Late Quaternary deep-sea ostracod taxonomy of the eastern North Atlantic Ocean

Abstract

Taxonomic revision and re-evaluation of the eastern North Atlantic deep-sea ostracods are conducted based on late Quaternary sediments from Ocean Drilling Program (ODP) Hole 982A, Rockall Plateau, eastern North Atlantic. Twenty-one genera and 51 species were examined and (re-)illustrated with highresolution scanning electron microscopy images. Six new species are described: Polycope lunaris, Argilloecia labri, Bythoceratina nuda, Cytheropteron colesoabyssorum, Cytheropteron colesopunctatum and Cytheropteron paramediotumidum. Excellent fossil ostracod preservation in this sediment core enabled us to provide a robust taxonomic baseline of the eastern North Atlantic deep-sea ostracods for application to palaeoceanographical, palaeoecological and biogeographical studies.

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Journal of Micropalaeontology Online First
10.1144/jmpaleo2013-022 , first published July 1, 2014; doiJournal of Micropalaeontology
Moriaki Yasuhara and Hisayo Okahashi
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Late Quaternary deep-sea ostracod taxonomy of the eastern North Atlantic
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Journal of Micropalaeontology. http://dx.doi.org/10.1144/jmpaleo2013-022
Published Online First
© 2014 The Micropalaeontological Society
INTRODUCTION
North Atlantic deep-sea ostracods have been well investigated
(e.g. Brady, 1880; Whatley & Coles, 1987; Coles & Whatley,
1989; Dingle & Lord, 1990; Cronin & Raymo, 1997; Cronin
et al., 1999; Didié et al., 2002; Yasuhara & Cronin, 2008;
Yasuhara et al., 2008, 2009a; Alvarez Zarikian, 2009; Yamaguchi
& Norris, 2012). However, detailed taxonomic studies using the
scanning electron microscope (SEM) are still limited and concen-
trated on the ostracod faunas from the lower bathyal and abyssal
zones (i.e. >2000 m water depth) (Whatley & Coles, 1987; Coles
& Whatley, 1989; Coles et al., 1994; Alvarez Zarikian, 2009).
Thus, little is known on the bathyal North Atlantic ostracod tax-
onomy, compared to well-investigated Mediterranean bathyal
fauna (Bonaduce et al., 1976; Colalongo & Pasini, 1980; Aiello
et al., 2000; Guernet, 2005; Aiello & Barra, 2010), even though
bathyal faunas are usually much more diverse than abyssal fau-
nas (e.g. Yasuhara et al., 2012). Furthermore, there is some taxo-
nomic confusion in North Atlantic ostracod taxonomic, faunal
and palaeoceanographic studies in which a same species has
often been called by several different names (see synonymy lists
in the present study).
Recently, Yasuhara et al. (2009b) conducted a comprehensive
taxonomic revision of western North Atlantic Quaternary
deep-sea ostracods using a sediment core taken from the upper
bathyal zone with high-resolution SEM images of 87 species and
a detailed literature survey. However, a comparable in-depth taxo-
nomic study has not been undertaken previously for the upper
bathyal zone of the eastern North Atlantic.
Ocean Drilling Program (ODP) Hole 982A gave us an ideal
opportunity to study eastern North Atlantic deep-sea ostracods from
the upper bathyal zone in detail, because its sediments have an
abundant, diverse and well-preserved ostracod fauna. Here we
investigate late Quaternary ODP 982A ostracod taxonomy using
high-resolution SEM images to reduce taxonomic confusion of
North Atlantic bathyal ostracods. In addition, we briefly discuss
similarity of bathyal ostracod faunas among the western and eastern
North Atlantic, the Mediterranean, and the western North Pacific.
MATERIALS AND METHODS
A total of 47 samples of ODP Hole 982A (57°30.992′N,
15°52.001′W, 1135.3 m water depth; Rockall Plateau, eastern
North Atlantic) covering the past 230 000 years and Marine
Isotope Stages (MIS) 1–7 (Venz et al., 1999) were examined for
ostracod taxonomy. The full information for the samples and
specimens used for the present study is shown in Tables 1 and 2.
Uncoated specimens were digitally imaged with a Philips XL-30
environmental SEM. High-resolution figures of ostracod SEM
images (Figs 2–16) are available at Dryad (http://datadryad.org/;
http://doi.org/10.5061/dryad.sc193. We follow the higher classifi-
cation scheme of the World Register of Marine Species (WoRMS:
http://www.marinespecies.org/) with certain modifications.
Repository. Figured specimens are deposited in the National
Museum of Natural History (Washington DC, catalogue numbers
USNM 603625–USNM 603760).
Abbreviations. LV, left valve; RV, right valve; L, length (mm);
H, height (mm).
SYSTEMATIC PALAEONTOLOGY
Class Ostracoda Latreille, 1802
Subclass Myodocopa Sars, 1866
Order Halocyprida Dana, 1853
Suborder Cladocopina Sars, 1866
Superfamily Polycopidea Sars, 1866
Family Polycopidae Sars, 1866
Genus Polycope Sars, 1866
Type species. Polycope orbicularis Sars, 1866
Remarks. We use the genus name Polycope in a broad sense fol-
lowing typical deep-sea ostracod taxonomy, but note that recent
zoological studies, for example Karanovic & Brandão (2012),
divide the genus into several separate genera based on soft parts,
which are not preserved in fossil ostracods.
Polycope martinezi (Karanovic & Brandão, 2012)
(Fig. 2A)
2001 Polycope sp. cf. P. arcys Joy & Clark; Didié & Bauch: 104,
pl. 1, fig. 28 (as erratum for Didié & Bauch, 2000).
Late Quaternary deep-sea ostracod taxonomy of the eastern North Atlantic Ocean
MORIAKI YASUHARA* & HISAYO OKAHASHI
School of Biological Sciences, Swire Institute of Marine Science, and Department of Earth Sciences,
University of Hong Kong, Kadoorie Biological Sciences Building, Pokfulam Road, Hong Kong SAR, China
*Corresponding author (e-mail: yasuhara@hku.hk)
ABSTRACT – Taxonomic revision and re-evaluation of the eastern North Atlantic deep-sea ostracods are
conducted based on late Quaternary sediments from Ocean Drilling Program (ODP) Hole 982A, Rockall Pla-
teau, eastern North Atlantic. Twenty-one genera and 51 species were examined and (re-)illustrated with high-
resolution scanning electron microscopy images. Six new species are described: Polycope lunaris, Argilloecia
labri, Bythoceratina nuda, Cytheropteron colesoabyssorum, Cytheropteron colesopunctatum and Cytherop-
teron paramediotumidum. Excellent fossil ostracod preservation in this sediment core enabled us to provide
a robust taxonomic baseline of the eastern North Atlantic deep-sea ostracods for application to palaeoceano-
graphical, palaeoecological and biogeographical studies.
KEYWORDS: deep-sea, Ostracoda, taxonomy, Quaternary, eastern North Atlantic, upper bathyal
research-articlePapersXXX10.1144/jmpaleo2013-022M. Yasuhara & H. OkahashiLate Quaternary deep-sea ostracods, North Atlantic
2014
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M. Yasuhara & H. Okahashi
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Table 1. Detailed information of the specimens used for the present study.
USNM No. Species T V Instar Hole Section Figure
603625 ODP982154poly Polycope martinezi L ? 982A 1/3/2–4 2A
603626 ODP982161 Polycope lunaris sp. nov. H L? A? 982A 1/3/42–44 2B
603627 ODP982162 Polycope lunaris sp. nov. P R? A? 982A 1/3/42–44 2C
603628 ODP982155poly Polycope cf. bireticulata L ? 982A 1/3/32–34 2D
603629 ODP982156poly Polycope cf. bireticulata R ? 982A 1/3/32–34 2E
603630 ODP982157poly Polycope orbicularis s.l. R ? 982A 1/3/112–114 2F
603631 ODP982158poly Polycope orbicularis s.l. L ? 982A 1/3/112–114 2G
603632 ODP982159poly Polycope vasfiensis L A? 982A 1/2/27–29 2H
603633 ODP982160poly Polycope vasfiensis R A? 982A 1/2/27–29 2I
603634 ODP982163poly Polycope reticulata L ? 982A 1/3/32–34 2J
603635 ODP982164poly Polycope reticulata R ? 982A 1/3/32–34 2K
603636 ODP982075 Cytherella robusta L J 982A 1/2/107–109 3A
603637 ODP982076 Cytherella robusta R J 982A 1/2/107–109 3B
603638 ODP982066 Bairdoppilata conformis L A 982A 1/2/67–69 3C, D
603639 ODP982067 Bairdoppilata conformis R A 982A 1/3/82–84 3E, F
603640 ODP982124 Macrocyprissa arcuata R A 982A 1/1/97–99 3G
603641 ODP982126 Macrocyprissa arcuata R A 982A 1/3/102–104 3H
603642 ODP982125 Macrocyprissa arcuata L A 982A 1/3/102–104 3I
603643 ODP982062 Argilloecia acuminata L A 982A 1/1/60–62 4A
603644 ODP982063 Argilloecia acuminata R A 982A 1/1/60–62 4B
603645 ODP982064 Argilloecia acuminata L A 982A 1/1/50–52 4C
603646 ODP982065 Argilloecia acuminata R A 982A 1/1/50–52 4D
603647 ODP982052 Argilloecia caju L A 982A 1/2/17–19 4E
603648 ODP982053 Argilloecia caju R A 982A 1/2/17–19 4F
603649 ODP982058 Argilloecia labri sp. nov. P L A 982A 1/3/102–104 4G
603650 ODP982059 Argilloecia labri sp. nov. P R A 982A 1/3/102–104 4H
603651 ODP982060 Argilloecia labri sp. nov. H L A 982A 1/1/50–52 4I
603652 ODP982061 Argilloecia labri sp. nov. P R A 982A 1/3/82–84 4J
603653 ODP982054 Argilloecia bensoni R A 982A 1/1/97–99 4K
603654 ODP982057 Argilloecia bensoni L A 982A 1/1/142–144 4L
603655 ODP982056 Argilloecia bensoni R A 982A 1/3/112–114 4M
603656 ODP982055 Argilloecia bensoni L A 982A 1/1/107–109 4N
603657 ODP982165prop Propontocypris acuminata L J? 982A 1/2/37–39 5A
603658 ODP982166prop Propontocypris sp. L J? 982A 1/1/60–62 5B
603659 ODP982072 Bythoceratina scaberrima L A 982A 1/2/67–69 5C
603660 ODP982073 Bythoceratina nuda sp. nov. H L A 982A 1/3/52–54 5D
603661 ODP982074 Bythoceratina nuda sp. nov. P R A 982A 1/3/42–44 5E
603662 ODP982167pseu Pseudocythere caudata L A? 982A 1/3/42–44 5F
603663 ODP982168pseu Pseudocythere caudata R A? 982A 1/3/42–44 5G
603664 ODP982131 Paijenborchella cymbula R A 982A 1/1/97–99 5H
603665 ODP982132 Paijenborchella cymbula L A 982A 1/1/40–42 5I
