Ammopemphix hemisphaericus sp. nov., a new attached agglutinated foraminifer from the Pleistocene of the Arctic Ocean, and the taxonomic status of the genus Ammopemphix Loeblich, 1952

Ammopemphix hemisphaericus sp. nov., a new attached agglutinated foraminifer from the Pleistocene... The new species Ammopemphix hemisphaericus sp. nov. is described from Pleistocene sediments of the Central Arctic Ocean. The species is a finely agglutinated monothalamous form with hemispherical shape and a terminal aperture located at the top of the test, often occurring in pseudocolonies. A. hemisphaericus sp. nov. attaches itself to inorganic or organic substrates, often growing attached to the inside surfaces of calcareous or agglutinated foraminiferal tests. Pseudocolonies of A. hemisphaericus sp. nov. may also attach themselves to previous generations of Ammopemphix individuals. Based on our study of numerous specimens of Ammopemphix from the Central Arctic Ocean, we propose modifications to the description of the genus. The genus may be pseudocolonial or consist of isolated individuals, and in the case of pseudocolonial forms, individual specimens may be arranged randomly or symmetrically. The genus Ammopemphix is found mostly in polar regions, and is here reported for the first time from the fossil record. Keywords Agglutinated foraminifera · Arctic Ocean · Pleistocene Introduction approximately 20 species of agglutinated foraminifera [13]. The taxonomy and biostratigraphy of the benthic foraminiferal The glacial environment of the deep Arctic Ocean yields spe- assemblages from the PS87 cores will be described separately. cies that are predisposed to life in cold and oligotrophic condi- The purpose of this paper is to describe an enigmatic new spe- tions. Because of profound changes in sea-ice cover, surface cies that lives attached to sediment particles, or most often to productivity, and sedimentation linked to glacial/interglacial fragments of other specimens of agglutinated foraminifera. cycles, the benthic foraminiferal and ostracode assemblages The genus Ammopemphix has been previously reported found in the glacial Arctic Ocean are much less abundant and from the Arctic and Antarctic seas [e.g., 3, 14, 15, 17, 24], diverse than in the interglacial faunas [2]. Glacial assemblages but specimens from the Lomonosov Ridge display morpho- reported from sediment cores are sparse and typically only logical characteristics that are not known among the previ- consist of a few species of agglutinated forms [6, 7, 13]. ously described species. In this study we describe a new During the summer of 2014, four long kastenlot cores and species of Ammopemphix and modify the current definition 36 gravity cores were collected on the Lomonosov Ridge of the genus. during Expedition 87 of the R/V Polarstern. The Pleisto- cene foraminiferal assemblages from these cores consist of Materials and methods * Anna Waśkowska The sediment cores from which the new species is described waskowsk@agh.edu.pl were collected from the Lomonosov Ridge in the Central Faculty of Geology, Geophysics and Environmental Arctic Ocean during Expedition PS87 of the R/V Polarstern Protection, AGH University of Science and Technology, al. in the summer of 2014 [12]. The new species is described Mickiewicza 30, 30-059 Kraków, Poland from Kastenlot Core PS87/30-1 (coordinates: 88° 39.72′N, Geosciences Department, College of Petroleum Engineering 61° 32,52′W, water depth 1276.8  m); Kastenlot Core and Geosciences, King Fahd University of Petroleum PS87/023-1 corecatcher (coordinates: 86° 38.23′ N, 44° and Minerals, PO Box 701, Dhahran 31261, Saudi Arabia Vol.:(0123456789) 1 3 0 80 N LOMONOSOV RIDGE 14 Page 2 of 10 arktos (2018) 4:14 53.98′ W, water depth 2444.8 m), Gravity Core PS87/100 Superfamily PSAMMOSPHAEROIDEA Haeckel, 1894 corecatcher (coordinates: 81° 21.42′ N, 142° 35.46′ E, Family LACUSTRINELLIDAE Mikhalevich, 1995 water depth 951  m) and Gravity Core PS87/108-1 core- Genus Ammopemphix Loeblich, 1952 catcher (coordinates: 81° 12.79′ N, 141° 10.70′ E, water Ammopemphix hemisphaericus Waśkowska & Kamin- depth 1439.6 m). All these core locations are on the crest ski, sp. nov. of the Lomonosov Ridge (Fig. 1). The sediment lithology (Fig. 3a–e; Fig. 4a–f; Fig. 5a–e) is brown and dark brown silty clay, with interbedded sandy Derivation of name From the hemispherical shape of the clay (Fig. 2). The studied agglutinated foraminifer-bearing test. interval in Core PS PS87/30-1 encompasses the middle Type specimens The holotype (EMRC 7/8x) and para- Pleistocene between MIS4 and MIS19 [21]. types (EMRC 7/8x – x) are archived in the collections of Sediment cores were sampled onboard ship, typically the European Micropalaeontological Reference Centre at every 10 cm, and samples were gently washed through a Micropress Europe, Krakow, Poland, in Cabinet 7, Drawer 8. 63 µm sieve under running water. Samples of larger volume Material More than 65 specimens, both as single indi- were collected from the core catchers. Residues were dried viduals or in pseudocolonies consisting of several to more at 60 °C, transferred into vials, and picked into cardboard than a dozen individuals. Specimens were recovered from slides under a binocular microscope. Specimens were pho- kastenlot Core PS87/030-1, Kastenlot Core PS87/023-1, tographed in the SEM Laboratory of the Faculty of Geology, corecatcher, Gravity Core PS87/100, corecatcher, and Grav- Geophysics and Environmental Protection, AGH University ity Core PS87/108-1, corecatcher. of Science and Technology, Kraków. Locality and horizon Lomonosov Ridge in the Central Arctic Ocean, Core PS87/30, 278–280 cm. Middle Pleisto- cene, MIS-10 according to Stein et al. [21]. Systematic micropalaeontology Description Test monothalamous, attached, pseudocolo- nial. The chamber is hemispherical, planoconvex, with a flat Order ASTRORHIZIDA Lankester, 1885 attachment surface and a convex dorsal side. The attachment Suborder HEMISPHAERAMMININAE Loblich & Tap- surface consists of a thin agglutinated wall with a smooth pan, 1961 inner surface. Individuals are sometimes found as isolated Fig. 1 The Lomonosov Ridge NOVAYA area with position of cores ZEMLYA containing Ammopemphix hemi- SEVERNAYA BARENTS ASIA ZEMLYA sphaericus sp. nov SEA NEW SIBERIAN ISL. FRANZ JOSEF ISL. EAST SIBERIAN SEA PS 87/100 PS 87/108 A RCTIC OCEA N GREENLAND SEA PS 87/030 BEAUFORT PS 87/023 SEA GREENLAND ELLESMERE ISL. 1 3 arktos (2018) 4:14 Page 3 of 10 14 Fig. 2 Chronology and lithol- PS 87/030-1 ogy of the type section Core PS87/30-1, after Stein et al. [21] with distribution of samples STRATIGRAPHY containing the holotype and paratypes Ammopemphix hemi- sphaericus sp. nov Age Age Marine Isotope LITHOLOGY Epoch Stage Stages (Ma) 0 m 0.13 SAMPLES 0.2 234-236 Holotype 278-280 295-297 0.4 302-304 322-324 332-334 404-406 437-439 0.6 457-459 476-478 536-538 583-585 0.77 interglacial periods silty clay glacial periods sand samples with Ammophemphix hemisphaerica nov. sp 1 3 PLEIS TO CENE MIDDLE LATE 18 MIS 16 14 MIS 12 MIS 10 8 MIS 6 MIS 4 MIS 19 MIS 17 