JMJournal of MicropalaeontologyJMJ. Micropalaeontol.2041-4978Copernicus PublicationsGöttingen, Germany10.5194/jm-37-25-2018“Live” (stained) benthic foraminiferal living depths, stable isotopes, and
taxonomy offshore South Georgia, Southern Ocean: implications for
calcification depthsImplications for calcification depthsDejardinRowanrowan.dejardin@nottingham.ac.ukKenderSevhttps://orcid.org/0000-0003-4216-3214AllenClaire S.https://orcid.org/0000-0002-0938-0551LengMelanie J.https://orcid.org/0000-0003-1115-5166SwannGeorge E. A.https://orcid.org/0000-0002-4750-9504PeckVictoria L.Centre for Environmental Geochemistry, School of Geography, University
of Nottingham, University Park, Nottingham, NG7 2RD, UKCamborne School of Mines, University of Exeter, Penryn, Cornwall TR10 9FE,
UKBritish Geological Survey, Keyworth, Nottingham NG12 5GG, UKBritish Antarctic Survey, High Cross, Madingley Road, Cambridge, CB3
0ET, UKNERC Isotope Geosciences Facilities, British Geological Survey,
Keyworth, Nottingham, NG12 5GG, UKRowan Dejardin (rowan.dejardin@nottingham.ac.uk)5January20183712571This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit https://creativecommons.org/licenses/by/4.0/This article is available from https://jm.copernicus.org/articles/37/25/2018/jm-37-25-2018.htmlThe full text article is available as a PDF file from https://jm.copernicus.org/articles/37/25/2018/jm-37-25-2018.pdf
It is widely held that benthic foraminifera exhibit species-specific
calcification depth preferences, with their tests recording sediment pore
water chemistry at that depth (i.e. stable isotope and trace metal
compositions). This assumed depth-habitat-specific pore water chemistry
relationship has been used to reconstruct various palaeoenvironmental
parameters, such as bottom water oxygenation. However, many deep-water
foraminiferal studies show wide intra-species variation in sediment living
depth but relatively narrow intra-species variation in stable isotope
composition. To investigate this depth-habitat–stable-isotope relationship on
the shelf, we analysed depth distribution and stable isotopes of “living” (Rose Bengal stained) benthic foraminifera from two box cores collected on
the South Georgia shelf (ranging from 250 to 300 m water depth). We provide a
comprehensive taxonomic analysis of the benthic fauna, comprising 79
taxonomic groupings. The fauna shows close affinities with shelf assemblages
from around Antarctica. We find “live” specimens of a number
of calcareous species from a range of depths in the sediment column. Stable isotope ratios
(δ13C and δ18O) were measured on stained specimens of
three species, Astrononion echolsi, Cassidulinoides porrectus, and Buccella sp. 1, at 1 cm depth intervals within the
downcore sediment sequences. In agreement with studies in deep-water
settings, we find no significant intra-species variability in either
δ13Cforam or δ18Oforam with sediment living depth on the
South Georgia shelf. Our findings add to the growing evidence that infaunal
benthic foraminiferal species calcify at a fixed depth. Given the wide range
of depths at which we find “living”, “infaunal” species, we speculate that
they may actually calcify predominantly at the sediment–seawater interface,
where carbonate ion concentration and organic carbon availability is at a
maximum.
Introduction
Benthic foraminifera live both on (epifaunal) and beneath (infaunal) the
sediment–seawater interface, and their assemblages and stable isotope and trace
element compositions are widely utilized as tools in the reconstruction of
past oceanographic conditions (see review in Jorissen et al., 2007). Stable
isotopes of all species typically exhibit inter-species offsets. In part this
offset is accounted for by “vital effects”, inter-species differences in
the fractionation of stable isotopes due to a range of biological factors
(see review in Ravelo and Hillaire-Marcel, 2007). Another consideration,
adopted in numerous studies, is that epifaunal and infaunal inter-species
isotopic offsets reflect, in part, pore water chemistry at the preferred
depth habitat of each species within the sediment (e.g.
Loubere et al., 1995; McCorkle et al., 1997; Theodor et al., 2016). The carbon isotope composition
of pore water dissolved inorganic carbon (δ13CDIC) has
been shown to become lighter (lower δ13C) with sediment depth
(e.g. McCorkle et al., 1985), due to the remineralization (by oxic
respiration or denitrification) of organic carbon. The difference between
bottom and pore water δ13CDIC has been shown to
increase with bottom water oxygenation (McCorkle and Emerson, 1988). Assuming
that infaunal species calcify at preferred depths within the sediment
hypothetically allows δ13CDIC gradients to be
reconstructed through the isotope analysis of paired epifaunal and deep-infaunal taxa (e.g. Cibicidoides wuellerstorfi and
Globobulimina spp.; Hoogakker et al., 2015). Similarly, the
difference in δ13C between epifaunal and shallow infaunal species
(e.g. Uvigerina spp.) has been used to reconstruct the intensity of
organic matter remineralization, a proxy for the flux of organic matter to
the sea floor (Zahn et al., 1986; McCorkle and Emerson, 1988; Schilman et al., 2003). Additionally, Elderfield et al. (2012) used the presence of a
constant offset between the δ13C data from paired epifaunal
(Cibicidoides wuellerstorfi) and infaunal (Uvigerina spp.)
foraminifera to negate the suggestion that a negative δ13C
excursion during Marine Isotope Stage 22 was due to productivity changes and
to support their hypothesis that the δ13C excursion was instead
related to global ocean change.
These applications rely on the previously stated assumption that benthic
foraminifera inhabit species-specific depth habitats and, more importantly,
calcify at that depth, thereby recording the pore water δ13CDIC gradient. This assumption is held despite the wide
range of sediment depths that individual infaunal species have been observed
to live at (e.g. McCorkle et al., 1997). Furthermore, isotope analysis of
“live” (Rose Bengal stained) foraminifera reveals a narrow range of isotope
values for individuals of the same species despite them having been recovered
from a wide range of depths beneath the sediment surface (McCorkle et al., 1990, 1997; Schmiedl et al., 2004; Fontanier et al., 2006b, 2008;
Theodor et al., 2016). A narrow intra-species range of δ13C composition of
“live” specimens recovered from a range of sediment depths argues against
individuals having calcified at the depths from which they were recovered
(since pore water δ13CDIC gradients are not reflected
in the foraminifera). More recent studies raise further questions about the
validity of the assumed calcification depths of infaunal species. For
example, it has been assumed that Mg / Ca data from infaunal foraminifera are
relatively insulated from the effect of bottom water [CO32-] changes
(Elderfield et al., 2006, 2010). However, Weldeab et al. (2016) have shown
that infaunal Globobulimina spp. are sensitive to changes in bottom
water [CO32-], implying that calcification may not occur exclusively
beneath the sediment surface. To extend previous studies of the deep ocean
(e.g. McCorkle et al., 1990, 1997; Tachikawa and Elderfield, 2002; Schmiedl
et al., 2003; Fontanier et al., 2008, 2006b) onto the shelf environment, and
test the assumption that infaunal species calcify at preferred depths and
that their δ13C composition is reflective of pore water
δ13CDIC (i.e. decreasing δ13CDIC
with increasing calcification depth), we measured the living depths and
isotope composition of three as yet unstudied species (Astrononion echolsi, Cassidulinoides porrectus, and Buccella sp. 1)
from the South Georgia shelf, South Atlantic. We also report the depth ranges
of all “living” (Rose Bengal stained) benthic foraminifera. To underpin
these and future analyses, we carried out a detailed taxonomic analysis of
benthic foraminifera from six surface samples, two box-core downcore
sequences, and two gravity cores, the first such study focused on South
Georgia since Earland (1933).
Previous benthic foraminiferal studies
Earland (1933) conducted a survey of the benthic foraminifera
of South Georgia, utilizing surface sediment samples collected from around
the island as part of the Discovery expeditions. Since this study,
there has been no further taxonomic work focussed on South Georgia, but a
number of benthic foraminiferal assemblage studies have been conducted at
other Southern Ocean and Antarctic locations. The Antarctic Peninsula is one
of the better-studied areas in the region, with foraminiferal research here
commencing in the first half of the 20th century (Earland, 1934; Cushman,
1945), followed by a number of brief US Antarctic Program taxonomic reports
(e.g. Lipps et al., 1972). In a more comprehensive study of the Antarctic
Peninsula, Ishman and Domack (1994) analysed a large number of surface
sediment samples from three areas on the western side of the peninsula
(Bransfield Strait, Marguerite Bay, and Palmer Archipelago). A detailed
assessment of the benthic foraminiferal assemblages of Admiralty Bay, King
George Island, and the South Shetland Islands has been conducted by
Majewski (2005, 2010), again on surface sediment samples. On the eastern side
of the Antarctic Peninsula, Majewski and Anderson (2009) examined the
palaeoclimatic implications of Holocene foraminiferal assemblages from the
Firth of Tay. Further east, Anderson (1975) identified 160 species in surface
samples distributed across the Weddell Sea, whilst Mackensen et al. (1990)
focussed on surface samples from the sea's eastern corner.
Map of South Georgia and sampling locations. The location of South
Georgia in relation to the Antarctic Peninsula and South America showing the
approximate location of the Antarctic Polar Front (APF – light blue) and the
Southern Antarctic Circumpolar Current Front (SACCF – dark blue). The inset
and map on the left show a topographical map of South Georgia and surrounding bathymetry with
the sampling locations indicated. The dark blue lines represent the location
and average position of the SACCF (contemporary variation near South Georgia
shown by the dashed lines, indicating most easterly and westerly
extents (Thorpe et al., 2002). The mean surface chlorophyll-α
concentration (mg m-2) between September and March is shown in green
with darker colours indicating higher concentration (SeaWiFS derived
satellite data, NASA; means calculated only for cells where 10 values were
obtained within a year; from Murphy et al., 2013).
Herb (1971) described the variation in benthic faunas across the Drake
Passage, illustrating the differences between the shelf environments of
southern South America and Antarctica. Mackensen et al. (1993) analysed a
similar transect of surface samples through the subantarctic zone between 0
and 10∘ E. Useful studies from Southern Ocean localities more
distant from South Georgia include Majewski's (2013) description of benthic
foraminifera from core-top samples around Pine Island and Ferrero Bays in the
Amundsen Sea, Violanti's (1996) analysis of live and dead foraminifera from
surface samples from Terra Nova Bay, Ross Sea, and Igarashi et al.'s (2001)
study of samples from sediment cores extending into the Holocene from
Lützow-Hom Bay, East Antarctica.
Oceanographic setting
The subantarctic island of South Georgia (54∘ S, 36∘ W)
is located at the northern edge of the Scotia Sea in the path of the
Antarctic Circumpolar Current (ACC), between the Antarctic Polar Front (APF)
and the Southern ACC Front (SACCF) (Fig. 1). South Georgia is one of the
largest islands in the subantarctic zone (SAZ), measuring approximately
180 × 40 km, and is separated from the deep ocean by a
50–150 km wide continental shelf. The shelf, typically within 200–250 m
water depth, is traversed by canyons that connect the island's fjords to the
continental slope. The properties of the waters of the shelf itself routinely
differ from those of the surrounding ocean, with on-shelf surface waters
typically being warmer (2–3 ∘C on shelf, 1.5–2 ∘C off
shelf) and fresher (salinity 33.4–33.7 psu on shelf, 33.6–33.7 psu off
shelf) (Brandon et al., 1999). These differences are caused by a number of
shelf-specific processes, such as freshwater run-off and topographically
steered circulation (Brandon et al., 1999, 2000; Meredith et al., 2005).
Sample locations.
Core no.LatitudeLongitudeWater depth (m)SampleBC656-53.78168-37.95855303Surface sampleBC660-53.78188-38.12847354Surface sampleBC667-54.42063-35.74152252Surface sampleBC671-54.20237-36.37451239Surface sampleBC737-53.99709-36.82012260Surface sampleBC737 (D)-53.99709-36.8201226015 cm sub-coreBC738-53.85504-37.65095287Surface sampleBC738 (D)-53.85504-37.6509528720 cm sub-coreGC666*-54.4206-35.74148253815 cm gravity coreGC673*-54.20237-36.37451238884 cm gravity core
* Samples with foram species not present in the
box cores are included in Sect. 6 and Supplement.
MethodsSample acquisition and preparation
Two box cores (BC737 and BC738) were collected during British Antarctic
Survey scientific cruise JR-15002, in November 2015 (see Table 1 and Fig. 1),
from which two sub-cores were collected (BC737 (D) and BC738 (D)). The box
cores were collected using the British Antarctic Survey (BAS) box corer,
deployed from the RRS James Clark Ross. The sub-cores were taken
from BC737 (15 cm) and BC738 (20 cm); these cores were cut into 1 cm
sections which were weighed to the nearest gram before being washed over a
63 µm sieve. The coarse fraction was then soaked in a Rose
Bengal / seawater solution (1 g : 1 L) for 24 h before being rinsed
with seawater from the underway seawater supply until the water ran clear
and dried at 40∘ C after a final rinse with MilliQ water. Rose
Bengal is a non-vital biological stain that is adsorbed onto any proteins,
whether or not they are alive, and can therefore stain necrotic as well as
healthy cytoplasm (Bernhard, 2000).
Benthic foraminiferal assemblage analysis
All “living” (stained) and dead foraminifera were dry picked from each of
the 1 cm samples between 0 and 10 cm, and only “living” (stained)
foraminifera were dry picked from each of the 1cm samples between 10 and
13 cm, from BC737 and BC738. Tests where all, or all but one or two,
chambers were well stained bright red were considered to be “living” at the
time of staining (e.g. Caulle et al., 2015; Fontanier et al., 2016; Wang et
al., 2016). For thick-walled agglutinated taxa, where the opaque test wall
concealed the extent of the staining, the tests were broken open allowing the
staining to be assessed. Tests with a pink tint or isolated spots of red were
not included, nor were tests where only a small number of isolated chambers
were stained. The average living depth (ALD) (Jorissen et al., 1995) was
calculated for the 10 taxa with at least five “live” specimens in one or
more samples. We also calculated the median living depth (MLD) (Theodor et al.,
2016) for these taxa. Differences between ALD and MLD are generally small but
can be significant for taxa with a wide downcore occurrence range (Theodor et
al., 2016). Foraminiferal abundances are reported as number per gram.
Downcore “live” : dead ratio and histograms showing downcore
abundance of “live” (stained) foraminifera. BC7373 (dark grey) and BC738
(light grey) downcore “live” abundances (numbers per gram):
(a) downcore “live” : dead ratio; (b) total calcareous taxa;
(c–f) calcareous taxa with five or more “live” specimens in at least
one sample; (g) total agglutinated taxa;
(h–m) agglutinated taxa with five or more “live” specimens in at
least one sample.
Stable isotope analysis
Oxygen and carbon isotope analyses were carried out on three species of
foraminifera from BC737 (Astrononion echolsi, Cassidulinoides porrectus, and Buccella sp. 1), from 10 samples (0–1 to
9–10 cm), and on two species from BC738 (Astrononion echolsi and
Cassidulinoides porrectus) from 6 samples (0–1 to 5–6 cm). All
specimens for isotope analysis were picked from the 125–250 µm
fraction to minimize the possible effects of ontogenetic change in
fractionation rates. These species were selected because their ALD/MLD
revealed different microhabitat preferences and because they were present in
high enough abundance to provide sufficient calcite to conduct the analyses.
Each isotope data point was measured on between 4 and 15 specimens from each
sample for A. echolsi and C. porrectus and on 3–5 specimens from each sample for the more heavily calcified
Buccella sp. 1. The measurements were carried out at the NERC
Isotope Geosciences Laboratory (NIGL), British Geological Survey (Keyworth,
UK) using an IsoPrime 100 dual-inlet mass spectrometer with Multiprep device.
Isotope values were calculated according to the Vienna Pee Dee Belemnite (VPDB) scale
in parts per thousand using a within-run laboratory standard (KCM) calibrated
to international NBS standards. Analytical reproducibility was 0.1 ‰
(1σ) for both δ18O and δ13C. To eliminate
contamination by any organic carbon remaining in the test (the presence of
the stain indicates that organic carbon is present within the tests), the
samples were plasma ashed prior to analysis (at NIGL using an Emitech K1050X
plasma asher). To test whether the plasma ashing process had any impact on
the isotope composition of the foraminiferal calcite, five standards (KCM)
were subjected to the same processing and analysed in parallel. In addition,
to test whether the Rose Bengal stain had any impact on the isotope
composition, five standards (KCM) mixed with Rose Bengal and five KCM
standards mixed with Rose Bengal and plasma ashed were also analysed.
ResultsBenthic foraminiferal assemblage results
A total of 1181 “live” (Rose Bengal stained) and 6505 dead specimens were
identified from the sub-core sample sequences, representing 62 (36 “living” and 60 dead) taxonomic groupings (See “Systematic taxonomy” section and the Supplement).
The total abundance for the upper 10 samples from the sub-cores ranges from
6 to 16 g-1 for BC737, with the highest abundance evident in the 2–3 cm sample, and from 6 to 13 g-1 for BC738, with the highest abundance
evident in the 0–1 cm sample. In 0–1 cm samples of the sub-cores, the
ratio of “live” to dead specimens is 0.42 for BC737 and 0.32 for BC738
(Fig. 2). In BC737 this ratio decreases to 0.14 in the 3–4 cm sample before
increasing to 0.37 in the 6–7 cm sample, mainly due to the high abundance
of “live” M. earlandi, and then decreasing to 0.08 in the
9–10 cm sample. In BC738 the “live” / dead ratio decreases steadily
throughout the core to 0.02 in the 0–10 cm sample, apart from a small
increase to 0.13 in the 6–7 cm sample. A small number of “live” taxa are
still present in the 12–13 cm sample of both cores (4 specimens in BC737
and one in BC738).
Histograms showing downcore abundance of dead foraminifera. BC7373
(dark grey) and BC738 (light grey) downcore “live” abundances no. g-1):
(a) total calcareous taxa; (b–e) calcareous taxa with five or
more “live” specimens in at least one sample; (f) total agglutinated taxa
excluding Miliammina spp.; (g–m) agglutinated taxa with five
or more “live” specimens in at least one sample (H. alba has not
been plotted in this figure as no dead specimens of this taxon were
recorded).
Average and median living depths.ALD (square symbols), calculated following Jorissen et al. (1995), and MLD (circular symbols), calculated following Theodor et
al. (2016), plotted for all taxa, all calcareous taxa, all agglutinated taxa,
and individually for the 10 taxa with five or more “live” specimens in at
least one sample. Error bars represent the standard deviation about the ALD
(1 SD). Closed symbols represent BC737 and open symbols represent BC738.
Note that ALD and MLD have not been calculated for F. fusiformis
and Buccella sp. 1 for BC738 as the “live” abundance of these taxa
in this core was too low.
The downcore variation in abundance of the 10 taxa with at least five
“live” specimens in one or more samples are shown in Fig. 2 (for comparison
the dead abundances of the same taxa are shown in Fig. 3). For six of these
10 taxa (A. echolsi, C. porrectus, F. fusiformis, Buccella sp. 1,
M. earlandi, and D. scoresbyi), live specimens were
recovered from a wide range of sediment depths, to at least 8 cm, in at least
one of the cores. The remaining four taxa (L. scitula, Reophax subdentaliniformis, G. rostrata, and Hippocrepinella alba) were found to be restricted to the upper 0 to 2 cm. The vertical
range in living depths for individual species is presented in Fig. 4 where
the error bars (1 SD) reveal a broad depth range of occurrences around the
ALD for all species that do not present a restricted epifaunal/shallow
infaunal microhabitat preference. There is a clear separation between the
calcareous taxa, which were all found living at a range of sediment
depths, and the agglutinated taxa, for which the majority were found
living in the upper 2 cm. The exceptions to this are M. earlandi and D. scoresbyi, which were recorded “living” at a range of
depths in BC737, as was R. subdentaliniformis in BC738. The
agreement between ALD and MLD is generally good, with a mean difference of
0.21 cm. However, in BC737 the difference between ALD and MLD for D. scoresbyi is 1.33 cm, with this difference due to the bimodal depth
distribution of “live” occurrences of this species. Whilst MLD and ALD are
generally close within the same box core, there is a systematic difference
between the living depths calculated for BC7373 and those calculated for
BC738. For four of the five taxa where the difference is > 0.5 cm, both
ALD and MLD are shallower in BC738, relative to BC737, with the exception
being R. subdentaliniformis, which is shallower in BC737.
The dead abundance of calcareous taxa in BC738 is stable throughout the core,
whilst this abundance peaks between 2 and 4 cm in BC737 before becoming
stable below this depth. This is in marked contrast to the dead abundance of
agglutinated taxa (excluding Miliammina spp.), which shows a steady
decline with depth in both cores such that, whilst agglutinated taxa are
equal to or exceed calcareous taxa in dead assemblages in the upper core,
calcareous taxa come to dominate dead assemblages as sediment depth
increases. This trend appears linked to the preservation of taxa with
epifaunal/shallow infaunal microhabitat preference as L. scitula,
R. subdentaliniformis, and G. rostrata all exhibit a steep
decline with depth in the number of dead specimens.