603666 ODP982133 Paijenborchella cymbula R A 982A 1/1/40–42 5J
603667 ODP982134 Paijenborchella cymbula L A 982A 1/1/40–42 5K
603668 ODP982048 Aversovalva hydrodynamica R A 982A 1/1/107–109 6A
603669 ODP982049 Aversovalva hydrodynamica L A 982A 1/1/107–109 6B
603670 ODP982050 Aversovalva hydrodynamica L A 982A 1/1/107–109 6C
603671 ODP982051 Aversovalva hydrodynamica R A 982A 1/1/117–119 6D
603672 ODP982042 Cytheropteron aielloi L A 982A 1/3/42–44 6E
603673 ODP982043 Cytheropteron aielloi R A 982A 1/3/42–44 6F
603674 ODP982044 Cytheropteron aielloi R A 982A 1/3/42–44 6G
603675 ODP982084 Cytheropteron alatum R A 982A 1/3/112–114 6H
603676 ODP982077 Cytheropteron colesoabyssorum sp. nov. H R A 982A 1/2/37–39 7A
603677 ODP982082 Cytheropteron colesopunctatum sp. nov. H R A 982A 1/1/30–32 7B
603678 ODP982083 Cytheropteron colesopunctatum sp. nov. P L A 982A 1/1/30–32 7C
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USNM No. Species T V Instar Hole Section Figure
603679 ODP982085 Cytheropteron didieae R A 982A 1/1/50–52 7D
603680 ODP982086 Cytheropteron didieae R A 982A 1/1/60–62 7E
603681 ODP982087 Cytheropteron didieae R A 982A 1/1/60–62 7F
603682 ODP982088 Cytheropteron didieae L A 982A 1/1/60–62 7G
603683 ODP982089 Cytheropteron didieae L A 982A 1/2/127–129 7H
603684 ODP982035 Cytheropteron fugu L A 982A 1/2/67–69 7I
603685 ODP982036 Cytheropteron fugu L A 982A 1/2/67–69 7J
603686 ODP982037 Cytheropteron fugu R A 982A 1/2/57–59 8A
603687 ODP982038 Cytheropteron fugu R A 982A 1/2/57–59 8B
603688 ODP982040 Cytheropteron omega L A 982A 1/4/12–14 8C
603689 ODP982039 Cytheropteron omega R A 982A 1/4/12–14 8D
603690 ODP982041 Cytheropteron omega R A 982A 1/3/132–134 8E
603691 ODP982045 Cytheropteron omega L A 982A 1/2/127–129 8F
603692 ODP982080 Cytheropteron inornatum L A 982A 1/1/97–99 8G
603693 ODP982079 Cytheropteron inornatum R A 982A 1/2/17–19 8H
603694 ODP982081 Cytheropteron inornatum R A 982A 1/1/97–99 8I
603695 ODP982078 Cytheropteron inornatum L A 982A 1/2/17–19 8J
603696 ODP982047 Cytheropteron massoni L A 982A 1/1/117–119 8K
603697 ODP982046 Cytheropteron massoni R A 982A 1/2/17–19 8L
603698 ODP982092 Cytheropteron paramediotumidum sp. nov. H L A 982A 1/2/107–109 9A
603699 ODP982093 Cytheropteron paramediotumidum sp. nov. P R A 982A 1/2/127–129 9B
603700 ODP982090 Cytheropteron demenocali L A 982A 1/1/137–139 9C
603701 ODP982091 Cytheropteron demenocali R A 982A 1/2/17–19 9D
603702 ODP982095 Cytheropteron pararhombiformis L A 982A 1/1/30–32 9E
603703 ODP982096 Cytheropteron pararhombiformis R A 982A 1/1/40–42 9F
603704 ODP982097 Cytheropteron paucipunctatum L A 982A 1/2/127–129 9G
603705 ODP982098 Cytheropteron paucipunctatum R A 982A 1/3/52–54 9H
603706 ODP982033 Cytheropteron perlaria L A 982A 1/2/17–19 10A
603707 ODP982034 Cytheropteron perlaria R A 982A 1/2/17–19 10B
603708 ODP982032 Cytheropteron pherozigzag L A 982A 1/1/142–144 10C
603709 ODP982031 Cytheropteron pherozigzag R A 982A 1/2/17–19 10D
603710 ODP982094 Cytheropteron pseudoalatum L A 982A 1/2/127–129 10E
603711 ODP982100 Eucytherura calabra L A 982A 1/1/70–72 10F
603712 ODP982101 Eucytherura calabra L A 982A 1/1/90–92 10G
603713 ODP982102 Eucytherura calabra R A 982A 1/1/107–109 10H
603714 ODP982004 Eucytherura multituberculata R A 982A 1/3/112–114 10I
603715 ODP982005 Eucytherura multituberculata L A 982A 1/3/92–94 10J
603716 ODP982001 Eucytherura tetrapteron L A 982A 1/3/112–114 11A
603717 ODP982002 Eucytherura tetrapteron R A 982A 1/3/112–114 11B
603718 ODP982003 Eucytherura tetrapteron L A 982A 1/3/112–114 11C
603719 ODP982104 Kangarina abyssicola L A 982A 1/1/137–139 11D
603720 ODP982103 Kangarina abyssicola R A 982A 1/1/70–72 11E
603721 ODP982136 Pedicythere atroposopetasi L A 982A 1/3/82–84 11F, G
603722 ODP982137 Pedicythere atroposopetasi R A 982A 1/3/82–84 11H, I
603723 ODP982138 Pedicythere atroposopetasi R A 982A 1/1/70–72 12A
603724 ODP982139 Pedicythere atroposopetasi L A 982A 1/3/92–94 12B
603725 ODP982146 Pedicythere atroposopetasi R A 982A 1/1/142–144 12C, D
603726 ODP982140 Pedicythere lachesisopetasi R A 982A 1/1/97–99 12E, F
603727 ODP982141 Pedicythere lachesisopetasi L A 982A 1/3/122–124 12G, H
603728 ODP982142 Pedicythere lachesisopetasi R A 982A 1/3/122–124 12I, J
603729 ODP982144 Pedicythere lachesisopetasi R A 982A 1/3/122–124 13A
603730 ODP982145 Pedicythere lachesisopetasi L A 982A 1/1/60–62 13B
603731 ODP982143 Pedicythere lachesisopetasi L A 982A 1/3/122–124 13C, D
603732 ODP982147 Pedicythere kennettopetasi L A 982A 1/3/72–74 13E, F
Table 1. (Continued)
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USNM No. Species T V Instar Hole Section Figure
603733 ODP982148 Pedicythere klothopetasi L A 982A 1/1/80–82 13G
603734 ODP982149pedi Pedicythere klothopetasi R A 982A 1/1/80–82 13H
603735 ODP982150 Pedicythere klothopetasi R A 982A 1/1/80–82 13I, J
603736 ODP982151 Pedicythere klothopetasi R A 982A 1/1/80–82 14A
603737 ODP982152 Pedicythere klothopetasi L A 982A 1/1/70–72 14B, C
603738 ODP982153 Pedicythere klothopetasi R A 982A 1/1/70–72 14D, E
603739 ODP982099 Eucythere triangula L A 982A 1/2/117–119 14F
603740 ODP982127-2 Cluthia sp. L A 982A 1/3/12–14 14G
603741 ODP982128 Cluthia sp. R A 982A 1/3/92–94 14H
603742 ODP982129-1 Cluthia sp. L A 982A 1/3/92–94 14I
603743 ODP982130 Cluthia sp. R A 982A 1/3/92–94 14J
603744 ODP982123 Loxoconchidea minima L A 982A 1/3/12–14 15A
603745 ODP982135 Paracytherois bondi R A 982A 1/1/80–82 15B, C
603746 ODP982169rock Arcacythere enigmatica L A 982A 1/2/77–79 15D
603747 ODP982170rock Arcacythere enigmatica R A 982A 1/2/77–79 15E
603748 ODP982171rock Arcacythere enigmatica L A 982A 1/2/77–79 15F
603749 ODP982172rock Arcacythere enigmatica R A 982A 1/2/77–79 15G
603750 ODP982173rock Arcacythere enigmatica R A 982A 1/2/67–69 15H
603751 ODP982174rock Arcacythere enigmatica L A 982A 1/3/92–94 15I
603752 ODP982027 Echinocythereis echinata R A 982A 1/1/142–144 16A
603753 ODP982028 Echinocythereis echinata R A 982A 1/1/142–144 16B
603754 ODP982029 Echinocythereis echinata L A 982A 1/3/72–74 16C
603755 ODP982030 Echinocythereis echinata L A 982A 1/3/72–74 16D
603756 ODP982022 Henryhowella asperrima L A 982A 1/1/137–139 16E
603757 ODP982023 Henryhowella asperrima R A 982A 1/1/137–139 16F
603758 ODP982026 Henryhowella asperrima L A 982A 1/1/0–2 16G
603759 ODP982024 Henryhowella asperrima R A 982A 1/1/142–144 16H, I
603760 ODP982025 Henryhowella asperrima L A 982A 1/1/107–109 16J, K
All specimens from late Quaternary sediments. Core samples are specified by standard ODP notation (core/section/interval). USNM, catalog number;
No., M.Y.’s personal catalog number. T, type (P, paratype; H, holotype); V, valve (L, left; R, right); A, adult; J, juvenile.
2009b Polycope arcys Joy & Clark; Yasuhara et al.: 881, pl. 1, fig. 6.
2012 Archypolycope martinezi Karanovic & Brandão: 348, figs
20–24.
Remarks. Polycope martinezi is very similar to the Arctic species
Polycope arcys, but distinguished by lacking obvious lateral
spines and having finer reticulation. P. martinezi was originally
reported from the equatorial Atlantic (Karanovic & Brandão,
2012) and is also known from the North Atlantic (Didié & Bauch,
2000, 2001; Yasuhara et al. 2009b).
Polycope lunaris sp. nov.
(Fig. 2B–C)
2001 Polycope sp. Didié & Bauch: 103, pl. 1, fig. 27 (as erratum
for Didié & Bauch, 2000).
Derivation of name. From Latin lunaris (adjective; nominative
singular; gender, neutral) = lunar.
Diagnosis. A small, heavily calcified Polycope species with well-
developed primary reticulation.
Holotype. LV, USNM 603626 (ODP982161) (Fig. 2B).
Paratype. RV, USNM 603627 (ODP982162).
Type locality and horizon. ODP 982A, 1/3/42–44.
Description. Carapace heavily calcified, small in size. Outline
rounded in lateral view. Lateral surface ornamented with well-
developed, rounded primary reticulation. Anteroventral ridge
thick, well developed and bearing no reticulation. Internal features
as for genus.
Dimensions. USNM 603626 (ODP982161) (Holotype), L = 0.378,
H = 0.343; USNM 603627 (ODP982162) (Paratype), L = 0.353,
H = 0.328.
Remarks. This species is distinctive from any other Polycope spe-
cies by having a heavily calcified carapace with well-developed,
rounded primary reticulation and thick anteroventral ridge.
Polycope cf. bireticulata Joy & Clark, 1977
(Fig. 2D–E)
2009b Polycope cf. bireticulata Joy & Clark; Yasuhara et al.:
881, pl. 1, figs 3 and 4.
Remarks. ODP 982A specimens have better developed
reticulation compared to the specimens from the western
North Atlantic (Yasuhara et al., 2009b), considered here to be
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intraspecific variation. This species is similar to P. bireticu-
lata, but the latter has a more evenly rounded outline and dif-
ferent alignment of muri (Yasuhara et al. in press b).
Polycope orbicularis s.l. Sars, 1866
(Fig. 2F–G)
2009b Polycope cf. orbicularis Sars; Yasuhara et al.: 881, pl. 1, fig. 5.
2009b Polycope orbicularis s.l. Sars; Yasuhara et al.: 881.
2009 Polycope orbicularis Sars; Alvarez Zarikian: 3, pl. P1, fig. 7.
Remarks. Polycope orbicularis s.l. is discussed in Yasuhara et al.
(2009b).
Polycope vasfiensis Sissingh, 1972
(Fig. 2H–I)
1972 Polycope vasfiensis Sissingh: 68, pl. 1, fig. 6.
1976 Polycope vasfiensis Sissingh; Bonaduce et al.: 18, pl. 1, figs
6–8, text-fig. 6.
2000 Polycope vasfiensis Sissingh; Aiello et al.: 85, pl. 1, fig. 1.
2009b Polycope vasfiensis Sissingh; Yasuhara et al.: 882, pl. 1,
figs 1–2.
Polycope reticulata Müller, 1894
(Fig. 2J–K)
1894 Polycope reticulata Müller: 235, pl. 7, figs 44, 49–50; pl. 8,
fig. 20.
non 1976 Polycope reticulata Müller; Bonaduce et al.: 14, pl. 2,
figs 9 and 10.