MIS 15 MIS 13 MIS 11 MIS 9 MIS 7 MIS 5 DEPTH 14 Page 4 of 10 arktos (2018) 4:14 Fig. 3 Holotype specimen of Ammopemphix hemisphaericus sp. nov. a pseudocolony with the holotype (upper right corner), d Ammopem- attached to a fragment of an agglutinated test (Core PS87/30 278- phix hemisphaericus sp. nov. holotype (EMRC-7/x), e broken speci- 280cm) (SEM photo). a General view of the pseudocolony, b posi- men of Ammopemphix hemisphaericus sp. nov.. showing the finely tion of individual photos (white squares) and outlines of individual agglutinated interior, Scale bar = 100 µm specimens (highlighted with black lines), c details of a fragment of 1 3 arktos (2018) 4:14 Page 5 of 10 14 specimens, but more often the species forms pseudocolonies oligotrophic glacial environment, when the central Arctic with individuals attached to one another, but without any was covered by thick ice. connection between the individual tests. Test wall aggluti- nated, thin, made of small (ca. 10 µm) mineral grains in a yellowish-orange organic cement. Grain size of agglutinated Discussion grains is variable, with finer grains used in the interior of the test and around the aperture. Aperture single, simple, The Pleistocene agglutinated foraminiferal rounded, oval, or subtriangular, situated on a elevated collar assemblage on the dorsal side of the test. Dimensions Holotype (Fig. 3c, d) is 90 µm in diameter, Ammopemphix hemisphaericus sp. nov. is most often found 40 µm high, with an aperture 5 µm in diameter. Paratypes in agglutinated foraminiferal assemblages that show elevated range from 20 µm to 155 µm in diameter. biodiversity. The Pleistocene assemblages are normally Remarks Ammopemphix hemisphaericus sp. nov. belongs dominated by Reticulophragmium pusillum (Brady), with to one of the more commonly occurring agglutinated Haplophragmoides arcticus Kaminski, Waskowska and foraminiferal species in the Arctic Pleistocene. It can be Chan, Trochammina lomonosovensis Evans and Kaminski, found attaching itself to crevices and depressions on clastic and the attached form Placopsinella aurantiaca Earland as and bioclastic mineral grains. It often occupies the interiors subdominant or accompanying species. In samples with A. of other agglutinated foraminifera such as Psammosphaera, hemisphaericus sp. nov. monothalamids are commonly rep- hemispherical fragments of Hemisphaerammina, or the resented by such forms as Psammosphaera, Saccammina, concave surfaces of miscellaneous fragments derived from Hemisphaerammina, as well as agglutinated tubes, including an undetermined agglutinated foraminifer. Specimens have Psammosiphonella, Rhizammina, and Hyperammina. The also been observed on the interior of the bilamellar wall of species Reophax davisii Paar, Pseudonodosinella nodu- a species that resembles Reophax davisii Parr, and on the losa (Brady), Hormosina sp., Alvelophragmium polarensis interior of a tubular species belonging to the genus Psam- O’Neill, Cystammina pauciloculata (Brady), and Glomo- mosiphonella (Fig. 4c). More rarely, it occurs attached to spira gordialis (Jones and Parker) are rare elements of the a calcareous benthic foraminifer, a specimen of Neoglobo- assemblage. Some specimens of A. hemisphaericus sp. nov. quadrina, a bivalve shell fragment, or an ice-rafted quartz are found attached to large, coarsely agglutinated fragments grain (Fig. 4d). Successive generations of A. hemisphaericus that show little or no curvature, which may be pieces of a sp. nov. can be observed on a single substrate particle— larger agglutinated foraminifer such as a xenophyophore, where younger forms attach themselves on top of (or inside) but because of their fragmentary nature their taxonomic previous generations of Ammopemphix (Fig. 5b, d, e). In all affinity is difficult to establish. In the upper two metres of cases the apertures of adjacent individuals open dorsally, the studied cores, the interglacial layers also contain cal- indicating there was no foramen or other type of connection careous benthic foraminifera dominated by Cassidulina and between the tests. Their habit of building new tests on/in abundant planktonic foraminifera belonging to the genus those of previous generations implies that juveniles did not Neogloboquadrina. disperse widely after reproduction took place, and is likely related to their choice of a suitable substrate. The microhabitat and ecology of Ammopemphix Concentrations of A. hemisphaericus sp. nov. form pseu- docolonies in which individuals display different stages of Pseudocolonies of A. hemisphaericus sp. nov. are found on development. Smaller hemispherical specimens are found clastic substrates that are mainly of biogenic origin. Most among larger specimens that display well-developed aper- occur inside other foraminiferal tests, which they apparently tures. Pseudocolonies may consist of several to more than a occupy after the death of the host organism. There is a pref- dozen individuals. erence for coarsely agglutinated fragments of other primi- Ammopemphix hemisphaericus sp. nov. is a delicate form tive agglutinated foraminifera. Such a preference suggests that is easily damaged. The aperture or the entire top of the a detritivore feeding strategy, perhaps taking advantage of specimen may be missing, and sometimes only the outline some residual organic matter (the organic cement) that was of basal attachment is seen on clastic grains. preserved between the grains of the host agglutinated fora- The occurrence of A. hemisphaericus sp. nov. is variable minifer. The attached species Ammolagena clavata (Jones at the Lomonosov Ridge sites. The species is more common and Parker) also displays such a substrate preference [22]. in the interglacial layers of the sediment cores. The glacial By settling on biogenic particles, A. hemisphaericus sp. layers only contain sporadic single occurrences, which testi- nov. may also be taking advantage of a bacterial flora that fies to the fact that its development was limited in the highly colonizes such grains and produces iron-manganese stain- ing of the dead foraminiferal tests. An association between 1 3 14 Page 6 of 10 arktos (2018) 4:14 iron-manganese reducing bacteria and foraminifera that the specimens of agglutinated foraminifera from the Pleis- occupy the interior of foraminiferal tests has been noted tocene of the Lomonosov Ridge are encrusted by an iron- by Gooday and Haynes [8] in the North Atlantic. Many of manganese coating. This dark coating is especially well 1 3 arktos (2018) 4:14 Page 7 of 10 14 ◂Fig. 4 Paratype specimens of Ammopemphix hemisphaericus sp. added that “any test symmetry or appearance of growth habit nov. a Ammopemphix hemisphaericus sp. nov. on the internal sur- we believe to be purely fortuitous”. In an emendation of the face of a coarsely agglutinated bioclast (Core PS87/30 476-478 cm), description of the species Haman [9] changed the descrip- b Ammopemphix hemisphaericus sp. nov. specimens on the internal tion to “unilocular”. In our view, a revised description of the surface of an agglutinated bioclast (Core PS87/30 