Challenging cosmopolitanism in the deep sea: The case of “Cythere acanthoderma Brady, 1880” (Crustacea, Ostracoda)

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www.sciencedirect.com Revue de micropaléontologie 56 (2013) 2–19

Original article

Challenging cosmopolitanism in the deep sea: The case of “Cythere acanthoderma Brady, 1880” (Crustacea, Ostracoda) Remise en question du cosmopolitisme en milieu marin profond : le cas de « Cythere acanthoderma Brady, 1880 » (Crustacea, Ostracoda) Simone Nunes Brandão ∗ Senckenberg am Meer, Deutsches Zentrum für Marine Biodiversitätsforschung (DZMB), Südstrand 44, 26382 Wilhelmshaven, Germany

Abstract Cythere acanthoderma Brady was described from Recent sediments collected from all world’s oceans during the HMS Challenger expedition in the 1870s. Subsequently, this species has been recorded in at least 27 publications, extending its stratigraphical range from the Recent back into the Upper Eocene, its zoogeographical range from Atlantic, Indian, Pacific, Arctic and Southern oceans, and its bathymetric range from the continental shelf (> 400 m) to deeper regions of the abyss (> 5000 m). Importantly, C. acanthoderma was erected the type species of the widespread and abundant, deep-sea genus Legitimocythere Coles and Whatley, 1989. For the present publication, all the specimens of Legitimocythere acanthoderma housed in the Natural History Museum in London were re-studied. Based on the type material and on previous publications, L. acanthoderma is revised, re-diagnosed and re-illustrated with scanning electron microscopy. Twenty-two specimens are, herein, as Henryhowella, Trachyleberididae gen. nov. and cf. Legitimocythere. Finally, comparisons based on illustrations from earlier publications show that only a few of the over 70 previous records (i.e., fossil and Recent samples) of L. acanthoderma could be confirmed; the remaining records either involved distinct species or genera or are dubious. As a consequence, the revised distribution of L. acanthoderma includes only Recent sediments from bathyal depths of the Subantarctic region of the Southern Ocean, and so it can no longer be considered to be cosmopolitan, eurybathic and to have existed for over 30 million years. © 2013 Elsevier Masson SAS. All rights reserved. Keywords: Geographical distribution; Biogeography; Cosmopolitanism; Legitimocythere acanthoderma; Taxonomic revision; Deep Sea

Résumé Cythere acanthoderma Brady a été décrite à partir des sédiments actuels de tous les océans du monde lors de l’expédition du HMS Challenger dans les années 1870. Depuis, 27 publications discutent de cette espèce, ce qui a permis d’accroître son extension stratigraphique de l’actuel jusqu’à l’Éocène supérieur, son extension zoogéographique de l’océan Atlantique, à l’Indien, au Pacifique, à l’Arctique et aux mers du Sud, et son extension bathymétrique du plateau continental (> 400 m) aux régions profondes des abysses (> 5000 m). C. acanthoderma est devenue l’espèce type du genre Legitimocythere Coles et Whatley, genre répandu, abondant, et habitant les eaux profondes. Pour la présente publication, tous les spécimens de L. acanthoderma conservés au musée d’histoire naturelle de Londres ont été réétudiés. En se basant sur le matériel type et sur les publications antérieures, L. acanthoderma est révisée, elle bénéficie d’un nouveau diagnostic et elle est nouvellement illustrée par microscopie électronique à balayage. Vingt-deux spécimens sont ici re-identifié comme Henryhowella, Trachyleberididae gen. nov. et cf. Legitimocythere. Enfin, les comparaisons basées sur les illustrations de publications antérieures montrent que seulement quelques-unes des plus de 70 occurrences précédentes (échantillons fossiles et récents) de L. acanthoderma ont pu être confirmées ; les enregistrements restants concernant soit des espèces ou genres distincts, soit étant douteux. Par conséquent, la répartition révisée des L. acanthoderma ne comprend que les sédiments récents de profondeurs bathyales de la région subantarctique de l’Océan Austral, et cette espèce ne peut donc plus être considérée comme cosmopolite, eurybathic et exister depuis plus de 30 millions d’années. © 2013 Elsevier Masson SAS. Tous droits réservés. Mots clés : Distribution géographique ; Biogéographie ; Cosmopolitanisme ; Legitimocythere acanthoderma ; Révision taxonomique ; Milieu marin profond

∗

Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. E-mail address: [email protected]

0035-1598/$ – see front matter © 2013 Elsevier Masson SAS. All rights reserved. http://dx.doi.org/10.1016/j.revmic.2012.11.001

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

1. Introduction Brady (1880) when studying the ostracods collected during the H.M.S. Challenger Expedition from 1873 to 1876 described “Cythere acanthoderma n. sp.” and 142 other novel species. Brady (1880) neither designated holotypes nor gave information on the specimens he had illustrated. Many of these new species, especially those from the deep-sea (e.g., Cythere dasyderma Brady, 1880, Cytheropteron mucronalatum Brady, 1880), he recorded from several distinct basins. However, subsequent taxonomic reviews have shown that of these cosmopolitan species were confused groups of distinct species (e.g., Maddocks, 1990; Jellinek et al., 2006; Brandão, 2008; Brandão and Yasuhara, in press; Brandão et al., in review). Brady (1880) recorded “C. acanthoderma n. sp.” from the Northern Atlantic (stations 64 and 73), the Southwestern Indian Ocean (station 146), the Indo-Pacific (station 191a), the Northern Central Pacific (station 246), and the Southeastern Pacific (stations 296 and 302) (Brady, 1880: pp. 104–105; Jellinek and Swanson, 2003: p. 33) (Fig. 1 and Table 1). The recorded depth ranges were also very wide, ranging from the shallow slope to deep abyssal depths (1061 to 5029 m). Moreover, Brady’s (1880: pl. 18, figs. 5a-e) drawings are schematic and it is not even possible to recognise which genera he had illustrated. This not only led to C. acanthoderma being recorded from all oceans, from the Upper Eocene to Recent, and from shelf to abyssal depths (Tables 1–3 and Fig. 1) but also to it being assigned to several genera, as noted by Jellinek and Swanson (2003): “Echinocythereis (Puri, 1954; Levinson, 1974), Henryhowella (Puri, 1971), Thalassocythere nomen nudum (Ducasse and Peypouquet, 1979; Malz, 1987), Trachyleberis (Ruan and Hao, 1989) and Legitimocythere (Coles and Whatley, 1989; Dingle and Lord, 1990)”. Below I discuss key publications on C. acanthoderma, for more detailed information see the taxonomic section and Tables 1–3. Bate (1963) reported on material of C. acanthoderma housed in the Natural History Museum (NHM) in London and in the Hancock Museum (Newcastle upon Tyne). Puri and Hulings (1976) designated as lectotype a single (unfortunately) juvenile left valve from the Southwestern Indian Ocean (Challenger station 146, 46◦ 46’S, 45◦ 31’E, 2515 m). They also re-sorted the sediment from the type locality (i.e., sample M-170, station 146) and found one complete carapace (NHM 1974.277) and one right valve (NHM 1974.276). But my re-examination of these latter three valves has shown that they actually belong to the genus Henryhowella (see taxonomic section). Coles and Whatley (1989) described the cosmopolitan, deepsea genus Legitimocythere and selected C. acanthoderma as its type species. But, as noted by Jellinek and Swanson (2003), they illustrated a distinct species from the Upper Oligocene sediments from the Northern Atlantic, which they misidentified as Legitimocythere acanthoderma. Dingle et al. (1990) subsequently assigned C. acanthoderma to the genus Trachyleberis and, based on the previous studies of Brady (1880), Benson et al. (1983) Ducasse and Peypouquet (1979) and Whatley and Coles (1987), stated that “Trachyleberis