δ13C vs. δ18O cross-plot. Cross-plot of carbon and
oxygen isotopic ratios for all samples analysed in this study.
Astrononion echolsi – circular symbols; Cassidulinoides porrectus – square symbols; Buccella sp. 1 – triangular symbols
(closed symbols – BC737; open symbols - BC738).
Isotopic data from Rose Bengal test (STD: laboratory standard)
Results from the test isotope samples (KCM plus Rose Bengal, KCM plasma
ashed, plasma ashed KCM plus Rose Bengal), run to establish whether these
processing methods could have any impact on the foraminiferal isotope
results, revealed that the values from the treated standards for both
δ13C and δ18O were within the normal “within-run” error
(Table 2).
Isotopic data plotted against sample depth. (a) Carbon and
oxygen isotopic values for species analysed in this study.
(b) Carbon and oxygen isotopic values for seven species analysed in
McCorkle et al. (1997). (c) Carbon and oxygen isotopic values for
species analysed in this study and in McCorkle et al. (1997), normalized by
subtracting the mean value for each species from the individual value. There
is no systematic variation in either δ13Cforam or
δ18Oforam with sample depth in any of these three
plots.
The stable isotope compositions of “live” specimens display a relatively
wide range of values, from -1.20 to +0.72 ‰ for δ13C
and from +2.55 to +3.56 ‰ for δ18O (Table 3). Mean
δ13C values for Astrononion echolsi, Cassidulinoides porrectus, and Buccella sp. 1 are -0.84 ± 0.15 ‰,
-0.34 ± 0.15 ‰, and +0.54 ± 0.16 ‰
respectively; mean δ18O values are +2.85 ± 0.20 ‰,
+3.40 ± 0.09 ‰, and +3.49 ± 0.06 ‰. When
plotted on a δ13C /δ18O cross-plot (Fig. 5), samples
from the same species cluster together, irrespective of the living depth.
When δ13C is plotted against depth, it is clear that there is no
systematic variation with depth (Fig. 6a). Similarly, no trends with depth
are observed in δ18O.
DiscussionBenthic foraminiferal distributions
In this study we identified 24 of the 345 species identified by
Earland (1933), including 13 of the 24 species identified as being most
typical of South Georgia assemblages, and 7 of the 14 species Earland
described as being particularly characteristic of the region. The taxa
identified as being most abundant by Earland (1933) include many of the more
abundant species identified here, such as M. earlandi. The majority
of species recorded in this study are present at a range of locations around
the Southern Ocean, but there are five species that may be endemic to South
Georgia (e.g. Labrospira sp. 1, G. rostrata,
Recurvoides sp. 1, Buccella sp. 1, and Buccella
sp. 2). In a comprehensive review of a range a data sources, encompassing a
wide diversity of taxa, Hogg et al. (2011) highlighted the significance of
endemism within the South Georgia shelf ecosystem and its importance to
global biodiversity, and the identification of potentially unidentified
species of foraminifera further adds to that diversity. The benthic
foraminiferal assemblage documented here suggests that, whilst there may be
partial isolation, the microfauna of South Georgia are connected to those of
the rest of the Southern Ocean. A total of 20 species identified in this
study were also recorded by Ishman and Domack (1994) in their study of the
Bellingshausen Sea margin; 22 of the 105 species identified by
Majewski (2005), from King George Island, are also recorded in this study, as
are 11 of the 26 species identified by Majewski and Anderson (2009) from the
Firth of Tay. The strong affinity with Antarctic Peninsula taxa suggests that
the position of South Georgia in the path of the ACC downstream of the
Antarctic Peninsula (AP) allows for a degree of microfaunal connectivity between
the two. The species that are commonly recorded from both the AP and South
Georgia include Portatrochammina antarctica weisneri,
Reophax subdentaliniformis, A. echolsi,
Spiroplectammina biformis, Nonionella bradii, N. iridea, C. parkerianus, C. porrectus, and F. fusiformis, many of which are among the more abundant taxa. A comparable
level of similarity is seen with studies of Weddell Sea benthic foraminifera
(Mackensen et al., 1990; Anderson, 1975).
There is less similarity between the South Georgia foraminiferal fauna and
those recorded from more remote areas of the Southern Ocean. For example,
only six of the taxa recorded here were also recorded by Herb (1971) from the
Drake Passage. However, this may be due to the open-ocean/deep-water nature
of these locations, rather than proximity, as there are greater levels of
connectivity between our sites and more distal locations on the Antarctic
shelf, including Pine Island Bay and Terra Nova Bay (Majewski, 2013;
Violanti,
1996).
The decreasing abundance of “live” specimens with depth within the sediment
(Fig. 2) reflects well-established gradients in organic carbon and oxygen
availability (see Jorissen et al., 2007). The shallower ALD/MLD observed in
BC738 across almost all taxa (Fig. 4) may indicate steeper oxygen and organic
carbon concentration gradients at this location. Variation in these gradients
between the two sites may be linked to differences in sedimentation rate or
carbon flux, with the latter potentially linked to BC738's closer proximity
to seasonal phytoplankton blooms occurring north-west of South Georgia. The
interruption in the decreasing “live” abundance trend seen in the
agglutinated taxa in BC737 is the result of a peak in “live” abundance of a
single species, M. earlandi, between 5 and 7 cm.
Dead abundances in both cores peak around the ALD/MLD for all taxa combined;
the decline in dead abundances below this depth can be attributed to the
physical and chemical breakdown of the tests. The decrease in dead abundance
is more pronounced in the agglutinated taxa, indicating that the organic matrix of
agglutinated tests has a greater susceptibility to taphonomic processes. An
exception to this is M. earlandi, whose dead abundance remains
stable in both sub-cores, likely due to its siliceous cement being more
resistant to chemical breakdown (Earland, 1933). This enhanced downcore loss of
agglutinated taxa, relative to calcareous taxa, illustrates the
well-recognized complication of fossil/dead assemblages not truly reflecting
living assemblages due to the loss of taxa with poor preservation potential
(e.g. Majewski, 2005; Jorissen et al., 2007; Stefanoudis et al., 2017).
Sediment living and calcification depths
Many of the most abundant infaunal taxa recorded in BC737 and BC738 do not
appear to have a clear preference for a specific living depth. However, it is
important to recognize the potential limitations of the Rose Bengal staining
technique in identifying truly living specimens. Where decay rates are slow,
Rose Bengal may stain foraminifera that have been dead for months to several
years (Corliss and Emerson, 1990). It is therefore important to apply criteria that are as
strict as possible when categorizing which specimens are living
and which are dead (e.g. Fontanier et al., 2016; Caulle et al., 2015) and to
bear this limitation in mind when considering live abundances arrived at
through Rose Bengal staining. It is also possible for specimens identified as
living at a certain depth interval to have been transported there after
death by bioturbation (see Bernhard, 2000, and references therein). If
bioturbation were controlling the distribution of specimens within our data,
we would expect to see all taxa affected such that it would not be possible
to identify any depth-restricted habitat preferences. The restricted
distribution of L. scitula, R. subdentaliniformis, and G. rostrata to epifaunal/very shallow infaunal environments contradicts
bioturbation being prevalent at our sites. Furthermore, where decay rates are
very slow, specimens that appear live at different depths within the
sediment column may be significantly older than the overlying live
assemblages. This is dependent on the sedimentation rate being high enough to
bury specimens to a significant depth before decay is sufficiently progressed
that staining with Rose Bengal will no longer occur, and such age control in
not available for the samples studied here. Whilst it is important to
recognize the limitations of Rose Bengal staining, it is simple, reliable,
and inexpensive to use and is currently the most appropriate technique to
apply when, as in this study, the samples are processed immediately upon
collection (Bernhard, 2000). In addition, Murray and Bowser (2000) have
proposed that, in oxygenated environments, dead tests are unlikely to retain
protoplasm due to predation being the most likely cause of death. Whilst we
did not directly measure the oxygenation of the sediments studied here, there
was no indication of anoxia or dysoxia from the colour and odour of the
sediments on recovery.
δ13C vs. MLD. Mean δ13Cforam values
for the three species analysed here plotted against MLD. A. echolsi
– circular symbols; C. porrectus – square symbols; Buccella sp. 1 – triangular symbols (closed symbols – BC737; open
symbols – BC738). Error bars: 1 SD.
As discussed in the Introduction, it is commonly assumed that foraminifera
calcify at their living depth and record the pore water δ13CDIC in their test isotopic composition or an isotopic
composition offset from that of pore water by species-specific vital
effects. Whilst downcore pore water δ13CDIC data are
not available for the cores studied here, records from various other locations
consistently report that pore water δ13CDIC decreases
(becomes more negative) with increased sediment depth (e.g. McCorkle et al.,
1997; Holsten et al., 2004; Loubere et al., 2011; Luo et al., 2013), before
stabilizing below the oxygen penetration depth. Furthermore, Gehlen et
al. (1999) have modelled how the interaction of oxic organic matter decay and
CaCO3 dissolution generate this gradient within marine sediments. We
therefore assume that a negative gradient of some kind will be present in
sediments from the South Georgia shelf. The observed increase in
δ13Cforam with increased MLD (Fig. 7) is the opposite
of the expected trend were each species calcifying at their respective MLDs.
However, there are multiple vital-effect factors that lead to different
inter-species δ13C offsets despite specimens calcifying from waters
with the same δ13CDIC (see review in Ravelo and
Hillaire-Marcel, 2007), and without prior knowledge of the impact of these
vital effects for each species, we cannot use this as evidence that these
species do not calcify at their MLD.
Assessing the intra-species isotope data from the three species analysed
here, we note that the depth–δ13Cforam relationship we would
anticipate, were the foraminifera calcifying with the pore water
δ13CDIC at the depth from which they were recovered,
does not hold true; i.e. intra-species δ13C does not decrease with
depth (Fig. 6a). The range of inter-species δ13C and δ18O compositions considerably exceeds the intra-species range (Fig. 5).
Other studies that have reported isotope data from live foraminifera at a
range of living depths have also noted limited variability in the
intra-species stable isotope composition. For example, Bolivina spissa, Bulimina aculeata, Cibicidoides wuellerstorfi, Uvigerina peregrina, Pleurostomella alternans, Globobulimina affinis, and
G. pacifica (McCorkle et al., 1997) from midlatitude continental
margins in the Atlantic and Pacific oceans show no systematic downcore
isotope variations (Fig. 6b), even though the isotopic composition of the
same species can vary significantly between locations (e.g. G. affinis) (for further examples, see McCorkle et al., 1990; Tachikawa and
Elderfield, 2002; Schmiedl et al., 2003; Fontanier et al., 2006b, 2008).
When we normalize our δ13Cforam data, and
the data of McCorkle et al. (1997), by subtracting the mean for each species
at each sample location, there is no trend away from that mean with depth
(Fig. 6c) even though all pore water δ13CDIC values in
those locations (where measured) exhibit a gradient.
If the individual foraminifera were calcifying at the depth at which they
were found living (as inferred by Fontanier et al., 2006b, and Theodor et al.,
2016, amongst others), we would expect our intra-species δ13C data,
and those of other studies (e.g. McCorkle et al., 1997), to reflect a
decreasing δ13CDIC gradient with depth, which they do
not. The consistency of intra-species isotope data (Fig. 6), despite their
broad range of living depths (Figs. 2 and 6), are good evidence that benthic
foraminifera calcify at a fixed depth location, and possibly within
a specific season, because they fail to capture the variability of pore water
δ13CDIC that their observed living depth would expose
them to.
Foraminiferal calcification strategies
Many have concluded that the observed inter-species differences in δ13C are due to species preferences for different depth microhabitats, and
their position on the δ13CDIC gradient, implying that
calcification occurs infaunally at a specific, restricted depth for each
species (e.g. McCorkle et al., 1990; Schmiedl et al., 2004; Theodor et al., 2016). To account for fixed intra-species δ13C compositions of
infaunal taxa requires calcification to only occur at certain times during
the organism's life when it is exposed to a fixed
δ13CDIC value. We discuss two previously proposed
calcification strategies of infaunal species – at a specific depth within the
sediment and within macro-/meio-faunal burrows – and propose a third, novel
strategy: calcification at the sediment–seawater interface.
In the first strategy, calcification only occurs at a specific depth within
the sediment and this depth relates to the preferred microhabitat of the
species: broadly speaking epifaunal, shallow infaunal, or deep infaunal.
There is evidence for this strategy in some species: for example Holsten et
al. (2004) found that the δ13Cforam in three species
(Bolivina argentea, B. subadvena, and Buliminella tenuata)
matched the pore water δ13CDIC at their maximum live
abundance depth. However, if the species studied here calcified at a
preferred depth within the sediment, we need to question what is
controlling/determining the narrow calcification depth range (in contrast to
the observed wide living depth range) and why it is apparently not the
ALD/MLD. We note that, on the South Georgia shelf, species with deeper
ALD/MLD have heavier δ13C compositions (Fig. 7), whilst pore waters
are assumed to change (e.g. become isotopically lighter) with depth.
The second strategy involves calcification in association with the burrows of
larger fauna, where voids within the sediment allow pore water isotopic
composition to be homogenous across a range of depths (Loubere et al., 1995;
Schmiedl et al., 2004), meaning that calcification within a burrow would
result in the same δ13Cforam composition regardless of
depth. Loubere et al. (2011) have proposed that all calcareous foraminifera
use this strategy, with the activities of macro-/meio-fauna allowing
foraminifera to choose micro-habitats with particular geochemical conditions
within the complex “bio-irrigation” systems the larger fauna create.
Our third hypothesis, that calcification occurs at the sediment–seawater
interface, is supported, in principal, by the presence of live specimens
of many of the infaunal calcareous taxa recorded in this study (and e.g.
McCorkle et al., 1997) within the top 2 cm of the sediment. Foraminifera are
known to be capable of migrating throughout the sediment column (e.g. Linke
and Lutze, 1993; Geslin et al., 2004) and the occurrence of infaunal species
at these shallow depths within the sediment demonstrates their ability to
migrate to the sediment–seawater interface. Weldeab et al. (2016) have
recorded the presence of a steep [CO32-] gradient between bottom
water and pore water and also within the sediment column. This
[CO32-] gradient suggests that the sediment–seawater interface may
provide the most energy-efficient location for calcification, due to the
greater availability of carbonate ions. In addition, food (phytodetrital
matter from the overlying water column) availability and/or type is greatest
at the sediment–seawater interface. Whilst from the stable isotope data on
living specimens published here and elsewhere, it is not possible to
prove that the species were calcifying at the sediment–seawater interface, we
conclude that this may be the simplest explanation for the stability of
inter-species isotopic composition.
Alternatively, there may be no requirement to calcify at a specific depth, or
within specific geochemical conditions, and foraminifera may be capable of
calcifying at a range of sediment depths and a corresponding range of pore
water δ13CDIC compositions. Calcification across a
range of sediment depths could result in foraminiferal calcite recording an
average of the range in pore water δ13CDIC
compositions, with any inter-species variation accounted for by
species-specific vital effects. The presence of “living” specimens of
the species studied here at a range of sediment depths and the absence of a
clear maximum “live” abundance depth are not inconsistent with this
calcification strategy.
Implications of sediment–seawater interface calcification for
palaeoenvironmental research
Were foraminiferal calcification only to occur at the sediment–seawater
interface, why do inter-species δ13C offsets vary on annual,
decadal, centennial, and longer timescales? As discussed above (see
Introduction), δ13C offsets between epifaunal and infaunal taxa,
assumed to be indicative of the pore water δ13CDIC
gradient, have been used to reconstruct past changes in organic matter flux
and sea floor oxygenation (e.g. McCorkle and Emerson, 1988; Zahn et al.,
1986; Schilman et al., 2003; Hoogakker et al., 2015). Considering our
hypothesis that infaunal species migrate to the sediment–seawater interface
to calcify, we propose that changes in epifaunal–infaunal offsets are
controlled by the timing of calcification; epifaunal and shallow infaunal
taxa calcify persistently, even during periods of relatively low
Corg availability (oligotrophic/mesotrophic conditions), and
infaunal taxa calcify only during periods of high Corg availability
(eutrophic conditions). Deeper infaunal taxa may be opportunistic,
potentially “hibernating” or at least not calcifying new chambers, at
depth during oligotrophic periods when food is scarce and migrating to the
sediment–seawater interface to feed and calcify during eutrophic periods.
Supporting this hypothesis, Ohga and Kitazato (1997) observed that, at their
study site in Sagami Bay, Japan, all foraminifera where found living in the
uppermost part of the sediment column during the peak of phytodetrital flux.
Kitazato et al. (2000) have proposed that deep-infaunal taxa may migrate to
the sediment–seawater interface to take advantage of phytodetrital input and
to reproduce. Furthermore, in laboratory experiments Nomaki et al. (2005)
recorded that many infaunal foraminifera, especially the taxa they classified
as shallow or intermediate infauna, migrated upwards through the sediment
column in response to an addition of food at the surface. This migration may
be quite rapid, with foraminifera capable of moving through several millimetres of
sediment in a day (e.g. Gross, 2000), although Witte et al. (2003) found that
foraminifera took 8 days to respond to a phytodetritus pulse. In a
simulated phytodetritus event Sweetman et al. (2009) found that the uptake of
added carbon, labelled with 13C, was confined to surface-living
specimens of the deep-infaunal Globobulimina turgida. However, we
note that Enge et al. (2011) did not record a significant response by deeper
infaunal taxa to a simulated phytodetritus pulse, although this may be due to
the short duration (4 days) of their experiment. It is also important to note
that it has been hypothesized that deep-infaunal taxa may have a preference
for degraded organic matter (Caralp, 1989; Fontanier et al., 2003), which if
correct may further delay their response to influxes of labile organic matter
from phytodetrital pulses. Whilst these studies are consistent with our
hypothesis that infaunal taxa migrate to the sediment–seawater interface
during eutrophic conditions and calcify, no studies have yet recorded when
calcification occurs.
In our scenario, infaunal δ13C records are therefore biased towards
eutrophic conditions (even if this is just a brief seasonal pulse of
Corg to the sea floor), while epifaunal/shallow infaunal δ13C records are more reflective of mean annual conditions at the sea
floor. Since deep-infaunal δ13C is biased towards periods of
enhanced Corg flux to the sea floor, downcore records from the
δ13C of these taxa are potentially less sensitive to variability
in Corg flux. The abundance of opportunistic infaunal species,
however, can increase dramatically by accelerating growth and/or reproduction
during eutrophic episodes (e.g. Gooday, 1988; Gooday and Hughes, 2002; Fontanier et al., 2006a; Peck et al., 2015) due to greater organic carbon
availability and their tolerance of lower oxygen concentrations
(Piña-Ochoa et al., 2010). Downcore epifaunal δ13C is
potentially a more sensitive indicator of variable Corg flux and
bottom water conditions (oxygenation). During periods of exceptionally high
productivity, epifaunal δ13C may converge with infaunal
δ13C records, reflecting a prolonged abundance of Corg at
the seafloor throughout the year.
Summary
In this study, we show that the diversity of benthic foraminifera does not
vary greatly at our study sites across the South Georgia shelf and that
there are similarities between the assemblages recorded here and those found
on the Antarctic Peninsula and elsewhere on the Antarctica shelf,
particularly among the more abundant taxa. We find that for most of the taxa
in this study, there is no strong preference for a specific living depth,
except for a small number of species that exhibit a preference for an
epifaunal/shallow infaunal microhabitat (e.g. Labrospira scitula and
Glaphyrammina rostrata).
We find little intra-species variation in δ13C of the tests of the
“living” benthic foraminiferal species analysed here (Astrononion echolsi, Cassidulinoides porrectus, Buccella sp. 1), despite their recovery from a range of depths (and assumed pore water isotopic
compositions) below the sea floor. The failure of δ13Cforam from depth-restricted samples to document the pore
water δ13CDIC gradient common to marine sediments
suggests that infaunal taxa calcify at a fixed δ13CDIC
and therefore depth (e.g. McCorkle et al., 1990; Schmiedl et al., 2004;
Theodor et al., 2016). Whilst we are not able to conclusively state that the
species studied here calcify at the sediment–seawater interface, this
hypothesis is consistent with (i) the presence of “live” individuals of
all the most abundant taxa in the upper 1–2 cm of sediment in this study
(and other studies), (ii) the documented ability of foraminifera to migrate
through the sediment column (e.g. Linke and Lutze, 1993; Geslin et al.,
2004), (iii) the absence of a strong living depth preference in this study
(and other studies), and (iv) the sediment–seawater interface being the most
energetically favourable location for calcification (greatest Corg
and [CO32-] availability).