302-304  cm), c Ammopemphix hemisphaericus sp. nov. inside a specimen of Psam- genus needs to emphasise its monothalamous and pseudo- mosiphonella sp. (Core PS87/30 536-538 cm), d Ammopemphix hem- colonial nature. isphaericus sp. nov. occupying a surface depression on the surface of Two previously described species of Ammopemphix, A. a quartz grain (Core PS87/30 278-279  cm), e Ammopemphix hemi- arctica (Cushman 1948a) and A. quadrupla (Wiesner, 1931), sphaericus sp. nov. between some quartz grains (Core PS87/30 536- 538  cm), f Ammopemphix hemisphaericus sp. nov. on the internal both consist of individuals that are arranged in a regular surface of a chamber of Hormosina sp. (Core PS87/30 404-406 cm). manner: they are of nearly identical dimensions, are sym- Scale bar = 100 µm metrical about a central point, and they occur in fours or multiples of four [3, 15, 17, 24]. Adjacent individuals share developed on specimens of Reticulophragmium pusillum and a straight suture between them. As a result, the pseudoc- Haplophragmoides arcticus, but it can also occur between olonies appear to have a radial symmetry. In our material the grains of the coarsely agglutinated fragments that are from the Lomonosov Ridge, such an arrangement of the colonized by A. hemisphaericus sp. nov. tests has been observed, but is rare and limited to speci- The glacial environment of the Arctic Ocean must have mens of approximately equal dimensions. Wiesner [24] did been difficult for the development of normal marine faunas. not mention symmetry in his original definition of his genus The cold temperatures and extremely oligotrophic condi- Urnula—this feature seems to have been introduced into the tions caused by the thick permanent ice cover must have description by Loeblich and Tappan [16]. Our specimens presented a barrier for certain benthic organisms, and fos- from the Lomonosov Ridge consist of individuals of dif- tered a benthic fauna that is much reduced in terms of both ferent sizes, and these are not neccessarily arranged on the abundance and number of species. The opportunistic species substrate with any sense of symmetry. The fact that the tests that were able to exist in such a restricted environment, such may be irregularly arranged, means that the statement about as A. hemisphaericus sp. nov., apparently took advantage of symmetry in the current description of the genus needs to marginal food resources such as the organic matter preserved be modified. We therefore propose the following emended in empty foraminiferal tests or in the intergranular spaces description of the genus Ammopemphix: within the wall of dead agglutinated foraminifera. Genus Ammopemphix Loeblich, 1952 Type species: Urnula quadrupla Wiesner, 1931 The taxonomical status of Ammopemphix Synonyms. Ammopemphix Loeblich, 1952, p. 82 (nom. subst. pro One of the diagnostic features of the genus Ammopemphix is Urnula Wiesner, 1931). its ability to construct a test that appears “multichambered”, Urnula Wiesner, 1931, p. 82 (non Urnula Claparede & as pointed out in the description of Ammopemphix lacustris Lachmann, 1857). by Haman & Marolt [10]. In his description of Urnula arc- Description Test attached, monothalamous, pseudocolo- tica, Cushman [3] stated that “chambers are apparently in nial. Chamber hemispherical, individual tests are nearly cir- communication with one another”, even though Cushman’s cular in outline; when attached to one another or hidden in a illustrations do not suggest there is any such connection crevice, their shape may be rounded-polygonal in outline or between chambers. Loeblich and Tappan [16] mentioned in conform to their substrate. Adjacent tests may be separated their description of the genus that “usually symmetrically by a straight suture between them. Wall finely agglutinated, arranged, with few chambers in single whorl, or with outer white to yellowish in colour, smoothly finished, attachment ring of chambers”, which implies a certain mode of coiling. wall thin, delicate, and translucent. Aperture a single small, Therefore there has been some confusion in the literature rounded opening at the apex of each test, may possess a concerning its unilocular vs. multilocular status. collar or finely agglutinated rim. Pleistocene–Holocene; In our opinion, the genus is better described as being Antarctic: Arctic. monothalamous and pseudocolonial as pointed out by Remarks Ammopemphix is much like the attached forms Haynes [11], who regarded the genus to be a synonym of the of the genus Sorosphaera, (e.g., Sorosphaera depressa Paleozoic (Silurian) genus Colonammina. Haman & Marolt Heron-Allen and Earland) differing in possessing a termi- [10] and Loeblich & Tappan [17] did not uphold this pro- nal aperture on top of each chamber. The type species seems posed synonymy on the grounds that Colonammina is only to display a type of radial symmetry, often consisting of known from the Paleozoic. Haman & Marolt [10] did regard four tests, with sutures between each test radiating outward Ammopemphix to be pseudocolonial, describing it as “a from a central point like on a hot-cross bun. However, other random agglomeration of single chambers”. These authors 1 3 14 Page 8 of 10 arktos (2018) 4:14 Fig. 5 Paratype specimens of Ammopemphix hemisphaericus sp. interior of a larger broken specimen (Core PS87/30 536-538  cm), c nov. a1 several Ammopemphix hemisphaericus nov. sp. on the inter- Ammopemphix hemisphaericus sp. nov. (Core PS87/30 278-279 cm), nal surface of an agglutinated bioclast (Core PS87/30 295–297  cm), d Ammopemphix hemisphaericus sp. nov., a smaller specimen nes- showing more than one generation of specimens, a2 a small unbro- tled inside a larger one (Core PS87/30 536–538  cm), e Ammopem- ken specimen wedged into a crevice between grains, b Ammopem- phix hemisphaericus sp. nov., (Core PS87/30 583–585 cm). Scale bar phix hemisphaericus sp. nov., three small specimens occupying the = 100 µm 1 3 arktos (2018) 4:14 Page 9 of 10 14 shipboard participation on Expedition PS87, and the current research species of Ammopemphix display a more irregular arrange- from CPG start-up funds. The manuscript benefitted from comments by ment. Three species of Ammopemphix have been decribed Jaroslaw Tyszka and two anonymous reviewers. We thank the Micro- to date press Europe Foundation for logistical support by housing the type Ammopemphix arctica (Cushman, 1948a), Ammopemphix specimens. quadrupla (Wiesner, 1931), and A. hemisphaericus sp. nov. Sample availability Microscope slides are deposited in the Euro- A fourth unnamed species was illustrated by Barbosa et al. pean Micropalaeontological Reference Centre, Micropress Europe, al. [1] from a Brazilian mangrove environment, which might Mickiewicza 30, Kraków, Poland. Viewing is by appointment during be better assigned to the genus Lacustrinella Loeblich and working hours. Tappan, 1987. Haman and Marolt [10] described the spe- cies Ammopemphix lacustris from the Lac des Allemands Author contributions AW made SEM photos and plates; MAK took part in the PS87 Expedition and collected