3

acanthoderma” occurs in the Atlantic, Indian and Pacific oceans, from the Miocene to the Quaternary and from 1600 to 3884 m water depth. Jellinek and Swanson (2003: pp. 33–34) discussed the taxonomic confusion prevailing with Legitimocythere acanthoderma, and provided comparisons between what they believed to be the “real” L. acanthoderma and the specimens studied by previous authors. However, they did not study any specimens of L. acanthoderma and until then only three optical microscope photos of the juvenile lectotype were available (Puri and Hulings, 1976: pl. 1, figs. 13–18). Therefore, it is not clear where they obtained the information about the adult valve morphology of L. acanthoderma from. Consequently, the validity of their comparisons must be considered doubtful. Mazzini, 2005 (fig. 23) studied the type material of L. acanthoderma and proposed new diagnostic characters for the genus Legitimocythere. She sorted again the sediment of the original sample of Challenger’s station 146 (i.e., sediment sample M-170) and found two more adult right valves and one adult left valve. She also provided SEMs of the external, internal and dorsal views of the juvenile lectotype and one of the re-sorted adult right valves, and compared the morphology of L. acanthoderma with other Legitimocythere species. The purpose of this study is to complement Mazzini (2005) work on L. acanthoderma and provide: • a diagnosis, an emended description L. acanthoderma ‘sensu stricto’; • detailed illustrations of valves, i.e. left and right valves’ internal and external views, hinge, muscle scars, external ornamentation of L. acanthoderma ‘sensu stricto’ (Plates 1, 2); • an extensive synonymy of L. acanthoderma ‘sensu stricto’; • a revised geographical and stratigraphical distribution for this important deep-sea species (Fig. 1; Tables 1–3); • a full list of specimens and slides (included in the syntypes of L. acanthoderma), but herein re-identified as Henryhowella sp., cf. Legitimocythere sp., “Taracythere” sp. nov. aff. “T.” dasyderma and Trachyleberididae sp. (Table 3). • SEM photos of the taxa misidentified by Brady (1880) as C. acanthoderma (Plate 3).

2. Material and methods The specimens studied herein are all included in the ostracod Challenger collection from the Natural History Museum, London (NHM), mostly from the Zoology Department (for details see Table 3). Four of the 11 NHM slides had no NHM collection number and, following the curator’s (Miranda Lowe) advice, I added a “SNB” number (e.g., SNB-1 020, SNB-1 022, SNB-1 040, SNB-1 056b). These slides have also belatedly received a NHM collection number, i.e., NHM 2012.1468 (SNB-1 020); NHM 2012.1469 (SNB-1 022); NHM 2012.1470 (SNB-1 051); NHM 2012.1471 (SNB-1 040); NHM 2012.1467 (SNB-1 056b).

4

Table 1 Summary of previous records of “Cythere acanthoderma sensu lato”. Author’s Id

Id herein

Locality

Ocean

Latitude

Longitude

Station

Depth (m)

Depth (fms)

Date

Age

Comments

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 54-5126

425

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1.

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 3-5051

457

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 4-5056

600

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 4-5057

613

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 50-5120

747

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 49-5117

915

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 48-5116

1110

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 7-5062

1404

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 7-5061

1406

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 46-5114

1458

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1.

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

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Table 1 (Continued) Author’s Id

Id herein

Locality

Ocean

Latitude

Longitude

Station

Depth (m)

Depth (fms)

Date

Age

Comments

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 8-5064

1587

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 8-5063

1701

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 9-5065

1902

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 43-5111

2295

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 11-5068

2306

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 60-5136

2322

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

Legitimocythere acanthoderma (Brady, 1880)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 42-5110

2725

1971

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 41-5109

2970

–

Recent/SubfossilThe geographic coordinates are approximate and were extracted from Fig. 1 1971 Recent/Subfossil

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 58-5133

3150

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 12-5069

3277

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 13-5107

3566

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

The geographic coordinates are approximate and were extracted from Fig. 1

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

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6

Table 1 (Continued) Author’s Id

Id herein

Locality

Ocean

Latitude

Longitude

Station

Depth (m)

Depth (fms)

Date

Age

Comments

Ayress et al., 2004

Legitimocythere acanthoderma (Brady)

? (no illustration)

off Kerguelen and Heard Islands

Indian/Southern Ocean

53◦ S

72◦ E

Eltanin cruise 47, Stat. 39-5071

3554

–

1971

Recent/Subfossil

The geographic coordinates are approximate and were extracted from Fig. 1

Benson, 1977

“Thalassocythere” acanthoderma (Brady) “Thalassocythere” acanthoderma (Brady) Cythere acanthoderma n. sp. Cythere acanthoderma n. sp. Cythere acanthoderma n. sp.

? (no illustration)

São Paulo Plateau

Southern Atlantic

28◦ 17.22’S

41◦ 05.28’W

DSDP leg 39 Site 356

3175

–

16–21. 11.1974

Lower Miocene

? (no illustration)

Rio Grande Rise

Southern Atlantic

30◦ 00.25’S

35◦ 33.59’W

DSDP leg 39 Site 357

2086

–

23–28. 11.1974

Lower Miocene

material lost

–

Northern Atlantic

35◦ 35’N

50◦ 27’W

Challenger, #64

5029

2750

Recent

Gray ooze.

Henryhowella sp.

–

Northern Atlantic

38◦ 30’N

31◦ 14’W

Challenger, #73

1829

1000

Recent

Globigerina ooze

Legitimocythere acanthoderma (Brady, 1880), Henryhowella sp., “Taracythere” sp. nov. aff. “T.” dasyderma cf. Legitimocythere sp. (see comments)

East of Prince Edward’s Island

Southwestern Indian Ocean

46◦ 46’S

45◦ 31’E

Challenger, #146

2515

1375

29.12. 1873

Recent

Type locality. Globigerina ooze. Bottom temperature 1.5◦ C

off Ki Islands

Indo-Pacific

5◦ 26’S

133◦ 19’E

Challenger, #191a

1061

580

24.09; 1874

Recent

This slide is probably mislabeled with the station number 192, which is not listed by Brady (1880: 12-30) in his station and species list. However, on p. 23, Brady states that the species Cythere acanthoderma was found on station 191a. Therefore, I conclude that the above mentioned slide contains specimens from the station 191a

Benson, 1977

Brady, 1880

Brady, 1880

Brady, 1880

Brady, 1880

Cythere acanthoderma n. sp.

Brady, 1880

Cythere acanthoderma n. sp. Cythere acanthoderma n. sp.

material lost

abyss

Northern Central Pacific

36◦ 10’N

178◦ 0’E

Challenger, #246

3749

2050

02.07. 1875

Recent

Gray ooze. Bottom temperature 1.3◦ C

material lost

off Chile

Southeastern Pacific

38◦ 6’S

88◦ 2’W

Challenger, #296

3338

1825

09.11. 1875

Recent

Type locality and single record of Taracythere dasyderma s.s. Red clay. Bottom temperature 1.2◦ C

Brady, 1880

Cythere acanthoderma n. sp

material either lost or mislabelled as station 300

off Chile

Southeastern

42◦ 3’S

82◦ 11’W

Challenger, #302

2652

1450

28.12. 1875

Recent

Globigerina ooze. Bottom temperature 1.5◦ C

Brady, 1887

Cythere acanthoderma (Brady)

? (no illustration)

–

–

–

–

–

–

–

–

Recent

Brady, 1880

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Publication

Table 1 (Continued) Publication

Author’s Id

Id herein

Brady and Norman, 1889

Cythere acanthoderma, Brady Cythere acanthoderma, Brady Cythere acanthoderma, Brady

Trachyleberididae spp.