We propose that epifaunal taxa live and calcify at the sediment–seawater
interface throughout a wide range of organic carbon availability scenarios
(oligotrophic to eutrophic). Infaunal species, however, may only calcify when
the concentration of organic carbon at the seafloor is high and they migrate
to the sediment–seawater interface to feed. In this scenario, temporal
(downcore) variability in the offsets between epifaunal and infaunal
δ13C would reflect changing fluxes of organic carbon to the
seafloor, causing variation in δ13CDIC between
oligotrophic and eutrophic conditions. Our hypothesis, therefore, has
important implications for the use of benthic foraminiferal δ13C
offsets as proxies of productivity and oxygenation and for the use of trace
metal ratios (Mg / Ca, B / Ca) in reconstructing pore water
temperature and Δ[CO32-]. However, we maintain that changes
in epifaunal–infaunal foraminiferal δ13C offsets through time are
likely to reflect the evolution of phytoplankton blooms and water mass
chemistry (cf. Hoogakker et al., 2015). This study further highlights the
need for a good understanding of foraminiferal ecology when utilizing
isotopic composition in interpreting palaeoenvironmental change. Further work is
needed to investigate the timing and depth of calcification in calcareous
benthic foraminifera and the test-forming depths for agglutinated benthic
foraminifera.
Systematic taxonomy
The following species and taxonomic groupings are arranged in taxonomic order
following the supra-generic classifications of Kaminski (2004) for the
agglutinated taxa and Loeblich and Tappan (1988) for the calcareous walled
taxa. Species identification is largely based on the works of Heron-Allen
and Earland (1932a), Earland (1933), Anderson (1975), Mackensen et
al. (1990), Ishman and Domack (1994), Violanti (1996), Igarashi et
al. (2001), and Majewski (2005, 2013). Eleven additional species have been
described, recorded in benthic foraminiferal assemblage work on sediment from
surface samples from other box cores and gravity cores collected from the
South Georgia shelf (Table 1 and Fig. 1) that were not present in the
sub-cores. Described species are illustrated in Figs. 8–14.
Material. Thirteen specimens from six samples.
Description. Included in this group are any
thick-walled tube fragments comprising coarsely agglutinated material with
rough external surface and very smooth internal cavity.
Family Hippocrepinellidae Loeblich & Tappan, 1984 emend.
Mikhalevich, 1995
Genus Hippocrepinella Heron-Allen & Earland, 1932b
Hippocrepinella alba Heron-Allen & Earland,
1932b(Fig. 8:2)
1932b Hippocrepinella alba Heron-Allen & Earland:
259, pl. 1, figs. 16–18.
1933 Hippocrepinella alba Heron-Allen & Earland;
Earland: 71, pl. 7, figs. 10–12.
Material. Twenty-eight specimens from seven samples.
Description. Unilocular, cylindrical to fusiform
test; smooth, thin, white wall constructed of very small particles; aperture
round at end of narrow neck, which may have a collar; may also have a
secondary basal aperture.
Occurrence. Originally described from the South
Atlantic (Heron-Allen and Earland, 1932b) and subsequently recorded from
South Georgia (Earland, 1933), both from surface samples.
Family Rhizamminidae Wiesner, 1931
Genus Rhizammina Brady, 1879
Rhizammina spp.(Fig. 8:3–5)
Material. Forty-nine specimens from nine samples.
Description. Included in this group are any elongate
tubular tests with thin, flexible walls; only broken tubes recovered in this
study; specimens recorded here mainly composed of diatomaceous material.
Family Saccamminidae Brady, 1884
Subfamily Saccammininae Brady, 1884
Genus Lagenammina Rhumbler, 1911
Lagenamminacf.sphaerica
Moreman, 1930(Fig. 8:6)
1930 Lagenammina sphaerica Moreman: 51, pl. 5,
fig. 15.
Material. Seventy specimens from 13 samples.
Description. Unilocular, near spherical test; thin
agglutinated wall; medium sand grains; rough surface; simple aperture
produced at the end of an elongate neck; neck approximately two fifths of
test length.
Remarks. As Lagenammina sphaerica Moreman
was described from Silurian deposits in the USA, this cannot be the same
species. However the morphological similarities are uncanny. The species
described here does bear some similarity to the extant Lagenammina arenulata (Skinner) that is common at other Antarctic localities, but this
species does not possess an elongate neck. Jones (1994) suggests that
specimens similar to L. arenulata (Skinner), but with an elongate
neck, belong to L. atlantica (Cushman). However, the original
description of this species reveals that it too does not possess an elongate
neck.
Genus Saccammina Carpenter, 1869
Saccammina spp.(Fig. 8:8)
Material. Four specimens from three samples.
Description. Included in this group are any forms
that have a globular, unilocular test, with a firm, fine to medium
agglutinated wall held together with an organic cement, and a round aperture
produced on a short neck.
Subfamily Thurammininae Miklukho-Maklay, 1963
Genus Astrammina Rhumbler in Wiesner, 1931
Astrammina rara Rhumbler in Wiesner, 1931(Fig. 8:9–11)
1931 Astrammina rara Rhumbler, in Wiesner: 78,
pl. 2, fig. 19.
1932b Armorella sphaerica Heron-Allen & Earland:
257, pl. 2, figs. 4–11.
1932b Pelosphaera cornuta Heron-Allen & Earland:
255, pl. 2, figs. 12–15.
1933 Armorella sphaerica Heron-Allen & Earland;
Earland: 65, pl. 7, figs. 16–23.
1933 Pelosphaera cornuta Heron-Allen & Earland;
Earland, 61, pl. 7, figs. 24–27.
Material. Twelve specimens from six samples.
Description. Large, near-spherical, unilocular test
with a number of narrow “arms” extending in various directions, often along
sponge spicules that are incorporated into the test wall. Wall firmly
agglutinated, comprising fine to coarse sand grains, sponge spicules, and
diatoms, with a grey to yellowish brown cement.
Occurrence. Originally described from the South
Atlantic (Heron-Allen and Earland, 1932b) and subsequently recorded from
South Georgia (Earland, 1933), both from surface samples.
Remarks. The synonymy of Astrammina rara Rhumbler, Armorella sphaerica Heron-Allen & Earland, and
Pelosphaera cornuta Heron-Allen & Earland was demonstrated by
DeLaca (1986) in an analysis of live Antarctic specimens.
1912 Psammosphaerabowmanni Heron-Allen
& Earland: pl. 5, figs. 5–6, pl. 6, fig. 5.
2010 Capsammina bowmanni (Heron-Allen & Earland);
Gooday et al.: 349, pl. 1, figs. A & B.
Material. Four specimens from three samples.
Description. Free, monothalamous test, comprising
several small flat grains of mica cemented by very fine cement that form an
irregular polyhedral chamber. Openings are visible where the flakes meet and
cement is absent, which may be the aperture or accidental.
Occurrence. Originally described from North Sea
dredgings (Heron-Allen and Earland, 1912), it was subsequently recorded in
the north-east Atlantic (Gooday et al., 2010).
Remarks. Heron-Allen and Earland (1912) were unable
to assign a definite oral aperture, recording openings similar to those
recorded here on some specimens but not all. Heron-Allen and Earland (1912)
also note that this species is difficult to describe, as no two specimens
look exactly alike. The flakes are arranged to approximately make an oval or
flask shape, and the gaps, the size of which may vary considerably, between
the flakes are then filled with cement, so that the cemented areas vary from
narrow infill to broad flat areas.
Psammosphaera fusca Schulze, 1875(Fig. 9:1–2)
1875 Psammosphaera fusca Schulze: 113, pl. 2,
fig. 8.
1994 Psammosphaera fusca Schulze; R. W. Jones: 31,
pl. 18, figs. 1–8.
2003 Psammosphaera fusca Schulze; Gaździcki &
Majewski: 7, fig. 3 (1–2).
Material. Forty-six specimens from 15 samples.
Description. Small spherical test; usually
unilocular, occasionally multiple chambers loosely joined. Agglutinated wall
comprising medium to coarse grains in a fine cement.
Occurrence. Recorded from several locations in the
Atlantic (Jones, 1994) and Antarctica (Gaździcki and Majewski, 2003),
all from Recent sediments.
Family Ammodiscidae Reuss, 1862
Subfamily Ammodiscinae Reuss, 1862
Genus Ammodiscus Reuss, 1862
Ammodiscusincertus (d'Orbigny,
1839a)(Fig. 9:5–6)
1839a Operculina incerta d'Orbigny: 49, pl. 6,
figs. 16–17.
1996 Ammodiscus incertus (d'Orbigny); Violanti: 53,
pl. 3, fig. 6.2001 Ammodiscus sp.; Igarashi et al.: 142, pl. 2,
fig. 5.
2013 Ammodiscus incertus (d'Orbigny); Majewski: 175,
fig. 3(4).
Material. Overall, 109 specimens from 22 samples.
Description. Small test (ca. 200 µm
diameter), generally flat but tending towards irregular coiling in the last
whorls. Up to seven whorls in microspheric form and up to six in
megalospheric form; sutures between whorls somewhat depressed;
proloculus
spherical and central. The agglutinated test wall comprises angular grains of
sand in a cement matrix that is a yellow-brown colour. Aperture a simple
opening at the end of the tubular chamber.
Occurrence. Recorded from a number of locations on
the Antarctic shelf (Violanti, 1996; Igarashi et al., 2001; Majewski, 2013),
from Recent sediments.
1933 Miliammina oblonga (Heron-Allen & Earland);
Earland: 92, pl. 3, fig. 17, pl. 5, figs. 1–5, 7–8.
1955 Miliammina earlandi Loeblich & Tappan: 12,
pl. 1, figs. 15–16.
1981 Miliammina earlandi Loeblich & Tappan; Milam
& Anderson: 304, pl. 2, fig. 4.
1996 Miliammina earlandi Loeblich & Tappan;
Violanti: 53, pl. 3, figs. 10–11.
2004 Miliammina earlandi Loeblich & Tappan;
Murray & Pudsey: 71, pl. 1, figs. 8–9.
Material. Overall, 3240 specimens from 34 samples.
Description. Tubular chambers with a rounded
peripheral edge, arranged in a quinqueloculine pattern; sutures slightly
depressed, increasing as test size increases. Wall comprises very small
mineral grains (whose composition is unclear) completely embedded in
siliceous cement (insoluble in HCl), smooth/polished surface. Very light grey
in colour, and large variation in test size. Crescentiform aperture is at the
terminal end of the final chamber, flush or with a slight neck. There is
considerable morphological variation within this species due to variation in
the roundness of the chambers.
Occurrence. Originally described from the South
Atlantic (Heron-Allen and Earland, 1930) and subsequently reported from
several locations of the Antarctic shelf (Milam and Anderson, 1981; Violanti,
1996; Murray and Pudsey, 2004), from Recent sediments.
Remarks. Whilst initially included in
Miliammina oblonga (Chapman) by Heron-Allen and Earland (1930), this
South Georgia form was later separated from Miliammina arenacea
(Chapman) by Earland (1933) due to its more ovate and rounded outline and
was renamed Miliamminaoblonga Heron & Earland. However,
Loeblich and Tappan (1955) recognized that this species name had previously
been assigned to Miliolina oblonga (Montagu), incorrectly placed in
Miliammina, and renamed it Miliammina earlandi (which we
use here). Earland (1933) considered M. earlandi to be the most
characteristic and commonest foraminifera of the South Georgia area.
Miliammina lata Heron-Allen & Earland, 1930(Fig. 9:4)
1930 Miliammina lata Heron-Allen & Earland: 43,
pl. 1, figs. 13–17.
1933 Miliammina lata Heron-Allen & Earland;
Earland: 93, pl. 3, fig. 17, p. 5, figs. 15–19.
1996 Miliammina lata Heron-Allen & Earland;
Violanti: 52, pl. 3, fig. 12.
2001 Miliammina lata Heron-Allen & Earland;
Igarashi et al.: 140, pl. 2, fig. 10.
2005 Miliammina lata Heron-Allen & Earland;
Majewski: 193, fig. 12(8).
2013 Miliammina lata Heron-Allen & Earland;
Majewski: 176, fig. 4(3).
Material. Sixty specimens from 13 samples.
Description. Quinqueloculine test; early chambers
often not visible; inflated, broadly rounded chambers, broadening at the
base. Slightly depressed sutures; thick, smooth, agglutinated wall comprising
very small particles in an excess of siliceous cement. Small crescentiform
aperture at the end of the final chamber.
Occurrence. Originally described from the South
Atlantic (Heron-Allen and Earland, 1930), it was subsequently recorded from
South Georgia (Earland, 1933) and a number of locations on the Antarctic
shelf (Igarashi et al., 2001; Violanti, 1996; Majewski, 2005, 2013), all from
Recent sediments.
Family Hormosinellidae Rauzer-Chernousova & Reytlinger,
1986
Genus Hormosinella Shchedrina, 1969
Hormosinella sp.(Fig. 9:3)
Material. Three fragmentary specimens from one sample.
Description. Large, uniserial test; ovate/fusiform
chambers separated by long, delicate, necks. Thin, agglutinated wall; rounded
terminal aperture on elongate neck.
Remarks. Only recorded here as fragmentary
specimens.
1881 Lituola (Reophax) guttifera Brady: 49, pl. 31,
figs 10–15.
1975 Reophax guttifer (Brady); Anderson: 74, pl. 1,
fig. 18.
1994 Hormosinella guttifera (Brady); R. W. Jones:
38, pl. 31, figs. 10–15.
2005 Hormosinelloides guttifer (Brady); Kaminski
& Gradstein: 249, pl. 46, figs. 1–8.
Material. Thirty-one specimens from four samples.
Description. Uniserial test comprising a series of
approximately three to five pyriform chambers. Thin, agglutinated wall with a fairly
rough finish. Simple, rounded aperture; terminal on an elongated neck.
Occurrence. Originally described from the South
Atlantic by Brady (1881) who recorded that it was “exceedingly rare”, it
was subsequently recorded from the Weddell Sea (Anderson, 1975), from Recent
sediments.
Remarks. Placed in the genus
Hormosinelloides by Kaminski and Gradstein (2005) because the
chambers attach near the base of the apertural neck of the preceding
chamber.
Family Reophacidae Cushman, 1927
Genus Nodulina Rhumbler, 1895
Nodulina dentaliniformis (Brady, 1881)(Fig. 9:11)
1881 Lituola (Reophax) dentaliniformis Brady: 49,
pl. 30, figs. 21–22.
1994 Reophax dentaliniformis (Brady); R. W. Jones:
37, pl. 30, figs. 21–22.
2017 Nodulina dentaliniformis (Brady); Kender &
Kaminski: pl. 8, fig. 1.
Material. Six specimens from one sample.
Description. Slender, elongate test with straight
axis; three to six chambers; final chamber significantly larger and tapered; wall
comprises sand grains in firm cement, with a smooth finish. Large, round
terminal aperture on a short tubular neck of finer grains.
Occurrence. Recorded from the South Pacific, the
North Atlantic (Brady, 1881; Jones, 1994), and the Bering Sea (Kender and
Kaminski, 2017).
Remarks.Nodulina dentaliformis (Brady) is
the type species of the genus Nodulina, which is distinguished from
Reophax by its more symmetrical, regular chambers, horizontal
sutures, and straighter axis.
1950 Reophax subdentaliniformis Parr: 269, pl. 4,
fig. 20.
1996 Nodulina subdentaliniformis (Parr); Violanti:
53, pl. 3, fig. 18.
2005 Nodulina subdentaliniformis (Parr); Majewski:
194, fig. 13(6–7).
Material. Overall, 457 specimens from 25 samples.
Description. Slender, elongate test with a slight
curvature; usually four to six chambers (may be difficult to distinguish); final
chamber significantly larger and tapered. Wall comprises poorly sorted sand
grains in firm cement, coarsely finished. Large, round terminal aperture on a
short neck of finer sand grains.
Occurrence. Recorded from Recent Antarctic sediments
(Parr, 1950; Violanti, 1996; Majewski, 2005).
Remarks. We have placed this species within the
genus Reophax on the basis of the curvature of the test.
Material. Seventeen specimens from 10 samples.
Description. Slender, elongate, slightly arched test
comprising two to four chambers; final chamber large, tapering, and fusiform.
Chambers increase rapidly in size; final chamber often comprising the
majority of the test; depressed sutures; wall comprises poorly sorted sand
grains in firm cement, coarsely finished. Large, round terminal aperture on a
short neck.
Occurrence. This species was originally described by
Earland (1933) from a number of stations near South Georgia. The description
was subsequently emended by Höglund (1947) using specimens from the North
Sea. Earland (1934) also reported its occurrence around the Falkland Islands, where it is widely distributed but rare at most localities. Kaminski and
Gradstein (2005) report that R. subfusiformis is reported from Recent
sediments at high-latitude stations throughout the Atlantic and Southern
Oceans.
Family Hormosinidae Haeckel, 1894
Subfamily Hormosininae Haeckel, 1894
Genus Pseudonodosinella Saidova, 1970
Pseudonodosinella sp.(Fig. 9:15)
Material. One specimen from one sample.
Description. Elongate, uniserial test; ovate
chambers that overlap much of the preceding chamber, including the aperture.
Firmly agglutinated, smoothly finished wall; terminal aperture.
1962 Haplophragmoides bradyi (Robertson) subsp.
niigatensis Uchio: 385, pl. 18, fig. 6.
1975 Haplophragmoides bradyi (Robertson); Anderson:
76, pl. 2, fig. 8.
Material. Seventeen specimens from six samples.
Description. Small, subglobose, involute test;
becomes slightly evolute and umbilicate in adult tests; rounded periphery,
slightly lobulate. Chambers low and broad, somewhat inflated; five in final
whorl, gradually increasing in size as added. Distinct, depressed, radiate
sutures. Finely agglutinated wall with smooth surface, yellowish colour.
Aperture an interiomarginal crescentic slit.
Occurrence. Originally described from Recent
sediments from the Sea of Japan (Uchio, 1962), it was subsequently recorded
from the Weddell Sea (Anderson, 1975).
Remarks.Haplophragmoides bradyiniigatensis Uchio differs from Haplophragmoides bradyi (Robertson) in having a less evolute and more inflated test.
1934 Haplophragmoides quadratus Earland: 88, pl. 3,
figs. 7–8.
1960 Haplophragmoides quadratus Earland; Uchio: 52,
pl. 1, fig. 17, pl. 5, fig. 14.
Material. Forty-three specimens from seven samples.
Description. Planispiral, compressed test with
rounded periphery; approximately square outline with rounded corners.
Chambers globose, slightly inflated; four in final whorl. Distinct, depressed
sutures. Aperture not clearly visible in the specimens examined here.
Occurrence. Originally described from Bellingshausen
Sea, Antarctica (Earland, 1932).
Remarks.Haplophragmoides quadratus Uchio
1960, described from the Pacific coast of the USA, appears to be the same
species and would therefore be a junior synonym.
Genus Labrospira Höglund, 1947
Labrospira scitula (Brady, 1881)(Fig. 9:16–18)
1881 Lituola (Haplopragmium) scitulum Brady: 50,
pl. 34, figs 11–13.
1933 Haplophragmoides scitulum (Brady); Earland: 78,
pl. 3, figs. 11–12.
1994 Veleroninoides scitulus (Brady); Jones: 41,
pl. 34, fig. 13.? 1996 Recurvoides contortus Violanti: 39, pl. 4,
fig. 11.
2017 Veleroninoides scitulus (Brady); Kender and
Kaminski: pl. 8, fig. 10.
Material. Overall, 328 specimens from 25 samples.
Description. Planispiral, partially evolute test,
comprising three whorls with 8–11 chambers in the final whorl. Some
specimens become slightly streptospiral in the final whorl; fairly deep
umbilicus on both sides. Sutures slightly curved to straight, radial, and
very slightly depressed. Smooth-surfaced, thick agglutinated wall comprising
coarse grains with yellow-brown cement; colour rapidly fades to white after
death. Aperture an elongate oval-shaped areal opening, close to the base of
the apertural face, with a fine-grained, raised lip surrounding the aperture.
Occurrence. Originally described from the North
Atlantic (Brady, 1881); also recorded from the Bering Sea (Kender and
Kaminski, 2017).
Remarks.Labrospira scitula (Brady) is
also often placed within the genus Veleroninoides. However, we
believe this designation to be incorrect as L. scitula (Brady) is
not fully evolute – a diagnostic characteristic of Veleroninoides.
Labrospira sp. 1(Fig. 10:3–4)
Material. Overall, 236 specimens from 26 samples.
Description. Planispiral, slightly evolute test; slightly inflated chambers; six in the final whorl. Small
umbilicus; straight, radial, slightly depressed sutures; smooth surface; thick
agglutinated wall comprising coarse grains with yellow-brown cement. Aperture
an areal, elongate slit, close to the base of the apertural face, with a very
fine-grained, raised lip surrounding the aperture.
Remarks. Our material may represent the same species
as that recorded by Murray and Pudsey (2004) as Haplophragmoides canariensis (d'Orbigny). We consider that this classification is incorrect,
however, due to the significant differences in apertural characteristics.