and prepared samples; Both on the Mississippi Delta. Loeblich and Tappan [17] desig- authors jointly wrote the manuscript. nated it the type species of their new genus Lacustrinella, which was reported to differ from Ammopemphix in possess- Compliance with ethical standards ing a more inflated test and a more prominent and elevated aperture. In their remarks to Lacustrinella, Loeblich and Conflict of interest Both authors declare no conflict of interest. Tappan mentioned that Ammopemphix is more flattened and has a wholly organic membrane (lacks an agglutinated Open Access This article is distributed under the terms of the Crea- wall) on the attachment surface. Our specimens from the tive Commons Attribution 4.0 International License (http://creat iveco mmons.or g/licenses/b y/4.0/), which permits unrestricted use, distribu- Arctic do not show these features. We follow the suggestion tion, and reproduction in any medium, provided you give appropriate by Loeblich and Tappan and separate the genera based on credit to the original author(s) and the source, provide a link to the their ecological preferences, as Lacustrinella only occurs Creative Commons license, and indicate if changes were made. in brackish shallow-water subtropical environments. Future investigations of the molecular biology of living specimens may ultimately resolve the issue of synonymy. The validity of Lacustrinella was upheld by Haman [9], who provided References an updated description of the type species regarding it as 1. Borbosa CF, Scott DB, Seoane JCS, Turcq BJ (2005) Foraminif- unilocular, not multilocular. eral zonations as baselines for quaternary sea-level fluctuations Wiesner [24] described the type species from a single sta- in south-southeast Brazilian mangroves and marshes. J Foramin tion, 385 m depth, off Antarctica. The test wall as comprised Res 35:22–43 of sand and mica flakes. Ammopemphix is known mostly 2. Cronin TM, Cronin MA (2015) Biological response to climate change in the Arctic Ocean: the view from the past. Arctos 1:4–18. from polar regions of the northern and southern hemisphere, https ://doi.org/10.1007/s4106 3-015-0019-3 and until now has only been reported from modern sedi- 3. Cushman JA (1948) Arctic Foraminifera. Cushman Lab Foramini- ments [e.g., 3, 4, 15, 17–20, 23, 24]. The genus has also been fer Res Spec Publ 23:1–79 reported from Brazilian mangroves [1], and from the Ria de 4. Cushman JA (1948) Foraminifera, their classification and eco - nomic use. Harvard University Press, Cambridge, pp 605 Vigo on the northwestern Iberian margin [5]. Our finding is 5. Diz P, Frances G, Costas S, Souto C, Alejo I (2004) Distribution the first report of the genus from the Pleistocene. of benthic foraminifera in coarse sediments, Ria de Vigo, NW Iberian Margin. J Foramin Res 34:258–275 6. Evans JR, Kaminski MA (1998) Pliocene and Pleistocene chron- ostratigraphy and palaeoenvironment of the Central Arctic Ocean, Conclusions using deep water agglutinated foraminifera. Micropaleontology 44:109–130 The new species A. hemisphaericus sp. nov. is described 7. Evans JR, Kaminski MA, Cronin TM.,Fütterer DK (1995) Pleis- from mid-Pleistocene sediments from the Lomonosov Ridge tocene agglutinated foraminifera from the Lomonosov Ridge and Amundsen Basin, Arctic Basin. Initial report on position in the Central Arctic Ocean. The species forms pseudoc- cores 2177-5 (KAL) and 2176-3 (KAL). Mar Micropaleontol olonies that are often found in the inner surface of empty 26:245–253 agglutinated foraminiferal tests and more rarely on inorganic 8. Gooday AJ, Haynes JR (1983) Abyssal foraminifers, including two sediment particles. The definition of the genus Ammopem- new genera, encrusting the interior of Bathisiphon rusticus tubes. Deep Sea Res 30:591–614 phix is here modified taking into account the morphological 9. Haman D, Marolt RE (1985) Ammopemphix lacustris n.sp. characteristics of the new species. (Foraminiferida) from Lac Des Allemands, Louisiana. Tulane stud Geol Paleontol 18:157–159 Acknowledgements A.W. thanks AGH statutory funds Grant No. 10. Haman D (1989) Ontogeny and morphological variability of the 11.11.140.005. MAK thanks Prof. Rudiger Stein (AWI) for the oppor- foraminiferid Lacustrinella lacustris (Haman and Marolt). Micro- tunity to participate on the ALEX Expedition (PS87-ARK-XXVIII/4); paleontology 35:375–378 as well as the Deans of the Science Faculty and the College of Petro- 11. Haynes JR (1981) Foraminifera. MacMillan, London, pp 433 leum Engineering and Geosciences (CPG), KFUPM for funding 1 3 14 Page 10 of 10 arktos (2018) 4:14 12. Kaboth S, Kaminski MA, Nam S, Zwick M, De Vernal A (2014) 20. Siciński J, Jażdżewski K, De Broyer C, Presler P, Ligowski R, Micropaleontology and Biostratigraphy. In: Stein R (ed) ARK- Nonato EF, Corbisier TN, Petti MAV, Brito TAS, Lavrado HP, XXVII/4, 05 August–08 October, 2014. Unpublished Cruise Błażewicz-Paszkowycz M, Pabis K, Jażdżewska A, Campos LS Report, Alfred Wegener Institute, Bremerhaven, pp 147–160 (2011) Admiralty Bay Benthos diversity—a census of a complex 13. Kaminski MA, Waśkowska A (2017) Reticulophragmium pusillum polar ecosystem. Deep Sea Res Part II 58:30–48 assemblages from the Middle Pleistocene of the Central Arctic 21. Stein R, Forwick M, Niessen F (2014) Lithostratigraphy and chro- Ocean—preliminary results. In: The Micropaleontological Society nology of PS87 sediment cores: preliminary results. In: Stein R Annual Conference, London, England, 15–16 Nov 2017 (ed) ARK-XXVII/4, 05 August–08 October, 2014. Unpublished 14. Loeblich AR (1952) Ammopemphis, new name for the recent cruise report, Alfred Wegener Institute, Bremerhaven, pp 125–143 foraminifera genus Urnula Wiesner. J Wash Acad Sci 43:82 22. Waśkowska A (2014) Distribution of the agglutinated foramini- 15. Loeblich AR, Tappan H (1952) Studies of Arctic foraminifera. fer Ammolagena clavata (Jones and Parker) in Western Tethyan Smithson Misc Collect 121:1–150 Upper Cretaceous and Paleogene deep-water deposits (Outer Car- 16. Loeblich AR, Tappan H (1964) Sarcodina chiefly “Thecamoebi- pathians, Poland). Micropaleontology 60:77–88 ans” and Foraminiferida. In: Moore RC (ed) Treatise on inverte- 23. Wollenburg J (1995) Benthische Foraminiferenfauna als Wasser- brate paleontology, part C, Protista. University of Kansas Press, masses-, Produktions- und Eisdriftanzeiger im Arktischen Ozean. Lawrence, pp 900 Berichte zur Polarforschung, 179, Alfred Wegener Institute for 17. Loeblich AR, Tappan H (1987) Foraminiferal genera and their Polar and Marine Research, Bremerhaven, pp 1–227 classification. Van Nostrand Reinhold, New York, pp 970 24. Wiesner H (1931) Die Foraminiferen der Deutsche Südpolar- 18. Majewski W (2010) Benthic foraminifera from West Antarctic Expedition 1901–1903. Deutsche Südpolar-Exped 20:53–165 fiord environments: An overview. Pol Polar Res 31:61–82 19. Majewski W (2005) Benthic foraminiferal communities: distri- bution and ecology in Admiralty Bay, King George Island, West Antarctica. Pol Polar Res 26:159–214 1 3 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png arktos Springer Journals