2013 Polycope reticulata Müller; Cabral & Loureiro: 137, pl. 1,
fig. 1.
Remarks. We consider that Polycope reticulata sensu Bonaduce
et al. (1976) is not conspecific with P. reticulata Müller, 1894
because the original sketch by Müller (1894, pl. 8, fig. 20) lacks
secondary reticulation and has a different primary reticulation
pattern and more inflated carapace.
Subclass Podocopa Müller, 1894
Order Platycopida Sars, 1866
Suborder Platycopina Sars, 1866
Superfamily Cytherelloidea Sars, 1866
Table 2. The list of ODP 982A samples used for the present study.
Core Section
Interval
(top: cm)
Interval
(bottom: cm)
MCD
(cm)
Age
(ka BP) N S
1 1 0 2 0 0.0 87 10
1 1 10 12 10 1.0 119 15
1 1 20 22 20 2.2 107 14
1 1 30 32 30 3.4 204 19
1 1 40 42 40 5.2 276 26
1 1 50 52 50 7.2 221 35
1 1 60 62 60 10.0 231 40
1 1 70 72 70 14.9 184 39
1 1 80 82 80 19.9 141 25
1 1 90 92 90 24.9 140 31
1 1 97 99 97 28.3 237 42
1 1 107 109 107 33.3 234 30
1 1 117 119 117 38.3 241 25
1 1 127 129 127 43.2 312 29
1 1 137 139 137 48.2 259 28
1 1 142 144 142 50.7 354 34
1 2 7 9 157 58.1 264 40
1 2 17 19 167 63.1 283 34
1 2 27 29 177 68.0 268 37
1 2 37 39 187 73.0 228 34
1 2 47 49 197 77.9 227 35
1 2 57 59 207 82.9 214 28
1 2 67 69 217 87.9 195 32
1 2 77 79 227 92.8 155 32
1 2 87 89 237 97.8 264 29
1 2 97 99 247 102.8 123 25
1 2 107 109 257 107.7 320 25
1 2 117 119 267 112.7 201 21
1 2 127 129 277 117.6 237 28
1 2 137 139 287 122.6 148 24
1 3 2 4 302 130.0 220 30
1 3 12 14 312 136.3 169 25
1 3 22 24 322 146.5 161 27
1 3 32 34 332 156.8 139 28
1 3 42 44 342 165.0 195 27
1 3 52 54 352 172.5 152 28
1 3 62 64 362 179.6 228 26
1 3 72 74 372 184.9 128 25
1 3 82 84 382 190.2 229 28
1 3 92 94 392 195.3 164 31
1 3 102 104 402 200.4 294 43
1 3 112 114 412 205.6 198 34
1 3 122 124 422 210.7 367 39
1 3 132 134 432 215.8 131 27
1 3 142 144 442 220.9 164 27
1 4 2 4 452 226.0 202 29
1 4 12 14 462 231.1 161 29
N, number of ostracode specimens per sample; S, number of ostracode
species per sample. Chronology from Venz et al. (1999)
: : : : :
1

1
1
1









2'3
:DWHUGHSWKP
Fig. 1. Locality map of ODP Hole 982A.
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M. Yasuhara & H. Okahashi
6
Family Cytherellidae Sars, 1866
Genus Cytherella Jones, 1849
Type species. Cytherina ovata Roemer, 1841
(designated by Ulrich, 1894)
Cytherella robusta Colalongo & Pasini, 1980
(Fig. 3A–B)
1979 Cytherella sp. 11 Ducasse & Peypouquet: pl. 1, figs 3–4.
1980 Cytherella robusta Colalongo & Pasini: 78, pl. 6, figs
4–10.
1996b Cytherella robusta Colalongo & Pasini; Aiello et al.: 184,
pl. 2, figs 4–5, 8–12.
2001 Cytherella serratula (Brady); Didié & Bauch: 104, pl. 1, fig. 5
(erratum for Didié & Bauch, 2000).
2001 Cytherella sp. 1 Didié & Bauch: 104, pl. 1, fig. 6 (erratum
for Didié & Bauch, 2000).
Fig. 2. Scanning electron microscope (SEM) images of Polycope species. A, Polycope martinezi (Karanovic & Brandão, 2012), USNM 603625
(ODP982154poly); LV from 1/3/2–4. B–C, Polycope lunaris sp. nov.: B, Holotype USNM 603626 (ODP982161); adult? LV? from 1/3/42–44;
C, Paratype USNM 603627 (ODP982162); adult? RV? from 1/3/42–44. D–E, Polycope cf. bireticulata Joy & Clark, 1977: D, USNM 603628
(ODP982155poly); LV from 1/3/32–34; E, USNM 603629 (ODP982156poly); RV from 1/3/32–34. F–G, Polycope orbicularis s.l. Sars, 1866: F,
USNM 603630 (ODP982157poly); RV from 1/3/112–114; G, USNM 603631 (ODP982158poly); LV from 1/3/112–114. H–I, Polycope vasfiensis
Sissingh, 1972: H, USNM 603632 (ODP982159poly); adult? LV from 1/2/27–29; I, USNM 603633 (ODP982160poly); adult? RV from 1/2/27–29.
J–K, Polycope reticulata Müller, 1894: J, USNM 603634 (ODP982163poly); LV from 1/3/32–34; K, USNM 603635 (ODP982164poly); RV from
1/3/32–34. All lateral views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic. Scale bars
represent 0.5 mm.
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Late Quaternary deep-sea ostracods, North Atlantic
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2001 Cytherella sp. 2 Didié & Bauch: 104, pl. 1, fig. 7 (erratum
for Didié & Bauch, 2000).
2009 Cytherella sp. Alvarez Zarikian: 7, pl. P10, fig. 5.
2009b Cytherella robusta s.l. Colalongo & Pasini; Yasuhara
et al.: 882, pl. 1, figs 7–12.
2013 Cytherella robusta Colalongo & Pasini; Cabral & Loureiro:
137, pl. 1, fig. 3.
Remarks. Juvenile specimens of Cytherella robusta are shown
here. As discussed in Yasuhara & Okahashi (in press), this
species has certain intraspecific variation. A comprehensive
synonymy list in Aiello et al. (1996b), Yasuhara et al. (2009b)
and Yasuhara & Okahashi (in press).
Order Podocopida Sars, 1866
Suborder Bairdiocopina Gründel, 1967
Superfamily Bairdioidea Sars, 1866
Family Bairdiidae Sars, 1866
Remarks. We follow Maddocks’ (1969) genus-level taxonomy
for this family.
Fig. 3. SEM images of Cytherella robusta Colalongo & Pasini, 1980, Bairdoppilata conformis (Terquem, 1878) and Macrocyprissa arcuata (Colalongo
& Pasini, 1980). A–B, Cytherella robusta Colalongo & Pasini, 1980: A, USNM 603636 (ODP982075); juvenile LV from 1/2/107–109; B, USNM
603637 (ODP982076); juvenile RV from 1/2/107–109. C–F, Bairdoppilata conformis (Terquem, 1878): C–D, USNM 603638 (ODP982066); adult LV
from 1/2/67–69; E–F, USNM 603639 (ODP982067); adult RV from 1/3/82–84. G–I, Macrocyprissa arcuata (Colalongo & Pasini, 1980): G, USNM
603640 (ODP982124); adult RV from 1/1/97–99; H, USNM 603641 (ODP982126); adult RV from 1/3/102–104; I, USNM 603642 (ODP982125);
adult LV from 1/3/102–104. A–C, E, G–I, lateral views; D, F, internal views. All specimens from late Quaternary section of ODP Hole 982A, Rockall
Plateau, eastern North Atlantic. Scale bars represent 1 mm.
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M. Yasuhara & H. Okahashi
8
Genus Bairdoppilata Coryell, Sample & Jennings (1935)
Type species. Bairdoppilata martyni Coryell, Sample &
Jennings (1935)
Bairdoppilata conformis (Terquem, 1878)
(Fig. 3C–F)
1878 Bairdia subdeltoidea var. conformis Terquem: 93, pl. 10,
fig. 17a–c.
1962 Bairdia conformis Terquem; Ruggieri: 13.
1976 Bairdia conformis Terquem; Bonaduce et al.: 22, pl. 6,
figs 5–10.
2000 Bairdoppilata conformis (Terquem); Aiello et al.: 85, pl. 1,
fig. 2.
2003 Bairdoppilata conformis (Terquem); Sciuto: 182, fig. 2a.
2008 Bairdoppilata conformis (Terquem); Faranda et al.: 300,
tab. 2.
2010 Bairdoppilata conformis (Terquem); Aiello & Barra: 406.
Remarks. A comprehensive synonymy list is in Aiello et al.
(2000) and supplemented herein. Our specimens are identical to
that shown in Aiello et al. (2000). Our specimens are also very
similar to the specimens in Terquem (1878) and Bonaduce et al.
(1976), but the latter specimens have a more heavily calcified car-
apace and slightly more upturned caudal process. Other authors
have also reported this species, but with a slightly different out-
line (Sciuto, 2003) or without images (Faranda et al., 2008; Aiello
& Barra, 2010). We consider all of these differences as intraspe-
cific variation and include them in Bairdoppilata conformis.
Slight differences in outlines may be due to intraspecific variation
in calcification. No SEM or microscopic image of type specimens
has been published.
Suborder Cypridocopina Jones, 1901
Superfamily Macrocypridoidea Müller, 1912
Family Macrocyprididae Müller, 1912
Genus Macrocyprissa Triebel, 1960
Type species. Bairdia cylindracea Bornemann, 1855
Macrocyprissa arcuata (Colalongo & Pasini, 1980)
(Fig. 3G–I)
1980 Paramacrocypris arcuata Colalongo & Pasini: 106, pl. 25,
figs 1–8.
1990 Macrocyprissa arcuata (Colalongo & Pasini); Maddocks:
85, figs 12.17–18, 13.17–18, 21.24, 23.24, 24.34, 28.29–30,
29.9, 35.19, 44.5, 47.6–7, 50.15–19, 51.9–13, 56.10, 24, 27,
57.11, 32, 58.18, 59.26, 33, 60.4, 63.3, 17, 64.23, 40, 51,
68.5, 72.1–3, 76.3, 78.9–10; pl. 30, figs 7–10, pl. 31, figs
7–11; pl. 67, figs 8–9; pl. 68, figs 6–14; pl. 69, fig. 1; pl. 70,
figs 1–4; pl. 80, fig. 7; pl. 91, figs 10–14; pl. 104, figs 1–5;
pl. 111, fig. 6.
1996 Macrocyprissa arcuata (Colalongo & Pasini); Coles et al.:
132, pl. 1, figs 10–11.
Remarks. A comprehensive synonymy list is given in Maddocks
(1990).
Superfamily Pontocypridoidea Müller, 1894
Family Pontocyprididae Müller, 1894
Genus Argilloecia Sars, 1866
Type species. Argilloecia cylindrica Sars, 1866
Argilloecia acuminata Müller, 1894
(Fig. 4A–D)
1894 Argilloecia acuminata Müller: 261, pl. 12, figs 1–2, 12–22.
1975 Argilloecia acuminata Müller; Breman: 82, pl. 2, fig. 21,
pl. 6, fig. 69.
1987 Argilloecia sp. 5 Whatley & Coles: 87, pl. 1, figs 19–20.
2004 Argilloecia acuminata Müller; Aiello & Szczechura: 16,
pl. 1, fig. 2.
2009b Argilloecia acuminata Müller; Yasuhara et al.: 886, pl. 3,
figs 1–2, 4–5.
2009 (part) Argilloecia sp. 2 Alvarez Zarikian: 7, pl. P8, fig. 4 (non 3).
Remarks. Comprehensive synonymy lists are given in Aiello &
Szczechura (2004), Yasuhara et al. (2009b) and supplemented herein.
Argilloecia caju Yasuhara, Okahashi & Cronin 2009
(Fig. 4E–F)
2009b Argilloecia caju Yasuhara, Okahashi & Cronin: 886, pl. 3,
figs 21–24.