Brady and Norman, 1889 Brady and Norman, 1889

Coles and Whatley, 1989 Coles and Whatley, 1989 Cronin, 1996

Cronin, 1996

Didié and Bauch, 2000; Didié, 2001 Dingle and Lord, 1990 Ducasse and Peypouquet, 1979 Ducasse and Peypouquet, 1979 Finger et al., 2007 Finger et al., 2007 Finger et al., 2007 Guernet, 1998

Latitude

Longitude

Station

Depth (m)

Depth (fms)

Date

Age

Northern Atlantic

56◦ 11’N

37◦ 41’W

Valorous 1875, Stat 12

2652

1450

–

Recent

–

Northeastern Atlantic

56◦ 11’N

10◦ 56’W

Porcupine 1869, Stat. 19

2498

1366

–

Recent

Trachyleberididae spp

–

Northern Atlantic

55◦ 10’N

25◦ 58’W

Valorous 1875, Stat 16

3264

1785

–

Recent

? (no illustration)

North of St. Michael, Azores

Northern Atlantic

–

–

–

2295

“Thalassocythere” acanthoderma (Brady, 1880) “Thalassocythere” acanthoderma (Brady, 1880) Legitimocythere acanthoderma

? (no illustration)

–

Northern Atlantic

–

–

–

–

–

–

Upper Oligocene to Recent

? (no illustration)

–

Pacific

–

–

–

–

–

–

Middle Miocene to Quaternary

cf. Taracythere sp.

Goban Spur

Northeastern Atlantic

49◦ 05.29’N

13◦ 05.89’W

DSDP leg 80 Site 549 Hole 549A

2513

–

06.1981

Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880) Legitimocythere Acanthoderma (Brady) Legitimocythere Acanthoderma (Brady) Thalassocythere acanthoderma (Brady) Legitimocythere acanthoderma (Brady, 1880) Thalassocythere acanthoderma (Brady) Thalassocythere acanthoderma (Brady) Legitimocythere acanthoderma (Brady) Legitimocythere acanthoderma (Brady) Legitimocythere acanthoderma (Brady) Legitimocythere acanthoderma

cf. Taracythere sp.

west flank of the Mid-Atlantic Ridge

Northern Atlantic

37◦ 46.2’N

37◦ 20.61’W

DSDP leg 82 Site 558 Hole 558

3754

–

03–11. 10.1981

Upper Oligocene and Upper Miocene (Brady, 1880) Lower-Middle Miocene

cf. Taracythere sp.

west flank of the Mid-Atlantic Ridge

Northern Atlantic

33◦ 38.53’N

43◦ 46.04’W

DSDP leg 82 Site 563 Hole 563

3786

–

24–28. 10.1981

Middle-Upper Miocene

Legitimocythere sp. (maybe L.

Western flank of the Mid-Atlantic Ridge

Northern Atlantic

41◦ 00.07’N

32◦ 57.44’W

DSDP leg 94 Site 607

34272

–

09–11. 07.1983

Pliocene acanthoderma)

Legitimocythere sp. (maybe L. acanthoderma) Legitimocythere sp. (maybe L. acanthoderma) Legitimocythere sp. (maybe L. acanthoderma) cf. Legitimocythere sp., not L.

middle eastern flank of the Mid-Atlantic Ridge Rockall Plateau

Northern Atlantic

53◦ 13.30’N

18◦ 53.21’W

DSDP leg 94 Site 610

24172

–

28–31. 07.1983

Pliocene

Northeastern Atlantic

53◦ 32.2’N

20◦ 17.3’W

M23414-9

2196

Rockall Plateau

Northeastern Atlantic

55◦ 09’N

22◦ 45’W

–

2931

–

–

Quaternary

Northwestern Atlantic

56◦ 08.31’N

23◦ 17.64’W

DSDP leg 48 Site 403

2301

–

20–24. 06.1976

cf. Legitimocythere sp., not L. acanthoderma cf. Legitimocythere sp.

Southwestern margin of Rockall Plateau Southwestern margin of Rockall Plateau Punta Perro, Navidad

Northwestern Atlantic

55◦ 20.18’N

22◦ 03.49’W

DSDP leg 48 Site 405

2958

–

29.06–4.07 1976

Upper Miocene to Quaternary acanthoderma Quaternary

Southwestern Pacific

–

–

–

cf. Legitimocythere sp.

Punta Alta, Navidad

Southwestern Pacific

–

–

–

cf. Legitimocythere sp.

Punta el Fraile, Arauco

Southwestern Pacific

–

–

–

cf. Legitimocythere sp., not L. acanthoderma

Gulf of Guinea

Equatorial eastern Atlantic

03◦ 35.025’N

02◦ 43.990’W

ODP leg 159 Site 960 Hole 960C

Lower bathyal or deepera Lower bathyal or deepera Lower bathyal or deepera 2046.3

Comments

Either a “Travailleur” or a “Talisman” expedition in the 19th Century

Quaternary

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Coles and Whatley, 1989

Ocean

Trachyleberididae spp.

Brady and Norman, 1889

Coles, Ayress and Whatley, 1990 Coles et al., 1990

Locality

Lower Pliocene Lower Pliocene Lower Pliocene –

1995

Middle Miocene to Pliocene

7

8

Table 1 (Continued) Author’s Id

Id herein

Locality

Ocean

Latitude

Longitude

Station

Depth (m)

Guernet and Moullade, 1994 Guernet and Moullade, 1994 Guernet and Moullade, 1994 Guernet and Moullade, 1994 Majoran and Dingle, 2001

Legitimocythere cf. acanthoderma

cf. Legitimocythere sp., not L

Equatorial eastern Atlantic

Equatorial eastern Atlantic

18◦ 04.63’N

21◦ 01.57’W

ODP leg 108 Site 659 Hole 659A

3355.5

Legitimocythere cf. acanthoderma

Equatorial eastern Atlantic

Equatorial eastern Atlantic

18◦ 04.63’N

21◦ 01.57’W

ODP leg 108 Site 659 Hole 659B

3286

Equatorial eastern Atlantic

Equatorial eastern Atlantic

0◦ 06.44’N

23◦ 13.65’W

ODP leg 108 Site 664 Hole 664D

3807

Equatorial eastern Atlantic

Equatorial eastern Atlantic

4◦ 34.15’N

21◦ 54.68’W

ODP leg 108 Site 667 Hole 667A

3529

northeastern slope of the Northeast Georgia Rise

Atlantic Sector of the Southern Ocean

51◦ 32.537’S

30◦ 40.619’W

ODP leg 114 Site 699 Hole 699A

3716

–

Malz, 1987

“Thalassocythere” Acanthoderma (Brady, 1880) “Thalassocythere” Acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880) Legitimocythere Acanthoderma (Brady, 1880) Legitimocythere Acanthoderma (Brady, 1880) Cythere acanthoderma Brady

cf. Legitimocythere sp., not L.-acanthoderma cf. Legitimocythere sp., not L. acanthoderma cf. Legitimocythere sp., not L. acanthoderma cf. Legitimocy-Upper Eocene there sp., not L. acanthoderma Legitimocythere, sp. not L. acanthoderma Legitimocythere sp., not L. acanthoderma Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880)

Gulf of Aden

Northeastern Indian Ocean

13◦ 07.7’N

47◦ 56.6’E

Meteor station M5-263

2195

Rockall Plateau

Northeastern Atlantic

56◦ 17.41’N

23◦ 31.69’W

DSDP leg 81 Site 554

South Tasman Rise

Pacific/Southern Ocean

49◦ 10.84’S

150◦ 10.13’W

South Tasman Rise

Pacific/Southern Ocean

48◦ 29.99’S

South Tasman Rise

Pacific/Southern Ocean

East of Prince Edward’s Island

Malz, 1990

Mazzini, 2005

Mazzini, 2005

Mazzini, 2005

Puri and Hulings, 1976

Legitimocythere cf. acanthoderma Legitimocythere cf. acanthoderma Legitimocythere acanthoderma (Brady)

Depth (fms)

11–12. 03.1986

Date

Age

09–11. 03.1986

Miocene to Pliocene acanthoderma

Miocene to Pliocene 01–02. 04.1986. 03.1986 08–12. 04.1986. 03.1986 20–26. 03.1987

Miocene to Pliocene

–

–

Recent

2574

–

27-28. 08.1981

Pleistocene

TASQWA BX140

1636

–

–

Subfossil

149◦ 6.75’E

TASQWA BX147

2177

–

–

Subfossil

47◦ 46.85’S

149◦ 23.73’W

TASQWA BX153

1874

–

–

Subfossil

Indian Sector of the Southern Ocean

46◦ 46’0’S

45◦ 31’0’E

Challenger, #146

2515

1375

29.12. 1873

Recent

Lectotype designation. Type locality. Globigerina ooze. Bottom temperature 1.5◦ C. Not types, valves re-sorted by Ilaria Mazzini (pers. comm.) from the sediment of Challenger station 146 (sediment sample M-170)

Miocene to Pliocene

Puri and Hulings, 1976

Cythere acanthoderma Brady

Henryhowella sp.