Ammopemphix hemisphaericus sp. nov., a new attached agglutinated foraminifer from the Pleistocene of the Arctic Ocean, and the taxonomic status of the genus Ammopemphix Loeblich, 1952

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Abstract

The new species Ammopemphix hemisphaericus sp. nov. is described from Pleistocene sediments of the Central Arctic Ocean. The species is a finely agglutinated monothalamous form with hemispherical shape and a terminal aperture located at the top of the test, often occurring in pseudocolonies. A. hemisphaericus sp. nov. attaches itself to inorganic or organic substrates, often growing attached to the inside surfaces of calcareous or agglutinated foraminiferal tests. Pseudocolonies of A. hemisphaericus sp. nov. may also attach themselves to previous generations of Ammopemphix individuals. Based on our study of numerous specimens of Ammopemphix from the Central Arctic Ocean, we propose modifications to the description of the genus. The genus may be pseudocolonial or consist of isolated individuals, and in the case of pseudocolonial forms, individual specimens may be arranged randomly or symmetrically. The genus Ammopemphix is found mostly in polar regions, and is here reported for the first time from the fossil record. Keywords Agglutinated foraminifera · Arctic Ocean · Pleistocene Introduction approximately 20 species of agglutinated foraminifera [13]. The taxonomy and biostratigraphy of the benthic foraminiferal The glacial environment of the deep Arctic Ocean yields spe- assemblages from the PS87 cores will be described separately. cies that are predisposed to life in cold and oligotrophic condi- The purpose of this paper is to describe an enigmatic new spe- tions. Because of profound changes in sea-ice cover, surface cies that lives attached to sediment particles, or most often to productivity, and sedimentation linked to glacial/interglacial fragments of other specimens of agglutinated foraminifera. cycles, the benthic foraminiferal and ostracode assemblages The genus Ammopemphix has been previously reported found in the glacial Arctic Ocean are much less abundant and from the Arctic and Antarctic seas [e.g., 3, 14, 15, 17, 24], diverse than in the interglacial faunas [2]. Glacial assemblages but specimens from the Lomonosov Ridge display morpho- reported from sediment cores are sparse and typically only logical characteristics that are not known among the previ- consist of a few species of agglutinated forms [6, 7, 13]. ously described species. In this study we describe a new During the summer of 2014, four long kastenlot cores and species of Ammopemphix and modify the current definition 36 gravity cores were collected on the Lomonosov Ridge of the genus. during Expedition 87 of the R/V Polarstern. The Pleisto- cene foraminiferal assemblages from these cores consist of Materials and methods * Anna Waśkowska The sediment cores from which the new species is described waskowsk@agh.edu.pl were collected from the Lomonosov Ridge in the Central Faculty of Geology, Geophysics and Environmental Arctic Ocean during Expedition PS87 of the R/V Polarstern Protection, AGH University of Science and Technology, al. in the summer of 2014 [12]. The new species is described Mickiewicza 30, 30-059 Kraków, Poland from Kastenlot Core PS87/30-1 (coordinates: 88° 39.72′N, Geosciences Department, College of Petroleum Engineering 61° 32,52′W, water depth 1276.8  m); Kastenlot Core and Geosciences, King Fahd University of Petroleum PS87/023-1 corecatcher (coordinates: 86° 38.23′ N, 44° and Minerals, PO Box 701, Dhahran 31261, Saudi Arabia Vol.:(0123456789) 1 3 0 80 N LOMONOSOV RIDGE 14 Page 2 of 10 arktos (2018) 4:14 53.98′ W, water depth 2444.8 m), Gravity Core PS87/100 Superfamily PSAMMOSPHAEROIDEA Haeckel, 1894 corecatcher (coordinates: 81° 21.42′ N, 142° 35.46′ E, Family LACUSTRINELLIDAE Mikhalevich, 1995 water depth 951  m) and Gravity Core PS87/108-1 core- Genus Ammopemphix Loeblich, 1952 catcher (coordinates: 81° 12.79′ N, 141° 10.70′ E, water Ammopemphix hemisphaericus Waśkowska & Kamin- depth 1439.6 m). All these core locations are on the crest ski, sp. nov. of the Lomonosov Ridge (Fig. 1). The sediment lithology (Fig. 3a–e; Fig. 4a–f; Fig. 5a–e) is brown and dark brown silty clay, with interbedded sandy Derivation of name From the hemispherical shape of the clay (Fig. 2). The studied agglutinated foraminifer-bearing test. interval in Core PS PS87/30-1 encompasses the middle Type specimens The holotype (EMRC 7/8x) and para- Pleistocene between MIS4 and MIS19 [21]. types (EMRC 7/8x – x) are archived in the collections of Sediment cores were sampled onboard ship, typically the European Micropalaeontological Reference Centre at every 10 cm, and samples were gently washed through a Micropress Europe, Krakow, Poland, in Cabinet 7, Drawer 8. 63 µm sieve under running water. Samples of larger volume Material More than 65 specimens, both as single indi- were collected from the core catchers. Residues were dried viduals or in pseudocolonies consisting of several to more at 60 °C, transferred into vials, and picked into cardboard than a dozen individuals. Specimens were recovered from slides under a binocular microscope. Specimens were pho- kastenlot Core PS87/030-1, Kastenlot Core PS87/023-1, tographed in the SEM Laboratory of the Faculty of Geology, corecatcher, Gravity Core PS87/100, corecatcher, and Grav- Geophysics and Environmental Protection, AGH University ity Core PS87/108-1, corecatcher. of Science and Technology, Kraków. Locality and horizon Lomonosov Ridge in the Central Arctic Ocean, Core PS87/30, 278–280 cm. Middle Pleisto- cene, MIS-10 according to Stein et al. [21]. Systematic micropalaeontology Description Test monothalamous, attached, pseudocolo- nial. The chamber is hemispherical, planoconvex, with a flat Order ASTRORHIZIDA Lankester, 1885 attachment surface and a convex dorsal side. The attachment Suborder HEMISPHAERAMMININAE Loblich & Tap- surface consists of a thin agglutinated wall with a smooth pan, 1961 inner surface. Individuals are sometimes found as isolated Fig. 1 The Lomonosov Ridge NOVAYA area with position of cores ZEMLYA containing Ammopemphix hemi- SEVERNAYA BARENTS ASIA ZEMLYA sphaericus sp. nov SEA NEW SIBERIAN ISL. FRANZ JOSEF ISL. EAST SIBERIAN SEA PS 87/100 PS 87/108 A RCTIC OCEA N GREENLAND SEA PS 87/030 BEAUFORT PS 87/023 SEA GREENLAND ELLESMERE ISL. 1 3 arktos (2018) 4:14 Page 3 of 10 14 Fig. 2 Chronology and lithol- PS 87/030-1 ogy of the type section Core PS87/30-1, after Stein et al. [21] with distribution of samples STRATIGRAPHY containing the holotype and paratypes Ammopemphix hemi- sphaericus sp. nov Age Age Marine Isotope LITHOLOGY Epoch Stage Stages (Ma) 0 m 0.13 SAMPLES 0.2 234-236 Holotype 278-280 295-297 0.4 302-304 322-324 332-334 404-406 437-439 0.6 457-459 476-478 536-538 583-585 0.77 interglacial periods silty clay glacial periods sand samples with Ammophemphix hemisphaerica nov. sp 1 3 PLEIS TO CENE MIDDLE LATE 18 MIS 16 14 MIS 12 MIS 10 8 MIS 6 MIS 4 MIS 19 MIS 17 MIS 15 