Family Discamminidae Mikhalevich, 1980
Genus Glaphyrammina Loeblich & Tappan, 1984
Glaphyrammina americana (Cushman, 1910)(Fig. 10:6)
1910 Ammobaculites americanus Cushman; 117,
figs. 184–185.
1932a Ammobaculites americanus Cushman; Heron-Allen
& Earland: 341, pl. 8, figs. 15–17.
1994 Glaphyrammina americana (Cushman); Jones: 40,
pl. 34, figs. 1–4.
Material. Six specimens from four samples.
Description. Very large broad and flattened test,
initially planispiral becoming uniserial; spiral section slightly involute; nine chambers in the outer whorl. Wall comprises well-cemented, poorly sorted sand
grains and occasional diatom frustules. Long, slit-like aperture extends the
width of the final chamber.
Occurrence. Originally described off the west coast
of Mexico (Cushman, 1910), later from the South Atlantic (Jones, 1994).
Glaphyrammina rostrata (Heron-Allen and
Earland, 1929)(Fig. 10:7)
1929 Ammobaculites rostratus Heron-Allen &
Earland; 326, pl. 2, figs. 14–17.
1933 Ammobaculites rostratus Heron-Allen &
Earland; Earland: 80, pl. 5, figs. 22–25.
Material. Overall, 278 specimens from 23 samples.
Description. Thin-walled, flattened, planispiral
test. Sutures often indistinct, especially in smaller specimens;
approximately six chambers in final whorl, partially evolute. In some specimens
the chambers begin to uncoil to have two uniserial chambers; where this
uncoiling does not occur the final chamber protrudes at an angle away from
the spiral. The test is constructed of fine sand with larger grains
distributed throughout this matrix. The final chamber terminates with a
narrowing apertural neck, described as “nipple-like” in Heron-Allen and
Earland (1929) that is much more neatly constructed, solely comprising fine
sand (this structure is very fragile and often partially or completely
absent).
Occurrence. This species has been described from the
South Georgia region (Heron-Allen and Earland, 1929; Earland, 1933).
Remarks. Originally described by Heron-Allen and
Earland (1929) within the genus Ammobaculites, here we place this
species in the genus Glaphyrammina due to the flattened nature of
the test. When the apertural neck is absent the specimens can resemble
Glaphyrammina americana but can be separated by the less distinct
sutures.
Material. Five specimens from three samples.
Description. Involute, subglobose test,
streptospirally coiled in early stage, becoming planispiral. Broad, low
chambers. Agglutinated wall, with a fairly rough surface. Simple, areal,
aperture, just above the base of the final chamber.
Remarks. These specimens are placed in the genus
Recurvoides on the basis of wall and apertural characteristics. Due
to their small size, it has not been possible to definitively identify the
coiling pattern.
Family Spiroplectamminidae Cushman, 1927
Subfamily Spiroplectammininae Cushman, 1927
Genus Spiroplectammina Cushman, 1927
Spiroplectammina biformis (Parker and Jones, 1865)(Fig. 10:13)
1865 Textularia agglutinans var. biformis Parker
& Jones: 370, pl. 15, figs. 23–24.
1932a Spiroplectammina biformis (Parker & Jones);
Heron-Allen & Earland: 347, pl. 8, figs. 27–31.
1994 Spiroplectammina biformis (Parker & Jones);
R. W. Jones: 50, pl. 45, figs. 25–27.
1994 Spiroplectammina biformis (Parker & Jones);
Ishman and Domack: 150, pl. 1, fig. 4.
2003 Spiroplectammina biformis (Parker & Jones);
Gaździcki & Majewski: 8, fig. 4 (5).
2005 Spiroplectammina biformis (Parker & Jones);
Majewski: 195, fig. 14 (3–5).
2009 Spiroplectammina biformis (Parker & Jones);
Majewski & Anderson: 138, fig. 3.
2013 Spiroplectammina biformis (Parker & Jones);
Rodrigues et al.: 214, fig. 3 (9).
Material. One specimen from one sample.
Description. Small test mainly comprising sand
grains with a small amount of cement. Initially planispiral, rapidly
uncoiling and becoming biserial; coiled section broader than biserial
section; biserial chambers are subquadrate. Aperture a low arch at the inner
margin of the final chamber.
Occurrence. Originally described from Recent
sediments from Greenland (Parker and Jones, 1865) and later from the Arctic
(Goes, 1894; Höglund, 1947). More recently this species has been
described from the Antarctic Peninsula (Ishman and Domack, 1994; Majewski,
2005, 2013).
Remarks. The type species of
Spiroplectammina. The large geographical separation between these
two areas suggests that the Antarctic form may be a different species but
they are morphologically very similar. Also recorded from the Cretaceous
Chalk and Gault of England by Parker and Jones (1865), who suggest it is an
abbreviated form of Textularia annectens, which is commonly found
within the Cretaceous.
1931 Pseudobolivina antarctica Wiesner: 99, pl. 21,
figs. 257–258.
1975 Pseudobolivina antarctica Wiesner; Anderson:
78, pl. 3, fig. 4.
1981 Pseudobolivina antarctica Wiesner; Milam &
Anderson: 305, pl. 3, fig. 7.
1990 Pseudobolivina antarctica Wiesner; Mackensen et
al.: 258, pl. 5, fig. 7.
1996 Pseudobolivina antarctica Wiesner; Violanti,
55, pl. 4, figs. 12–13.
Material. Twenty-three specimens from 11 samples.
Description. Very small biserial test, tapered at
the base; chambers rapidly enlarging as they are added to give broad maximum
width. Wall comprises fine sand with little cement such that the surface is
fairly rough. Sutures clearly visible and slightly depressed. Aperture
loop-shaped extending up the face of the final chamber.
Occurrence. Originally described from Antarctica
(Wiesner, 1931) and subsequently recorded in Recent sediments from a number
of locations on the Antarctic shelf (Anderson, 1975; Milam and Anderson,
1981; Mackensen et al., 1990; Violanti, 1996).
Remarks.Pseudobolivina antarctica is also
described in Igarashi et al. (2001) and Majewski (2013). However, the
slit-like aperture shown by these authors does not agree with the type
description, suggesting it has been incorrectly assigned (these specimens may
be Textularia antarctica).
1950 Trochammina wiesneri Parr: 279, pl. 5, fig. 14.
1975 Portatrochammina wiesneri (Parr); Anderson: 92,
pl. 3, fig. 5.
1988 Portatrochammina antarctica wiesneri (Parr);
Brönnimann & Whittaker: 68, figs. 25D–F, 26H–K, 27D–I.
1994 Portatrochammina antarctica (Parr); Ishman &
Domack: 150, pl. 1, fig. 5.
1996 Portatrochammina antarctica (Parr); Violanti:
55, pl. 4, figs. 14–15.
2001 Portatrochammina antarctica antarctica (Parr);
Igarashi et al.: 145, pl. 4, fig. 6.
2004 Portatrochammina antarctica (Parr); Murray &
Pudsey: 71, pl. 1, figs. 14–16.
2009 Portatrochammina antarctica (Parr); Majewski
& Anderson: 138, fig. 3.
2013 Portatrochammina antarctica (Parr); Majewski:
178, fig. 6(4).
Material. Overall, 383 specimens from 31 samples.
Description. Low, trochospiral test of approximately
17 chambers, slightly convex on the spiral side and shallowly concave on the
umbilical side; three whorls with five chambers in the final whorl.
Well-defined, moderately depressed sutures producing slightly lobate outline.
Periphery sub-acute in edge view. Aperture covered by a large umbilical flap
that covers much of the umbilical depression (diagnostic of this genus); parts of the apertural flaps of preceding chambers also visible.
Agglutinated, imperforate wall, coarser on spiral side. Yellowish-brown
colour that is markedly darker (almost reddish) in earlier chambers.
Occurrence. Originally described from Antarctica
(Parr, 1950) and subsequently recorded from around Antarctica (Anderson,
1975; Brönnimann and Whittaker, 1988; Ishman and Domack, 1994; Violanti,
1996; Igarashi et al., 2001; Murray and Pudsey, 2004; Majewski and Anderson, 2009; Majewski, 2013).
Remarks.Portatrochammina antarctica wiesneri (Parr) can be distinguished from Portatrochammina antarctica antarctica (Parr) by its flatter, smoother test and subacute
peripheral margin.
1988 Deuterammina scoresbyi Brönnimann &
Whittaker: 116, fig. 43(A–J).
Material. Overall, 207 specimens from 26 samples.
Description. Low trochospiral test, becoming
planispiral in the final whorl; spiral side slightly depressed centrally;
deep wide umbilical depression. Sutures distinct, straight, radial, slightly
depressed; 20 chambers including the proloculus arranged in three whorls;
six to seven slightly inflated chambers in the final whorl. Primary aperture
interiomarginal, sub-peripheral loop-shape that widens towards the spiral
side with a lip; secondary aperture at umbilical tip of chamber, may have a
lip; both apertures present on every chamber. Agglutinated, imperforate wall.
Occurrence. Originally described from the
Bellingshausen Sea and the South Sandwich Islands (Brönnimann and Whittaker,
1988).
Remarks. Brönnimann and Whittaker (1988) describe
the wall as being almost entirely composed of barite spherules/ovoids. This
is not the case in the specimens described here, although some ovoids were
observed. Additionally the specimens described here have six to seven chambers in the
final whorl, rather than the eight described in Brönnimann and Whittaker (1988).
Material. Overall, 182 specimens from 21 samples.
Description. Low trochospiral test; globular
chambers that rapidly increase in size; four in final whorl. Depressed
sutures; smoothly finished, finely agglutinated wall; areal elongate slit aperture
located on the umbilical side but not in the umbilicus.
Family Cyclamminidae Marie, 1941
Subfamily Cyclammininae Marie, 1941
Genus Cyclammina Brady, 1879
Cyclammina cf. contorta Pearcey, 1914(Fig. 11:2)
1914 Cyclammina contorta Pearcey: 1009, pl. 2,
figs. 5–7.
Material. One specimen from one sample.
Description. Very large, planispiral, involute,
somewhat compressed test; rounded periphery; very slightly lobulate margin.
Numerous chambers (ca. 15) in the final whorl; sutures straight. Finely
agglutinated, surface smooth, slightly iridescent. Aperture appears to be an
interiomarginal, equatorial low arch, although this is poorly preserved on
the only specimen found in this study. Apertural face of the final chambers
much more coarsely agglutinated with a rough surface; no evidence of additional apertures.
Occurrence. Cyclammina contorta Pearcey was
originally described from the Weddell Sea, Antarctica.
Remarks. The species described here differs only in
its sutural characteristics, being straighter and less dark in colour.
Family Textulariidae Ehrenberg, 1838
Subfamily Textulariinae Ehrenberg, 1838
Genus Textularia Defrance, 1824
Textularia earlandi Parker, 1952(Fig. 11:3–4)
1933 Textularia tenuissima Earland: 95, pl. 3,
figs. 21–30.
1952 Textularia earlandi Parker: 458 (see synonymy).
1996 Textularia earlandi Parker; Violanti: 57,
pl. 5, figs. 13–14.
2010 Textularia tenuissima Earland; Majewski: 65,
fig. 2(2).
Material. Forty-nine specimens from 18 samples.
Description. Very small, delicate, elongate test.
Initial chambers coiled in both microspheric (tapered end) and megalospheric
(rounded end) forms, becoming biserial. Numerous subquadrate, rounded
chambers that enlarge slightly towards the terminal end. Distinct, slightly
depressed sutures. Aperture a low interiomarginal arch on the face of the
final chamber. Fairly smooth wall comprising fine grains with little cement
visible.
Occurrence. Originally described from South Georgia
(Earland, 1933); subsequently recorded from the Ross Sea (Violanti, 1996) and
the Antarctic Peninsula (Majewski, 2010).
Remarks. Originally described from South Georgia as
Textularia tenuissima Earland, this was however preoccupied by
Textularia tenuissima Häusler 1881. Therefore, Parker (1952)
proposed the new name Textularia earlandi. Morphologically it is
very similar to Textularia elegans Lacroix, found in the
Mediterranean.
Family Cornuspiridae Schultze, 1854
Subfamily Cornuspirinae Schultze, 1854
Genus Cornuspira Schultze, 1854
Cornuspira antarctica Rhumbler in Wiesner, 1931(Fig. 11:6–7)
1931 Cornuspira antarctica Rhumbler in Wiesner: 101,
pl. 14, figs. 164–166.
Material. Two specimens from two samples.
Description. Discoidal test comprising two chambers,
the globular proloculus and the planispirally enrolled, undivided, near-evolute second chamber. Calcareous, imperforate, porcellaneous wall with
transverse growth lines. Aperture simple opening at end of tube.
Occurrence. Originally described from Antarctica
(Wiesner, 1931).
Family Hauerinidae Schwager, 1876
Subfamily Hauerininae Schwager, 1876
Genus Quinqueloculina d'Orbigny, 1826
Quinqueloculina cf. occidentalis Bailey, 1851(Fig. 11:5 and 11:8–9)
1851 Quinqueloculina occidentalis Bailey: 13,
figs. 46–48.
2013 Quinqueloculina sp. Majewski: 179, fig. 7 (6).
2013 Quinqueloculina seminula (Linnaeus); Rodrigues
et al.: 214, fig. 3 (6–7).
Material. Twenty-one specimens from seven samples.
Description. Small, elongate, subovate test,
quinqueloculine chamber arrangement, rounded periphery, long, narrow chambers
that rapidly increase in size as added. Calcareous, smooth, thin,
porcellaneous wall; distinct, somewhat depressed sutures; aperture a terminal
high arch with a small tooth.
Occurrence. Quinqueloculina occidentalis Bailey was originally described from the North Atlantic. Similar specimens
have been recorded from Antarctica (Majewski, 2013; Rodrigues et al., 2013)
but never previously assigned to Q. occidentalis Bailey.
Remarks. The specimens described here appear quite
similar to Quinqueloculina occidentalis Bailey, although the very
brief original description hinders any certainty in this designation. The
specimens also share a number of features with Triloculinella antarctica Kennett, but the absence of a broad apertural flap indicates that
this species does not belong within the genus Triloculinella; its
placement within this genus by Majewski (2005) is therefore incorrect.
1758 Serpula seminulum Linnaeus: 786, pl. 2, fig. 1.
1932a Miliolina seminulum (Linnaeus); Heron-Allen
& Earland: 313, pl. 6, figs. 25–40.
1981 Quinqueloculinaseminulum (Linnaeus);
Milam and Anderson: 307, pl. 5, fig. 5.
1994 Quinqueloculinaseminulum (Linnaeus);
Jones: 21, pl. 5, fig. 6, text figs. 2–3.
2013 Quinqueloculinaseminulum (Linnaeus);
Rodrigues et al.: 214, fig. 3 (6–7).
Material. Two specimens from two samples.
Description. Slightly compressed ovate test, rounded
at base, more angular at apertural end, rounded triangular in section.
Quinqueloculine coiling; five chambers visible in final whorl, four from one side
and three from the other; more convex on the four-chambered side. Calcareous,
imperforate, porcellaneous wall; smooth surface. Large, terminal,
horseshoe-shaped aperture with a small bifid tooth.
Occurrence. Recorded from the North Atlantic in the
Challenger material (Jones, 1994) and from Antarctica by Milam and
Anderson (1981).
1884 Miliolina circularis var. sublineata
Brady: 169, pl. 4, fig. 7.
1994 Triloculinella sublineata (Brady); R. W. Jones:
20, pl. 4, fig. 7.
Material. One specimen from one sample.
Description. Test near-circular in outline;
compressed, rounded section. Quinqueloculine coiling; five chambers visible in
final whorl; calcareous, imperforate, porcellaneous wall; terminal, low,
broad, arch-shaped aperture. Surface ornamented with very delicate,
longitudinal striae and large pits.
Remarks. The absence of an apertural flap in the
specimen described here, and in the image of Jones (1994), plus the presence
of five chambers in the final whorl, leads us to place it within
Quinqueloculina. This specimen differs from the specimen figured in
Brady (1884) and Jones (1994) in the presence of large pits that cover its
surface.
Genus Pyrgo Defrance, 1824
Pyrgo cf. subpisumParr, 1950(Fig. 11:14)
1950 Pyrgosubpisum Parr: 297, pl. 7,
figs. 5–6.
Material. One specimen from one sample.
Description. Large, subglobular test, near-circular
from front, rounded periphery; two chambers visible in final whorl
(biloculine). Calcareous, imperforate, porcellaneous wall; large, ovate,
terminal aperture, surrounded by a thickened, raised rim; unusual large tooth
with three lobes.
Occurrence. Pyrgo subpisum Parr was
described from a number of locations in the southern Indian Ocean.
Remarks. The specimen described here is very similar
to Pyrgo subpisum Parr, differing only in the unusual tooth
characteristics.
Pyrgo patagonica (d'Orbigny, 1839b)(Fig. 11:13)
1839b Biloculina patagonica d'Orbigny: 65, pl. 3,
figs. 15–17.
1932a Biloculina patagonica d'Orbigny; Heron-Allen
& Earland: 311, pl. 6, figs. 4–6.
2004 Pyrgo patagonicum (d'Orbigny); Mikhalevich:
188, pl. 3, figs. 6–7.
Material. One specimen from one sample.
Description. Large, subglobular test, ovate from
front, rounded periphery; two inflated chambers visible in final whorl
(biloculine). Calcareous, imperforate, porcellaneous, think wall, with a
smooth, shiny surface; medium, round, terminal aperture, surrounded by a
small, thickened, raised rim; bifid tooth.
Occurrence. Originally described from the Patagonian
coast (d'Orbigny, 1839b).
Genus Triloculina d'Orbigny, 1826
Triloculina sp.(Fig. 11:15)
Material. Twenty-three specimens from 11 samples.
Description. Ovate text, triangular in section with
rounded corners; triloculine, three chambers visible in final whorl. Chambers
slightly inflated; sutures somewhat depressed; calcareous, imperforate,
porcellaneous wall; smooth surface. Terminal, rounded aperture; no tooth
visible.
Family Nodosariidae Ehrenberg, 1838
Subfamily Nodosariinae Ehrenberg, 1838
Genus Laevidentalina Loeblich & Tappan, 1986
Laevidentalina communis (d'Orbigny, 1826)(Fig. 11:17–18)
1826 Nodosaria (Dentalina) communis d'Orbigny: 254 (no type figure).
1996 Dentalina communis (d'Orbigny); Violanti: 59,
pl. 6, fig. 11.
2005 Dentalina communis (d'Orbigny); Majewski: 201,
fig. 20(2–3).
2013 Laevidentalina communis (d'Orbigny); Majewski:
181, fig. 9(2).
Material. Two specimens from two samples.
Description. Elongate, slightly arcuate test; fusiform, apiculate proloculus, followed by a number of uniserially arranged
chambers; straight, horizontal sutures. Calcareous, hyaline, finely perforate
wall; smooth surface; multilamellar, terminal aperture damaged in our
specimens.
Occurrence. Recorded from Antarctica in Recent
sediments (Violanti, 1996; Majewski, 2005, 2013)
Remarks. Placed within the genus
Laevidentalina due to the absence of costae.
1866 Nodosaria (Nodosaria) calomorpha Reuss: 129,
pl. 1, figs. 15–19.
1933 Nodosaria calomorpha Reuss; Earland: 117,
pl. 4, fig. 19.
1994 Glandulonodosaria calomorpha (Reuss); Jones:
72, pl. 61, figs. 23–26.
Material. Six specimens from four samples.
Description. Small, elongate, slightly arcuate test
comprising two to four sausage-shaped chambers. Simple, round, terminal aperture.
Thin, hyaline, calcareous wall, with smooth surface; sutures straight and
horizontal.
Occurrence. Reported from South Georgia (Earland,
1933) and the South Atlantic (Jones, 1994).
Genus Nodosaria Lamarck, 1812
Nodosaria simplex Silvestri, 1872(Fig. 12:3–4)
1872 Nodosaria simplex Silvestri: 95, pl. 11,
figs. 268–272.
1994 Nodosaria simplex Silvestri; Jones: 73, pl. 62,
figs. 4–5.
Material. Two specimens from two samples.
Description. Small, slightly elongate test
comprising two spherical/ovate chambers of approximately the same size;
straight, depressed suture between the chambers. Both chambers narrow
terminally: the upper towards the aperture, whilst the lower becomes apicate.
Calcareous, hyaline, perforate wall, smooth and unornamented. Terminal,
rounded aperture with radiating grooves.
1950 Lenticulina (Robulus) asterizans Parr: 322,
pl. 11, figs. 9–10.
Material. One specimen from one sample.
Description. Planispiral, involute, lenticular test;
narrow rounded keel on periphery; eight chambers in final whorl. Limbate, flush
sutures; triangular, slightly rounded apertural face; calcareous, hyaline,
smooth wall. Aperture damaged in the specimen described here.
