? (ZMH K-35501); D, holotype adult male (SNB 0739, ZMH K-40820); E, paratype adult male (SNB 0022); F, paratype adult female (SNB 0056); G, paratype adult female (SNB 0741) (ZMH K-40837). Macroscapha walterae sp. nov.: K, holotype adult male (SNB 0745, ZMH K-41484); L, paratype adult male (SNB 0051); M, paratype adult male (SNB 0750); N, paratype adult female (SNB 0048) (ZMH K-40818). Scale bar = 500 Mm.

Figure 38. External view of left valves of Macroscapha inaequata Maddocks, 1990, Macroscapha scotia sp. nov.; and Macroscapha walterae sp. nov. Macroscapha inaequata Maddocks, 1990: A, adult female (SNB 0752, ZMH K-40821); B, adult male (SNB 0578-DNA 285. ZMH K-41487); H, (SNB 0052); I, (SNB 0053); J, adult female (SNB 0054) (ZMH 40819). Macroscapha scotia sp. nov.: C, paratype A (ZMH K-35501); D, holotype adult male (SNB 0739, ZMH K-40820); E, paratype adult male (SNB 0022); F, paratype adult female (SNB 0056); G, paratype adult female (SNB 0741) (ZMH K-40837). Macroscapha walterae sp. nov.: K, holotype adult male (SNB 0745, ZMH K-41484); L, paratype adult male (SNB 0051); M, paratype adult male (SNB 0750); N, paratype adult female (SNB 0048) (ZMH K-40818). Scale bar = 500 Mm.

Figure 39. Appendages V, VI, and VII and furcae of Macroscapha inaequata Maddocks, 1990, Macroscapha rehmi sp. nov., Macroscapha scotia sp. nov., Macroscapha turbida (Müller, 1908), and Macroscapha walterae sp. nov. Macroscapha scotia sp. nov.: A, paratype adult female (SNB 0024); L, M, paratype adult male (SNB 0738), ZMH K-40837. Macroscapha rehmi sp. nov.: B, paratype adult female (SNB 0028); N, O, paratype adult male (SNB 0743); ZMH K-40836. Macroscapha turbida (Müller, 1908): C, adult female (SNB 0348-DNA 85); I, adult male (SNB 0355); R, adult male (SNB 0351-DNA 88); S, adult male (SNB 0347-DNA 84) (ZMH K-41476); T, U, paralectotype (SNB 0747, ZMB 13133). Macroscapha inaequata Maddocks, 1990: D, adult female (SNB 0752, ZMH K-40821); H, adult male (SNB 0753, ZMH K-40822). Macroscapha walterae sp. nov.: E, paratype adult female (SNB 0611-DNA 318); F, J, K, paratype adult female (SNB 0462-DNA 169); G, P, (A-1) male paratype (SNB 0612-DNA 319); Q, paratype (A-1) male (SNB 0613-DNA 320) (ZMH K-41485). A–E, female appendage V; F, appendage VI; G–I, K, furca; J, appendage VII; L–O, R–U, male appendage V; P, Q, (A-1) appendage V. Scale bars = 100 Mm.

Figure 47. Right valves and Zenker’s organ of species of the informal group ‘Macroscapha tensa-opaca’. Macroscapha cactus sp. nov.: A, P, holotype adult male (SNB 0041, ZMH K-41483); B, paratype adult male (SNB 0764); C, paratype adult male (SNB 0766) (ZMH K-40824). Macroscapha falcis sp. nov.: D, holotype adult female (SNB 0854, ZMH K-41497); E, paratype adult male (SNB 0471-DNA 178); F, paratype adult male (SNB 0493-DNA 200); Q, adult male (SNB 0104-DNA 7), (ZMH K-41496). Macroscapha solecavai sp. nov.: G, paratype adult female (SNB 0566-DNA273); H, paratype adult male (SNB 0563-DNA 271); I, paratype adult male (SNB 0561-DNA 268) (ZMH K-41499); R, adult male (SNB 0518-DNA 225), (ZMH K-41498). Macroscapha sp. nov. 1: J, A (ZMH K-33305). Macroscapha sp. nov. 2: L, A (ZMH K-34505a). Macroscapha sp. nov. 3: K, A? (ZMH K-33786). Macroscapha sp.: M, A (ZMH K-34500); N, A (ZMH K-35373).?Macroscapha sp.: O, subfossil (ZMH K-33970). A–O, right valve external view; P–R, Zenker’s organ. Scale bars = 500 Mm.

Figure 59. Geographical distribution and height ¥ length scatter plot of valves of Macroscapha turbida informal group. Previous records of Mh. turbida from the northern Scotia Sea and from the Pacific Sector of the Southern Ocean are herein considered dubious because they were based on a much wider species morphological concept, which includes at least the three new species described herein – Macroscapha rehmi, Macroscapha scotia, and Macroscapha subhemispherica. Mh. subhemis. – Mh. subhemispherica sp. nov.; Mh. spp. Mad. – dubious records of Mh. turbida by Maddocks (1990); Mh. spp. Hart. – dubious records of Mh. turbida by Hartmann (1987, 1989b).