East of Prince Edward’s Island

Indian Sector of the Southern Ocean

46◦ 46’0»S

45◦ 31’0»E

Challenger, #146

2515

1375

29.12. 1873

Recent

Steineck et al., 1988

“Thalassocythere” acanthoderma

off southern Hawaii

tropical central Pacific

–

–

DSDP leg 85

deep abyssal

–

1982

SylvesterBradley and Benson, 1971 Whatley, 1993

‘Cythere’ acanthoderma Brady Legitimocythere acanthoderma “Thalassocythere” acanthoderma (Brady)

cf. Legitimocythere, not L. acanthoderma Legitimocythere sp., not L. acanthoderma ? (no illustration)

Gulf of Mexico

Northwestern Atlantic

–

–

–

–

–

–

Upper Miocene to Quaternary (Brady, 1880) Recent

North Atlantic

Northern Atlantic

–

–

–

–

–

–

Lower Oligocene

Northern Atlantic

37◦ 20.32’N

35◦ 29.99’W

DSDP leg 94 Site 606

3022b

–

2–4.07. 1983

Lower Pliocene to Upper Quaternary

Whatley and Coles, 1987

cf. Legitimocythere sp.

Western flank of the Mid-Atlantic Ridge

Comments

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Publication

Table 1 (Continued) Author’s Id

Id herein

Locality

Ocean

Latitude

Longitude

Station

Depth (m)

Depth (fms)

Date

Age

Whatley and Coles, 1987

“Thalassocythere” acanthoderma (Brady) “Thalassocythere” acanthoderma (Brady) “Thalassocythere” acanthoderma (Brady) “Thalassocythere” acanthoderma (Brady) “Thalassocythere” acanthoderma (Brady) Legitimocythere acanthoderma Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady)

cf. Legitimocythere sp.

Western flank of the Mid-Atlantic Ridge

Northern Atlantic

41◦ 00.07’N

32◦ 57.44’W

DSDP leg 94 Site 607

3427b

–

9–11.07. 1983

Upper Miocene to Upper Quaternary

cf. Legitimocythere sp.

Northern Atlantic

42◦ 50.21’N

23◦ 20.25’W

DSDP leg 94 Site 608

3526b

–

13–17. 07.1983

Upper Pliocene to Upper Quaternary

Northern Atlantic

49◦ 52.67’N

24◦ 14.29’W

DSDP leg 94 Site 609

3884b

–

21–23. 07.1983

Lower Pliocene to Upper Quaternary

Northern Atlantic

53◦ 13.30’N

18◦ 53.21’W

DSDP leg 94 Site 610

2417b

–

28–31. 07.1983

Upper Miocene to Upper Quaternary

cf. Legitimocythere sp.

Southern flank of the King’s Trough tectonic complex middle eastern flank of the Mid-Atlantic Ridge middle eastern flank of the Mid-Atlantic Ridge southeastern flank of Gardar Ridg

Northern Atlantic

52◦ 50.47’N

30◦ 19.58’W

DSDP leg 94 Site 611

3195b

–

6–7.08. 1983

Upper Quaternary

? (no illustration)

North Atlantic

Northern Atlantic

–

–

–

–

–

–

? (no illustration)

off Kap Norvegica, Eastern Weddell Sea Meteor Rise

Southern Ocean

70◦ 36.80’S

13◦ 57.80’W

RV Polarstern, ARK I/2, PS1003-2

2796

–

1983

Upper Oligocene to Recent Late Quaternary

Southwestern Atlantic/Southern Ocean Northwestern Pacific

46◦ 52.8’S

7◦ 25.3’ E

ODP leg 114 Site 704 Hole 704A

2532

–

25–27. 04.1987

Quaternary

25◦ 12’N

123◦ 07’E

Core 255

1575

–

–

Upper Pleistocene to Upper Quaternary

Whatley and Coles, 1987 Whatley and Coles, 1987 Whatley and Coles, 1987 Whatley and Coles, 1987 Whatley and Coles, 1991 Whatley and Roberts, 1999 Yasuhara et al., 2009 Zhao et al., 2004

cf. Legitimocythere sp. cf. Legitimocythere sp.

Legitimocythere acanthoderma (Brady, 1880) cf. Legitimocythere, not L. acanthoderma

Okinawa Trough

Comments

560 cm gravity core

DSDP: Deep-Sea Drilling Project; ODP: Ocean Drilling Program. * In the NHM collection there is a MP slide (NHM 2012.1469, SNB-1 022) putatively mislabeled with the station number 192, which is not listed by Brady (1880: 12-30) in his station and species list. However, on p. 23, Brady states that the species Cythere acanthoderma was found on station 191a. Therefore, I conclude that the above mentioned slide contains specimens from the station 191a. a Depth estimated by the authors for the moment of deposition of the sediments which form the stratus. b Present day water depth.

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Publication

9

10

Table 2 Specimens assigned to “Cythere acanthoderma sensu lato” by subsequent authors. Id herein

Locality

Ocean

Latitude

Longitude

Station

Vessel

Depth (m)

Depth (fms)

Age

Brady (1866)

Cythere scabra, Von Münster (1830)

Trachyleberididae

Atlantic

Atlantic

–

–

–

–

3749,048

2050

Recent

Brady (1866)

Cythere scabra, Von Münster (1830)

Trachyleberididae

Abrolhos, Brazil

Atlantic

–

–

–

–

–

?

Recent

Brady (1866)

Cythere scabra, Von Münster (1830)

Trachyleberididae

Mediterranean Sea

–

–

–

–

658,3694

360

Recent

Dingle et al. (1990)

Trachyleberis sp. 3017

Southeastern Atlantic

–

–

–

–

Abyss

–

Quaternary

Tressler (1941)

Cythere ericea

Jellinek and Swanson, 2003: L. acanthoderma

cf. Legitimocythere sp., not L acanthoderma Legitimocythere sp. not L. acanthoderma

Creta, Mediterranean Sea off Southwestern Africa

Atlantic

49◦ 32’00”N

29◦ 21’00”W

Western Union cable ship Lord Kelvin

3250

–

?

Tressler (1941)

Cythere ericea

Jellinek and Swanson, 2003: acanthoderma

Legitimocythere sp. not L.L. acanthoderma

Atlantic

49◦ 37’00”N

13◦ 34’00”W

Core 7, sample H-series 48 (0-7 cm below core top) Core 12, sample H-series 130 (183-191 cm below core top)

Western Union cable ship Lord Kelvin

3230

–

?

Puri and Hulings, 1976: Cythere acanthoderma Puri and Hulings, 1976: Cythere acanthoderma Puri and Hulings, 1976: Cythere acanthoderma Jellinek and Swanson, 2003: L. acanthoderma

Faraday Hills, Mid-Atlantic Ridge, North Atlantic off Lands End, England, Northwestern Atlantic

Comments

Piggot deep-sea core sampler

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Additional specimens assigned to Legitimocythere acanthoderma Publication Publications author’s Subsequent Id authors’ Id

Table 3 Specimens studied herein. 1, specimens herein belong to the syntype series of Legitimocythere acanthoderma but are herein assigned to distinct taxa. Legitimocythere acanthoderma sensu stricto. Slide nr.

Previous slide nr.

Type

Material

Previously labelled as

Identification by Brady

Identification by other authors

Locality

Type of slide / vial

SEMs

Legitimocythere acanthoderma (Brady, 1880) Legitimocythere acanthoderma (Brady, 1880)

80.38.48.A.1

80.38.48.A

lectotype

1 LV

Cythere acanthoderma

Cythere acanthoderma

Cythere acanthoderma

Challenger, #146

white paper

SB212

2012.1467 (SNB-1 056b)

no number

no

1 LV, 2 coated RV

Cythere acanthoderma

-

Mazzini: Cythere acanthoderma

Challenger, #146

Squared, paper MP

SB953SB955, SB1000-1002, SB1010-1015, SB1043SB1050, SB1468SB1473

Henryhowella sp.