MIS 13 MIS 11 MIS 9 MIS 7 MIS 5 DEPTH 14 Page 4 of 10 arktos (2018) 4:14 Fig. 3 Holotype specimen of Ammopemphix hemisphaericus sp. nov. a pseudocolony with the holotype (upper right corner), d Ammopem- attached to a fragment of an agglutinated test (Core PS87/30 278- phix hemisphaericus sp. nov. holotype (EMRC-7/x), e broken speci- 280cm) (SEM photo). a General view of the pseudocolony, b posi- men of Ammopemphix hemisphaericus sp. nov.. showing the finely tion of individual photos (white squares) and outlines of individual agglutinated interior, Scale bar = 100 µm specimens (highlighted with black lines), c details of a fragment of 1 3 arktos (2018) 4:14 Page 5 of 10 14 specimens, but more often the species forms pseudocolonies oligotrophic glacial environment, when the central Arctic with individuals attached to one another, but without any was covered by thick ice. connection between the individual tests. Test wall aggluti- nated, thin, made of small (ca. 10 µm) mineral grains in a yellowish-orange organic cement. Grain size of agglutinated Discussion grains is variable, with finer grains used in the interior of the test and around the aperture. Aperture single, simple, The Pleistocene agglutinated foraminiferal rounded, oval, or subtriangular, situated on a elevated collar assemblage on the dorsal side of the test. Dimensions Holotype (Fig. 3c, d) is 90 µm in diameter, Ammopemphix hemisphaericus sp. nov. is most often found 40 µm high, with an aperture 5 µm in diameter. Paratypes in agglutinated foraminiferal assemblages that show elevated range from 20 µm to 155 µm in diameter. biodiversity. The Pleistocene assemblages are normally Remarks Ammopemphix hemisphaericus sp. nov. belongs dominated by Reticulophragmium pusillum (Brady), with to one of the more commonly occurring agglutinated Haplophragmoides arcticus Kaminski, Waskowska and foraminiferal species in the Arctic Pleistocene. It can be Chan, Trochammina lomonosovensis Evans and Kaminski, found attaching itself to crevices and depressions on clastic and the attached form Placopsinella aurantiaca Earland as and bioclastic mineral grains. It often occupies the interiors subdominant or accompanying species. In samples with A. of other agglutinated foraminifera such as Psammosphaera, hemisphaericus sp. nov. monothalamids are commonly rep- hemispherical fragments of Hemisphaerammina, or the resented by such forms as Psammosphaera, Saccammina, concave surfaces of miscellaneous fragments derived from Hemisphaerammina, as well as agglutinated tubes, including an undetermined agglutinated foraminifer. Specimens have Psammosiphonella, Rhizammina, and Hyperammina. The also been observed on the interior of the bilamellar wall of species Reophax davisii Paar, Pseudonodosinella nodu- a species that resembles Reophax davisii Parr, and on the losa (Brady), Hormosina sp., Alvelophragmium polarensis interior of a tubular species belonging to the genus Psam- O’Neill, Cystammina pauciloculata (Brady), and Glomo- mosiphonella (Fig. 4c). More rarely, it occurs attached to spira gordialis (Jones and Parker) are rare elements of the a calcareous benthic foraminifer, a specimen of Neoglobo- assemblage. Some specimens of A. hemisphaericus sp. nov. quadrina, a bivalve shell fragment, or an ice-rafted quartz are found attached to large, coarsely agglutinated fragments grain (Fig. 4d). Successive generations of A. hemisphaericus that show little or no curvature, which may be pieces of a sp. nov. can be observed on a single substrate particle— larger agglutinated foraminifer such as a xenophyophore, where younger forms attach themselves on top of (or inside) but because of their fragmentary nature their taxonomic previous generations of Ammopemphix (Fig. 5b, d, e). In all affinity is difficult to establish. In the upper two metres of cases the apertures of adjacent individuals open dorsally, the studied cores, the interglacial layers also contain cal- indicating there was no foramen or other type of connection careous benthic foraminifera dominated by Cassidulina and between the tests. Their habit of building new tests on/in abundant planktonic foraminifera belonging to the genus those of previous generations implies that juveniles did not Neogloboquadrina. disperse widely after reproduction took place, and is likely related to their choice of a suitable substrate. The microhabitat and ecology of Ammopemphix Concentrations of A. hemisphaericus sp. nov. form pseu- docolonies in which individuals display different stages of Pseudocolonies of A. hemisphaericus sp. nov. are found on development. Smaller hemispherical specimens are found clastic substrates that are mainly of biogenic origin. Most among larger specimens that display well-developed aper- occur inside other foraminiferal tests, which they apparently tures. Pseudocolonies may consist of several to more than a occupy after the death of the host organism. There is a pref- dozen individuals. erence for coarsely agglutinated fragments of other primi- Ammopemphix hemisphaericus sp. nov. is a delicate form tive agglutinated foraminifera. Such a preference suggests that is easily damaged. The aperture or the entire top of the a detritivore feeding strategy, perhaps taking advantage of specimen may be missing, and sometimes only the outline some residual organic matter (the organic cement) that was of basal attachment is seen on clastic grains. preserved between the grains of the host agglutinated fora- The occurrence of A. hemisphaericus sp. nov. is variable minifer. The attached species Ammolagena clavata (Jones at the Lomonosov Ridge sites. The species is more common and Parker) also displays such a substrate preference [22]. in the interglacial layers of the sediment cores. The glacial By settling on biogenic particles, A. hemisphaericus sp. layers only contain sporadic single occurrences, which testi- nov. may also be taking advantage of a bacterial flora that fies to the fact that its development was limited in the highly colonizes such grains and produces iron-manganese stain- ing of the dead foraminiferal tests. An association between 1 3 14 Page 6 of 10 arktos (2018) 4:14 iron-manganese reducing bacteria and foraminifera that the specimens of agglutinated foraminifera from the Pleis- occupy the interior of foraminiferal tests has been noted tocene of the Lomonosov Ridge are encrusted by an iron- by Gooday and Haynes [8] in the North Atlantic. Many of manganese coating. This dark coating is especially well 1 3 arktos (2018) 4:14 Page 7 of 10 14 ◂Fig. 4 Paratype specimens of Ammopemphix hemisphaericus sp. added that “any test symmetry or appearance of growth habit nov. a Ammopemphix hemisphaericus sp. nov. on the internal sur- we believe to be purely fortuitous”. In an emendation of the face of a coarsely agglutinated bioclast (Core PS87/30 476-478 cm), description of the species Haman [9] changed the descrip- b Ammopemphix hemisphaericus sp. nov. specimens on the internal tion to “unilocular”. In our view, a revised description of the surface of an agglutinated bioclast (Core PS87/30 