Remarks. Argilloecia caju is similar to Pacific species A. viriosa
Hao, 1988 (in Ruan & Hao, 1988), but distinguished by having a
much more slender outline. A. caju was originally reported from
the western North Atlantic (Yasuhara et al., 2009b) and is here
confirmed in the eastern North Atlantic.
Argilloecia labri sp. nov.
(Fig. 4G–J)
1987 Argilloecia sp. 4 Whatley & Coles: 86, pl. 1, figs 17–18.
2000 Argilloecia sp. 2 Didié & Bauch: 116, pl. 3, figs 3–4.
Derivation of name. From Latin labri (noun, genitive singu-
lar) = lip.
Diagnosis. A small, moderately calcified Argilloecia species with
lip-shaped outline.
Holotype. LV, USNM 603651 (ODP982060) (Fig. 4I).
Paratypes. LV, USNM 603649 (ODP982058); RV, USNM
603650 (ODP982059); RV, USNM 603652 (ODP982061).
Type locality and horizon. ODP 982A, 1/1/50–52.
Description. Carapace moderately calcified, small, highest at mid-
length. Outline trapezoidal in lateral view; anterior margin rounded
and upturned; posterior margin acuminate and slightly upturned;
dorsal margin arched; ventral margin slightly sinuous. Anterodorsal
and posterodorsal margins obtuse-angular. RV strongly overlaps
LV. Lateral surface smooth. Internal features as for genus.
Dimensions. USNM 603651 (ODP982060) (Holotype), L = 0.461,
H = 0.202;
USNM 603650 (ODP982059) (Paratype), L = 0.497, H = 0.223.
Remarks. This species is similar to the Pacific species Argilloecia
spicata Hao, 1988 (in Ruan & Hao, 1988), but distinguished by
having a much more upturned posterior margin.
Argilloecia bensoni Barra, Aiello & Bonaduce, 1996
(Fig. 4K–N)
1996 Argilloecia bensoni Barra, Aiello & Bonaduce: 129, pl. 2,
figs 3–4; pl. 3, figs 1–3.
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Late Quaternary deep-sea ostracods, North Atlantic
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Remarks. This species was originally reported from Mediterranean
Plio-Pleistocene strata.
Genus Propontocypris Sylvester-Bradley, 1947
Type species. Pontocypris trigonella
Sars, 1866
Propontocypris acuminata (Müller, 1894)
(Fig. 5A)
1894 Erythrocypris acuminata Müller: 259, pl. 11, figs 5–6, 16–
18, 40–42; pl. 28, figs 23, 30; pl. 38, figs 47–48.
1976 Pontocypris acuminata (Müller); Bonaduce et al.: 25, pl. 9,
figs 1–2.
2000 Pontocypris sp. Didié & Bauch: 116, pl. 4, fig. 17.
Fig. 4. SEM images of Argilloecia species. A–D, Argilloecia acuminata Müller, 1894: A, USNM 603643 (ODP982062); adult LV from 1/1/60–62;
B, USNM 603644 (ODP982063); adult RV from 1/1/60–62; C, USNM 603645 (ODP982064); adult LV from 1/1/50–52; D, USNM 603646
(ODP982065); adult RV from 1/1/50–52. E–F, Argilloecia caju Yasuhara, Okahashi & Cronin, 2009: E, USNM 603647 (ODP982052); adult
LV from 1/2/17–19; F, USNM 603648 (ODP982053); adult RV from 1/2/17–19. G–J, Argilloecia labri sp. nov.: G, Paratype USNM 603649
(ODP982058); adult LV from 1/3/102–104; H, Paratype USNM 603650 (ODP982059); adult RV from 1/3/102–104; I, Holotype USNM 603651
(ODP982060); adult LV from 1/1/50–52; J, Paratype USNM 603652 (ODP982061); adult RV from 1/3/82–84. K–N, Argilloecia bensoni Barra,
Aiello & Bonaduce, 1996: K, USNM 603653 (ODP982054); adult RV from 1/1/97–99; L, USNM 603654 (ODP982057); adult LV from 1/1/142–
144; M, USNM 603655 (ODP982056); adult RV from 1/3/112–114; N, USNM 603656 (ODP982055); adult LV from 1/1/107–109. C–F, I–J, M–N,
lateral views; A–B, G–H, K–L, internal views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North
Atlantic. Scale bar represents 1 mm.
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M. Yasuhara & H. Okahashi
10
Propontocypris sp.
(Fig. 5B)
?2009 Propontocypris trigonella Sars; Alvarez Zarikian: 7, pl. P8,
fig. 10.
Suborder Cytherocopina Gründel, 1967
Superfamily Cytheroidea Baird, 1850
Family Bythocytheridae Sars, 1866
Genus Bythoceratina Hornibrook, 1952
Type species. Bythoceratina mestayerae, Hornibrook, 1952
Bythoceratina scaberrima (Brady, 1886)
(Fig. 5C)
1886 Cytherura scaberrima Brady: 198, pl. 14, figs 10–11.
Fig. 5. SEM images of Propontocypris, Bythoceratina, Pseudocythere and Paijenborchella species. A, Propontocypris acuminata (Müller, 1894),
USNM 603657 (ODP982165prop); juvenile? LV from 1/2/37–39. B, Propontocypris sp., USNM 603658 (ODP982166prop); juvenile? LV from
1/1/60–62. C, Bythoceratina scaberrima (Brady, 1886), USNM 603659 (ODP982072); adult LV from 1/2/67–69. D–E, Bythoceratina nuda sp. nov.:
D, Holotype USNM 603660 (ODP982073); adult LV from 1/3/52–54; E, Paratype USNM 603661 (ODP982074); adult RV from 1/3/42–44. F–G,
Pseudocythere caudata Sars, 1866:. F, USNM 603662 (ODP982167pseu); adult? LV from 1/3/42–44; G, USNM 603663 (ODP982168pseu); adult?
RV from 1/3/42–44. H–K, Paijenborchella cymbula Ruggieri, 1950: H, USNM 603664 (ODP982131); adult RV from 1/1/97–99; I, USNM 603665
(ODP982132); adult LV from 1/1/40–42; J, USNM 603666 (ODP982133); adult RV from 1/1/40–42; K, USNM 603667 (ODP982134); adult LV from
1/1/40–42. A–J, lateral views; K, internal view. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic.
Scale bars represent 0.5 mm.
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Late Quaternary deep-sea ostracods, North Atlantic
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1980 Bythoceratina scaberrima mediterranea Colalongo & Pasini:
72, pl. 1, fig. 9; pl. 4, figs 9–10.
2001 Bythoceratina scaberrima (Brady); Didié & Bauch: pl. 1,
fig. 29 (erratum for Didié & Bauch, 2000).
2005 Retibythere scaberrima (Brady); Guernet: 109.
Remarks. Comprehensive synonymy list and detailed discussion
in Guernet (2005) and Yasuhara et al. (in press b).
Bythoceratina nuda sp. nov.
(Fig. 5D–E)
Derivation of name. From Latin nuda (adjective, nominative sin-
gular, gender feminine or neuter) = stripped, with reference to its
carapace without any spine or reticulation.
Diagnosis. A large, moderately calcified Bythoceratina species
without spines or reticulation.
Holotype. LV, USNM 603660 (ODP982073) (Fig. 5D)
Paratype. RV, USNM 603661 (ODP982074).
Type locality and horizon. ODP 982A, 1/3/52–54.
Description. Carapace moderately calcified, large, highest at
anterodorsal corner (= anterior cardinal angle). Outline parallelo-
gram-like in lateral view; anterior margin rounded; caudal pro-
cess upturned; dorsal margin sinuous; ventral margin slightly
curved. Anterodorsal margin prominent; posterodorsal margin
slightly angular. Lateral surface smooth. A ventrolateral ridge
well developed, reaching to anterior margin; thin dorsolateral
ridge present. A median sulcus present, but very shallow. Internal
features as for genus.
Dimensions. USNM 603660 (ODP982073) (Holotype), L = 0.920,
H = 0.462; USNM 603661 (ODP982074) (Paratype), L = 0.948,
H = 0.471.
Remarks. This species is distinguished from any other
Bythoceratina species by its lack of spines and reticulation.
Genus Pseudocythere Sars, 1866
Type species. Pseudocythere caudata Sars, 1866
Pseudocythere caudata Sars, 1866
(Fig. 5F–G)
1866 Pseudocythere caudata Sars: 88.
1926 Pseudocythere caudata Sars; Sars: 239, pl. 109, fig. 2a–k.
2009b Pseudocythere caudata Sars; Yasuhara et al.: 892, pl. 4,
figs 7–12.
Remarks. We think that this species has considerable intraspecific
variation. A comprehensive synonymy list and detailed discussion
are given in Yasuhara et al. (in press b).
Family Cytheridae Baird, 1850
Genus Paijenborchella Kingma, 1948
Type species. Paijenborchella iocosa Kingma, 1948
Paijenborchella cymbula Ruggieri, 1950
(Fig. 5H–K)
1950 Paijenborchella cymbula Ruggieri: 60, 1 unnumbered fig.
on p. 61.
1973 Paijenborchella (Eopaijenborchella) malaiensis cymbula
Ruggieri; Doruk: 161, pls 1.30.162, 1.30.164.
2000 Paijenborchella malaiensis cymbula Ruggieri; Aiello et al.:
93, pl. 2, fig. 12.
2005 Paijenborchella cymbula Ruggieri; Guernet: 107.
Remarks. A comprehensive synonymy list can be found in Aiello
et al. (2000) and Guernet (2005). To our knowledge, this is the
first well-illustrated record (SEM images) of this species from the
Atlantic. This species was recently reported from the Iberian
Margin at IODP Site U1387 (Expedition 339 Scientists, 2013).
Family Cytheruridae Müller, 1894
Genus Aversovalva Hornibrook, 1952
Type species. Cytheropteron
(Aversovalva) aureum Hornibrook, 1952
Remarks. Several authors have considered Aversovalva Hornibrook,
1952 as a subgenus (Aiello et al., 1996a) or junior synonym
(Whatley & Masson, 1979; Horne & Whittaker, 1988) of
Cytheropteron Sars, 1866. However, in our opinion, differences in
hingement and outline including a straight median hinge element,
highly developed smooth and rounded terminal teeth clearly distinct
from the median hinge element, and parallelogram-like outline in
Aversovalva (e.g. Figs 6C–D) are sufficient to separate Aversovalva
from Cytheropteron. Most recent deep-sea ostracod papers have
considered Aversovalva as an independent genus (e.g. see synonymy
of Aversovalva hydrodynamica below), supporting our decision.
Aversovalva hydrodynamica Whatley & Coles, 1987
(Fig. 6A–D)
1987 Aversovalva hydrodynamica Whatley & Coles: 69, pl. 3,
figs 10–11.
1996 Aversovalva hydrodynamica Whatley & Coles; Coles et al.:
150, pl. 3, fig. 17.
1988 (part) Aversovalva sp. 2 Whatley & Ayress: 742, pl. 2, fig.
1a (non 1b).
?2001 Aversovalva sp. cf. A. hydrodynamica Didié & Bauch: 103,
pl. 1, fig. 12 (as erratum for Didié & Bauch, 2000).
2009 Aversovalva hydrodynamica Whatley & Coles; Alvarez
Zarikian: 3, pl. P3, fig. 7.
Remarks. Very similar, but slightly different species are reported
from the western North Atlantic as Aversovalva sp. 1 and A. cf.
hydrodynamica (Yasuhara et al., 2009b). A Pliocene Mediterranean
species Aversovalva denticulatum (Aiello, Barra & Bonaduce,
1996) shows strong affinity to A. hydrodynamica Whatley &
Coles, 1987, but the former has a more triangular outline. A.
hydrodynamica is also similar to A. consueta (Dall’Antonia, 2003),
but the latter has thicker and more downward-extended alae and
weaker reticulation. Although Coles et al. (1990, 1996) suggested
a global distribution for this species, reliable records with SEM
images are restricted in the eastern North Atlantic.