80.38.48.A.2 (SNB-1 023) 1974.276, 1974.277

80.38.48.A

1

1 RV

no

1 LV, 2 RV

Cythere acanthoderma -

Cythere acanthoderma -

Challenger, #146 Challenger, #146

white paperMP white plastic MP

SB0084

1974.276, 1974.277

Cythere acanthoderma Cythere acanthoderma

Henryhowella sp.

2012.1468 (SNB-1 020)

no number

no

2 LV, 7 RV

Cythere acanthoderma

Cythere acanthoderma

-

Challenger, #73

Henryhowella sp.

1961.12.4.50

1961.12.4.50

1

3 LV, 2 RV

Cythere acanthoderma

Cythere acanthoderma

cf. Legitimocythere sp.

2012.1469 (SNB-1 022)

New slide by SNB

1

1 RV

Cythere acanthoderma

Cythere acanthoderma

unknown author: Cythereis scraba (Münster) –

“Taracythere” sp. nov. aff. “T.” dasyderma Trachyleberididae sp.

80.38.49

80.38.49

1

2 LV

Cythere acanthoderma

Cythere acanthoderma

2012.1470 (SNB-1 051)

New slide by SNB

1

1 LV

Cythere acanthoderma, dasyderma dictyon

1900-3-6-234

1900-3-6-234

no

1911.11.8, M. 3308 2012.1471 (SNB-1 040)

1911.11.8, M. 3308 no number

no

5 LV, 4 RV, 1 RLV 9V

no

1 LV

Cythere acanthoderma Cythere acanthoderma Cythere acanthoderma

Henryhowella sp.

Trachyleberididae spp. Trachyleberididae spp. Trachyleberididae spp.

Comments

This valve was picked for Mazzini’s publication (2005) (pers. comm.). The slide is labelled “Cythere acanthoderma Brady (1+2), Challenger, n. 146, D. 1375”. One of the RV is probably the one illustrated by Mazzini (2005: fig. 23.C,D,F). Otherwise, the collection number given by this last author (i.e., OS 16001) is not written on the slide. Before SNB mislabeled this slide with “SNB-1 055”

SB001SB005, SB293-SB295

These valves was picked in the early 70s for Puri and Hulings’ (1976) publication, therefore it is not a type specimen

White plastic MP

SB0053-0069

These valves was picked in the early 70s for Puri and Hulings’ (1976) publication, therefore it is not a type specimen

Challenger, #300

Brady’s original MP

SB0046SB0051

Challenger, #191a

Black plastic MP by SNB

SB458

-

Challenger, #146

Brady’s original MP

SB959-SB963

Cythere acanthoderma

–

Challenger, #192

white paper MP by SNB

SNB0052

–

Cythere acanthoderma Cythere acanthoderma Cythere acanthoderma

Valorous 1875, Stat 12 Porcupine 1869, Stat. 19 Valorous 1875, Stat 16

Norman’s original MP Norman’s original MP Norman’s original MP

SB0060SB0064

– –

This MP slide is putatively mislabeled with the station number 192, which is not listed by Brady (1880: 12-30) in his station and species list. However, on p. 23, Brady states that the species Cythere acanthoderma was found on station 191a. Therefore, I conclude that this slide (i.e., NHM 2012.1469 SNB-1 022) contains specimens From the station 191a

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Identification herein

Before this RV was on the slide NHM 80.38.71A. 48A.75 (new number added “SNB-1 021”). This slide (i.e., NHM 2012.1470 SNB-1 051) was added to the NHM collection by SNB

LV, left valve; MP, micropalaeontological slide; RLV, closed carapace (i.e., articulated left and right valves); RV, right valve; SNB-1, slide number added by SNB to the Challenger ostracod collection in the NHM, mostly for slides previously without NHM collection number. SEMs, name of the digital files of the photos taken by SNB with Zeiss environmental SEM LEO 1455VP in the Natural History Museum, London. These photos are available from the author. 11

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S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

Fig. 1. Geographical distribution of Legitimocythere acanthoderma (Brady, 1880), and previous misidentifications of “Cythere acanthoderma” (see Table 1 for references). The map was made with the Ocean Data View 4 (Schlitzer, 2012).

Valves (mostly uncoated) were digitally imaged by SNB with Zeiss environmental SEM LEO 1455VP in the Natural History Museum, London. Abbreviations: DSDP: Deep-Sea Drilling Project; LV: left valve; MP: micropalaeontological slide; NHM: Zoology collection of the Natural History Museum, London; ODP: Ocean Drilling Program; RLV: closed carapace (i.e., articulated left and right valves); RV: right valve; SEM: Scanning Electron Microscope; SNB-1: slide number added by SNB to the Challenger ostracod collection in the NHM, mostly for slides previously without NHM collection number. 3. Results and discussion: challenging the deep-sea cosmopolitanism According to the present taxonomic revision (below), the distribution of L. acanthoderma ‘sensu stricto’, is restricted to Recent sediments from bathyal depths (1636 to 2931 m) of the Subantarctic region of the Southern Ocean (Fig. 1). This replaces the previous distribution from all oceans, and from geological epochs ranging from the Upper Eocene to the Recent, and bathymetrically from shelf to the abyss (341 to 3754 m) (for references see Table 1 and synonymy below). Most of the previous records of C. acanthoderma sensu lato have involved specimens of different species and even different genera. The specimens collected at the Challenger stations 73 (East of Prince Edward’s Island, Indian Sector of the Southern Ocean) and 146 (Northern Atlantic) clearly belong to Henryhowella. These specimens have a more sub-quadrate outline and much shorter spines than L. acanthoderma sensu stricto (Plates 1 and 3). The fossil specimens recorded from the Northern Atlantic by Coles and Whatley (1989: pl. 4, fig. 9) probably belong to Taracythere, since their anterior margins are more

broadly rounded than their posterior margins; and they have far fewer spines than is typical of Legitimocythere. Other specimens studied by other authors (e.g., Malz, 1987) do belong to the genus Legitimocythere, but are not co-specific with L. acanthoderma sensu stricto, they (for example) have more elongate carapaces (in lateral view), and longer and fewer spines (Malz, 1987: fig. 2b). Finally, an important inconsistency relates to the exceptionally wide ecological niche of L. acanthoderma as derived from all the former records (Table 1). Generally ostracods are important proxies for palaeoenvironmental reconstructions (e.g., Martens and Horne, 2000), because of their sensitivity to physico-chemical characteristics (temperature, salinity, oxygen content) of the water masses in the environments in which they live. The earlier studies of C. acanthoderma sensu lato imply an exceptionally broad range of environmental conditions ranging from starkly contrasting bathymetric zones, salinities, in situ temperatures, trophic regimes, etc. For example, Ayress et al. (2004) recorded L. acanthoderma from relatively shallow and eutrophic zones (Kletou and Hall-Spencer, 2012) around Kerguelen and Heard Islands, in the Subantarctic region of the Indian sector of the Southern Ocean. Several other authors recorded the same nominal species from nutrient poor abyssal basins in the Northern Atlantic (Brady, 1880; Brady and Norman, 1889; Coles and Whatley, 1989; Cronin, 1996; Didié and Bauch, 2000; Didié, 2001; Dingle and Lord, 1990; Ducasse and Peypouquet, 1979; Malz, 1990; Sylvester-Bradley and Benson, 1971; Whatley, 1993; Whatley and Coles, 1987, 1991) and in the extremely oligothrophic abyssal areas in the Pacific (Brady, 1880; Coles et al., 1990; Steineck et al., 1988). In order to describe accurately the ecological preferences (or ecological niche) of a specific taxon, it is necessary to have records, based on a broad geographical and temporal

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

13

Plate 1. Lectotype and “topotypes” of Legitimocythere acanthoderma (Brady, 1880). Photos: SNB and John Whittaker. Copyright: Natural History Museum, London. First numbers indicate single valves (i.e. Figs. 3.1 and 3.2 are distinct views of the same valve). Challenger station 146, 46◦ 46 S, 45◦ 31 E, 2515 m (= 1375 fathoms), 29.12.1873, Globigerina ooze, bottom temperature 1◦ 5 C. “Topotypes” sorted by I. Mazzini (personal communication) from the dried sediments of the sample from the Challenger Station 146 (NHM 2012.1467, SNB-1 056b). Figs. 1, 3.1. Right valve, external view; Fig. 2. Left valve, external view; Fig. 3.2. Right valve, internal view Lectotype. Fig. 4. (NHM 80.38.48.A.1), left valve: Fig. 4.1. External view; Fig. 4.2. External view; Fig. 4.3. Dorsal view; Fig. 4.4. Adductor muscle scars, internal view.

