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NEW THALASSIOSIRA SPECIES (BACILLARIOPHYTA) FROM HOLOCENE SEDIMENTS OF THE BUNGER OASIS, EAST ANTARCTICA PDF

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Polish Botanical Journal 55(1): 135–147, 2010 NEW THALASSIOSIRA SPECIES (BACILLARIOPHYTA) FROM HOLOCENE SEDIMENTS OF THE BUNGER OASIS, EAST ANTARCTICA* RINAT M. GOGOREV, ZINAIDA V. PUSHINA, MARTIN MELLES & SERGEY R. VERKULICH Abstract. Studies of Holocene marine sediments from the Bunger Oasis (East Antarctica) collected during the joint Russian-German Expedition in 1993–1994 revealed several dominant diatom species. This paper focuses on the morphology of Thalassiosira species studied by light and scanning electron microscopy. One of them, Thalassiosira longifultoportulata Gogorev & Pushina sp. nov., characterized by relatively large valves, several central fultoportulae, one ring of marginal fultoportulae and a single rimoporula, is described here as new for science. The unique morphological feature of T. longifultoportulata are the very long external tubes of marginal fultoportulae. Based on some morphological features (especially number and location of subcentral fultoportulae clusters) another new species Thalassiosira bungerensis Gogorev & Pushina sp. nov. is described. Key words: diatoms, morphology, new taxa, Thalassiosira, sediment cores, Holocene, Bunger Hills, East Antarctica Rinat M. Gogorev, Department of Algology, Komarov Botanical Institute, Russian Academy of Sciences, Prof. Popov Street, 2, St. Petersburg 197376, Russia; e-mail: [email protected] Zinaida V. Pushina, Russian Research Institute for Oceanology and Mineral Resourses, Angliyskiy Street, 1, St. Petersburg 190121, Russia; e-mail: [email protected] Martin Melles, Universität zu Köln, Mathematisch-Naturwissenschaftliche Fakultät, Institut für Geologie und Mineralogie, Zülpicher Str., 49a, D-50674, Köln, Germany; e-mail:[email protected] Sergey R. Verkulich, Arctic and Antarctic Research Institute, Department of Polar Geography, Bering Street, 199226, St. Pe- tersburg, Russia; e-mail: [email protected] INTRODUCTION In 1993–1994, a joint Russian-German scientifi c are separated from the Southern Hills by marine expedition undertook coring of lacustrine and basin inlets (Edisto Channel, Kakapon Bay, Rybiy marine sediment in the Bunger Oasis. This paper Khvost Bay), which have a hydraulic connection considers two new taxa found in bottom sediments to the open ocean beneath the fl oating glaciers and from the marine basins in the Bunger Oasis. the Shackleton Ice Shelf. The Bunger Oasis (66°16′S, 100°45′E; Fig. 1) The climate in the Southern Hills is character- forms one of the largest ice-free areas in East Ant- ized by average monthly temperatures of up to arctica, located to the east of the Scott/Denman 1.8°C in summer down to –20.6°C in winter, and Glacier system. To the north the Oasis is bordered by precipitation of ca 200 mm/yr and potential by the Shackleton Ice Shelf, to the east by the evaporation of 400–600 mm/yr (Rusin 1961). East Antarctic Ice Sheet, to the south by the Apfel The waters of marine basins on the Bunger Glacier, and to the west by the Edisto Glacier. Oasis are NaCl type (Klokov et al. 1989), with The southern part of the Oasis (Southern Hills) is Si content of 1–10 mg Si/l. Recently the waters a large land area of ca 280 km2 with a great number of the Rybiy Khvost Bay show extremely high of freshwater lakes. To the north, numerous islands phytoplankton production. The chlorophyll a concentration is 1–10 mg/l in this bay. In Rybiy Khvost Bay, salinity varies from 9‰under the * Dedicated to Dr. Kurt Krammer on the occasion of his 85th ice cover to 18.6‰ at 20 m depth. The highly birthday 136 POLISH BOTANICAL JOURNAL 55(1). 2010 40 30 20 10 0 10 20 30 40 in sediment cores from the lakes (Roberts et al. 2000; Verkulich et al. 2002). 