Genus Cytheropteron Sars, 1866
Type species. Cythere latissima Norman, 1865 (designated by
Brady & Norman, 1889; see Horne & Whittaker (1988) for
details and lectotype).
Remarks. We agree with Horne & Whittaker (1988) and consider
Kobayashiina Hanai, 1957b and Lobosocytheropteron Ishizaki &
Gunther, 1974 as junior synonyms of Cytheropteron.
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M. Yasuhara & H. Okahashi
12
Cytheropteron aielloi Yasuhara, Okahashi & Cronin, 2009
(Fig. 6E–G)
1996 Cytheropteron sedovi Schneider; Whatley et al.: 19, pl. 2,
figs 15–17.
1998 Cytheropteron sedovi Schneider; Whatley et al.: 21, pl. 2,
figs 11–12.
2009 Cytheropteron aielloi Yasuhara, Okahashi & Cronin: 898,
pl. 10, figs 3–6.
Remarks. This species is known not only from the North Atlantic
proper but also from the Nordic seas.
Cytheropteron alatum Sars, 1866
(Fig. 6H)
1866 Cytheropteron alatum Sars: 81.
1926 Cytheropteron alatum Sars; Sars: 225, pl. 104, fig. 1.
1993 Cytheropteron alatum Sars; Penney: figs 4n–o.
1996 Cytheropteron vespertilio (Reuss); Coles et al.: 136, pl. 3,
fig. 9.
1998 Cytheropteron alatum Sars; Freiwald & Mostafawi: 260,
pl. 59, fig. 7.
2000 Cytheropteron alatum Sars; Didié & Bauch: pl. 2, fig. 6.
Fig. 6. SEM images of Aversovalva and Cytheropteron species. A–D, Aversovalva hydrodynamica Whatley & Coles, 1987: A, USNM 603668 (ODP982048);
adult RV from 1/1/107–109; B, USNM 603669 (ODP982049); adult LV from 1/1/107–109; C, USNM 603670 (ODP982050); adult LV from 1/1/107–109;
D, USNM 603671 (ODP982051); adult RV from 1/1/117–119. E–G, Cytheropteron aielloi Yasuhara, Okahashi & Cronin, 2009: E, USNM 603672
(ODP982042); adult LV from 1/3/42–44; F, USNM 603673 (ODP982043); adult RV from 1/3/42–44; G, USNM 603674 (ODP982044); adult RV from
1/3/42–44; H, Cytheropteron alatum Sars, 1866, USNM 603675 (ODP982084); adult RV from 1/3/112–114. A–B, E–F, H, lateral views; C–D, G, internal
views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic. Scale bar represents 0.5 mm.
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Late Quaternary deep-sea ostracods, North Atlantic
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Remarks. Reliable occurrence records of this species with SEM
image(s) or sketches are known only from the eastern North
Atlantic as listed in the synonymy list above. Detailed discussion
of this species can be found in Yasuhara et al. (in press b).
Cytheropteron colesoabyssorum sp. nov.
(Fig. 7A)
1996 Cytheropteron cf. abyssorum Brady; Coles et al.: 136, pl. 3,
figs 12–13.
Derivation of name. In honour of Graham P. Coles for his con-
tribution to deep-sea ostracod research; and with reference to its
similarity to Cytheropteron abyssorum as indicated by him.
Diagnosis. A large, moderately calcified Cytheropteron species
with finely punctate carapace, upturned caudal process, and rela-
tively rounded outline.
Holotype. RV, USNM 603676 (ODP982077) (Fig. 7A).
Type locality and horizon. ODP 982A, 1/2/37–39.
Description. Carapace moderately calcified, large, highest at mid-
length. Outline rhomboidal and rounded in lateral view; anterior
margin evenly rounded; caudal process strongly upturned; dorsal
margin arched; ventral margin slightly curved; alae well devel-
oped, almost reaching to anterior margin and slightly extended
below ventral margin; median sulcus present on alae; thin dorso-
lateral ridge present along dorsal margin. Anterodorsal and pos-
terodorsal margins slightly angular. Lateral surface finely punctate.
Internal features as for genus.
Dimensions. USNM 603676 (ODP982077) (Holotype), L = 0.489,
H = 0.301.
Remarks. Cytheropteron colesoabyssorum sp. nov. is similar to
C. abyssorum Brady, 1880 (see Passlow & Ayress, 1994) in cer-
tain aspects, such as presence of a median sulcus on alae and
punctate carapace, but easily distinguished by having much finer
punctation covering entire carapace, thinner and longer alae, and
upturned caudal process and presence of dorsolateral ridge.
C. colesoabyssorum sp. nov. is similar to C. cf. tenuialatum of
Coles et al. (1996), but distinguished by having a dorsolateral
ridge and more strongly upturned caudal process, and by the
absence of primary reticulation on the caudal process.
Cytheropteron colesopunctatum sp. nov.
(Fig. 7B–C)
1996 (part) Cytheropteron gr. punctatum Brady; Coles et al.: 136,
pl. 3, figs 7–8 (non 5–6).
in press a Cytheropteron sp. Yasuhara et al.: fig. 6.7–8.
Derivation of name. In honour of Graham P. Coles for his con-
tribution to deep-sea ostracod research; and with reference to its
similarity to Cytheropteron punctatum as indicated by him.
Diagnosis. A large, moderately calcified Cytheropteron species
with coarsely punctate carapace and horizontally long, well-devel-
oped alae.
Holotype. RV, USNM 603677 (ODP982082) (Fig. 7B).
Paratype. LV, USNM 603678 (ODP982083).
Type locality and horizon. ODP 982A, 1/1/30–32.
Description. Carapace moderately calcified, large, highest at mid-
length. Outline rhomboidal or almond-like in lateral view; anterior
margin evenly rounded; caudal process prominent and upturned;
dorsal margin arched and slightly sinuous; ventral margin curved;
alae curved, horizontally long and well-developed, reaching to
anterior margin and extending slightly below ventral margin, with
a small spine at apex. Anterodorsal margin slightly angular; pos-
terodorsal margin weakly angular. Lateral surface coarsely punc-
tate; primary and secondary reticulation developed in posterior
one third. Internal features as for genus.
Dimensions. USNM 603677 (ODP982082) (Holotype), L = 0.514,
H = 0.306; USNM 603678 (ODP982083) (Paratype), L = 0.545,
H = 0.338.
Remarks. Cytheropteron colesopunctatum sp. nov. is similar to
C. punctatum Brady, 1868 in certain aspects, such as punctate
carapace and general outline, but is distinguished by having
smoothly curved and horizontally longer alae reaching to anterior
margin and well-developed primary and secondary reticulation in
posterior one-third. C. punctatum has sinuous and horizontally
shorter alae and no or only poorly developed reticulation, accord-
ing to the sketches and SEM images shown in Brady (1868), Sars
(1928) and Whatley & Masson (1979). C. colesopunctatum sp.
nov. is similar to C. paracarolinae Zhao et al., 2000 (see Zhao
et al., 2000; Hou & Gou, 2007), but the latter is larger, lacks
punctation in anterior one-third, and has more slender outline and
stronger caudal process.
Cytheropteron didieae Yasuhara, Okahashi & Cronin 2009
(Fig. 7D–H)
2009 Cytheropteron didieae Yasuhara, Okahashi & Cronin: 900,
pl. 6, figs 5–6, 8–9, 11–12.
Remarks. This is the first record of the species from the eastern
North Atlantic.
Cytheropteron fugu Yasuhara, Okahashi & Cronin, 2009
(Figs. 7I–J, 8A–B)
2009 Cytheropteron fugu Yasuhara, Okahashi & Cronin: 902,
pl. 7, figs 1–6.
Remarks. This is the first record of the species from the eastern
North Atlantic.
Cytheropteron inornatum Brady & Robertson, 1872
(Fig. 8G–J)
1872 Cytheropteron inornatum Brady & Robertson: 61, pl. 2, figs
1–3.
1979 Cytheropteron inornatum Brady & Robertson; Whatley &
Masson: 238, pl. 3, figs 1–3, 5–7.
1981 Cytheropteron hanaii Ishizaki: 55, pl. 11, figs 11–12; pl. 12,
figs 1–4; pl. 13, figs 8–9; pl. 14, fig. 3.
1988 Cytheropteron hanaii Ishizaki; Wang et al.: 265, pl. 52, figs
17–18.
1989 Cytheropteron inornatum Brady & Robertson; Athersuch
et al.: 226, fig. 95; pl. 8, fig. 4.
1992 Cytheropteron hanaii Ishizaki; Ikeya & Suzuki: 126, pl. 4,
fig. 4.
1996 Cytheropteron inornatum Brady & Robertson; Coles et al.:
135, pl. 2, fig. 15.
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M. Yasuhara & H. Okahashi
14
2000 Cytheropteron hanaii Ishizaki; Zhao et al.: 262, pl. 3, figs
8–9.
2007 Cytheropteron hanaii Ishizaki; Hou & Gou: 294, pl. 119,
figs 14–15; pl. 122, figs 11–12.
2009 Cytheropteron sp. g Yasuhara, Okahashi & Cronin: 908, pl.
6, fig. 14.
2012 Cytheropteron hanaii Ishizaki; Tanaka et al.: 10, pl. 1, fig. 13.
Remarks. Detailed comparison with similar species such as
C. fraudulentum Aiello, Barra & Bonaduce 1996 and C. sulcatum
Bonaduce, Ciampo & Masoli, 1976 is found in Aiello et al.
(1996a) and Athersuch et al. (1989). In our opinion, C. hanaii
Ishizaki, 1981 (see Ishizaki, 1981; Zhao et al., 2000; Hou & Gou,
2007) is a junior synonym of C. inornatum Brady & Robertson,
1872.
Fig. 7. SEM images of Cytheropteron species. A, Cytheropteron colesoabyssorum sp. nov., Holotype USNM 603676 (ODP982077); adult RV from
1/2/37–39. B–C, Cytheropteron colesopunctatum sp. nov.: B, Holotype USNM 603677 (ODP982082); adult RV from 1/1/30–32; C, Paratype USNM
603678 (ODP982083); adult LV from 1/1/30–32. D–H, Cytheropteron didieae Yasuhara, Okahashi & Cronin, 2009: D, USNM 603679 (ODP982085);
adult RV from 1/1/50–52; E, USNM 603680 (ODP982086); adult RV from 1/1/60–62; F, USNM 603681 (ODP982087); adult RV from 1/1/60–62;
G, USNM 603682 (ODP982088); adult LV from 1/1/60–62; H, USNM 603683 (ODP982089); adult LV from 1/2/127–129. I–J, Cytheropteron fugu
Yasuhara, Okahashi & Cronin, 2009: I, USNM 603684 (ODP982035); adult LV from 1/2/67–69; J, USNM 603685 (ODP982036); adult LV from
1/2/67–69. A–E, G, I, lateral views; F, H, J, internal views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern
North Atlantic. Scale bar represents 0.5 mm.
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Cytheropteron massoni Whatley & Coles, 1987
(Fig. 8K–L)
1987 Cytheropteron massoni Whatley & Coles: 63, pl. 2, figs 15–17.
2000 Cytheropteron massoni Whatley & Coles; Didié & Bauch:
113, pl. 2, fig. 11.
2009b Cytheropteron massoni Whatley & Coles; Yasuhara et al.:
904, p. 6, figs 7, 10, 13.
Remarks. C. massoni Whatley & Coles, 1987 is known from
both the eastern and western North Atlantic.
Cytheropteron omega Aiello, Barra & Bonaduce, 1996
(Fig. 8C–F)
1987 (part) Cytheropteron syntomoalatum Whatley & Coles: pl. 2,
fig. 27 (non pl. 2, figs 25–26, 28–29).