Occurrence. Originally described from Antarctica
(Parr, 1950).
Subfamily Marginulininae Wedekind, 1937
Genus Astacolus de Montfort, 1808
? Astacolus sp.(Fig. 11:16)
Material. One specimen from one sample.
Description. Elongate, flattened test; low, broad
chambers added on a slightly curved axis; strongly oblique, curved sutures;
rounded periphery; smooth, perforate, calcareous wall; terminal, radiate
aperture.
Family Lagenidae Reuss, 1862
Genus Lagena Walker & Jacob in Kanmacher, 1798
Lagena substriata Williamson, 1848(Fig. 12:6–7)
1848 Lagena substriata Williamson: 15, pl. 2,
fig. 12.
1994 Lagena substriata Williamson; Loeblich &
Tappan: 79, pl. 138, figs. 1–5.
2001 Lagena substriata elegantula R. W. Jones;
Igarashi et al.: 148, pl. 7, fig. 5.
2009 Lagena weisneri Parr; Majewski & Anderson:
139, fig. 4.
Material. Four specimens from three samples.
Description. Unilocular, ovate test that may be
significantly elongated. Calcareous, hyaline wall ornamented with numerous
delicate, parallel, longitudinal costae. Simple aperture with phialine lip at
the end of a long neck. Some of the costae continue up the neck, either
straight up or spiralling around the neck. Some specimens also present a
honeycomb pattern on the base of the test.
Occurrence. Originally described from the Recent of
England (Williamson, 1848), this species has also been described from the
Timor Sea (Loeblich and Tappan, 1994) and Antarctica (Igarashi et al., 2001;
Majewski and Anderson, 2009).
Lagena multilatera McCulloch, 1977(Fig. 12:8)
1977 Lagena multilatera McCulloch: 40, pl. 50,
fig. 5.
1994 Lagena multilatera McCulloch; Jones: 65,
pl. 58, figs. 2–3, 7–8, 22–24.
Material. Five specimens from three samples.
Description. Unilocular, fusiform test. Calcareous,
hyaline wall; approximately eight longitudinal costae, high, narrow,
tapering, some of which continue onto neck. Small, round, terminal aperture
produced on a long delicate neck.
Occurrence. McCulloch (1977) described this species
from deep water off Bikini Atoll, Pacific Ocean; subsequently recorded from
the South Pacific and Southern Ocean (Jones, 1994).
Material. Eighteen specimens from five samples.
Description. Fusiform, elongate, unilocular test
that slowly tapers at the base. Hyaline, calcareous, unornamented wall.
Simple aperture with a phialine lip, at the end of a long, tapering neck.
Occurrence. Originally described from the Miocene of
Italy (Seguenza, 1862); subsequently recorded from Recent sediments from
Antarctica (Majewski, 2005), the Southern Ocean (Jones, 1994), and the Bering
Sea (Setoyama and Kaminski, 2015).
Remarks. Whilst Jones (1994) retained this species in
Procerolagena, as he considered Hyalinonetrion to be a
junior synonym of Procerolagena, here we place it in
Hyalinonetrion due to the absence of any longitudinal ornamentation
(Loeblich and Tappan, 1988).
Genus Procerolagena Puri, 1954
Procerolagena distoma (Parker & Jones M. S. in Brady,
1864)(Fig. 12:10)
1864 Lagena distoma Parker & Jones in Brady: 467,
pl. 48, fig. 6.
? 1981 Lagena sulcata var. distomapolita Milam &
Anderson: 308, pl. 6, fig. 2.
1994 Hyalinonetrion distoma (Parker & Jones);
Loeblich & Tappan: 77, pl. 137, fig. 9.
Material. Thirteen specimens from nine samples.
Description. Unilocular, elongate test with nearly
parallel sides in the central part of the test which tapers towards apiculate
ends. Calcareous wall ornamented by delicate longitudinal striae.
Occurrence. Originally described from the Shetlands
(Brady, 1864); subsequently recorded from Antarctica (Milam and Anderson,
1981) and the Timor Sea (Loeblich and Tappan, 1994).
Remarks. Loeblich and Tappan (1994) assign this
species to the genus Hyalinonetrion. However, this designation is in
conflict with the genus description in Loeblich and Tappan (1988) defining
Hyalinonetrion as unornamented.
Genus Pygmaeoseistron Patterson & Richardson, 1987
1913 Lagena hispidula Cushman: 14, pl. 5,
figs. 2–3.
2013 Pygmaeoseistron hispidulum (Cushman 1913);
Majewski: 180, fig. 8(11).
Material. Two specimens from two samples.
Description. Unilocular flask-shaped/ovoid test that
rapidly narrows to a tubular neck (delicate and often broken, as in figured
specimen), evenly covered in fine hispid ornamentation.
Occurrence. Originally described from the North
Pacific (Cushman, 1913); also recorded from the Amundsen Sea (Majewski,
2013).
Pygmaeoseistron sp.(Fig. 12:12)
Material. One specimen from one sample.
Description. Unilocular, ovoid test with a narrow,
tubular neck; sporadic pustulose ornamentation.
Remarks. Bears some similarity to
Pygmaeoseistron islandicum (Jones, 1994).
Family Ellipsolagenidae A. Silvestri, 1923
Subfamily Oolininae Loeblich & Tappan, 1961
Genus Favulina Patterson & Richardson, 1987
Favulina squamosa (Montagu, 1803)(Fig. 12:15)
1803 Vermiculum squamosum Montagu: 526, pl. 14,
fig. 2.
1994 Oolina squamosa (Montagu); Jones: 66, pl. 58,
figs. 28–32.
2015 Oolina squamosa (Montagu); Setoyama &
Kaminski: 23, figs. 7(14), 8(31).
Material. Two specimens from two samples.
Description. Unilocular, ovate test, circular in
section, narrowing towards the apertural end. Calcareous, hyaline wall,
ornamented with roughly perpendicular vertical costae. Horizontal costae join
the vertical to form loosely hexagonal shapes. Round aperture, terminal on a
short neck with a rounded lip.
Occurrence. Originally described from Britain
(Montagu, 1803); also recorded from the South Pacific (Jones, 1994) and the
Bering Sea (Setoyama and Kaminski, 2015).
Remarks. The specimens figured here most closely
resemble the specimen from Plate 58, fig. 32, of Jones (1994) from the
Challenger material that has previously been variously assigned to
Lagena hexagona (Williamson) by Brady (1881) and Thalmann (1932)
and Oolina hexagona (Williamson) by Barker (1960), Hermelin (1989),
and Van Marle (1991). The ornamentation on our specimens forms roughly
perpendicular lines, a feature not seen in Favulina hexagona (Williamson), and it has therefore been assigned to Oolina squamosa (Montagu), following Knight (1986), which is then assigned to the genus
Favulina following Clark et al. (1994).
Genus Oolina d'Orbigny, 1839b
Oolina globosa (Montagu, 1803)(Fig. 12:16–17)
1803 Vermiculum globosum Montagu: 523, pl. 1,
fig. 8.
1994 Oolina globosa (Montagu); Jones: 62, pl. 56,
figs. 1–3, 15–16.
Material. Two specimens from two samples.
Description. Unilocular, tear-drop shaped test; may
have apiculate base. Calcareous, hyaline, smooth wall. Terminal, simple,
round aperture, with an entosolenian tube that extends over half the length
of the chamber and flares dramatically at the end.
Occurrence. Recorded from the Southern Ocean (Jones,
1994).
Remarks. Following Jones (1994), specimens with an
apiculate base are not considered to be distinct from Oolina globosa
(Montagu).
Oolina spp.(Fig. 12:13)
Material. Eight specimens from four samples.
Description. Included in this group any unilocular,
ovate tests with calcareous, hyaline, smooth walls, an apiculate base, and a
rounded terminal aperture that have not been assigned to Oolina globosa (Montagu).
1865 Lagena sulcata Walker & Jones var.
striatopunctata Parker & Jones: 350, pl. 13, figs. 25–27.
2001 Vasicostella striatopunctata (Parker &
Jones); Igarashi et al.: 148, pl. 8, fig. 5.
2013 Vasicostella sp. Majewski: 180, fig. 8 (8).
Material. One specimen from one sample.
Description. Unilocular, flask-like, compressed
test; carinate periphery. Calcareous, hyaline wall, ornamented with strong
longitudinal costae that appear striped, perpendicular to their length, due
to differences in the optical properties of their calcite (only visible with
light microscopy). Rounded, terminal aperture, with a slight lip and a short,
centrally placed entosolenian tube.
Occurrence. Parker and Jones (1865) originally
described this species from Greenland, the Red Sea, the South Atlantic, and
the Indian Ocean. Igarashi et al. (2001) subsequently described it from
Antarctica.
Remarks. Le Coze and Hayward (2017) places Lagena striatopunctata Parker & Jones within the genus Cushmanina. However, we believe that, based on the type figures, this classification is incorrect.
The compressed nature of the test and the lack of an elongate neck suggest
that this species should be placed within the genus Vasicostella.
The specimen described here and that of Igarashi et al. (2001) also reveal
the carinate nature of this species, further supporting this classification.
Subfamily Ellipsolageninae A. Silvestri, 1923
Genus Fissurina Reuss, 1850
Fissurina subformosa Parr, 1950(Fig. 12:19)
1950 Fissurina subformosa Parr: 313, pl. 9, fig. 9.
1996 Lagena subformosa (Parr); Violanti: 43, pl. 7,
fig. 3.
Material. Three specimens from three samples.
Description. Small, pyriform test, slightly
compressed, tapering towards a long neck; keel extends from the neck around
the test, with two tabulated supplementary keels that create a “crimped”
appearance. Circular, terminal aperture; straight entosolenian tube extends
centrally to approximately one third the length of the chamber.
Occurrence. Previously recorded from Antarctica
(Parr, 1950; Violanti, 1996).
Remarks. This species can be distinguished from
F. formosa (Lagena Formosa Schwager) by the supplementary keels.
Fissurina sp. 1(Fig. 12:14)
Material. Seven specimens from four samples.
Description. Unilocular, pyriform test, with pointed
protuberance towards basal end; circular section. Smooth, hyaline, calcareous
wall; terminal, oval aperture within a slightly depressed fissure at the apex
of the test.
Fissurina sp. 2(Fig. 12:20)
Material. Thirty-one specimens from 15 samples.
Description. Elongate, compressed test, broadening
towards the base; rounded ends; base somewhat produced; oval section. Smooth,
hyaline, calcareous wall; terminal, slit-like aperture within a significantly
depressed fissure at the apex of the test.
Remarks. Bears some similarity to the Oligocene
Fissurina oblonga Reuss and to Parafissurina lateralis
(Cushman). May have previously been recorded from the Weddell Sea by
Anderson (1975) as Parafissurina lateralis (Cushman).
1931 Parafissurina fusiformis Wiesner: 126, pl. 24,
fig. j.
1996 Parafissurina fusiformis Wiesner; Violanti, 62,
pl. 8, fig. 8.
2001 Parafissurina fusiformis Wiesner; Igarashi:
151, pl. 9, fig. 5.
2005 Parafissurina fusiformis Wiesner; Majewski:
203, fig. 22(1–2).
Material. Thirty-seven specimens from 14 samples.
Description. Small test, ovate in front view; slightly compressed, rounded periphery. Aperture an elongate slit protected
by a protruding hood, with a large lip opposite the hood. Entosolenian tube
extends centrally along inner surface of chamber, flaring at the end.
Occurrence. Originally described from high southern
latitudes (Wiesner, 1931) and later from other Antarctic localities
(Violanti, 1996; Majewski, 2005).
Family Bolivinidae Glaessner, 1937
Genus Bolivinellina Saidova, 1975
Bolivinellina earlandi (Parr, 1950)(Fig. 12:22)
1950 Bolivina earlandi Parr: 339, pl. 12, fig. 16.
1996 Bolivina pseudopunctata Höglund; Violanti:
65, pl. 9, fig. 1.
2001 Bolivinellina earlandi (Parr); Igarashi et al.:
153, pl. 10, fig. 5.
2013 Bolivinellina earlandi (Parr); Majewski: 181,
fig. 9(4).
Material. Seventy specimens from 19 samples (for
Bolivinellina spp.).
Description. Elongate test, biserial throughout,
oval in section. Chambers up to 12 in number, slightly inflated; strongly
curved, slightly depressed sutures. Wall calcareous, hyaline, smooth; finely
perforate in the lower half of the chamber, imperforate in the upper half.
Narrow, comma-shaped aperture starting at the inner margin of the final
chamber.
Occurrence. Parr (1950) described this species as
widely distributed around Antarctica.
Remarks. Following Igarashi et al. (2001) and
Majewski (2013), we have placed this species in the genus
Bolivinellina due the lack of perforation in the upper half of the
chambers.
1947 Bolivina pseudopunctata Höglund: 273,
pl. 24, fig. 5, pl. 32, figs. 23–24.
2005 Bolivina pseudopunctata Höglund; Majewski:
203, fig. 22(6–7).
2013 Bolivinellina pseudopunctata (Höglund);
Majewski: 181, fig. 9(3).
Material. Seventy specimens from 19 samples (for
Bolivinellina spp.).
Description. Elongate, gently tapering test,
biserial from a large globular proloculus, oval in section. Chambers
distinct; up to 23 in number; slightly inflated; sutures strongly curved,
slightly depressed. Wall calcareous, hyaline, smooth, finely perforate in the
lower half of the chamber, imperforate in the upper half. Oval-shaped
aperture with a double-folded tongue, starting at the inner margin of the
final chamber.
Occurrence. Originally described from the Gullmar
Fjord, Sweden (Höglund, 1947), this species has subsequently been
described from Antarctica (Majewski, 2005, 2013).
Remarks. Following Majewski (2013) we have placed
this species in the genus Bolivinellina due the lack of perforation
in the upper half of the chambers. B. pseudopunctata can be
distinguished from B. earlandi by the greater number of chambers, an
oval-shaped aperture with a double-folded tongue, and a large globular
proloculus.
Material. Overall, 713 specimens from 32 samples.
Description. Crozier-shaped test, initially
planispiral, rapidly uncoiling to become biserial; number of chambers in
biserial section quite variable; biserial section cylindrical, straight or
arcuate. Chambers subglobose, inflated, with small overlap at the periphery;
sutures slightly depressed. Thick, calcareous, perforate, hyaline
bi-multilamellar wall. Interiomarginal, loop-shaped aperture; subterminal.
Occurrence. Originally described from the South
Pacific (Brady, 1881) and subsequently described from various locations
around Antarctica (Heron-Allen and Earland, 1932a; Anderson, 1975; Igarashi et al., 2001; Majewski, 2005;
Majewski and Anderson, 2009; Rodrigues et al., 2013).
Cassidulinoides porrectus (Heron-Allen and Earland,
1932a)(Fig. 13:1–2)
1932a Cassidulina crassa d'Orbigny var.
porrecta Heron-Allen & Earland, p. 358, pl. 9, figs. 34–37.
1981 Cassidulinoides porrecta (Heron-Allen &
Earland); Milam & Anderson: 308, pl. 6, fig. 6.
1996 Cassidulinoides porrectus (Heron-Allen &
Earland): Violanti: 65, pl. 9, figs. 2–3.
2001 Cassidulinoides porrectus (Heron-Allen &
Earland); Igarashi et al.: 153, pl. 10, fig. 11.
2005 Cassidulinoides porrectus (Heron-Allen &
Earland); Majewski: 204, fig. 23(3).
2013 Cassidulinoides porrectus (Heron-Allen &
Earland); Rodrigues et al.: p. 214, fig. 3 (3).
Material. Overall, 1452 specimens from 37 samples.
Description. Test small, planispirally enrolled in
early stages, uncoiling to become biserial; commonly 2–4 chambers in the
biserial part but can be as many as 10. Chambers subglobose, inflated, with
small overlap at the periphery; sutures slightly depressed. Thick,
calcareous, perforate, hyaline wall, covered in distinctive large pores.
Interiomarginal, loop-shaped aperture, subterminal.
Occurrence. Originally described from various
subantarctic locations (Heron-Allen and Earland, 1932a), subsequently in
numerous additional Antarctic studies (Milam and Anderson, 1981; Violanti, 1996; Igarashi et al., 2001;
Majewski, 2005; Rodrigues et al., 2013).
1967 Globocassidulina crassa (d'Orbigny)
subsp. rossensis Kennett 134, pl. 11, figs. 4–6.
1981 Globocassidulina crassa rossensis Kennett;
Milam & Anderson: 312, pl. 10, fig. 1.
1993 Globocassidulina rossensis Kennett; Mackensen
et al.: 56, pl. 2, figs. 7–8.
1996 Globocassidulina rossensis Kennett; Violanti:
p. 65, pl. 9, fig. 7.
2006 Globocassidulina rossensis Kennett; Hromic et
al.: 124, pl. 1, fig. 5.
Material. Overall, 727 specimens from 32 samples.
Description. Oval to circular, involute, slightly
compressed test; rounded, slightly lobulate, planispiral coil of biserial,
inflated chambers that gradually increase in size; four to five pairs in the last
whorl. Thin, smooth, calcareous wall. Aperture a narrow, interiomarginal slit
with an aerial branch that extends perpendicular to the main slit
approximately halfway up the apertural face, surrounded by a narrow rim.
Occurrence. Originally described from the Ross Sea
(Kennett, 1967); subsequently described from other Antarctic locations (Milam
and Anderson, 1981; Mackensen et al., 1993; Violanti, 1996; Hromic et al.,
2006).
Remarks. Distinguished from other subspecies of
Globocassidulina crassa (d'Orbigny) by an aerial branch that extends
from the main part of the aperture.
Subfamily Ehrenbergininae Cushman, 1927
Genus Ehrenbergina Reuss, 1850
Ehrenbergina glabra Heron-Allen & Earland, 1922
1922 Ehrenbergina hystrix var. glabra Heron-Allen
& Earland: 140, pl. 5, figs. 1–6.
1981 Ehrenbergina glabra Heron-Allen & Earland;
Milam & Anderson: 311, pl. 9, fig. 7.
1990 Ehrenbergina glabra Heron-Allen & Earland;
Mackensen et al.: 254, pl. 1, figs. 5–6.
1996 Ehrenbergina glabra Heron-Allen & Earland;
Violanti: 65, pl. 9, fig. 9.
2001 Ehrenbergina glabra Heron-Allen & Earland;
Igarashi et al.: 154, pl. 11, fig. 1.
2013 Ehrenbergina glabra Heron-Allen & Earland;
Majewski: 182, fig. 10(13).
Material. Two specimens from one sample.
Description. Biserial test, initially enrolled
rapidly uncoiling; lenticular in section with a convex dorsal side; flatter
ventral side with a median furrow. Chambers are low and broad with
considerable overlap at the peripheral midline. Spinose lateral margins; sutures curved, flush. Calcareous, smooth wall. Curved, elongate, slit-like
aperture that runs parallel to the peripheral margin, perpendicular to the
base of the apertural face.
Occurrence. Recorded from a number of locations
around Antarctica (Milam and Anderson, 1981; Mackensen et al., 1990;
Violanti, 1996; Igarashi et al., 2001; Majewski, 2013).
Remarks. Heron-Allen and Earland (1922)
distinguished this species from E. hystrix Brady by the absence of
spines on the early chambers, the more flattened oral face, the more marginal
position of the aperture, and the near absence of striations around the
aperture.
Family Buliminidae Jones in Griffith and Henfrey, 1875
Genus Bulimina d'Orbigny, 1826
Bulimina aculeata d'Orbigny, 1826(Fig. 13:3–4)
1826 Bulimina aculeata d'Orbigny: 269 (no type
figure).
1990 Bulimina aculeata d'Orbigny; Mackensen et al.:
255, pl. 2, figs. 1–3.
1993 Bulimina aculeata d'Orbigny; Mackensen et al.:
54, pl. 1, figs. 3–4.
1994 Bulimina aculeata d'Orbigny; Ishman &
Domack: 152, pl. 2, fig. 8.
2001 Bulimina aculeata d'Orbigny; Igarashi et al.:
154, pl. 11, fig. 4.
2013 Bulimina aculeata d'Orbigny; Majewski: 181,
fig. 9(8–9).
Material. Fifty-nine specimens from 20 samples.
Description. Elongate, triserial test, strongly
tapered towards base. Globular, inflated chambers rapidly increase in size; distinct, depressed sutures. Smooth, finely perforate, calcareous wall. Early
chambers ornamented with long, tapering spines that extend dorsally; fragile
and often partially or completely broken. Loop-shaped aperture extends up the
face of the final chamber from the basal margin; well-developed, elevated
rim.
Occurrence. Originally described from the Adriatic
(d'Orbigny, 1826) but more recently from various Antarctic localities
(Mackensen et al., 1990, 1993; Ishman and Domack, 1994; Igarashi et al.,
2001; Majewski, 2013).
Bulimina subteres Brady, 1881
1881 Bulimina subteres Brady: 55, pl. 50,
figs. 17–18.