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Plate 2. “Topotypes” of Legitimocythere acanthoderma (Brady, 1880). Photos: SNB and John Whittaker. Copyright: Natural History Museum, London. First numbers indicate single valves (i.e. Figs. 1.1 to 1.4 are distinct views of the same valve). Challenger station 146, 46◦ 46 S, 45◦ 31 E, 2515 m (= 1375 fathoms), 29.12.1873, Globigerina ooze, bottom temperature 1◦ 5 C. “Topotype” sorted by I. Mazzini (personal communication) from the sediments of the sample from the Challenger Station 146 (NHM 2012.1467, SNB-1 056b). Figs. 1.1–1.4. Right valve; Figs. 1.1, 1.3. Details of anterior and posterior hinge teeth; Fig. 1.2. Internal view; Fig. 1.4. Dorsal view; Fig. 2. Left valve; Fig. 2.1. Internal view; Figs. 2.2–2.5. Dorsal view; Figs. 2.3–2.5. Details of spines and hinge.

S.N. Brandão / Revue de micropaléontologie 56 (2013) 2–19

15

Plate 3. Challenger specimens misidentified by Brady (1880). Photos: SNB. Copyright: Natural History Museum, London. Figs. 1, 2. Henryhowella sp., left valve external view; 1, Challenger station 73, 38◦ 30 N, 31◦ 14 W., 1829 m (1000 fathoms), Globigerina ooze; 2, Challenger station 146, 46◦ 46 S, 45◦ 31 E, 2515 m (= 1375 fathoms), 29.12.1873, Globigerina ooze, bottom temperature 1◦ 5 C. Fig. 3. Trachyleberididae sp., left valve external view, Challenger station 191a*, Depth 129 fathoms, off Ki Islands. * In the NHM collection there is a MP slide (NHM 2012.1469, SNB-1 022) putatively mislabeled with the station number 192, which is not listed by Brady (1880: 12-30) in his station and species list. However, on page 23, Brady states that the species Cythere acanthoderma was found on station 191a. Therefore, I conclude that the above mentioned slide contains specimens from the station 191a.

(e.g. monthly, seasonal, annual) range of samples. As shown in Table 1, only six among the 76 previously published records of C. acanthoderma sensu lato (i.e., 76 distinct samples where C. acanthoderma sensu lato was found) could be confirmed as certain. These six records originated from five studies (Ayress et al., 2004; Brady, 1880; Mazzini, 2005; Puri and Hulings, 1976; Yasuhara et al., 2009), and only one of which (the first) provides in situ physico-chemical measurements of the water masses overlying the sea floor. However, these physico-chemical measurements may not be truely representative of the environment in which Legitimocythere acanthoderma lives, since they were taken at least a few tens of meters above the sea floor and not in/on the sediment. However the benthic boundary layer (BBL) that overlying the sediment and is both isothermal and isohaline, varies in thickness from 10-100+ metres depending on the eddy structure of the overlying water column. Ayress et al. (2004: pl. 1, figs. 6-7; Table 1) recorded L. acanthoderma from 22 stations off Kerguelen and Heard Islands. They figure two valves collected from the station 42-5110, where the in situ temperature and the salinity were 0.95 ◦ C and 34.728 psu, respectively. In this revision, all the records from the remaining 21 stations have

been considered dubious, because these authors routinely use a morphological species concept, which is much wider than the one adopted herein. However, if all these 22 records from the vicinities of the Kerguelen and Heard Islands are considered certain, the bathymetric range of L. acanthoderma would be significantly extended (i.e., from 425 to 3584 m depth), compared to the bathyal range (1636 to 2931 m) stated in the first paragraph of this section. In addition, if all these 22 records are considered valid, the physico-chemical ranges of L. acanthoderma are greatly extended: temperature from 0.11 to 2.26 ◦ C, salinity from 34.435 to 34.767 psu, and oxygen concentrations (based on data from only 4 stations) from 4.89 to 5.34 ml/l; and phosphate concentrations (also available for only 4 stations) from 0.85 to 2.2 ␮mol/l. In conclusion, the ecological niche of L. acanthoderma s.s. would include: • relatively cool waters, but not as cold as the polar bottom waters, like Antarctic Bottom Water, but colder than other bathyal regions in temperate and tropical latitudes (e.g., North Pacific Intermediate Water, Red Sea Intermediate Water, Mediterranean Intermediate Water);

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• middle salinity values in relation to other intermediate water masses; • well oxygenated environments.

4. Taxonomy Remarks: In order to clarify the biogeography of L. acanthoderma ‘sensu stricto’, I include in the taxonomic sections below all specimens contained in the syntype series of “C. acanthoderma”, i.e. the following taxa: cf. Legitimocythere sp., Henryhowella spp. and Trachyleberididae gen. and sp. nov. Finally, the specimens identified by Brady and Norman (1889) as C. acanthoderma, which cannot be identified beyond the family level (i.e., Trachyleberididae spp.) are also listed below. Higher classification based on Horne et al. (2002). Classe OSTRACODA Latreille, 1806 Subclasse PODOCOPA Müller, 1894 Order PODOCOPIDA Sars, 1866 Suborder CYTHEROCOPINA Baird, 1850 Superfamily CYTHEROIDEA Baird, 1850 Family TRACHYLEBERIDIDAE Sylvester-Bradley, 1948 Genus Legitimocythere Coles and Whatley, 1989 Type species: C. acanthoderma Brady, 1880. Additional species: Cythereis bermudezi Bold, 1946 (assigned to Legitimocythere by Coles and Whatley, 1989); Trachyleberis bermudezi crebripustulosa Bold, 1966; Legitimocythere aculeata Jellinek and Swanson, 2003; Legitimocythere castanea Jellinek and Swanson, 2003; Legitimocythere geniculata Mazzini, 2005; Acanthocythereis? presequenta Benson, 1977 (assigned to Legitimocythere by Coles and Whatley, 1989). Legitimocythere acanthoderma (Brady, 1880) (Plates 1, 2, Fig. 1, Tables 1–3) 1880. in part C. acanthoderma Brady, pp. 104–105,?non pl. 18, figs. 5a-e. 1887.? Cythere acanthoderma (Brady), 165. 1889. not C. acanthoderma Brady – Brady and Norman, p. 151. 1889.? not C. acanthoderma Brady – Brady and Norman, p. 244. 1963. in part C. acanthoderma – Bate, p. 80, tab. 1. 1971. not ‘Cythere’ acanthoderma Brady – SylvesterBradley and Benson, p. 263, not fig. 16. 1976. in part C. acanthoderma – Puri and Hulings, pp. 267268, pl. 11.16–18. 1977.? “Thalassocythere” acanthoderma – Benson, p. 869. 1979. not Thalassocythere acanthoderma – Ducasse and Peypouquet, not figs 4, 6, not pl. 3, fig. 4. 1983. not “Thalassocythere” acanthoderma – Benson et al., pl. 2, fig. 9. 1987. not “Thalassocythere” acanthoderma (Brady) – Malz, fig. 2.b. 1987.? “Thalassocythere” acanthoderma (Brady) – Whatley and Coles,?figs. 1, 2B, 3-12B;?pl. 6, figs. 1-2; tab. 9.