Fig. 8. SEM images of Cytheropteron species. A–B, Cytheropteron fugu Yasuhara, Okahashi & Cronin, 2009: A, USNM 603686 (ODP982037); adult
RV from 1/2/57–59; B, USNM 603687 (ODP982038); adult RV from 1/2/57–59. C–F, Cytheropteron omega Aiello, Barra & Bonaduce, 1996: C,
USNM 603688 (ODP982040); adult LV from 1/4/12–14; D, USNM 603689 (ODP982039); adult RV from 1/4/12–14; E, USNM 603690 (ODP982041);
adult RV from 1/3/132–134; F, USNM 603691 (ODP982045); adult LV from 1/2/127–129. G–J, Cytheropteron inornatum Brady & Robertson,
1872: G, USNM 603692 (ODP982080); adult LV from 1/1/97–99; H, USNM 603693 (ODP982079); adult RV from 1/2/17–19; I, USNM 603694
(ODP982081); adult RV from 1/1/97–99; J, USNM 603695 (ODP982078); adult LV from 1/2/17–19. K–L, Cytheropteron massoni Whatley & Coles,
1987: K, USNM 603696 (ODP982047); adult LV from 1/1/117–119; L, USNM 603697 (ODP982046); adult RV from 1/2/17–19. A, C–D, G–H, K–L,
lateral views; B, E–F, I–J, internal views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic.
Scale bar represents 0.5 mm.
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M. Yasuhara & H. Okahashi
16
1996 Cytheropteron omega Aiello, Barra & Bonaduce: 170, pl. 2,
figs 7–9.
Remarks. Detailed comparison with similar species such as
C. garganicum Bonaduce, Ciampo & Masoli, 1976 can be found
in Aiello et al. (1996a). Our specimens have relatively weakly
developed dorsal ridges compared to the type specimens and thus
the ‘upside-down omega’ structure is unclear, but otherwise iden-
tical. We consider this difference as intraspecific variation. Well-
preserved specimens shown here indicate that there are two spines
at the apex of alae. A paratype specimen of C. syntomoalatum of
Whatley & Coles (1987, pl. 2, fig. 27) is not conspecific with
C. syntomoalatum Whatley & Coles, 1987 and is considered here
and by Aiello et al. (1996a) as C. omega Aiello, Barra &
Bonaduce, 1996, although this specimen has only one spine at the
apex of the alae and a slightly more slender outline.
Cytheropteron paramediotumidum sp. nov.
(Fig. 9A–B)
1996 (part) Cytheropteron gr. punctatum Brady; Coles et al.: 136,
pl. 3, figs 5–6 (non 7–8).
Derivation of name. With reference to its similarity to
Cytheropteron mediotumidum.
Diagnosis. A large, moderately calcified Cytheropteron species
with finely punctate carapace and straight-sided alae.
Holotype. LV, USNM 603698 (ODP982092) (Fig. 9A).
Paratype. RV, USNM 603699 (ODP982093).
Type locality and horizon. ODP 982A, 1/2/107–109.
Description. Carapace moderately calcified, large, highest at mid-
length. Outline subrhomboidal in lateral view; anterior margin
rounded; caudal process moderately prominent; dorsal margin
arched; ventral margin slightly curved; alae straight, thin and hori-
zontally long, almost reaching to anterior margin, and extending
slightly below ventral margin; a small subcentral depression pre-
sent on alae. Anterodorsal margin slightly angular; posterodorsal
margin weakly prominent. Lateral surface finely punctate in pos-
terior two-thirds; primary and secondary reticulation weakly
developed in posterior one-third. Internal features as for genus.
Dimensions. USNM 603698 (ODP982092) (Holotype), L = 0.638,
H = 0.389; USNM 603699 (ODP982093) (Paratype), L = 0.671,
H = 0.431.
Remarks. Cytheropteron paramediotumidum sp. nov. is very sim-
ilar to C. mediotumidum Zhao, Whatley & Zhou, 2000, but distin-
guished by having a less upturned caudal process, finer punctation,
less distinct primary reticulation and straight-sided alae. C. para-
mediotumidum sp. nov. is also similar to C. colesopunctatum sp.
nov., but the latter has more prominent caudal processes, coarser
and more distinct punctation, and curved-sided alae.
Cytheropteron demenocali Yasuhara, Okahashi & Cronin, 2009
(Fig. 9C–D)
?2000 Cytheropteron porterae Whatley & Coles; Didié & Bauch:
113, pl. 2, fig. 20 (non figs 19 and 21).
2009 Cytheropteron demenocali Yasuhara, Okahashi & Cronin:
900, pl. 9, figs 1–10.
Remarks. Our specimens have two spines (instead of one) at the
apex of alae and slightly more ventrally-extended alae, but are
otherwise identical. We consider these differences as intraspecific
variation, at least for now.
Cytheropteron pararhombiformis Zhao, Whatley & Zhou, 2000
(Fig. 9E–F)
1988 Cytheropteron rhombiformis Chen; Ruan & Hao: 283,
pl. 47, figs 18–20.
1996 Cytheropteron sp. Coles et al.: 136, pl. 3, fig. 16.
2000 Cytheropteron pararhombiformis Zhao, Whatley & Zhou:
275, pl. 4, figs 5–8.
2007 Cytheropteron pararhombiforme Zhao, Whatley & Zhou;
Hou & Gou: 301, pl. 123, figs 5–8.
Remarks. Cytheropteron pararhombiformis Zhao, Whatley &
Zhou, 2000 is very similar to C. zinzulusae Bonaduce, Ciampo &
Masoli, 1976, but distinguished by having irregular and coarser
punctation and a less upturned caudal process. This species was
originally reported from the western North Pacific.
Cytheropteron paucipunctatum Whatley & Coles, 1987
(Fig. 9G–H)
1987 Cytheropteron paucipunctatum Whatley & Coles: 63, pl. 2,
figs 18–20.
1988 Cytheropteron sp. 1 Whatley & Ayress: 740, pl. 1, fig. 6a–b.
Remarks. Our specimens have weakly developed primary reticu-
lation in the posterior one-third, but otherwise are identical to
C. paucipunctatum Whatley & Coles, 1987. We consider this dif-
ference as intraspecific variation. This species is known only from
the eastern North Atlantic.
Cytheropteron perlaria Hao, 1988 (in Ruan & Hao, 1988)
(Fig. 10A–B)
1988 Cytheropteron perlaria Hao (in Ruan & Hao, 1988): 280,
pl. 47, figs 4–9.
1996 Cytheropteron testudo Sars; Coles et al.: 136, pl. 3,
figs 10–11.
1999 Cytheropteron perlaria Hao; Swanson & Ayress: 155, pl. 1,
figs 7–13; pl. 2, figs 1–3.
non 2004 Cytheropteron perlaria Hao; Ayress et al.: 29, pl. 3,
figs 7–8.
2006 Cytheropteron perlaria Hao; Stepanova: S163, pl. 3, figs
8–10.
2007 Cytheropteron testudo Sars; Hou & Gou: 290, pl. 120, figs
9–10.
2009 Cytheropteron perlaria Hao; Alvarez Zarikian: 4, pl. P3,
figs 1–2.
2009b Cytheropteron perlaria Hao; Yasuhara et al.: 904, pl. 7,
figs 12–13.
2011 Cytheropteron perlaria Hao; Zhao et al.: 27, pl. 1, fig. 26.
Remarks. This species is known from the Atlantic, Arctic and
Pacific oceans.
Cytheropteron pherozigzag Whatley, Ayress & Downing, 1986
(Fig. 10C–D)
1986 Cytheropteron pherozigzag Whatley, Ayress & Downing:
32, pl. 1, figs 6–20.
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Late Quaternary deep-sea ostracods, North Atlantic
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1988 Cytheropteron pherozigzag Whatley, Ayress & Downing;
Whatley & Ayress: pl. 2, fig. 3a–b.
1996 Cytheropteron pherozigzag Whatley, Ayress & Downing;
Zhao & Zheng: 72, pl. 2, fig. 3.
2000 Cytheropteron pherozigzag Whatley, Ayress & Downing;
Zhao et al.: 263, pl. 1, fig. 20.
2005 Cytheropteron pherozigzag Whatley, Ayress & Downing;
Zhao: 39, pl. 2, fig. 15.
2007 Lobosocytheropteron pherozigzag (Whatley, Ayress &
Downing); Hou & Gou: 309, pl. 125, fig. 17.
2009b Cytheropteron pherozigzag Whatley, Ayress & Downing;
Yasuhara et al.: 906, pl. 5, figs 6–8, 10.
Remarks. This species is known from the eastern and western
North Atlantic and northwestern Pacific oceans.
Fig. 9. SEM images of Cytheropteron species. A–B, Cytheropteron paramediotumidum sp. nov.: A, Holotype USNM 603698 (ODP982092); adult
LV from 1/2/107–109; B, Paratype USNM 603699 (ODP982093); adult RV from 1/2/127–129. C–D, Cytheropteron demenocali Yasuhara, Okahashi
& Cronin, 2009: C, USNM 603700 (ODP982090); adult LV from 1/1/137–139; D, USNM 603701 (ODP982091); adult RV from 1/2/17–19. E–F,
Cytheropteron pararhombiformis Zhao, Whatley & Zhou, 2000: E, USNM 603702 (ODP982095); adult LV from 1/1/30–32; F, USNM 603703
(ODP982096); adult RV from 1/1/40–42. G–H, Cytheropteron paucipunctatum Whatley & Coles, 1987: G, USNM 603704 (ODP982097); adult LV
from 1/2/127–129; H, USNM 603705 (ODP982098); adult RV from 1/3/52–54. All lateral views. All specimens from late Quaternary section of ODP
Hole 982A, Rockall Plateau, eastern North Atlantic. Scale bars represent 0.5 mm.
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M. Yasuhara & H. Okahashi
18
Cytheropteron pseudoalatum Colalongo & Pasini, 1980
(Fig. 10E)
1980 Cytheropteron pseudoalatum Colalongo & Pasini: 92, pl. 8,
fig. 8; pl. 9, figs 1–5.
1996a Cytheropteron pseudoalatum Colalongo & Pasini; Aiello
et al.: 171, pl. 2, figs 1–3; pl. 3, figs 1–2.
Remarks. This species was originally reported from the
Mediterranean. This is the first record from the North Atlantic.
Genus Eucytherura Müller, 1894
Type species. Cythere complexa Brady, 1867 (designated by
Alexander, 1936).
Remarks. We agree with Ayress et al. (1995) and consider
Typhlocythere Bonaduce, Ciampo & Masoli, 1976, Typhloeucytherura
Colalongo & Pasini, 1980 and Parahemingwayella Dingle, 1984 as
junior synonyms of Eucytherura Müller, 1894.
Eucytherura calabra (Colalongo & Pasini, 1980)
(Fig. 10F–H)
1980 Typhloeucytherura calabra Colalongo & Pasini: 122, pl. 20,
figs 1–8; pl. 21, figs 1–2.
1987 Eucytherura calabra (Colalongo & Pasini); Whatley &
Coles: pl. 3, figs 14–16.
1988 Eucytherura sp. 1; Ruan & Hao: 291, pl. 49, fig. 18.
1988 Eucytherura calabra (Colalongo & Pasini); Whatley &
Ayress: pl. 1, fig. 9a–b.
1995 Eucytherura calabra (Colalongo & Pasini); Ayress et al.:
211, fig. 3A–D.
1996 Eucytherura calabra (Colalongo & Pasini); Coles et al.:
136, pl. 3, fig. 18.
1996 Eucytherura calabra (Colalongo & Pasini); Zhao & Zheng:
72, pl. 2, fig. 36.
2001 Eucytherura calabra (Colalongo & Pasini); Didié & Bauch
(as erratum of Didié & Bauch, 2000): 103, pl. 1, figs 9–10.
Remarks. Eucytherura calabra (Colalongo & Pasini, 1980) is
similar to E. spinicorona Yasuhara, Okahashi & Cronin 2009, but
the former has a curved dorsal margin and rectangular outline.
E. calabra is known from the Atlantic, Mediterranean and Pacific.
Eucytherura multituberculata Ayress, Whatley,
Downing & Millson, 1995
(Fig. 10I–J)
1983 ?Tuberculocythere sp. Cronin: 107, pl. 6, fig. A.