Figure 60. Details of hemipenis of Macroscapha rehmi sp. nov., Macroscapha scotia sp. nov., and Macroscapha turbida (Müller, 1908). Macroscapha turbida (Müller, 1908): A–D, adult male (SNB 0344-DNA 81, ZMH K-41476). Macroscapha rehmi sp. nov.: E–H, paratype adult male (SNB 0025, ZMH K-40836). Macroscapha scotia sp. nov.: I–L, holotype adult male (SNB 0739, ZMH K-40820). A, entire hemipenis; B, F, J, conical, perforated process of copulatory rod; C, G, K, distal, short process of copulatory rod; D, H, L, lamelliform structure of copulatory rod; E, I, entire copulatory rod. Scale bars: A, E, I = 100 Mm; B–D, F–H, J–L = 50 Mm.

Figure 61. Hemipenis of Macroscapha rehmi sp. nov., Macroscapha scotia sp. nov., and Macroscapha turbida (Müller, 1908). Macroscapha turbida (Müller, 1908): A, B, paralectotype adult male (SNB 0747, ZMB 13133); C, adult male (SNB 0344-DNA 81); D, (SNB 0347-DNA 84), (ZMH K-41476). Macroscapha rehmi sp. nov.: E, holotype adult male (SNB 0744, ZMH K-41482); F, paratype adult male (SNB 0025); G, paratype adult male (SNB 0058); H, paratype adult male (SNB 0026) (ZMH K-40836); I, paratype adult male (SNB 0742, ZMH K-40836). Macroscapha scotia sp. nov.: J, paratype adult male (SNB 0022); K, paratype adult male (SNB 0027); L, paratype adult male (SNB 0738); M, paratype adult male (SNB 0739) (ZMH K-40837). Scale bar = 100 Mm.

Figure 62. External view of right valves of Macroscapha rehmi sp. nov., Macroscapha subhemispherica sp. nov., Macroscapha turbida (Müller, 1908), and Macroscapha spp. (from the Mh. turbida group). Macroscapha turbida (Müller, 1908): A, paralectotype adult male (SNB 0747, ZMB 13133); B, adult male (SNB 0343-DNA 80); C, adult male (SNB 0352); D, adult male (SNB 0355); E, adult female (SNB 0356) (ZMH K-41476). Macroscapha rehmi sp. nov.: G, holotype adult male (SNB 0744, ZMH K-41482); H, paratype adult male (SNB 0743); I, paratype adult male (SNB 0059) (ZMH K-40836). Macroscapha subhemispherica sp. nov.: J, paratype A (ZMH K-34830); K, L, paratype A (ZMH K-34835). Macroscapha spp.: F, M, adult female? (ZMH K-35502b). Scale bar = 500 Mm.

Figure 63. External view of left valves of Macroscapha rehmi sp. nov., Macroscapha subhemispherica sp. nov., Macroscapha turbida (Müller, 1908), and Macroscapha spp. (from the Mh. turbida group). Macroscapha turbida (Müller, 1908): A, paralectotype adult male (SNB 0747, ZMB 13133); B, adult male (SNB 0343-DNA 80); C, adult male (SNB 0352); D, adult male (SNB 0355); E, adult female (SNB 0356) (ZMH K-41476). Macroscapha rehmi sp. nov.: G, holotype adult male (SNB 0744, ZMH K-41482); H, paratype adult male (SNB 0743); I, paratype adult male (SNB 0059) (ZMH K-40836). Macroscapha subhemispherica sp. nov.: J, paratype A (ZMH K-34830); K, L, paratype A (ZMH K-34835). Macroscapha spp.: F, M, adult female? (ZMH K-35502b). Scale bar = 500 Mm.

Figure 64. Dorsal, ventral, and internal views of Macroscapha inaequata (Maddocks, 1990), Macroscapha rehmi sp. nov., Macroscapha scotia sp. nov., Macroscapha turbida (Müller, 1908), and Macroscapha walterae sp. nov. Macroscapha walterae sp. nov.: A, subfossil paratype (ZMH K-41486); I, J, paratype adult male (SNB 0051, ZMH K-40818). Macroscapha turbida (Müller, 1908): B, C, adult male; D, adult female (ZMH K-41476). Macroscapha scotia sp. nov.: E, paratype adult female, F, paratype adult male (ZMH K-33192); M, N, paratype adult male (SNB 0022, ZMH K-40837). Macroscapha inaequata (Maddocks, 1990): G, H, adult female (SNB 0054, ZMH K-40819). Macroscapha rehmi sp. nov.: K, L, paratype adult female (SNB 0059, ZMH K-40836). A, B, D–F, right and left valve (RLV) dorsal view; C, RLV ventral view; G, I, K, M, LV internal view; H, J, L, N, RV internal view. Scale bar = 500 Mm. The arrows point towards the anterior of the animal.

Figure 65. Soft parts of Macroscapha scotia sp. nov. A–D, paratype adult male (SNB 0022, ZMH 40837). A, furca; B, C, male appendage V; D, appendage VI. Scale bars = 100 Mm.