302-304  cm), c Ammopemphix hemisphaericus sp. nov. inside a specimen of Psam- genus needs to emphasise its monothalamous and pseudo- mosiphonella sp. (Core PS87/30 536-538 cm), d Ammopemphix hem- colonial nature. isphaericus sp. nov. occupying a surface depression on the surface of Two previously described species of Ammopemphix, A. a quartz grain (Core PS87/30 278-279  cm), e Ammopemphix hemi- arctica (Cushman 1948a) and A. quadrupla (Wiesner, 1931), sphaericus sp. nov. between some quartz grains (Core PS87/30 536- 538  cm), f Ammopemphix hemisphaericus sp. nov. on the internal both consist of individuals that are arranged in a regular surface of a chamber of Hormosina sp. (Core PS87/30 404-406 cm). manner: they are of nearly identical dimensions, are sym- Scale bar = 100 µm metrical about a central point, and they occur in fours or multiples of four [3, 15, 17, 24]. Adjacent individuals share developed on specimens of Reticulophragmium pusillum and a straight suture between them. As a result, the pseudoc- Haplophragmoides arcticus, but it can also occur between olonies appear to have a radial symmetry. In our material the grains of the coarsely agglutinated fragments that are from the Lomonosov Ridge, such an arrangement of the colonized by A. hemisphaericus sp. nov. tests has been observed, but is rare and limited to speci- The glacial environment of the Arctic Ocean must have mens of approximately equal dimensions. Wiesner [24] did been difficult for the development of normal marine faunas. not mention symmetry in his original definition of his genus The cold temperatures and extremely oligotrophic condi- Urnula—this feature seems to have been introduced into the tions caused by the thick permanent ice cover must have description by Loeblich and Tappan [16]. Our specimens presented a barrier for certain benthic organisms, and fos- from the Lomonosov Ridge consist of individuals of dif- tered a benthic fauna that is much reduced in terms of both ferent sizes, and these are not neccessarily arranged on the abundance and number of species. The opportunistic species substrate with any sense of symmetry. The fact that the tests that were able to exist in such a restricted environment, such may be irregularly arranged, means that the statement about as A. hemisphaericus sp. nov., apparently took advantage of symmetry in the current description of the genus needs to marginal food resources such as the organic matter preserved be modified. We therefore propose the following emended in empty foraminiferal tests or in the intergranular spaces description of the genus Ammopemphix: within the wall of dead agglutinated foraminifera. Genus Ammopemphix Loeblich, 1952 Type species: Urnula quadrupla Wiesner, 1931 The taxonomical status of Ammopemphix Synonyms. Ammopemphix Loeblich, 1952, p. 82 (nom. subst. pro One of the diagnostic features of the genus Ammopemphix is Urnula Wiesner, 1931). its ability to construct a test that appears “multichambered”, Urnula Wiesner, 1931, p. 82 (non Urnula Claparede & as pointed out in the description of Ammopemphix lacustris Lachmann, 1857). by Haman & Marolt [10]. In his description of Urnula arc- Description Test attached, monothalamous, pseudocolo- tica, Cushman [3] stated that “chambers are apparently in nial. Chamber hemispherical, individual tests are nearly cir- communication with one another”, even though Cushman’s cular in outline; when attached to one another or hidden in a illustrations do not suggest there is any such connection crevice, their shape may be rounded-polygonal in outline or between chambers. Loeblich and Tappan [16] mentioned in conform to their substrate. Adjacent tests may be separated their description of the genus that “usually symmetrically by a straight suture between them. Wall finely agglutinated, arranged, with few chambers in single whorl, or with outer white to yellowish in colour, smoothly finished, attachment ring of chambers”, which implies a certain mode of coiling. wall thin, delicate, and translucent. Aperture a single small, Therefore there has been some confusion in the literature rounded opening at the apex of each test, may possess a concerning its unilocular vs. multilocular status. collar or finely agglutinated rim. Pleistocene–Holocene; In our opinion, the genus is better described as being Antarctic: Arctic. monothalamous and pseudocolonial as pointed out by Remarks Ammopemphix is much like the attached forms Haynes [11], who regarded the genus to be a synonym of the of the genus Sorosphaera, (e.g., Sorosphaera depressa Paleozoic (Silurian) genus Colonammina. Haman & Marolt Heron-Allen and Earland) differing in possessing a termi- [10] and Loeblich & Tappan [17] did not uphold this pro- nal aperture on top of each chamber. The type species seems posed synonymy on the grounds that Colonammina is only to display a type of radial symmetry, often consisting of known from the Paleozoic. Haman & Marolt [10] did regard four tests, with sutures between each test radiating outward Ammopemphix to be pseudocolonial, describing it as “a from a central point like on a hot-cross bun. However, other random agglomeration of single chambers”. These authors 1 3 14 Page 8 of 10 arktos (2018) 4:14 Fig. 5 Paratype specimens of Ammopemphix hemisphaericus sp. interior of a larger broken specimen (Core PS87/30 536-538  cm), c nov. a1 several Ammopemphix hemisphaericus nov. sp. on the inter- Ammopemphix hemisphaericus sp. nov. (Core PS87/30 278-279 cm), nal surface of an agglutinated bioclast (Core PS87/30 295–297  cm), d Ammopemphix hemisphaericus sp. nov., a smaller specimen nes- showing more than one generation of specimens, a2 a small unbro- tled inside a larger one (Core PS87/30 536–538  cm), e Ammopem- ken specimen wedged into a crevice between grains, b Ammopem- phix hemisphaericus sp. nov., (Core PS87/30 583–585 cm). Scale bar phix hemisphaericus sp. nov., three small specimens occupying the = 100 µm 1 3 arktos (2018) 4:14 Page 9 of 10 14 shipboard participation on Expedition PS87, and the current research species of Ammopemphix display a more irregular arrange- from CPG start-up funds. The manuscript benefitted from comments by ment. Three species of Ammopemphix have been decribed Jaroslaw Tyszka and two anonymous reviewers. We thank the Micro- to date press Europe Foundation for logistical support by housing the type Ammopemphix arctica (Cushman, 1948a), Ammopemphix specimens. quadrupla (Wiesner, 1931), and A. hemisphaericus sp. nov. Sample availability Microscope slides are deposited in the Euro- A fourth unnamed species was illustrated by Barbosa et al. pean Micropalaeontological Reference Centre, Micropress Europe, al. [1] from a Brazilian mangrove environment, which might Mickiewicza 30, Kraków, Poland. Viewing is by appointment during be better assigned to the genus Lacustrinella Loeblich and working hours. Tappan, 1987. Haman and Marolt [10] described the spe- cies Ammopemphix lacustris from the Lac des Allemands Author contributions AW made SEM photos and plates; MAK took part in the PS87 Expedition and collected and