1987 Eucytherura sp. 2 Whatley & Coles: 90, pl. 3, fig. 18.
1995 Eucytherura multituberculata Ayress, Whatley, Downing &
Millson: 213, fig. 5A–E.
Remarks. This species is known both from the western and east-
ern North Atlantic.
Eucytherura tetrapteron (Bonaduce, Ciampo & Masoli, 1976)
(Fig. 11A–C)
1976 ?Cytheropteron tetrapteron Bonaduce, Ciampo & Masoli:
99, pl. 47, fig 1–7.
1980 Cytheropteron? tetrapteron Bonaduce, Ciampo & Masoli;
Ciampo: 19, pl. 3, fig. 5.
1980 Tuberculocythere tetrapteron (Bonaduce, Ciampo & Masoli);
Colalongo & Pasini: 120, pl. 34 fig. 2.
1985 Tuberculocythere tetrapteron (Bonaduce, Ciampo &
Masoli); Moncharmont-Zei et al.: 28, pl. 1, fig. 1.
1994 Parahemingwayella tetrapteron (Bonaduce, Ciampo &
Masoli); Malz & Jellinek: 28, pl. 7, figs 37–40.
2000 Parahemingwayella tetrapteron (Bonaduce, Ciampo &
Masoli); Aiello et al.: 94, pl. 3, fig. 9.
2001 Eucytherura sp. Didié & Bauch (as erratum of Didié &
Bauch, 2000): 103, pl. 1, fig. 11.
2005 Parahemingwayella tetrapteron (Bonaduce, Ciampo &
Masoli); Guernet: 108.
Remarks. Eucytherura tetrapteron (Bonaduce, Ciampo & Masoli,
1976) is very similar to E. downingae (Coles & Whatley, 1989), but
the latter has more strongly developed primary and secondary reticu-
lation and bears two spines on the anterodorsal margin (the former
bears a continuous frill). Although these differences are subtle and
may be a result of intraspecific variation in calcification, we consider
these two as separate species at least for now. E. tetrapteron is
known from the Mediterranean and the eastern North Atlantic.
Genus Kangarina Coryell & Fields, 1937
Type species. Kangarina quellita Coryell & Fields, 1937
Kangarina abyssicola (Müller, 1894)
(Fig. 11D–E)
1894 Cytheropteron abyssicolum Müller: 302, pl. 20, figs 5, 11;
pl. 21, figs 6–9.
1952 Cytheropteron (Kangarina) abyssicolum Müller; Ruggieri:
77, pl. 6, fig. 9.
1953 Kangarina abyssicola coarctata Ruggieri: 53, figs 16, 16a.
1953 Kangarina abyssicola (Müller); Ruggieri: 53, figs 15, 15a.
1972 Kangarina septentrionalis Neale: 33, pl. 1, figs 1–8.
1976 Kangarina abyssicola (Müller); Bonaduce et al.: 84, pl. 17,
fig. 16.
1980 Kangarina abyssicola (Müller); Colalongo & Pasini: 58,
pl. 22, fig. 2.
1988 Kangarina abyssicola (Müller); Guernet & Fourcade: 145,
pl. 4, fig. 12.
non 1993 Kangarina abyssicola (Müller); Witte: 43, pl. 9,
figs 25–26.
1996 Kangarina abyssicola (Müller); Coles et al.: 135, pl. 2,
figs 10–11.
2004 Kangarina? abyssicola (Müller); Aiello & Szczechura: 53,
pl. 8, fig. 15.
2005 Kangarina abyssicola (Müller); Guernet: 103.
2005 Kangarina coarctata Ruggieri; Guernet: 103.
2009b Kangarina cf. abyssicola (Müller); Yasuhara et al.: 914,
p. 14, fig. 13.
2010 Kangarina abyssicola (Müller); Aiello & Barra: 412.
Remarks. We consider Kangarina coarctata Ruggieri, 1953 and
K. septentrionalis Neale, 1972 as junior synonyms of K. abyssicola
(Müller, 1894). K. abyssicola is known from the Mediterranean and
the eastern and western North Atlantic.
Genus Pedicythere Eagar, 1965
Type species. Pedicythere tessae Eagar, 1965
Remarks. We found four Pedicythere species in ODP 982A, all
of which also occur in the western North Atlantic (Yasuhara
et al., 2009b).
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Pedicythere atroposopetasi Yasuhara, Okahashi & Cronin, 2009
(Figs 11F–I, 12A–D)
?2000 Pedicythere sp. B Guernet & Bellier: 270, pl. 5, fig. 3.
2009 Pedicythere atroposopetasi Yasuhara, Okahashi & Cronin:
914, pl. 15, figs 1–13.
Pedicythere kennettopetasi Yasuhara, Okahashi & Cronin, 2009
(Fig. 13E–F)
?2000 (part) Pedicythere sp. A Guernet & Bellier: 270, pl. 5, fig. 2
(non fig. 1).
2009 Pedicythere kennettopetasi Yasuhara, Okahashi & Cronin:
916, pl. 16, figs 1–10.
Pedicythere klothopetasi Yasuhara, Okahashi & Cronin, 2009
(Figs 13G–J, 14A–E)
2009 Pedicythere klothopetasi Yasuhara, Okahashi & Cronin:
916, pl. 15, figs 14–21.
Fig. 10. SEM images of Cytheropteron and Eucytherura species. A–B, Cytheropteron perlaria Hao, 1988: A, USNM 603706 (ODP982033); adult LV
from 1/2/17–19; B, USNM 603707 (ODP982034); adult RV from 1/2/17–19. C–D, Cytheropteron pherozigzag Whatley, Ayress & Downing, 1986: C,
USNM 603708 (ODP982032); adult LV from 1/1/142–144; D, USNM 603709 (ODP982031); adult RV from 1/2/17–19. E, Cytheropteron pseudoalatum
Colalongo & Pasini, 1980, USNM 603710 (ODP982094); adult LV from 1/2/127–129. F–H, Eucytherura calabra (Colalongo & Pasini, 1980): F,
USNM 603711 (ODP982100); adult LV from 1/1/70–72; G, USNM 603712 (ODP982101); adult LV from 1/1/90–92; H, USNM 603713 (ODP982102);
adult RV from 1/1/107–109. I–J, Eucytherura multituberculata Ayress, Whatley, Downing & Millson, 1995: I, USNM 603714 (ODP982004); adult
RV from 1/3/112–114; J, USNM 603715 (ODP982005); adult LV from 1/3/92–94. A–F, H–J, lateral views; G, internal view. All specimens from late
Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic. Scale bars represent 0.5 mm.
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M. Yasuhara & H. Okahashi
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Pedicythere lachesisopetasi Yasuhara, Okahashi & Cronin, 2009
(Figs 12E–J, 13A–D)
1983 Pedicythere sp. A Cronin: 110, pl. 4H.
2008 Pedicythere sp. Bergue & Coimbra: 130, pl. 6, fig. 13.
2009 Pedicythere lachesisopetasi Yasuhara, Okahashi & Cronin:
918, pl. 16, figs 11–21.
Family Eucytheridae Puri, 1954
Genus Eucythere Brady, 1868
Type species. Cythere declivis Norman, 1867 (designated by
Brady & Norman, 1889; see Horne & Whittaker (1985) for
details and lectotype).
Eucythere triangula Whatley & Coles, 1987
(Fig. 14F)
1987 Eucythere triangula Whatley & Coles: 74, pl. 4, figs 16–18.
2000 Eucythere triangula Whatley & Coles; Didié & Bauch: 114,
pl. 3, fig. 21.
Fig. 11. SEM images of Eucytherura, Kangarina and Pedicythere species. A–C, Eucytherura tetrapteron (Bonaduce, Ciampo & Masoli, 1976):
A, USNM 603716 (ODP982001); adult LV from 1/3/112–114; B, USNM 603717 (ODP982002); adult RV from 1/3/112–114; C, USNM 603718
(ODP982003); adult LV from 1/3/112–114. D–E, Kangarina abyssicola (Müller, 1894): D, USNM 603719 (ODP982104); adult LV from 1/1/137–139;
E, USNM 603720 (ODP982103); adult RV from 1/1/70–72. F–I, Pedicythere atroposopetasi Yasuhara, Okahashi & Cronin, 2009: F–G, USNM 603721
(ODP982136); adult LV from 1/3/82–84; H–I, USNM 603722 (ODP982137); adult RV from 1/3/82–84. A–B, D–F, H, lateral views; C, internal view;
G, I, oblique views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic. Scale bar represents
0.5 mm.
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2009 Eucythere triangula Whatley & Coles; Alvarez Zarikian: 4,
pl. P6, fig. 4.
2009b Eucythere triangula Whatley & Coles; Yasuhara et al.:
920, pl. 17, figs 2–7.
Remarks. This species is known both from the eastern and west-
ern North Atlantic.
Family Krithidae Mandelstam, 1958 (in Bubikyan, 1958)
Genus Krithe Brady, Crosskey & Robertson, 1874
Type species. Ilyobates praetexta Sars, 1866
Remarks. Krithe is one of the most abundant genera in this
core, representing on average 20% of the total fauna. We
followed the taxonomy of Coles et al. (1994). Krithe in this
core is mainly composed of K. dolichodeira van den Bold,
1946, K. ayressi Coles et al., 1994 and K. minima Coles et al.,
1994.
Family Leptocytheridae Hanai, 1957
Genus Cluthia Neale, 1973
Type species. Cythere cluthae Brady, Crosskey &
Robertson, 1874
Fig. 12. SEM images of Pedicythere species. A–D, Pedicythere atroposopetasi Yasuhara, Okahashi & Cronin, 2009: A, USNM 603723 (ODP982138);
adult RV from 1/1/70–72; B, USNM 603724 (ODP982139); adult LV from 1/3/92–94; C–D, USNM 603725 (ODP982146); adult RV from 1/1/142–
144. E–J, Pedicythere lachesisopetasi Yasuhara, Okahashi & Cronin, 2009: E–F, USNM 603726 (ODP982140); adult RV from 1/1/97–99; G–H,
USNM 603727 (ODP982141); adult LV from 1/3/122–124; I–J, USNM 603728 (ODP982142); adult RV from 1/3/122–124. C, E, G, I, lateral views;
A–B, internal views; D, F, H, J, oblique views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic.
Scale bar represents 0.5 mm.
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M. Yasuhara & H. Okahashi
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Cluthia sp.
(Fig. 14G–J)
1998 Nannocythere sp. Whatley, Eynon & Moguilevsky: 23, pl.
3, figs 9–10.
?2000 Nannocythere sp. Didié & Bauch: 111, pl. 4, fig. 25.
Remarks. This species is formally described as new in Yasuhara
et al. (in press b).
Family Loxoconchidae Sars, 1926
Genus Loxoconchidea Bonaduce, Ciampo & Masoli, 1976
Type species. Loxoconchidea minima Bonaduce, Ciampo &
Masoli, 1976
Loxoconchidea minima Bonaduce, Ciampo & Masoli, 1976
(Fig. 15A)
1976 Loxoconchidea minima Bonaduce, Ciampo & Masoli: 112,
pl. 59, figs 1–7, text-fig. 43.
Fig. 13. SEM images of Pedicythere species. A–D, Pedicythere lachesisopetasi Yasuhara, Okahashi & Cronin, 2009: A, USNM 603729
(ODP982144); adult RV from 1/3/122–124; B, USNM 603730 (ODP982145); adult LV from 1/1/60–62; C–D, USNM 603731 (ODP982143); adult
LV from 1/3/122–124. E–F, Pedicythere kennettopetasi Yasuhara, Okahashi & Cronin, 2009, USNM 603732 (ODP982147); adult LV from 1/3/72–
74. G–J, Pedicythere klothopetasi Yasuhara, Okahashi & Cronin, 2009: G, USNM 603733 (ODP982148); adult LV from 1/1/80–82; H, USNM
603734 (ODP982149pedi); adult RV from 1/1/80–82; I–J, USNM 603735 (ODP982150); adult RV from 1/1/80–82. C, E, H, J, lateral views; A–B,
G, internal views; D, F, I, oblique views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic.