Material. Three specimens from three samples.
Description. Elongate, triserial, ovate test,
rounded at the distal end and tapering to a point at the basal end; slightly
inflated chambers; depressed sutures; curved, slit-like aperture extending up
the face of the terminal chamber.
Occurrence. Originally described from the Recent of
Fiji and the Caribbean (Brady, 1881).
Bulimina sp. 1(Fig. 13:9–10)
Material. Twenty-four specimens from 11 samples.
Description. Near spherical test, wider than high;
only the three chambers of the final whorl visible. Inflated, globular
chambers; depressed sutures. Calcareous, smooth, perforate wall; loop-shaped
aperture extending from the base up the face of the final chamber, with a
raised rim.
Family Uvigerinidae Haeckel, 1894
Subfamily Angulogerininae Galloway, 1933
Genus Trifarina Cushman, 1923
Trifarina earlandi (Parr, 1950)(Fig. 13:8)
1950 Angulogerina earlandi Parr: 341, pl. 12,
fig. 21.
2001 Angulogerina earlandi Parr; Igarashi et al.:
154, pl. 11, fig. 7.
2005 Angulogerina earlandi Parr; Majewski: 203,
fig. 22(8–9).
2013 Angulogerina earlandi Parr; Majewski: 181,
fig. 9(13–14).
Material. Overall, 204 specimens from 22 samples.
Description. Elongate, fusiform test, triserial,
becoming uniserial; triangular cross section becoming rectilinear with
rounded to sub-acute corners. Base rounded; slightly lobulate margins.
Numerous inflated chambers ornamented with narrow, high costae that generally
do not cross the sutures; in some specimens final chambers lack
ornamentation. Terminal, ovate aperture, with internal hemicylindrical
tooth plate, produced on a neck with a phialine lip.
Occurrence. Recorded from a number of locations
around Antarctica (Parr, 1950; Igarashi et al., 2001; Majewski, 2005, 2013).
Remarks. Parr (1950) placed this species within
Angulogerina. However, Loeblich and Tappan (1988) define
Angulogerina as triserial throughout, whilst this species is
triserial, becoming uniserial, diagnostic of Trifarina. The high
narrow costae of Trifarina earlandi (Parr) distinguish it from the
otherwise similar T. angulosa (Williamson).
1930 Virgulina fusiformis Cushman: 45, pl. 8,
fig. 8.
1981 Fursenkoina fusiformis (Cushman); Milam &
Anderson: 311, pl. 9, fig. 6.
1994 Fursenkoina fusiformis (Cushman); Ishman &
Domack: 152, pl. 2, fig. 7.
1996 Fursenkoina fusiformis (Cushman); Violanti: 65,
pl. 9, figs. 14–15.
2005 Fursenkoina fusiformis (Cushman); Majewski:
204, fig. 23(9–12).
2009 Fursenkoina fusiformis (Cushman); Majewski &
Anderson: 139, fig. 4.
Material. Overall, 1260 specimens from 34 samples.
Description. Narrow, elongate test, ovate in
section. High, narrow, slightly inflated chambers, biserial, twisted around
the test axis. Oblique, depressed sutures; calcareous, hyaline, finely
perforate wall, with smooth surface. Narrow, elongate aperture; basal part of
the aperture may be closed in adult specimens, leaving an areal comma-shaped
opening.
Occurrence. Recorded from a range of Antarctic
locations (Milam and Anderson, 1981; Ishman and Domack, 1994; Violanti, 1996;
Majewski, 2005; Majewski and Anderson, 2009).
Remarks. Very similar to Stainforthia fusiformis (Williamson), a species common in the high, northern latitudes.
F. fusiformis (Cushman) can be distinguished from this species by
the shape of the aperture and by being biserial throughout. F. fusiformis (Cushman) is common throughout the high, southern latitudes.
Occurrence. Originally described from the Falkland
Islands (d'Orbigny, 1839b), subsequently from various locations on the
Antarctic shelf (Heron-Allen and Earland, 1932a; Anderson, 1975; Igarashi et al., 2001; Majewski, 2005, 2013;
Majewski and Anderson, 2009).
Remarks. Chambers less inflated and alar projections
more angular than those of Rosalina globularis d'Orbigny, with which
it may sometimes be confused.
Family Cibicididae Cushman, 1927
Subfamily Cibicidinae Cushman, 1927
Genus Cibicides de Montfort, 1808
Cibicides sp. 1(Fig. 13:11–12)
Material. Four specimens from four samples.
Description. Trochospiral, planoconvex test; flat,
slightly involute spiral side with somewhat thickened sutures; strongly
convex, completely involute umbilical side. Sutures flush except around final
chamber; rounded periphery; calcareous wall, coarsely perforate on spiral
side with five large pores in each chamber; pores in earlier chambers filled
by lamellar thickening; umbilical side finely perforate; keel and apertural
face imperforate. Low, interiomarginal, equatorial aperture extending along
the spiral side suture and slightly onto the umbilical side, with a narrow
lip.
Cibicides sp. 2
Material. One specimen from one sample.
Description. Trochospiral, planoconvex test; flat,
slightly involute spiral side with somewhat thickened sutures; strongly
convex, completely involute umbilical side. Calcareous wall, coarsely
perforate on spiral side; umbilical side finely perforate; keel and apertural
face imperforate; aperture not preserved.
Family Nonionidae Schultze, 1854
Subfamily Nonioninae Schultze, 1854
Genus Nonionella Cushman, 1926
Nonionella bradii (Chapman,
1916)(Fig. 14:2–4)
1916 Nonionina scapha (Fichtel & Moll) var.
bradii Chapman: 71, pl. 5, fig. 42.
1990 Nonionella bradii (Chapman); Mackensen et al.:
254, pl. 1, fig. 4.
1994 Nonionella bradii (Chapman); Ishman &
Domack: 152, pl. 2, figs. 10 & 12.
1994 Nonionella bradii (Chapman); Jones: 108, pl.
109, fig. 16.
1996 Nonionella bradii (Chapman); Violanti: 67, pl.
10, figs. 8, 13, 17.
2001 Nonionella bradii (Chapman); Igarashi et al.:
155, pl. 12, fig. 5.
2005 Nonionella bradii (Chapman); Majewski: 206,
fig. 25(4–5).
2009 Nonionella bradii (Chapman); Majewski &
Anderson: 139, fig. 4.
Material. Sixty-seven specimens from 17 samples.
Description. Low trochospiral, compressed test with
rounded periphery; involute umbilical side; partially evolute spiral side.
Low, broad chambers that widen towards the periphery; 9–10 in the final
whorl; chambers rapidly enlarge in the final whorl to produce a flared test.
Flap-like projections from each chamber overhang the umbilicus. Slightly
curved, narrow, depressed sutures. Calcareous, hyaline, finely perforate wall
with a smooth surface. Aperture a low arch, interiomarginal, equatorial.
Occurrence. First described from the Ross Sea
(Chapman, 1916); subsequently reported from numerous other Antarctic
localities (Mackensen et al., 1990; Ishman and Domack, 1994; Jones, 1994; Violanti, 1996; Igarashi et al., 2001; Majewski, 2005; Majewski and Anderson, 2009).
Nonionella iridea Heron-Allen & Earland, 1932a(Fig. 14:1 and 5)
1932a Nonionella iridea Heron-Allen & Earland:
438, pl. 16, figs. 14–16.
1990 Nonionella iridea Heron-Allen & Earland;
Mackensen et al.: 254, pl. 1, figs. 7–9.
1994 Nonionella iridea Heron-Allen & Earland;
Ishman & Domack: 152, pl. 2, fig. 11.
1996 Nonionella iridea Heron-Allen & Earland;
Violanti: 67, pl. 10, figs. 9, 14, 18.
2001 Nonionella iridea Heron-Allen & Earland;
Igarashi et al.: 155, pl. 12, fig. 6.
2005 Nonionella iridea Heron-Allen & Earland;
Majewski: 206, fig. 25(2–3).
2009 Nonionella iridea Heron-Allen & Earland;
Majewski & Anderson: 139, fig. 4.
2013 Nonionella iridea Heron-Allen & Earland;
Majewski: 182, fig. 10(8).
Material. Overall, 323 specimens from 28 samples.
Description. Low trochospiral, compressed test, with
rounded periphery; involute umbilical side; partially evolute spiral side.
Low, broad, somewhat inflated chambers that widen towards the periphery; six to
seven in the final whorl. Chambers enlarge in the final whorl to produce a slightly
flared test; flap-like projections from each chamber overhang the umbilicus
and overlap each other. Slightly curved, narrow, depressed sutures.
Calcareous, hyaline, finely perforate wall with a smooth surface. Aperture a
low arch; interiomarginal, equatorial, supplementary apertures develop in
later chambers along the sutures from the umbilicus approximately halfway to
the periphery (not visible in all specimens).
Occurrence. Originally described from the
subantarctic, in particular South Georgia and the Falkland Islands
(Heron-Allen and Earland, 1932a); subsequently reported from locations around
Antarctica (Mackensen et al., 1990; Ishman and Domack, 1994; Violanti, 1996;
Igarashi et al., 2001; Majewski, 2005, 2013; Majewski and Anderson, 2009).
Subfamily Astrononioninae Saidova, 1981
Genus Astrononion Cushman and Edwards, 1937
Astrononion echolsi Kennett, 1967(Fig. 14:6–8)
1967 Astrononion echolsi Kennett: 134, pl. 11,
figs. 7–8.
1981 Astrononion echolsi Kennett; Milam &
Anderson: 313, pl. 11, fig. 3.
1994 Astrononion echolsi Kennett; Ishman &
Domack: 152, pl. 2, fig. 3.
1996 Astrononion echolsi Kennett; Violanti: 67,
pl. 10, fig. 16.
2001 Astrononion echolsi Kennett; Igarashi et al.:
155, pl. 12, fig. 11.
2005 Astrononion echolsi Kennett; Majewski: 206,
fig. 25(6–7).
2009 Astrononion echolsi Kennett; Majewski &
Anderson: 139, fig. 4
2013 Astrononion echolsi Kennett; Majewski: 182,
fig. 10(7).
Material. Overall, 1216 specimens from 39 samples.
Description. Involute, planispiral, slightly
compressed test, with a rounded periphery. Size of chambers gently increases
as they are added; eight to nine in the final whorl; somewhat inflated in some
specimens but more commonly not. Slightly limbate, curved sutures; slightly
depressed to flush. Indistinct, tube-like supplementary chambers extend from
the umbilical region, only visible as sunken pits on the sutural lines.
Calcareous, hyaline, finely perforate, smooth wall, often with slightly
yellow colour. Primary aperture a low, interiomarginal, equatorial arch.
Occurrence. Abundant in the Ross Sea and Scotia Sea,
where it was first described (Kennett, 1967); also recorded from various
other Antarctic locations (Milam and Anderson, 1981; Ishman and Domack, 1994;
Violanti, 1996; Igarashi et al., 2001; Majewski, 2005, 2013; Majewski and
Anderson, 2009).
Subfamily Pulleniinae Schwager, 1877
Genus Pullenia Parker & Jones in Carpenter et al., 1862
1839b Nonionina subcarinata d'Orbigny: 28, pl. 5,
figs. 23–24.
1932a Pullenia subcarinata (d'Orbigny); Heron-Allen
& Earland: 403, pl. 13, figs. 14–18.
1981 Pullenia subcarinata (d'Orbigny); Milam &
Anderson: 313, pl. 11, fig. 5.
1993 Pullenia subcarinata (d'Orbigny); Mackensen et
al.: 58, pl. 3, figs. 8–9.
? 1994 Pullenia quinqueloba (Reuss); Jones: 92,
pl. 84, figs. 14–15.
1996 Pullenia subcarinata (d'Orbigny); Violanti: 48,
pl. 10, fig. 19.
Material. Thirty-nine specimens from 18 samples.
Description. Involute, planispiral, slightly
compressed test with a rounded periphery. Chambers slightly inflated, giving
the test a slightly lobulate outline; five to six in the final whorl; face of final
chamber has a somewhat triangular shape. Near-radial, slightly depressed
sutures. Calcareous, smooth, very finely perforate wall. Aperture a low,
interiomarginal, equatorial, crescentic slit, bordered on the upper margin by
a raised lip.
Occurrence. Originally described from recent
deposits from the Falkland Islands (d'Orbigny, 1839b); subsequently recorded
from around Antarctica (Heron-Allen and Earland, 1932a; Milam
and Anderson, 1981; Mackensen et al., 1993; Jones, 1994; Violanti, 1996).
Remarks.Pullenia quinqueloba (Reuss) is
morphologically very similar to P. subcarinata (d'Orbigny), and
Jones (1994) assigns the Challenger specimens to P. quinqueloba (Reuss). However, Pullenia quinqueloba (Reuss) was originally
described from German Eocene deposits, and we therefore assign our specimens
to P. subcarinata (d'Orbigny), which was originally described from
Recent deposits from the Falkland Islands.
Family Chilostomellidae Brady, 1881
Subfamily Chilostomellinae Brady, 1881
Genus Chilostomella Reuss, 1849
Chilostomella sp. 1(Fig. 14:13)
Material. One specimen from one sample.
Description. Ovoid, planispiral, involute test;
near-circular cross section. Two strongly embracing, inflated chambers in
final whorls; final chamber significantly larger. Calcareous, finely
perforate wall; smooth surface; equatorial, interiomarginal, narrow
slit-shaped aperture, with a raised margin. Distinctive spine-like structure
on basal end.
Family Trichohyalidae Saidova, 1981
Genus Buccella Andersen, 1952
Buccella sp. 1(Fig. 14:14–16)
Material. Overall, 241 specimens from 31 samples.
Description. Large, biconvex, carinate, trochospiral
test; spiral side low convex and evolute; umbilical side higher and involute.
Numerous rhomboidal chambers with curved sides, slightly inflated on the
umbilical side; three and a half whorls; 8–10 chambers in the final whorl.
Sutures on spiral side flush, slightly limbate, swept back towards periphery,
straighter and depressed on the umbilical side. Umbilical region and sutural
fissures on umbilical side filled with a mixture of fine and coarse granules;
spiral side surface smooth, finely perforate on the chamber faces; periphery
very slightly lobate, with a distinct keel. Aperture not visible in any of
the specimens examined; may be covered by the abundant granules.
Remarks. Openings at the end of the sutural fissures
that are common in members of this genus are not visible in any of the
specimens examined here.
Buccella sp. 2(Fig. 14:11–12)
Material. Twenty-five specimens from 12 samples.
Description. Test small for this genus;
trochospiral, biconvex; spiral side flatter and evolute; umbilical side
convex and involute. Chambers almost semi-circular; two whorls; six to seven chambers
in the final whorl; sutures limbate, flush, slightly curved on the spiral
side, very depressed on the umbilical side. Umbilical region and sutural
fissures on umbilical side filled with granules; spiral side surface smooth,
finely perforate on the chamber faces; periphery slightly lobate and
thickened but not carinate. Aperture not visible in any of the specimens
examined and may be covered by the abundant granules.
Remarks. Bears some similarity to
Epistominella exigua (Brady), especially in light microscope.
Genus Neogloboquadrina Bandy, Frerichs & Vincent, 1967
Neogloboquadrina pachyderma (Ehrenberg, 1861)
1861 Aristerospira pachyderma Ehrenberg: 276, 277,
303.
1932a Globigerina pachyderma Ehrenberg; Heron-Allen
& Earland: 401, pl. 13, figs. 9–13.
1975 Globigerina pachyderma Ehrenberg; Anderson: 90,
pl. 9, fig. 4.
1996 Neogloboquadrina pachyderma (Ehrenberg);
Violanti: 62, pl. 8, figs. 17 and 21.
2001 Neogloboquadrina pachyderma (Ehrenberg);
Igarashi et al.: 152, pl. 10, figs. 8–9.
Material. Overall, 209 specimens from 27 samples.
Description. Globular, low trochospiral test; globular chambers that rapidly enlarge as added; five to six in the final whorl.
Straight to slightly curved, depressed, radial sutures; rounded periphery; open umbilical region. Calcareous, perforate wall; interiomarginal,
extra-umbilical to umbilical aperture with bordering lip (may be absent in
later stages).
All the data analysed in this paper are available in
the tables provided within the paper or in the Supplement.
The Supplement related to this article is available online at https://doi.org/10.5194/jm-37-25-2018-supplement.
The authors declare that they have no conflict of
interest.
Acknowledgements
This study forms part of a doctoral project carried out by Rowan Dejardin,
funded by the Centre for Environmental Geochemistry (University of Nottingham
and the British Geological Survey) and supported by BGS-University Funding
Initiative (BUFI). We thank the officers, crew, and scientific party on board
the RRS James Clark Ross during scientific cruises JR257 and
JC15002. Participation in cruise JR15002 by Rowan Dejardin was funded by the
British Antarctic Survey Collaborative Gearing Scheme. We thank Hilary Sloane
and Jack Lacey (BGS) for analytical assistance and Tony Milodowski and
Gren Turner for their assistance generating the SEM images. We also thank the
editor, Laia Alegret, one anonymous reviewer, and Wojciech Majewski for their
insightful comments, which helped to strengthen this paper. This project
was supported by NERC Isotope Geoscience Facility grant IP/1495/1114
(RCUK). Edited by: Laia Alegret Reviewed by:
Wojciech Majewski and one anonymous referee
References
Alekseychik-Mitskevich, L. S.: Towards the classification of the
foraminiferal family Haplophragmiidae, Trudy Vsesoyuzznogo neftyanogo
Nauchnoissledovatel'skogo Geologorazvedochnogo Instituta, 343, 12–44, 1973.
Andersen, H. V.: Buccella, a new genus of the rotalid foraminifera, Journal
of the Washington Academy of Sciences, 42, 143–151, 1952.
Anderson, J. B.: Ecology and distribution of foraminifera in the Weddell Sea
of Antarctica, Micropalaeontology, 21, 69–96, 1975.
Bailey, J. W.: Microscopical examination of soundings made by the U.S. Coast
Survey of the Atlantic coast of the U.S., Smithsonian Contributions, 2,
1–48, 1851.Bandy, O. L., Frerichs, W. E., and Vincent, E.: Origin, development, and
geologic significance of Neogloboquadrina Bandy, Frerichs, and
Vincent, gen. nov., Contributions from the Cushman Foundation for
Foraminiferal Research, 18, 152–157, 1967.
Barker, R. W.: Taxonomic notes on the species figured by H.B. Brady in his
report of the foraminifera dredged by HMS Challenger during the years
1873–1876, American Association of Petroleum Geologists Special Publication,
9, 10238, 2–240, 1960.
Bernhard, J. M.: Distinguishing live from dead foraminifera: Methods review
and proper applications, Micropaleontology, 46, 38–46, 2000.Brady, H. B.: Contributions to the knowledge of the foraminifera. – On the
rhizopodal fauna of the Shetlands, Transactions of the Linnaean Society, 24,
463–476, 10.1111/j.1096-3642.1863.tb00170.x, 1864.
Brady, H. B.: Notes on some of the Reticularian Rhizopoda of the
“Challenger” Expedition. I. On new or little known arenaceous types, Q. J.
Microsc. Sci., 19, 20–62, 1879.
Brady, H. B.: Notes on some of the Reticularian Rhizopoda of the
“Challenger” Expedition. Part III, Q. J. Microsc. Sci., 21, 31–71, 1881.
Brady, H. B.: Report on the scientific results of the voyage of H.M.S.
Challenger during the years 1873–76, in: Zoology of the Challenger
Expedition, edited by: Thompson, C. W. and Murray, J., London, Neill,
Edinburgh,
1884.Brandon, M. A., Murphy, E. J., Whitehouse, M. J., Trathan, P. N., Murray, A.
W. A., Bone, D. G., and Priddle, J.: The shelf break front to the east of the
sub-Antarctic island of South Georgia, Cont. Shelf Res., 19, 799–819,
10.1016/s0278-4343(98)00112-5, 1999.
Brandon, M. A., Murphy, E. J., Trathan, P. N., and Bone, D. G.: Physical
oceanographic conditions to the northwest of the sub-Antarctic Island of
South Georgia, J. Geophys. Res.-Oceans, 105, 23983–23996,
10.1029/2000jc900098, 2000.
Brönnimann, P.: Two new genera of Recent Trochamminidae (Foraminiferida),
Achives des Sciences, Geneve, 29, 215–218, 1976.
Brönnimann, P. and Beurlen, G.: Recent benthonic foraminifera from Brasil.
Morphology and ecology. Part I, Achives des Sciences, Geneve, 30, 77–89,
1977.Brönnimann, P. and Whittaker, J. E.: The Trochamminacea of the Discovery
Reports, British Museum, London, 1988.
Brönnimann, P., Zaninetti, L., and Whittaker, J. E.: On the classification of
the Trochamminacea (Foraminiferida), J. Foramin. Res., 13, 202–218, 1983.Caralp, M. H.: Abundance of Bulimina exilis and Melonis barleeanum – relationship to the quality of marine organic-matter,
Geo-Marine Letters, 9, 37–43, 10.1007/bf02262816, 1989.