1988. not “Thalassocythere” acanthoderma – Steineck et al., p. 602, not pl. 1.10. 1988.? “Thalassocythere” acanthoderma – Ayress, p. 836,?pl. 32, fig. 16. 1988 not Trachyleberis acanthoderma (Brady) – Ruan and Hao, p. 356, pl. 64, figs. 22–25. 1989 not Trachyleberis acanthoderma (Brady) – Ruan, p. 126, pl. 23, fig. 18. 1989. not L. acanthoderma (Brady, 1880) – Coles and Whatley, pp. 100-101, not pl. 4, fig. 9. 1990.? “Thalassocythere” acanthoderma (Brady, 1880) – Coles, Ayress and Whatley,?tab. 2. 1990.?not L. acanthoderma (Brady, 1880) – Dingle and Lord,?not fig. 2.11. 1990. not “Thalassocythere” acanthoderma (Brady, 1880) – Malz, not fig. 8.5–8.7. 1991.? L. acanthoderma (Brady, 1880) – Whatley and Coles,?fig. 4,? appendix 2. 1993.? L. acanthoderma – Whatley,?tab. 1. 1994. not Legitimocythere cf. acanthoderma (Brady, 1880) – Guernet and Moullade, 266, not pl. 3, figs. 12-13, 15. 1996.? L. acanthoderma – Cronin,?pl. 1, fig. 7,?tab. 1. 1998. not L. acanthoderma – Guernet, not pl. 2, fig. 2; not tab. 3–5. 1999.? L. acanthoderma (Brady, 1880) – Whatley and Roberts, 13. 2000.? Thalassocythere acanthoderma (Brady) – Didié and Bauch,? tab. 2,?non pl. 3, fig. 15, non pl. 3, fig. 16. 2001.? Thalassocythere acanthoderma (Brady) – Didié, p. 108, fig. 2.5,?non pl. 3, fig. 15, non pl. 3, fig. 16. 2001. not L. acanthoderma (Brady) – Majoran and Dingle, p. 214, not pl. 1, fig. 15. 2004. L. acanthoderma (Brady) – Ayress et al., pl. 1, fig. 6–7. 2004. not L. acanthoderma (Brady) – Zhao et al., not fig. 2.18. 2005. L. acanthoderma (Brady, 1880) – Mazzini, pp. 42-44, figs. 22.A-22.J, 23,?fig. 22.K-22.L. 2007. non L. acanthoderma (Brady) – Finger et al., fig. 14.C. 2009. L. acanthoderma (Brady, 1880) – Yasuhara et al., p. 922, figs. 5.5, fig. 11.1–11.6, tab. 1. Material examined: Lectotype: 1 juvenile LV on a white, paper MP labelled “173, Cythere acanthoderma Brady, H.S. Puri 7/67, Lectoholotype, Foto SB212, Challenger, No. 146, D. 1375, from trawl, 80.38.48A.1”. This valve was illustrated (with an optic camera) and designated as the lectotype by Puri and Hulings (1976: pl. 11, figs. 16–18). Mazzini (2005: fig. 23.A-C) provided SEM photos of this specimen. This lectotype is herein illustrated in Plate 1, Figs. 4(1-4.4). Type locality (from Brady, 1880: 16): Challenger station 146, 46◦ 46 S, 45◦ 31 E, 2515 m (= 1375 fathoms), Globigerina ooze, bottom temperature 1.5 ◦ C, 29th December 1873. Specimens (not types) later picked from the dried sediment (sample M-170) collected at the type locality (i.e., Challenger station 146): 1 LV, 2 coated RV on a squared micropaleontological slide previously without NHM collection number and labelled “C. acanthoderma Brady (1 + 2), Challenger, n. 146, D. 1375; SNB-1 056b”. This slide was labelled by Ilaria Mazzini (personal communication), who also illustrated

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one of the RV (Mazzini, 2005: fig. 23, D–F). However, the collection number given by Mazzini (i.e., OS 16001, a number from the Ostracoda collection within the Micropalaeontology collections in the Palaeontology Department of the NHM) is not written on this slide. I initially mislabeled this slide with the number SNB1 055, but it has been renumbered and labelled SNB-1 056b, and now bears the collection number NHM 2012.1467. These specimens are illustrated in Plate 1, Figs. 1–3 (1–3.2); Plate 2. Measurements: Lectotype (NHM 80.38.48.A.1): LV height 0.61 mm; length 0.99 mm. “Topotypes” (NHM 2012.1467, SNB-1 056b): LV height 0.66 mm; length 1.13 mm; LV height 0.68 mm; length 1.14 mm; RV height 0.70 mm; length 1.15 mm. Diagnosis: Suboval lateral outline, with anterior margin only slightly broader than posterior. Lateral surface covered by numerous spines of diverse forms and sizes. Conical, long spines almost exclusively restricted to marginal area and to ventral ridge. The longest conical spines located in the ventral ridge and posterior cardinal angle. Other spines mostly short, verrucose and multi-furcated. Apex of spines connected by a low ridge, forming a slight reticulation on valve surface, especially in the postero-ventral area. Description (modified after Mazzini, 2005): External view. Surface covered by small, verrucose tuberclelike (i.e., multi-furcate) spines, often furcate. Spines apex mostly jointed by a faint ridge forming a slight reticulation, which is weaker in the antero-dorsal area. Posterior with a thick marginal rim, with long, conical spines; one large, conical spine on dorsal edge. Anterior margin with a less pronounced rim and slightly thicker, conical marginal spines. Rows of small, often multifurcate tubercles on the inner margin on both rims. On the anterior, tubercles are larger and multi-furcate towards the ventral margin, where they fuse with the ventral ridge. Ventral ridge with six long, conical spines of which the posterior-most is the better developed. Dorsal margin with shorter, mostly conical spines. Around five spines on the sub-central tubercle. At an almost equal distance between the anterior margin and the sub-central tubercle, two larger and more pointed multi-furcate spines occur. Both tubercle-like and conical spines with simple pores. Ventral margin with a single row of spatulate spines. In dorsal view, the posterior rim is clearly separated from the domicilial swelling by a pronounced indentation; maximum width on the posterior half of the length. Internal view. Ventral margin concave around oral region; dorsal margin straight. Anterior margin more broadly rounded than posterior. Fused zone moderately wide, selvage robust. Marginal pore-canals straight with medial swelling, often paired; 29 to 32 anteriorly and 21 to 26 posteriorly. Hinge holamphidont, with smooth anterior, posterior and medial elements. Hinge of RV with large and stepped anterior tooth; large and sub-spherical posterior tooth; median groove widens at extremities. Anterior socket of hinge of left valve with a small adjacent tooth; median bar, smooth stout and thickened at both ends; fine crenulation present in the posterior half of the dorsal part of the median bar. Central adductor scars arranged in a row: two ventral-most scars small and almost fused; the upper median scar elongate; and dorsal-most scar inclined at 45◦ approximately with respect to the others. The