Scale bar represents 0.5 mm.
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2000 Loxoconchidea minima Bonaduce, Ciampo & Masoli; Aiello
et al.: 97, pl. 3, fog. 10.
2004 Loxoconchidea minima Bonaduce, Ciampo & Masoli; Aiello
& Szczechura: 35, pl. 7, figs 1–3.
2006 Loxoconchidea minima Bonaduce, Ciampo & Masoli;
Bergue et al.: 206, fig. 6E.
2008 Loxoconchidea minima Bonaduce, Ciampo & Masoli;
Bergue & Coimbra: 115, pl. 1, fig. 16.
2009b Loxoconchidea minima Bonaduce, Ciampo & Masoli;
Yasuhara et al.: 920, pl. 17, figs 8–11.
Remarks. A comprehensive synonymy list can be found in Aiello
& Szczechura (2004) and Yasuhara et al. (2009b). This species is
known from the Atlantic and Mediterranean regions.
Family Paradoxostomatidae Brady & Norman, 1889
Genus Paracytherois Müller, 1894
Type species. Paracytherois striata Müller, 1894 [designated by
Howe, 1955 (he considered this species a junior synonym of
Paradoxostoma flexuosum Brady (1868) (sic: correctly, Bythocythere?
flexuosa Brady, (1867)); see Ellis & Messina Catalogue].
Fig. 14. SEM images of Pedicythere, Eucythere and Cluthia species. A–E, Pedicythere klothopetasi Yasuhara, Okahashi & Cronin, 2009: A, USNM
603736 (ODP982151); adult RV from 1/1/80–82; B–C, USNM 603737 (ODP982152); adult LV from 1/1/70–72; D–E, USNM 603738 (ODP982153);
adult RV from 1/1/70–72. F, Eucythere triangula Whatley & Coles, 1987, USNM 603739 (ODP982099); adult LV from 1/2/117–119. G–J, Cluthia sp.: G,
USNM 603740 (ODP982127-2); adult LV from 1/3/12–14; H, USNM 603741 (ODP982128); adult RV from 1/3/92–94; I, USNM 603742 (ODP982129-1);
adult LV from 1/3/92–94; J, USNM 603743 (ODP982130); adult RV from 1/3/92–94. C–D, F–H, lateral views; A, I–J, internal views; B, E, oblique views.
All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic. Scale bars represent 0.5 mm.
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M. Yasuhara & H. Okahashi
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Paracytherois bondi Yasuhara, Okahashi & Cronin, 2009
(Fig. 15B–C)
2009b Paracytherois bondi Yasuhara, Okahashi & Cronin: 924,
pl. 19, figs 5–10, 15 (?12).
Remarks. This species was known only from the western North
Atlantic, but this record confirms its presence in the eastern North
Atlantic.
Family Rockalliidae Whatley, Uffenorde, Harlow, Downing &
Kesler, 1982
Genus Arcacythere Hornibrook, 1952
Type species. Arcacythere chapmani Hornibrook, 1952
Remarks. We agree with Ayress (1991) in considering Rockallia
Whatley, Frame & Whittaker, 1978 as a junior synonym of
Arcacythere Hornibrook, 1952. See Yasuhara & Okahashi (in
press) for detailed discussion.
Fig. 15. SEM images of Loxoconchidea, Paracytherois, and Arcacythere species. A, Loxoconchidea minima Bonaduce, Ciampo & Masoli, 1976, USNM
603744 (ODP982123); adult LV from 1/3/12–14. B–C, Paracytherois bondi Yasuhara, Okahashi & Cronin, 2009: USNM 603745 (ODP982135); adult
RV from 1/1/80–82. D–I, Arcacythere enigmatica (Whatley, Frame & Whittaker, 1978): D, USNM 603746 (ODP982169rock); adult LV from 1/2/77–
79; E, USNM 603747 (ODP982170rock); adult RV from 1/2/77–79; F, USNM 603748 (ODP982171rock); adult LV from 1/2/77–79; G, USNM 603749
(ODP982172rock); adult RV from 1/2/77–79; H, USNM 603750 (ODP982173rock); adult RV from 1/2/67–69; I, USNM 603751 (ODP982174rock);
adult LV from 1/3/92–94. A–G, lateral views; H–I, internal views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau,
eastern North Atlantic. Scale bars represent 0.5 mm for A–B, D–I and 0.1 mm for C.
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Arcacythere enigmatica (Whatley, Frame & Whittaker, 1978)
(Fig. 15D–I)
1978 Rockallia enigmatica Whatley, Frame & Whittaker: 137, pls
5-138, 5-140, 5-142, 5-144; text-fig. 1.
1979 Indet. Gen. 3 Ducasse & Peypouquet: pl. 5, fig. 9.
1982 Rockallia enigmatica Whatley, Frame & Whittaker; Whatley
et al.: 3, pl. 1, figs 1, 4.
1987 Rockallia enigmatica Whatley, Frame & Whittaker; Whatley
& Coles: 80, pl. 2, figs 3–4.
1987 Rockallia sp. Whatley & Coles: 89, pl. 2, fig. 5.
1988 Rockallia enigmatica Whatley, Frame & Whittaker; Ruan &
Hao: 377, pl. 70, figs 2–4.
1988 Rockallia inceptiocelata Whatley, Uffenorde, Harlow,
Downing & Kesler; Ruan & Hao: 377, pl. 70, figs 5–7.
1990 Rockallia enigmatica Whatley, Frame & Whittaker; Malz:
143, fig. 4.2.
2000 Rockallia enigmatica Whatley, Frame & Whittaker; Didié &
Bauch: 116, pl. 3, figs 13–14.
Fig. 16. SEM images of Echinocythereis and Henryhowella species. A–D, Echinocythereis echinata (Sars, 1866): A, USNM 603752 (ODP982027);
adult RV from 1/1/142–144; B, USNM 603753 (ODP982028); adult RV from 1/1/142–144; C, USNM 603754 (ODP982029); adult LV from 1/3/72–74;
D, USNM 603755 (ODP982030); adult LV from 1/3/72–74. E–K, Henryhowella asperrima (Reuss, 1850): E, USNM 603756 (ODP982022); adult LV
from 1/1/137–139; F, USNM 603757 (ODP982023); adult RV from 1/1/137–139; G, USNM 603758 (ODP982026); adult LV from 1/1/0–2;
H–I, USNM 603759 (ODP982024); adult RV from 1/1/142–144; J–K, USNM 603760 (ODP982025); adult LV from 1/1/107–109. A, C, E–G, I, K,
lateral views; B, D, H, J, internal views. All specimens from late Quaternary section of ODP Hole 982A, Rockall Plateau, eastern North Atlantic. Scale
bar represents 1 mm.
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M. Yasuhara & H. Okahashi
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2003 Rockallia enigmatica Whatley, Frame & Whittaker; Cronin
& Dwyer: 263, pl. 2, fig. n.
2005 Rockallia enigmatica Whatley, Frame & Whittaker; Mazzini:
86, figs 50P, 51B.
2007 Rockallia enigmatica Whatley, Frame & Whittaker; Hou &
Gou: 509, pl. 198, figs 1–4.
2007 Rockallia inceptiocelata Whatley, Uffenorde, Harlow,
Downing & Kesler; Hou & Gou: 509, pl. 198, figs 5, 8 (?6–7).
2009 Rockallia enigmatica Whatley, Frame & Whittaker; Alvarez
Zarikian: 5, pl. P9, fig. 5.
Remarks. This species is known from the eastern North Atlantic
and northwestern Pacific oceans.
Family Trachyleberididae Sylvester-Bradley, 1948
Remarks. Ambocythere, Buntonia and Pennyella occur in this
core, but we will discuss these genera elsewhere.
Genus Echinocythereis Puri, 1954
Type species. Cythere margaritifera Brady, 1870 [=Cythereis
garretti Howe & McGuirt, 1935 (in Howe & graduate students,
1935); see Hazel (1967)].
Echinocythereis echinata (Sars, 1866)
(Fig. 16A–D)
1866 Cythereis echinata Sars: 44.
1880 Cythere irpex Brady: 107, pl. 17, fig. 2a–d.
?1967 Echinocythereis echinata (Sars); Hazel: 37, pl. 6, figs 10–11.
1976 Cythere irpex Brady; Puri & Hulings: 278, pl. 11, figs 1–9.
1990 Echinocythereis whatleyi Dingle, Lord & Boomer: 303, figs
35B–F, 36E–G, I–J.
2000 Echinocythereis echinata (Sars); Barra & Bonaduce: 214, pl.
1, figs 1–10; text-fig. 1.
2004 Echinocythereis echinata (Sars); Ayress et al.: 35, pl. 3,
fig. 9.
2009 Echinocythereis echinata (Sars); Alvarez Zarikian: 6, pl. P9,
figs 3–4.
2009b Echinocythereis echinata (Sars); Yasuhara et al.: 926,
pl. 21, figs 6–9.
Remarks. A comprehensive synonymy and detailed discussion
are found in Yasuhara et al. (2009b) and references therein. This
species is known from the Atlantic and Southern Oceans.
Genus Henryhowella Puri, 1957
Type species. Cythere evax Ulrich & Bassler, 1904
Henryhowella asperrima (Reuss, 1850)
(Fig. 16E–K)
1850 Cypridina asperrima Reuss: 74, pl. 10, fig. 5a–b.
2005 Henryhowella asperrima (Reuss); Mazzini: 50, figs 26A–I, 27B.
2009 Henryhowella dasyderma (Brady); Alvarez Zarikian: 6,
pl. 9, figs 6–8.
2009b Henryhowella cf. asperrima (Reuss); Yasuhara et al.: 926,
pl. 20, fig. 7; pl. 21, figs 1–4.
2010 Henryhowella asperrima (Reuss); Bergue & Govindan: 751,
fig. 3.14.
2011 Henryhowella asperrima (Reuss); Pirkenseer & Berger: 54,
pl. 7, figs 6a–6c, 7a–7c; pl. 8, figs 1a–1c, 2a–2c, 3a–3c.
Remarks. To be discussed in detail elsewhere.
DISCUSSION
ODP 982A late Quaternary ostracod assemblages show a strong
affinity to the western North Atlantic and Mediterranean bathyal
faunas and, to a lesser extent, to the western North Pacific bathyal
fauna. There are many common or closely related species among
these regions, as shown in the Systematic Palaeontology section
above. The strong affinity among the western North Atlantic, the
eastern North Atlantic and the Mediterranean faunas is under-
standable, given their proximity and bathyal-depth connections
(e.g. Strait of Gibraltar and Greenland–Iceland–Faeroe Ridge).
However, an affinity between the North Atlantic and the North
Pacific is more difficult to explain. Two possibilities are pro-
posed: (1) many bathyal ostracod species are cosmopolitan; or (2)
they were able to migrate through the Bering Strait despite its
present-day shallow depth (<50 m). We will need further modern
and palaeo-biogeographical studies to evaluate these hypotheses.
ACKNOWLEDGEMENTS
We thank D. A. Hodell for kindly loaning ODP 982A samples; L.
M. Y. Wong and C. Sanford for continuous support; S. Whittaker
for help with SEM imaging; G. Hunt for help in depositing type and
figured specimens; C. Alvarez Zarikian and an anonymous reviewer
for valuable comments. Samples used for this research were pro-
vided by the Integrated Ocean Drilling Program (IODP). This work
was supported by the Seed Funding Programme for Basic Research
of the University of Hong Kong (project codes: 201105159002;
201210159043), Hung Hing Ying Physical Sciences Research Fund
2012–13, Smithsonian Postdoctoral Fellowship, and Smithsonian
Marine Science Network Postdoctoral Fellowship (to M.Y.).
Manuscript received 8 September 2013
Manuscript accepted 15 January 2014
Scientific Editing by Alan Lord.
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