Carpenter, W. B.: On the rhizopodal fauna of the deep sea, Proceedings of the
Royal Society of London, 18, 59–62, 1869.
Carpenter, W. B., Parker, W. K., and Jones, T. R.: Introduction to the study
of the Foraminifera, Ray Society, London, 1862.Caulle, C., Mojtahid, M., Gooday, A. J., Jorissen, F. J., and Kitazato, H.:
Living (Rose-Bengal-stained) benthic foraminiferal faunas along a strong
bottom-water oxygen gradient on the Indian margin (Arabian Sea),
Biogeosciences, 12, 5005–5019, 10.5194/bg-12-5005-2015,
2015.
Chapman, F.: Report on the Foraminifera and Ostracoda from elevated deposits
on the shores of the Ross Sea, British Antarctic Expedition 1907–9 under the
command of Sir E. H. Shackleton, C.V.O.: Reports on the scientific
investigations, Geology, Vol. II – Contributions to the palaeontology and
petrology of South Victoria Land, edited by: Benson, W. N., Chapman, F.,
Cohen, F., Cotton, L. A., Hedley, C., Jensen, H. I., Mawson, D., Skeats, E.
W., Thomson, J. A., Walkom, A. B., and Woolnough, W. G., William Heinemann,
London, 1916.
Chapman, F., Parr, W. J., and Collins, A. C.: Tertiary Foraminifera of
Victoria, Australia. The Balcombian deposits of Port Phillip, Journal of the
Linnaean Society, 38, 553–576, 1934.
Clark, F. E., Patterson, R. T., and Fishbein, E.: Distribution of Holocene
benthic foraminifera from the tropical southwest Pacific Ocean, J. Foramin.
Res., 24, 241–267, 1994.Corliss, B. H. and Emerson, S.: Distribution of Rose-Bengal stained deep-sea
benthic foraminifera from the Nova Scotian continental-margin and Gulf of
Maine, Deep-Sea Res. Pt. A, 37, 381–400, 10.1016/0198-0149(90)90015-n,
1990.
Cushman, J. A.: A monograph of the Foraminifera of the North Pacific Ocean.
Part I. Astrorhizidae and Lituolidae, Bulletin of the United States National
Museum, 71, 1–134, 1910.
Cushman, J. A.: A monograph of the foraminifera of the North Pacific Ocean.
Pt. III. Lagenidae, Bulletin of the United States National Museum, 71,
1–125, 1913.
Cushman, J. A.: The foraminifera of the Atlantic Ocean, Part 4. Lagenidae,
Bulletin United States National Museum, 104, 1–228, 1923.
Cushman, J. A.: Foraminifera of the typical Monerey of California, Cushman
Foundation for Foraminiferal Research Special Publication, 2, 53–69, 1926.
Cushman, J. A.: An outline of a re-classification of the foraminifera,
Contributions from the Cushman laboratory for foraminiferal research, 3,
1–105, 1927.
Cushman, J. A.: The foraminifera of the Atlantic Ocean, Part 7, Nonionidae,
Camerinidae, Peneroplidae and Alveolinellidae, Bulletin United States
National Museum, 104, 1–79, 1930.
Cushman, J. A.: Foraminifera, their classification and economic use Cushman
Lab. Foram. Res., Special Publications Cushman Laboratory for Foraminiferal
Research, 4, 1–349, 1933.
Cushman, J. A.: Foraminifera of the United States Antarctic Service
Expedition 1939–1941, Proceedings, American Philosophical Society, 89,
285–288, 1945.Cushman, J. A. and Edwards, P. G.: Astrononion a new genus of the
foraminifera and its species, Contributions from the Cushman laboratory for
foraminiferal research, 13, 29–36, 1937.
d'Orbigny, A.: Tableau méthodique de la classe des Céphalopodes, Ann.
Sci. Nat., 7, 245–314, 1826.
d'Orbigny, A.: Foraminifères, in: Histoire Physique, Politique et
Naturelle de l'île de Cuba, edited by: de la Sagra, R., A. Bertrand,
Paris, 1–224, 1839a.
d'Orbigny, A.: Voyage dans l'Amérique Méridionale (le Brésil, la
République orientale de l'Uruquay, la République Argentine, la
Patagonie, la République du Chili, la République de Bolivia, la
République du Pérou) éxécuté pendant les années 1826,
1827, 1832 et 1833 in: Foraminifères, edited by: Levrault, S., Bertrand,
Paris, 1–86, 1839b.
Defrance, J. L. M.: Dictionnaire des Sciences Naturelles, F. G. Levrault,
Strasbourg, 1824.DeLaca, T. E.: The morphology and ecology of Astrammina rara, J.
Foramin. Res., 16, 216–233, 1986.
Earland, A.: Foraminifera, Part II, South Georgia, London, 27–138, 1933.
Earland, A.: Foraminifera. Part III. The Falklands sector of the Antarctic
(excluding South Georgia), London, 1–208, 1934.
Echols, R. J.: Distribution of foraminifera in sediments of the Scotia Sea
area, Antarctic waters, in: Antarctic Oceanography I, edited by: Reid, J. L.,
American Geophysical Union, Washington, D.C., 93–168, 1971.
Ehrenberg, C. G.: Uber die Bildung der Kreidefelsen und des Kreidemergels
durch unsichtbare Organismen, Physikalische Abhandlungen der Koniglichen
Akademie der Wissenschaften zu Berlin, 1838, 59–147, 1838.
Ehrenberg, C. G.: Elemente des tiefen Meeresgrundes im Mexikanishen
Golfstrome bei Florida: uber die Tiefgrund-Verhaltnisse des Oceans am
eingange der Davisstrasse und bei Island, Monatsbericht der Koniglichen
Preussichen Akademie der Wissenschaften zu Berlin, 1861, 275–315, 1861.Elderfield, H., Yu, J., Anand, P., Kiefer, T., and Nyland, B.: Calibrations
for benthic foraminiferal Mg / Ca paleothermometry and the carbonate ion
hypothesis, Earth Planet. Sc. Lett., 250, 633–649,
10.1016/j.epsl.2006.07.041, 2006.Elderfield, H., Greaves, M., Barker, S., Hall, I. R., Tripati, A., Ferretti,
P., Crowhurst, S., Booth, L., and Daunt, C.: A record of bottom water
temperature and seawater δ18O for the Southern Ocean over the past
440 kyr based on Mg / Ca of benthic foraminiferal Uvigerina spp,
Quaternary Sci. Rev., 29, 160–169, 10.1016/j.quascirev.2009.07.013,
2010.Elderfield, H., Ferretti, P., Greaves, M., Crowhurst, S., McCave, I. N.,
Hodell, D., and Piotrowski, A. M.: Evolution of Ocean Temperature and Ice
Volume Through the Mid-Pleistocene Climate Transition, Science, 337,
704–709, 10.1126/science.1221294, 2012.Enge, A. J., Nomaki, H., Ogawa, N. O., Witte, U., Moeseneder, M. M., Lavik,
G., Ohkouchi, N., Kitazato, H., Kucera, M., and Heinz, P.: Response of the
benthic foraminiferal community to a simulated short-term phytodetritus pulse
in the abyssal North Pacific, Mar. Ecol.-Prog. Ser., 438, 129–142,
10.3354/meps09298, 2011.Fontanier, C., Jorissen, F. J., Chaillou, G., David, C., Anschutz, P., and
Lafon, V.: Seasonal and interannual variability of benthic foraminiferal
faunas at 550 m depth in the Bay of Biscay, Deep-Sea Res. Pt. I, 50,
457–494, 10.1016/s0967-0637(02)00167-x, 2003.Fontanier, C., Jorissen, F., Anschutz, P., and Chaillou, G.: Seasonal
variability of benthic foraminiferal faunas at 1000 m depth in the Bay of
Biscay, J. Foramin. Res., 36, 61–76, 10.2113/36.1.61, 2006a.Fontanier, C., Mackensen, A., Jorissen, F. J., Anschutz, P., Licari, L., and
Griveaud, C.: Stable oxygen and carbon isotopes of live benthic foraminifera
from the Bay of Biscay: Microhabitat impact and seasonal variability, Mar.
Micropaleontol., 58, 159–183, 10.1016/j.marmicro.2005.09.004, 2006b.Fontanier, C., Jorissen, F. J., Michel, E., Cortijo, E., Vidal, L., and
Anschutz, P.: Stable oxygen and carbon isotopes of live (stained) benthic
foraminifera from Cap-Ferret Canyon (Bay of Biscay), J. Foramin. Res., 38,
39–51, 10.2113/gsjfr.38.1.39, 2008.Fontanier, C., Garnier, E., Brandily, C., Dennielou, B., Bichon, S., Gayet,
N., Eugene, T., Rovere, M., Grémare, A., and Deflandre, B.: Living
(stained) benthic foraminifera from the Mozambique Channel (eastern Africa):
Exploring ecology of deep-sea unicellular meiofauna, Deep-Sea Res. Pt. I,
115, 159–174, 10.1016/j.dsr.2016.06.007, 2016.
Frerichs, W. E.: Recent arenaceous foraminifers from Gulf of Mexico,
Palaeontological Contributions, University of Kansas, 46, 1–2, 1969.
Galloway, J. J.: A manual of Foraminifera, Principia Press, Bloomington,
1933.
Gaździcki, A. and Majewski, W.: Recent foraminifera from Goulden Cove of
King George Island, Antarctica, Pol. Polar Res., 24, 3–12, 2003.Gehlen, M., Mucci, A., and Boudreau, B.: Modelling the distribution of stable
carbon isotopes in porewaters of deep-sea sediments, Geochim. Cosmochim. Ac.,
63, 2763–2773, 10.1016/s0016-7037(99)00214-8, 1999.Geslin, E., Heinz, P., Jorissen, F., and Hemleben, C.: Migratory responses of
deep-sea benthic foraminifera to variable oxygen conditions: laboratory
investigations, Mar. Micropaleontol., 53, 227–243,
10.1016/j.marmicro.2004.05.010, 2004.
Glaessner, M. F.: Die Entfaltung der Foraminiferenfamilie Buliminidae,
Problemy Palaeontologii, Palaeontologicheskaya Laboratoriya Moskovskogo
Gosudarstvennogo Universiteta, 2–3, 411–422, 1937.
Goes, A.: A synopsis of the Arctic and Scandinavian Recent marine
Foraminifera hitherto discovered, Kongl. svenka vetenskaps-Akademiens
handlinger 25, 1–127, 1894.Gooday, A. J.: A response by benthic foraminifera to the deposition of
phytodetritus in the deep-sea, Nature, 332, 70–73, 10.1038/332070a0,
1988.Gooday, A. J. and Hughes, J. A.: Foraminifera associated with phytodetritus
deposits at a bathyal site in the northern Rockall Trough (NE Atlantic):
seasonal contrasts and a comparison of stained and dead assemblages, Mar.
Micropaleontol., 46, 83–110, 10.1016/s0377-8398(02)00050-6, 2002.
Gooday, A. J., da Silva, A. A., Koho, K. A., Lecroq, B., and Pearce, R. B.:
The “mica sandwich”; a remarkable new genus of Foraminifera (Protista,
Rhizaria) from the Nazare Canyon (Portuguese margin, NE Atlantic),
Micropaleontology, 56, 345–357, 2010.
Griffith, J. W. and Henfrey, A.: The Micrographic Dictionary, van Voorst,
London, 1875.Gross, O.: Influence of temperature, oxygen and food availability on the
migrational activity of bathyal benthic foraminifera: evidence by microcosm
experiments, Hydrobiologia, 426, 123–137, 10.1023/a:1003930831220, 2000.
Haeckel, E.: Systematische Phylogenie. Entwurf eines Naturlichen Systems der
Organismen auf Grund ihrer Stammesgeschichte. Theil 1, Systematische
Phylogenie der Protisten und Pflanzen, Georg Reimer, Berlin, 1894.
Hayward, B. W., Kawagata, S., Sabaa, A. T., Grenfell, H. R., van Kerckhoven,
L., Johnson, K., and Thomas, E.: The last global extinction (Mid-Pleistocene)
of deep-sea benthic foraminifera (Chrysalogoniidae, Ellipsoidinidae,
Glandulonodosariidae, Plectofrondiculariidae, Pleursostomellidae,
Stilostomellidae), their Late Cretaceous-Cenozoic history and taxonomy,
Cushman Foundation for Foraminiferal Research Special Publication, 43,
1–410,
2012.
Heron-Allen, E. and Earland, A.: On some Foraminifera from the North Sea,
etc., dredged by the Fisheries Cruiser “Goldseeker” (International North
Sea Investigations – Scotland). I. On some new Astrorhizidae and their
shell-structure, Journal of the Royal Microscopical Society, 1912, 382–389,
1912.
Heron-Allen, E. and Earland, A.: Les Foraminifères des “Sables Rouges”
du golfe de Ajaccio (Côte Nord), Bulletin de la Société des
sciences historiques et naturelles de la Corse, 42, 109–149, 1922.Heron-Allen, E. and Earland, A.: XXVII. – Some new Foraminifera from the
South Atlantic, Journal of the Royal Microscopical Society, 49, 324–334,
10.1111/j.1365-2818.1929.tb00787.x, 1929.
Heron-Allen, E. and Earland, A.: Some new foraminifera from the South
Atlantic, Part 3., Journal of the Royal Microscopical Society, 50, 38—45,
1930.
Heron-Allen, E. and Earland, A.: Foraminifera. Part 1. The ice-free area of
the Falkland Islands and adjacent seas, Cambridge University Press, London, 291–460,
1932a.
Heron-Allen, E. and Earland, A.: Some new foraminifera from the South
Atlantic, IV, Journal of the Royal Microscopical Society, 52, 253–261,
1932b.
Herb, R.: Distribution of Recent Benthonic Foraminifera in the Drake Passage,
in: Biology of the Antarctic Seas IV, American Geophysical Union, 251–300,
1971.
Hermelin, J. O. R.: Pliocene benthic foraminifera from the Ontong-Java
plateau (western equatorial Pacific Ocean): faunal response to changing
paleoenvironments, Cushman Foundation for Foraminiferal Research Special
Publication, 26, 1–143, 1989.Hogg, O. T., Barnes, D. K. A., and Griffiths, H. J.: Highly Diverse, Poorly
Studied and Uniquely Threatened by Climate Change: An Assessment of Marine
Biodiversity on South Georgia's Continental Shelf, Plos One, 6, e19795,
10.1371/journal.pone.0019795, 2011.
Höglund, H.: Foraminifera in the Gullmar Fjord and the Skagerak,
Zoologiska Bidrag fran Uuppsala, 26, 1–328, 1947.Holsten, J., Stott, L., and Berelson, W.: Reconstructing benthic carbon
oxidation rates using δ13C of benthic foraminifers, Mar.
Micropaleontol., 53, 117–132, 10.1016/j.marmicro.2004.05.006, 2004.Hoogakker, B. A. A., Elderfield, H., Schmiedl, G., McCave, I. N., and
Rickaby, R. E. M.: Glacial-interglacial changes in bottom-water oxygen
content on the Portuguese margin, Nat. Geosci., 8, 40–43,
10.1038/ngeo2317, 2015.Hromic, T., Ishman, S., and Silva, N.: Benthic foraminiferal distributions in
Chilean fjords: 47∘ S to 54∘ S, Mar. Micropaleontol., 59,
115–134, 10.1016/j.marmicro.2006.02.001, 2006.Igarashi, A., Numanami, H., Tsuchiya, Y., and Fukucki, M.: Bathymetric
distribution of fossil foraminifera within marine sediment cores from the
eastern part of Lutzow-Holm Bay, East Antarctica, and its paleoceanographic
implications, Mar. Micropaleontol., 42, 125–162,
10.1016/s0377-8398(01)00004-4, 2001.Ishman, S. E. and Domack, E. W.: Oceanographic controls on benthic
foraminifers from the Bellingshausen margin of the Antarctic Peninsula, Mar.
Micropaleontol., 24, 119–155, 10.1016/0377-8398(94)90019-1, 1994.
Jones, A. F.: The Micrographic Dictionary, edited by: Griffith, J. W. and
Henfrey, A., van Voorst, London, 1875.
Jones, R. W.: A revised classification of the unilocular Nodosariida and
Buliminida (Foraminifera), Revista Espanola de Micropaleontologia, 16,
91–160, 1984.
Jones, R. W.: The Challenger Foraminifera, Oxford University Press, Oxford,
1994.Jorissen, F. J., deStigter, H. C., and Widmark, J. G. V.: A conceptual model
explaining benthic foraminiferal microhabitats, Mar. Micropaleontol., 26,
3–15, 10.1016/0377-8398(95)00047-x, 1995.
Jorissen, F. J., Fontanier, C., and Thomas, E.: Paleoceanographical Proxies
Based on Deep-Sea Benthic Foraminiferal Assemblage Characteristics, in:
Proxies in Late Cenozoic Paleoceanography, edited by: Hillaire-Marcel, C.,
and de Vernal, A., Developments in Marine Geology, Elsevier, Amsterdam,
263–325, 2007.
Kaminski, M. A.: The Year 2000 Classification of the Agglutinated
Foraminifera, in: Proceedings of the Sixth International Workshop on
Agglutinated Foraminifera (Prague, Czech Republic, 1–7 September 2001),
edited by: Kaminski, M. A. and Bubik, M., Grzybowski Foundation Special
Publication, London, UK, 237–255, 2004.
Kaminski, M. A. and Gradstein, F. M.: Atlas of Paleogene Cosmopolitan
deep-water agglutinated foraminifera, Grzybowski Foundation Special
Publication, Krakow, 2005.
Kanmacher, F.: Adam's Essays on the Microscope: the Second Edition, with
Considerable Additions and Improvements, Dillon & Keating, London, 1798.Kender, S. and Kaminski, M. A.: Modern deep-water agglutinated foraminifera from IODP Expedition 323,
Bering Sea: ecological and taxonomic implications, J. Micropalaeontology, 10.1144/jmpaleo2016-026, 2017.
Kennett, J. P.: New Foraminifera from the Ross Sea, Antarctica, Contributions
from the Cushman Foundation for Foraminiferal Research, 18, 133–135, 1967.Kitazato, H., Shirayama, Y., Nakatsuka, T., Fujiwara, S., Shimanaga, M.,
Kato, Y., Okada, Y., Kanda, J., Yamaoka, A., Masuzawa, T., and Suzuki, K.:
Seasonal phytodetritus deposition and responses of bathyal benthic
foraminiferal populations in Sagami Bay, Japan: preliminary results from
“Project Sagami 1996–1999”, Mar. Micropaleontol., 40, 135–149,
10.1016/s0377-8398(00)00036-0, 2000.
Knight, R.: Apertural characteristics of certain unilocular foraminifera:
methods of study, nomenclature and taxonomic significance, J.
Micropaleontol., 5, 37–47, 1986.
Lamarck, J. B.: Suite des mémoires sur les fossiles des environs de
Paris, Annales du Muséum d'Histoire Naturelle, 5, 105–115, 179-188,
1804.
Lamarck, J. B.: Extrait du course de zoologie du Museum d'Histoire Naturelle,
sur les animaux sans vertebres, d'Hautel, Paris, 1812.Le Coze, F. and Hayward, B.: Lagena striatopunctata Parker & Jones, 1865,
in: World Foraminifera Database, edited by: Hayward, B. W., Cedhagen, T.,
Kaminski, M., and Gross, O., avaiable at:
http://www.marinespecies.org/foraminifera./aphia.php?p=taxdetails&id=525593
(last access: 12 July 2017), 2017.Linke, P. and Lutze, G. F.: Microhabitat preferences of benthic foraminifera
– a static concept or a dynamic adaptation to optimize food acquisition,
Mar. Micropaleontol., 20, 215–234, 10.1016/0377-8398(93)90034-u, 1993.
Linnaeus, C.: Systema Naturae per Regnia tria Naturae, secundum classes,
ordines, genera, species, cum characteribus, differentiis, synonymis, locis,
Stockholm, 823 pp., 1758.
Lipps, J. H., DeLaca, T. E., Krebs, W. N., and Stockton, W.: Shallow-water
foraminifera studies, Antarctic Peninsula, 1971–1972, Foraminiferal ecology,
Antarctic Peninsula, Antarctic, Journal of the United States, 7, 82–83,
1972.
Loeblich, A. R. and Tappan, H.: Revision of some recent foraminiferal genera,
Smithsonian Miscellaneous Collections, 128, 1–37, 1955.
Loeblich, A. R. and Tappan, H.: Suprageneric classification of the
Rhizopodea, J. Paleontol., 35, 245–330, 1961.
Loeblich, A. R. and Tappan, H.: Some new proteinaceous and agglutinated
genera of Foraminiferida, J. Paleontol., 58, 1158–1163, 1984.