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frontal scar is oval with the main axis vertical. Fulcral point prominent. Revised stratigraphic, geographic and bathymetric occurrence (Table 1): Recent. Subantarctic region of the Southern Ocean, 1636 to 2931 m. Comparisons with other species assigned to Legitimocythere: Based on Bold’s (1946: pl. 10, fig. 18) schematic drawings, L. acanthoderma differs from L. bermudezi bermudezi and L. bermudezi crebripustulosa in the following characters: in lateral view, the last species shows anterior margin conspicuously more broadly rounded than the posterior margin; in dorsal view, the first species is wider posteriorly, while in the first species both anterior and posterior are similarly wide. Legitimocythere aculeata shows more oval lateral outline and longer and more numerous spines than L. acanthoderma (Jellinek and Swanson, 2003). In lateral view, the dorsal margin of the left valve of L. castanea and L. geniculata are conspicuously concave, while in L. acanthoderma it is fairly straight (Jellinek and Swanson, 2003; Mazzini, 2005). L. acanthoderma differs from L. presequenta in the following characters: in lateral view, the last species shows anterior margin conspicuously more broadly rounded than the posterior margin; the first species shows more robust but fewer spines than L. acanthoderma (Benson, 1977: pl. 2, fig. 5). Twenty-two subfossils (i.e., empty valves) paralectotypes of L. acanthoderma (Brady, 1880), are herein, reassigned to distinct taxa (see details in the taxonomic sections below and in Table 3): • 1 RV of cf. Legitimocythere sp. on the MP NHM 2012.1469 (SNB-1 022); • 1 RV of Henryhowella spp. on the MP NHM 80.38.48A.2 (SNB-1 023); • 1 LV and 2 RV of Henryhowella spp. on the MP NHM 1974.276-277; • 7 RV, 2 LV of Henryhowella spp. on the MP NHM 2012.1468 (SNB-1 020); • 2 RV, 3 LV of Henryhowella spp. on the MP NHM 1961.12.4.50; • 2 (possibly juvenile) LV of “Taracythere” sp. nov. on the MP NHM 80.38.49; • 1 LV of Trachyleberididae spp. on the MP NHM 2012.1470 (SNB-1 051). The following taxonomic sections are on the taxa of specimens previously included in the syntype series of “Cythere acanthoderma”. cf. Legitimocythere sp. (Tables 1 and 3) 1880. in part C. acanthoderma n. sp. Brady, pp. 104–105. Material examined: 1 RV on a MP labelled “Henryhowella cf. acanthoderma, Challenger #192, Depth 129 fathms., off Ki Islands, G.S. Brady 26/9/74, This valve was on slide with original Nr. = 80.38.71A.48A.75 (which is now labelled

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SNB-1 021). SNB-1 022”. NHM collection number 2012. 1469. Remarks: This right valve could not be further identified because it is decalcified. Genus Henryhowella Puri, 1957 Henryhowella spp. (Plate 3, Figs. 1, 2, Tables 1 and 3) 1880. in part C. acanthoderma n. sp. Brady, pp. 104–105. Material examined: 1 RV on a MP labelled “173, Henryhowella, Brady, Cythere acanthoderma Brady, H.S. Puri 7/967, Foto SB0084, Lectoparatype, 173, 28 T, SNB-1 023, “Challenger” No. 146, D. 1375, from trawl, 80.38.48A.2”. This specimen was studied by H.S. Puri in 1967 but it was not listed in Puri and Hulings (1976). 1 LV and 2 RV on a MP labelled “Cythere acanthoderma”, 1974.276-277, topotypes, 1974.276, 1974.277, Sediment sample M-170 (Stat. 146, 1375 fathoms, Lat. 46◦ 46 S, Long. 45◦ 31 E). Fotos SB0001-0005, ‘Challenger’ 29.12.1873”. These specimens were studied by H.S. Puri and listed in Puri and Hulings (1976). These specimens are herein illustrated in Plate 3, Fig. 2. 7 RV, 2 LV on a MP slide labelled “Recent, Challenger, St. 73, Lat 38◦ 30 N., Long 31◦ 14 W., 1000 fathoms, SNB-1 020, M95, fotos SB0053-0069, Henryhowella, Cythere acanthoderma Brady, n. sp.”. These specimens seem to have been studied by H.S. Puri but it was not listed in Puri and Hulings (1976). One left valve is herein illustrated in Plate 3, Fig. 1. 2 RV, 3 LV on a MP slide labelled “Cythere acanthoderma Brady, 1961.12.4.50, Cythereis scraba (Münster), Henryhowella, fotos SB0046-0051, “Challenger”, No. 300, Depth 1375 fathoms, G.S. Brady, 179”. Remarks: These eighteen valves could not be identified to species level because their surfaces are eroded (Plate 3, Figs. 1, 2). Genus Trachyleberididae gen. nov. Trachyleberididae gen. nov. and sp. nov. 1880. in part C. acanthoderma n. sp. Brady, pp. 104-105. in press. Trachyleberididae gen. nov. and sp. nov. Brandão and Yasuhara, pl. 2, figs. a-b. Material examined: 2 (possibly juvenile) LV on a MP originally used by Brady and labelled “Cythere dasy acanthoderma Brady, 80.38.49, Type, “CHALLENGER”, No. 146, depth 1375 fathoms, from trawl, G. S. Brady, 173”. These valves were illustrated by Brandão and Yasuhara, in press (pl. 2, figs. A-b). Remarks: See Brandão and Yasuhara (in press) for taxonomic diagnosis, illustrations and remarks on this material. Trachyleberididae spp. (Plate 3, Fig. 3, Tables 1 and 3) 1889. C. acanthoderma Brady - Brady and Norman, p. 244. Material examined: 1 LV was on a MP slide originally used by Brady (now labelled 80.38.71A.48A.75), is now on a new

white MP slide added to the NHM collection by myself and labelled NHM 2012.1470 “Challenger # [sic] 192, Depth 129 fathoms, off Ki Islands, (SNB-1 051), this valve was on slide 80.38.71A.48A.75”. This slide is probably mislabeled with the station number 192, which is not listed by Brady (1880: 12-30) in his station and species list. However, on page 23, Brady states that the species C. acanthoderma was found on station 191a. Therefore, I conclude that the above mentioned slide contains specimens from the station 191a. This valve is herein illustrated in Plate 3, Fig. 3. 4 RV, 5 LV, 1 RLV on a MP slide labelled “C. acanthoderma G.S. Brady, 1900-3-6-234, Valorous 1875, Stat 12, Lat 56◦ 11 N, Long 37◦ 41 W, 1450 fathoms, 177”. 5 RV, 4 LV on a MP slide labelled “C. acanthoderma Brady, NORMAN COLLECTION, 1911.11.8, M. 3308, Porcupine 1869, Stat. 19, Lat 56◦ 11 N, Long 10◦ 56 W, 1366 fathoms, 173”. 1 LV on a MP slide labelled “C. acanthoderma Brady, Valorous 1875, Stat 16, 1785 fathoms, 173, SNB-1 040”. Remarks: These valves could not be further identified because they are decalcified and covered by sediment and glue. Acknowledgements Miranda Lowe (NHM, London) carefully assisted me during five visits to the NHM and also kindly provided access to and permission for studying the Challenger and other ostracod specimens housed in the NHM. Martin V. Angel suggested revisions to the English. “This research received support from the SYNTHESYS Project http://www.synthesys.info/ which is financed by the European Community Research Infrastructure Action under the FP7 Integrating Activities Programme.” The author thanks the Alexander von Humboldt Foundation for the PostDoctoral Fellowship; and Angelika Brandt (ZMH, Universität Hamburg) and Pedro Martínez Arbizú (DZMB, Senckenberg Institute) for mentorship. Ilaria Mazzini and John E. Whittaker provided information and SEM photos from Challenger specimens taken from John E. Whittaker. Vincent Perrier translated the title and abstract into French. Cristianini T. Bergue, Moriaki Yasuhara and Dayou Zhai provided publications. This study was also financially supported by the Encyclopedia of Life and the Hansische Universitäts-Stiftung. Constructive remarks by Taniel Danelian, Thomas Cronin (USGS) and an anonymous reviewer are gratefully acknowledged. References Ayress, M.A., De Deckker, P., Coles, G.P., 2004. A taxonomic and distributional survey of marine benthonic Ostracoda off Kerguelen and Heard Islands, South Indian Ocean. Journal of Micropalaeontology 23, 15–38. Bate, R.H., 1963. The Ostracoda collected during the Voyage of H.M.S Challenger. Micropaleontology 9, 79–84. Benson, R.H., 1977. The Cenozoic ostracode faunas of the São Paulo Plateau and the Rio Grande Rise (DSDP leg. 39, Sites 356 and 357). In: Supko, P.R., Perch-Nielsen, K. (Eds.), Initial Reports of the Deep Sea Drilling Project 39. U.S Government Printing Office, Washington, pp. 869–883. Benson, R.H., DelGrosso, R.M., Steineck, P.L., 1983. Ostracode distribution and biofacies. Newfoundland continental slope and rise. Micropaleontology 29, 430–453.

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