prepared samples; Both on the Mississippi Delta. Loeblich and Tappan [17] desig- authors jointly wrote the manuscript. nated it the type species of their new genus Lacustrinella, which was reported to differ from Ammopemphix in possess- Compliance with ethical standards ing a more inflated test and a more prominent and elevated aperture. In their remarks to Lacustrinella, Loeblich and Conflict of interest Both authors declare no conflict of interest. Tappan mentioned that Ammopemphix is more flattened and has a wholly organic membrane (lacks an agglutinated Open Access This article is distributed under the terms of the Crea- wall) on the attachment surface. Our specimens from the tive Commons Attribution 4.0 International License (http://creat iveco mmons.or g/licenses/b y/4.0/), which permits unrestricted use, distribu- Arctic do not show these features. We follow the suggestion tion, and reproduction in any medium, provided you give appropriate by Loeblich and Tappan and separate the genera based on credit to the original author(s) and the source, provide a link to the their ecological preferences, as Lacustrinella only occurs Creative Commons license, and indicate if changes were made. in brackish shallow-water subtropical environments. Future investigations of the molecular biology of living specimens may ultimately resolve the issue of synonymy. The validity of Lacustrinella was upheld by Haman [9], who provided References an updated description of the type species regarding it as 1. Borbosa CF, Scott DB, Seoane JCS, Turcq BJ (2005) Foraminif- unilocular, not multilocular. eral zonations as baselines for quaternary sea-level fluctuations Wiesner [24] described the type species from a single sta- in south-southeast Brazilian mangroves and marshes. J Foramin tion, 385 m depth, off Antarctica. The test wall as comprised Res 35:22–43 of sand and mica flakes. Ammopemphix is known mostly 2. Cronin TM, Cronin MA (2015) Biological response to climate change in the Arctic Ocean: the view from the past. Arctos 1:4–18. from polar regions of the northern and southern hemisphere, https ://doi.org/10.1007/s4106 3-015-0019-3 and until now has only been reported from modern sedi- 3. Cushman JA (1948) Arctic Foraminifera. Cushman Lab Foramini- ments [e.g., 3, 4, 15, 17–20, 23, 24]. The genus has also been fer Res Spec Publ 23:1–79 reported from Brazilian mangroves [1], and from the Ria de 4. Cushman JA (1948) Foraminifera, their classification and eco - nomic use. Harvard University Press, Cambridge, pp 605 Vigo on the northwestern Iberian margin [5]. Our finding is 5. Diz P, Frances G, Costas S, Souto C, Alejo I (2004) Distribution the first report of the genus from the Pleistocene. of benthic foraminifera in coarse sediments, Ria de Vigo, NW Iberian Margin. J Foramin Res 34:258–275 6. Evans JR, Kaminski MA (1998) Pliocene and Pleistocene chron- ostratigraphy and palaeoenvironment of the Central Arctic Ocean, Conclusions using deep water agglutinated foraminifera. Micropaleontology 44:109–130 The new species A. hemisphaericus sp. nov. is described 7. Evans JR, Kaminski MA, Cronin TM.,Fütterer DK (1995) Pleis- from mid-Pleistocene sediments from the Lomonosov Ridge tocene agglutinated foraminifera from the Lomonosov Ridge and Amundsen Basin, Arctic Basin. Initial report on position in the Central Arctic Ocean. The species forms pseudoc- cores 2177-5 (KAL) and 2176-3 (KAL). Mar Micropaleontol olonies that are often found in the inner surface of empty 26:245–253 agglutinated foraminiferal tests and more rarely on inorganic 8. Gooday AJ, Haynes JR (1983) Abyssal foraminifers, including two sediment particles. The definition of the genus Ammopem- new genera, encrusting the interior of Bathisiphon rusticus tubes. Deep Sea Res 30:591–614 phix is here modified taking into account the morphological 9. Haman D, Marolt RE (1985) Ammopemphix lacustris n.sp. characteristics of the new species. (Foraminiferida) from Lac Des Allemands, Louisiana. Tulane stud Geol Paleontol 18:157–159 Acknowledgements A.W. thanks AGH statutory funds Grant No. 10. Haman D (1989) Ontogeny and morphological variability of the 11.11.140.005. MAK thanks Prof. Rudiger Stein (AWI) for the oppor- foraminiferid Lacustrinella lacustris (Haman and Marolt). Micro- tunity to participate on the ALEX Expedition (PS87-ARK-XXVIII/4); paleontology 35:375–378 as well as the Deans of the Science Faculty and the College of Petro- 11. Haynes JR (1981) Foraminifera. MacMillan, London, pp 433 leum Engineering and Geosciences (CPG), KFUPM for funding 1 3 14 Page 10 of 10 arktos (2018) 4:14 12. Kaboth S, Kaminski MA, Nam S, Zwick M, De Vernal A (2014) 20. Siciński J, Jażdżewski K, De Broyer C, Presler P, Ligowski R, Micropaleontology and Biostratigraphy. In: Stein R (ed) ARK- Nonato EF, Corbisier TN, Petti MAV, Brito TAS, Lavrado HP, XXVII/4, 05 August–08 October, 2014. Unpublished Cruise Błażewicz-Paszkowycz M, Pabis K, Jażdżewska A, Campos LS Report, Alfred Wegener Institute, Bremerhaven, pp 147–160 (2011) Admiralty Bay Benthos diversity—a census of a complex 13. Kaminski MA, Waśkowska A (2017) Reticulophragmium pusillum polar ecosystem. Deep Sea Res Part II 58:30–48 assemblages from the Middle Pleistocene of the Central Arctic 21. Stein R, Forwick M, Niessen F (2014) Lithostratigraphy and chro- Ocean—preliminary results. In: The Micropaleontological Society nology of PS87 sediment cores: preliminary results. In: Stein R Annual Conference, London, England, 15–16 Nov 2017 (ed) ARK-XXVII/4, 05 August–08 October, 2014. Unpublished 14. Loeblich AR (1952) Ammopemphis, new name for the recent cruise report, Alfred Wegener Institute, Bremerhaven, pp 125–143 foraminifera genus Urnula Wiesner. J Wash Acad Sci 43:82 22. Waśkowska A (2014) Distribution of the agglutinated foramini- 15. Loeblich AR, Tappan H (1952) Studies of Arctic foraminifera. fer Ammolagena clavata (Jones and Parker) in Western Tethyan Smithson Misc Collect 121:1–150 Upper Cretaceous and Paleogene deep-water deposits (Outer Car- 16. Loeblich AR, Tappan H (1964) Sarcodina chiefly “Thecamoebi- pathians, Poland). Micropaleontology 60:77–88 ans” and Foraminiferida. In: Moore RC (ed) Treatise on inverte- 23. Wollenburg J (1995) Benthische Foraminiferenfauna als Wasser- brate paleontology, part C, Protista. University of Kansas Press, masses-, Produktions- und Eisdriftanzeiger im Arktischen Ozean. Lawrence, pp 900 Berichte zur Polarforschung, 179, Alfred Wegener Institute for 17. Loeblich AR, Tappan H (1987) Foraminiferal genera and their Polar and Marine Research, Bremerhaven, pp 1–227 classification. Van Nostrand Reinhold, New York, pp 970 24. Wiesner H (1931) Die Foraminiferen der Deutsche Südpolar- 18. Majewski W (2010) Benthic foraminifera from West Antarctic Expedition 1901–1903. Deutsche Südpolar-Exped 20:53–165 fiord environments: An overview. Pol Polar Res 31:61–82 19. Majewski W (2005) Benthic foraminiferal communities: distri- bution and ecology in Admiralty Bay, King George Island, West Antarctica. Pol Polar Res 26:159–214 1 3

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