Loeblich, A. R. and Tappan, H.: Some new and redefined genera and families of
Textulariina, Fusulinina, Involutinina and Miliolina (Foraminiferida), J.
Foramin. Res., 16, 334–346, 1986.
Loeblich, A. R. and Tappan, H.: Foraminiferal genera and their
classification, Van Nostrand Reinhold, New York, 1988.
Loeblich, A. R. and Tappan, H.: Foraminifera of the Sahul Shelf and Timor
Sea, Cushman Foundation for Foraminiferal Research Special Publication, 31,
1–661,
1994.Loubere, P., Meyers, P., and Gary, A.: Benthic foraminiferal microhabitat
selection, carbon isotope values, and association with larger animals – a
test with Uvigerina peregrina, J. Foramin. Res., 25, 83–95, 1995.Loubere, P., Jacobsen, B., Klitgaard Kristensen, D., Husum, K., Jernas, P.,
and Richaud, M.: The structure of benthic environments and the paleochemical
record of foraminifera, Deep-Sea Res. Pt. I, 58, 535–545,
10.1016/j.dsr.2011.02.011, 2011.Luo, M., Chen, L. Y., Wang, S. H., Yan, W., Wang, H. B., and Chen, D. F.:
Pockmark activity inferred from pore water geochemistry in shallow sediments
of the pockmark field in southwestern Xisha Uplift, northwestern South China
Sea, Mar. Petrol. Geol., 48, 247–259, 10.1016/j.marpetgeo.2013.08.018,
2013.Mackensen, A., Grobe, H., Kuhn, G., and Futterer, D. K.: Benthic
Foraminiferal Assemblages from the Eastern Weddell Sea Between 68 and
73∘ S – Distribution, Ecology and Fossilisation Potential, Mar.
Micropaleontol., 16, 241–283, 10.1016/0377-8398(90)90006-8, 1990.Mackensen, A., Futterer, D. K., Grobe, H., and Schmiedl, G.: Benthic
Foraminiferal Assemblages from the Eastern South-Atlantic Polar Front Region
Between 35 and 57∘ S – Distribution, Ecology and Fossilization
Potential, Mar. Micropaleontol., 22, 33–69,
10.1016/0377-8398(93)90003-g, 1993.
Majewski, W.: Benthic foraminiferal communities: distribution and ecology in
Admiralty Bay, King George Island, West Antarctica, Pol. Polar Res., 26,
159–214, 2005.Majewski, W.: Benthic foraminifera from West Antarctic fiord environments: An
overview, Pol. Polar Res., 31, 61–82, 10.4202/ppres.2010.05, 2010.Majewski, W.: Benthic foraminifera from Pine Island and Ferrero bays,
Amundsen Sea, Pol. Polar Res., 34, 169–200, 10.2478/popore-2013-0012,
2013.Majewski, W. and Anderson, J. B.: Holocene foraminiferal assemblages from
Firth of Tay, Antarctic Peninsula: Paleoclimate implications, Mar.
Micropaleontol., 73, 135–147, 10.1016/j.marmicro.2009.08.003, 2009.
Marie, P.: La foraminifères de la Craie à Belemnitella mucronata du
Bassin de Paris, Mémoires de Museum Nationale d'Histoire Naturelle, 12,
1–296, 1941.Maync, W.: Critical taxonomic study and nomenclatural revision of the
Lituolidae based upon the prototype of the family, Lituola nautiloidea Lamarck 1804, Contributions from the Cushman Foundation for
Foraminiferal Research, 3, 35–56, 1952.McCorkle, D. C. and Emerson, S. R.: The relationship between pore water
carbon isotopic composition and bottom water oxygen concentration, Geochim.
Cosmochim. Ac., 52, 1169–1178, 10.1016/0016-7037(88)90270-0, 1988.McCorkle, D. C., Emerson, S. R., and Quay, P. D.: Stable carbon isotopes in
marine porewaters, Earth Planet. Sc. Lett., 74, 13–26,
10.1016/0012-821x(85)90162-1, 1985.McCorkle, D. C., Keigwin, L. D., Corliss, B. H., and Emerson, S. R.: The
influence of microhabitats on the carbon isotopic composition of deep-sea
benthic foraminifera, Paleoceanography, 5, 161–185,
10.1029/PA005i002p00161, 1990.McCorkle, D. C., Corliss, B. H., and Farnham, C. A.: Vertical distributions
and stable isotopic compositions of live (stained) benthic foraminifera from
the North Carolina and California continental margins, Deep-Sea Res. Pt. I,
44, 983–1024, 10.1016/S0967-0637(97)00004-6, 1997.
McCulloch, I.: Qualitative observations on Recent foraminiferal tests with
emphasis on the eastern Pacific, University of Southern California, Los
Angeles, 1977.Meredith, M. P., Brandon, M. A., Murphy, E. J., Trathan, P. N., Thorpe, S.
E., Bone, D. G., Chernyshkov, P. P., and Sushin, V. A.: Variability in
hydrographic conditions to the east and northwest of South Georgia,
1996–2001, J. Marine Syst., 53, 143–167, 10.1016/j.jmarsys.2004.05.005,
2005.
Mikhalevich, V. I.: Systematics and evolution of forminifera in the light of
new data on their cytology and ultrastructure, Trudy Zoologicheskogo
Instituta Akademiya Nauk SSSR, 94, 42–61, 1980.
Mikhalevich, V. I.: A new classification of the class Astrorhizata,
Zoosystematica Rossica, 3, 161–174, 1995.Mikhalevich, V. I.: The general aspects of the distribution of Antarctic
foraminifera, Micropaleontology, 50, 179–194,
10.1661/0026-2803(2004)050[0179:moeoti]2.0.co;2, 2004.
Miklukho-Maklay, A. D.: Upper Palaeozoic of Central Asia, Leningradskiy
Universitet, Lenigrad, 1963.
Milam, R. W. and Anderson, J. B.: Distribution and Ecology of Recent
Benthonic Foraminifera of the Adelie-George-V Continental-Shelf and Slope,
Antarctica, Mar. Micropaleontol., 6, 297–325, 1981.
Montagu, G.: Testacea Britannica or Natural History of British Shells,
Marine, Land, and Fresh-Water, Including the Most Minute, J. S. Hollis,
Romsey, England, 1803.
Montfort, P. D. d.: Conchyliologie systématique et classification
méthodique des coquilles, F. Schoell, Paris, 676 pp., 1808.
Moreman, W. L.: Arenaceous foraminifera from Ordovician and Silurian
limestones of Oklahoma, J. Paleontol., 4, 42–59, 1930.Murphy, E. J., Hofmann, E. E., Watkins, J. L., Johnston, N. M., Pinones, A.,
Ballerini, T., Hill, S. L., Trathan, P. N., Tarling, G. A., Cavanagh, R. A.,
Young, E. F., Thorpe, S. E., and Fretwell, P.: Comparison of the structure
and function of Southern Ocean regional ecosystems: The Antarctic Peninsula
and South Georgia, J. Marine Syst., 109, 22–42,
10.1016/j.jmarsys.2012.03.011, 2013.Murray, J. W. and Bowser, S. S.: Mortality, protoplasm decay rate, and
reliability of staining techniques to recognize “living” foraminifera: A
review, J. Foramin. Res., 30, 66–70, 10.2113/0300066, 2000.Murray, J. W. and Pudsey, C. J.: Living (stained) and dead foraminifera from
the newly ice-free Larsen Ice Shelf, Weddell Sea, Antarctica: ecology and
taphonomy, Mar. Micropaleontol., 53, 67–81,
10.1016/j.marmicro.2004.04.001, 2004.Nomaki, H., Heinz, P., Hemleben, C., and Kitazato, H.: Behavior and response
of deep-sea benthic foraminifera to freshly supplied organic matter: A
laboratory feeding experiment in microcosm environments, J. Foramin. Res.,
35, 103–113, 10.2113/35.2.103, 2005.Ohga, T. and Kitazato, H.: Seasonal changes in bathyal foraminiferal
populations in response to the flux of organic matter (Sagami Bay, Japan),
Terra Nova, 9, 33–37, 10.1046/j.1365-3121.1997.d01-6.x, 1997.
Parker, F. L.: Foraminiferal distribution in the Long Island Sound –
Buzzards Bay area, Bulletin of the Museum of Comparative Zoology, Harvard,
106, 428–473, 1952.
Parker, W. K. and Jones, T. R.: On some foraminifera from the North Atlantic
and Arctic Oceans, including Davis Straits and Baffin's Bay, Philos. T. R.
Soc., 155, 325–441, 1865.Parr, W. J.: On Torrresina, a new genus of the foraminifera from
eastern Australia, Journal of the Royal Microscopical Society, 64, 129–135,
1947.
Parr, W. J.: Foraminifera, in: Reports B.A.N.Z. Antarctic Research Expedition 1929–1931, 232–392, 1950.
Patterson, R. T. and Richardson, R. H.: A taxonomic revision of the
unilocular Foraminifera, J. Foramin. Res., 17, 212–226, 1987.
Pearcey, F. G.: Foraminifera of the Scottish National Antarctic Expedition,
T. Roy. Soc. Edin., 49, 991–1044, 1914.Peck, V. L., Allen, C. S., Kender, S., McClymont, E. L., and Hodgson, D.:
Oceanographic variability on the West Antarctic Peninsula during the Holocene
and the influence of upper circumpolar deep water, Quaternary Sci. Rev., 119,
54–65, 10.1016/j.quascirev.2015.04.002, 2015.Piña-Ochoa, E., Høgslund, S., Geslin, E., Cedhagen, T., Revsbech, N.
P., Nielsen, L. P., Schweizer, M., Jorissen, F., Rysgaard, S., and
Risgaard-Petersen, N.: Widespread occurrence of nitrate storage and
denitrification among Foraminifera and Gromiida, P. Natl. Acad. Sci. USA,
107, 1148–1153, 10.1073/pnas.0908440107, 2010.
Puri, H. S.: Contribution to the study of the Miocene of teh Florida
panhandle, Bulletin Florida State Geological Survey, 36, 1–345, 1954.
Rauzer-Chernousova, D. M. and Reytlinger, E. A.: On the suprageneric
systematics of the Order Hormosinida (Foraminifera), Paleontologicheskiy
Zhurnal, 1986, 15–20, 1986.
Ravelo, A. C. and Hillaire-Marcel, C.: The use of oxygen and carbon isotopes
of foraminifera in paleoceanography, in: Proxies in Late Cenozoic
Paleoceanography, edited by: Hillaire-Marcel, C. and De Vernal, A.,
Developments in Marine Geology, Elsevier, Amsterdam, 735–764, 2007.
Reuss, A. E.: Uber zwei neue Arten von Foraminiferen aus dem Tegel van Baden
und Mollersdorf, in: Bericht uder die Mittheilungen Freunden der
Naturwissenschaften in Wien, edited by: Czjzek, J., 50–56, 1849.
Reuss, A. E.: Neues Foraminiferen aus den Schichten des Osterreichischen
Tertiarbeckens, Denkschriften der Akademie des Wissenschaften, 1, 365–390,
1850.
Reuss, A. E.: Die Foraminiferen der westphälischen Kreideformation,
Sitzungsberichte der mathematisch-naturwissenschaflichen Classe der
kaiserlichen Akademie der Wissenschaften, 40, 147–238, 1860.
Reuss, A. E.: Entwurf einer systematischen Zusammenstellung der
Foraminiferen, Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften
in Wien. Mathematicsch-Naturwissenschaftliche Classe, 1861, 355–396, 1862.
Reuss, A. E.: Die Foraminiferen, Anthozoen und Bryozoen des deutschen
Septarienthones, Denkschriften der Akademie des Wissenschaften, Wien, 25,
117–214, 1866.
Rhumbler, L.: Entwurf eines naturlichen systems der Thalamophoren,
Nachrichten von der Gesellschaft der Wissenschaften zu Gottingen, Math-Physik
Klasse, 1895, 51–98, 1895.
Rhumbler, L.: Die Foraminiferen (Thalamophoren) der Plankton-Expedition,
Erster Teil, Die allgemein Organizationsverhaltnisse der Foraminiferen,
Ergebnisse der Plankton-Expedition der Humboldt-Stiftung Kiel u Liepzig, 3,
1–331, 1911.
Rodrigues, A. R., Eichler, P. P. B., and Eichler, B. B.: Foraminifera in two
inlets fed by a tidewater glacier, King George Island, Antarctic Peninsula,
J. Foramin. Res., 43, 209–220, 2013.
Saidova, K. M.: Bentosnye Foraminifery rayona kurilo-Kamchatskogo zheloba (po
materianlam 39-go reysa e/s “Vityaz”) [Benthic foraminifera in the
Kurile-Kamchatka region based on the data of the 39th cruise of the R/V
“Vityaz.”], Trudy Instituta Okeanologii, 86, 134–161, 1970.
Saidova, K. M.: Bentosniye foraminifery Tikhogo Okeana, P.P. Shirshov
Institute of Oceanology, Academy of Sciences of the USSR, Moscow, 1975.
Saidova, K. M.: O sovremennom sostoyanii sistemy nadvidovykh taksonov
Kaynozoyskikh bentosnykh foraminifer [On an up-to-date system of
supraspecific taxonomy of Cenozoic benthonic foraminifera], Akademiya Nauk
SSSR, Moscow, 1981.
Sars, M.: Fortsatte bemerkinger over det dyriske livs udbredning i havets
dybder, Förhandlinger i Videnskabsselskabet i Kristiania, 1869, 246–275,
1869.Schilman, B., Almogi-Labin, A., Bar-Matthews, M., and Luz, B.: Late Holocene
productivity and hydrographic variability in the eastern Mediterranean
inferred from benthic foraminiferal stable isotopes, Paleoceanography, 18,
10.1029/2002pa000813, 2003.Schmiedl, G., Mitschele, A., Beck, S., Emeis, K. C., Hemleben, C., Schulz,
H., Sperling, M., and Weldeab, S.: Benthic foraminiferal record of ecosystem
variability in the eastern Mediterranean Sea during times of sapropel S-5 and
S-6 deposition, Palaeogeogr. Palaeocl., 190, 139–164,
10.1016/s0031-0182(02)00603-x, 2003.Schmiedl, G., Pfeilsticker, M., Hemleben, C., and Mackensen, A.:
Environmental and biological effects on the stable isotope composition of
recent deep-sea benthic foraminifera, from the western Mediterranean Sea,
Mar. Micropaleontol., 51, 129–152, 10.1016/j.marmicro.2003.10.001, 2004.
Schultze, M. J. S.: Über den Organismus der Polythalamien
(Foraminiferen), nebst Bemerkungen über die Rhizopoden im allgemeinen,
Leipzig, Ingelmann, 1854.
Schulze, F. E.: Rhizopodenstudien. III., Archiv für Mikroskopische
Anatomie 11, 94–139, 1875.
Schwager, C.: Saggio di una classificazione dei foraminiferi avuto riguardo
alle loro famiglie naturali, Bolletino R. Comitato Geologico d'Italia, 7,
475–485, 1876.
Schwager, C.: Quadro del proposto sistema di classificazione dei foraminiferi
con guscio, Bolletino R. Comitato Geologico d'Italia, 8, 18–27, 1877.
Seguenza, G.: Dei terreni Terziarii del distretto di Messina: Parte II –
Descrizione dei foraminiferi monotalamici delle marne Mioceniche del
distretto di Messina, T. Capra, Messina, 1862.
Setoyama, E. and Kaminski, M. A.: Neogene benthic foraminifera from the
southern Bering Sea (IODP Expedition 323), Palaeontologia Electronica,
18.2.38A, 1–30, 2015.
Shchedrina, Z. G.: O nekotorykh izmeneniyakh v sisteme semeytv Astrorhizidae
i Reophacidae (Foraminifera) [On some changes in the systematics of the
families Astrorhizidae and Reopacidae (Foraminifera)], Voprosy
Mikropaleontologii, 11, 157–170, 1969.
Silvestri, A.: Lo stipite della Elissoforme e le sue affinita, Memorie della
Pontificia Accademia della Scienze, Nouvi Lincei, 6, 231–270, 1923.
Silvestri, O.: Saggio di studi sulla fauna microscopia fossile appartenente
al terreno subappenino italiano. Mem. I – monografia delle Nodosarie,
Academia Gioenia Scienze Naturali Catania 7, 108, 1872.Stefanoudis, P. V., Bett, B. J., and Gooday, A. J.: Relationship between
“live” and dead benthic foraminiferal assemblages in the abyssal NE
Atlantic, Deep-Sea Res. Pt. I, 121, 190–201, 10.1016/j.dsr.2017.01.014,
2017.Sweetman, A. K., Sommer, S., Pfannkuche, O., and Witte, U.: Retarded response
by macrofauna-size foraminifera to phytodetritus in a deep Norwegian fjord,
J. Foramin. Res., 39, 15–22, 10.2113/gsjfr.39.1.15, 2009.Tachikawa, K. and Elderfield, H.: Microhabitat effects on Cd / Ca and
δ13C of benthic foraminifera, Earth Planet. Sc. Lett., 202,
607–624, 10.1016/S0012-821X(02)00796-3, 2002.Thalmann, H. E.: Nomenclator (Um- und Neubennungen) zu den Tafeln 1 bis 115
in H. B. Brady's Werk uber die Foraminferen der
Challenger-Expedition, London 1884, Eclogae Geologicae Helvetiae,
25, 293–312, 1932.Theodor, M., Schmiedl, G., and Mackensen, A.: Stable isotope composition of
deep-sea benthic foraminifera under contrasting trophic conditions in the
western Mediterranean Sea, Mar. Micropaleontol., 124, 16–28,
10.1016/j.marmicro.2016.02.001, 2016.Thorpe, S. E., Heywood, K. J., Brandon, M. A., and Stevens, D. P.:
Variability of the southern Antarctic Circumpolar Current front north of
South Georgia, J. Marine Syst., 37, 87–105,
10.1016/s0924-7963(02)00197-5, 2002.
Uchio, T.: Ecology of living benthonic foraminifera from the San Diego,
California area, Special Publications Cushman Laboratory for Foraminiferal
Research, 5, 1–72, 1960.
Uchio, T.: Influence of the River Shinano on foraminifera and sediment grain
size distribution, Publications of the Seto Marine Biological Laboratory, 10,
363–392, 1962.
Van Marle, L.: Eastern Indonesia Late Cenozoic smaller benthic foraminifera,
Verhandeling Koninklijke Nederlandse Akademie van Wetenschappen, Afdeling
Antuurkunde, Eerste Reeks, 34, 1–328, 1991.
Violanti, D.: Taxonomy and distribution of recent benthic foraminifers from
Terra Nova Bay (Ross Sea, Antarctica), oceanographic campaign 1987/1988,
Palaeontographica Italica, 83, 25–71, 1996.Voloshinova, N. A.: Progress in micropalaeontology in the work of studying
the inner structure of Foraminifera, Trudy Pervogo Seminara po Mikrofaune,
48–87, 1960.
Wang, F., Gao, M., Liu, J., Pei, S., Li, C., Mei, X., and Yang, S.:
Distribution and environmental significance of live and dead benthic
foraminiferal assemblages in surface sediments of Laizhou Bay, Bohai Sea,
Mar. Micropaleontol., 123, 1–14, 10.1016/j.marmicro.2015.12.006, 2016.
Wedekind, P. R.: Einfuhrung in die Grundlagen der historischen Geologie. Band
II. Mikrobiostratigraphie der Korallen- und Foraminiferenzeit, Ferdinand
Enke, Stuttgart, 1937.Weldeab, S., Arce, A., and Kasten, S.: Mg / Ca-Delta
CO3pore2-(water)-temperature calibration for Globobulimina spp.: A sensitive
paleothermometer for deep-sea temperature reconstruction, Earth Planet. Sc.
Lett., 438, 95–102, 10.1016/j.epsl.2016.01.009, 2016.
Wiesner, H.: Die Foraminiferen der Deutschen Südpolar Expedition
1901–1903. Deutschen Südpolar Expedition, Berlin (Zoology), 20, 53–165,
1931.Williamson, W. C.: On the Recent British species of the genus
Lagena, Annals and Magazine of Natural History, 2, 1–20, 1848.
Witte, U., Wenzhofer, F., Sommer, S., Boetius, A., Heinz, P., Aberle, N.,
Sand, M., Cremer, A., Abraham, W. R., Jorgensen, B. B., and Pfannkuche, O.:
In situ experimental evidence of the fate of a phytodetritus pulse at the
abyssal sea floor, Nature, 424, 763–766, 2003.Zahn, R., Winn, K., and Sarnthein, M.: Benthic foraminiferal δ13C
and accumulation rates of organic carbon: Uvigerina peregrina group
and Cibicidoides wuellerstorfi, Paleoceanography, 1, 27–42, 1986.
Zheng, S. and Fu, Z.: Fauna Sinica, Phylum Granuloreticulosa, Class
Foraminiferea, Agglutinated Foraminifera, Science Press, Beijing, 2001.