50 50 60 60 MATERIAL AND METHODS Weddell Sea 70 70 Marine and lacustrine sediment sequences were sam- 80 80 pled by gravity and piston corers from a platform (3.5 × 2.7 m), which was run either with 4 infl atable rubber 90 70 Mirniy 90 pontoons on open water or with 4 runners on lake ice. Bunger 100 100 The gravity corer was employed for sampling undis- turbed near-surface sediments. Long sediment sequences 110 110 were recovered with the piston corer (Melles et al. Ross Sea 120 1994b). Sediment cores PG 1165 (246 cm), PG 1169 120 (190 cm), PG 1172 (265 cm), PG 1173 (1278 cm), PG 70 130 1179 (100 cm), PG 1180 (1202 cm) were sectioned on 130 site at 0.2 cm intervals. Sampling locations are shown in 140 150 160 170 180 170 160 150 140 Figure 1. The samples were processed for LM and SEM by standard procedures (Jousé et al. 1969); 300 diatom 30′ 40′ 50′ 101° E 10′ valves of each sample were counted at 1000×. Stratigraphic information was obtained by radio- Shackleton carbon dating of the total organic carbon of marine 66° S Ice Shelf and lacustrine sediments (Melles et al. 1994a, b, 1997). Channel Measurements were performed by accelerator mass 05′ Edisto soploegcytr oamnde tHryis taot ryth oe f RAerst,e aOrxchfo rLda. bFourlalt omrayr ifnoer eAnrvcihroane-- ments with biogenic sedimentation were established at 8540±80 BP (Bolshiyanov et al. 1991; Melles et al. 1994a, b, 1997; Kulbe et al. 2001). The lithology, 10′ Kakapon Bay Edisto radiocarbon chronology, granulometry, chemistry and 1179 Glacier distribution of diatoms were analyzed. Analyses of the Rybiy Khvost Bay 1172 sediment composition revealed the environmental his- 1165 Ice 15′ 1173 Sheet tory of the water basins, providing information about 1169 1180 the evolution of the Hills. Figurnoe Lake 0 5 10 km 20′ RESULTS Fig. 1. Sampling sites and geographical setting of Bunger Oasis, The deglaciation on the Bunger Oasis started close East Antarctica. to the Pleistocene/Holocene boundary with rapid ice decay, at least in the Southern Hills. The diatom mineralized water in Rybiy Khvost Bay is of fl ora from sediments of the marine basins of the marine origin. Bunger Oasis is here determined for the fi rst time. Modern diatom assemblages in lakes of the About 100 diatom species and one silicofl agellate Bunger Oasis were investigated by Gibson et al. Dictyocha speculum were identifi ed in marine di- (2006). There are no data on modern marine dia- atom-rich sediments. The most species-rich genera toms in this region. Fossil diatoms were studied are Fragilariopsis (11 species), Navicula (8; 4 un- and analyzed for diatom composition and abun- determined), Thalassiosira (7), Cocconeis (5). The dance in sediment cores from the marine basins taxonomic composition of diatoms found is more (Bolshiyanov et al. 1991; Melles et al. 1997; Kulbe or less similar to modern marine assemblages of et al. 2001; Pushina et al. 2004; Pushina 2008) and near-shore Antarctic waters. Chaetoceros resting R. M. GOGOREV ET AL.: NEW THALASSIOSIRA SPECIES FROM EAST ANTARCTICA 137 EEccoollooggiiccaall ggrroouuppss %% ttoottaall ddiiaattoommss ss ssii (())ccmmtteepphhDD llyyooggiittLLhhoo ))ll..,,((rrccyyaaBBPPggeeAA lliittttrrvvccccaaeeoonneennaaoonnVVlliitt((rr))ssnnppeeggeeddeennmmmmmm iiiittttrrccccccccuubbaannaaooeeaannSS iiiitttt--rrrrccppeennaaeenneewwOO llIItt--rrcceeeeaaeedd iillttttrrLLooaa ttrrccsshhaaeeooeeooCCsspppp.. TThhaallaassssiioossiirraa bbuunnggeerreenn iillii..rrccsshhggaaTT iillii..ttrrccssxxcchhaaeeppeeaaTT..rrvvggaa ii..ttrrrrhhaaggaaaaeeTTmm ..lliillllffttttttrraaaaTThhoonngguuooppoouu iijjlllliiiittsskknnoooonneeeeaannEEmmmm iiiittttrrccccccuuuuppaaaannaaaaEEmm iilliiiittrrrrrrssssccaaggaaooppuuaaFF llii..rrccyyssnndduuFF llssccppeeuuuuDDmmiiccttyyoocchhaa lliillttrrssccaaaaaauuaaaaPP 00 11000000%% 335500 11000000 220000 440000 11000000 330000 9900 330000 4400 440000 7700 7700 440000 110000 440000 55 00 00 110000 11000000 220000 22000000 330000 440000 33000000 550000 44000000 660000 55000000 770000 66000000 880000 990000 77000000 11000000 88000000 11110000 11220000 99000000 11229900 SSaapprrooppeell llaammiinnaatteedd ccllaayyeeyy ssiilltt MMoorraaiinnee//ttiillll Fig. 2. Lithology and Holocene variations of diatoms in sediment core PG 1173 of Rybiy Khvost Bay (Bunger Oasis, East Antarctica). spores identifi ed as Chaetoceros neglectus Karsten most stable and warm Holocene paleoceanographic were the most abundant. Diatom assemblages are conditions in the Bunger Oasis were established dominated by Chaetoceros spp., mostly from else- at 4300–1800 BP. where in the sediments of marine bays (Pushina The fi rst data on the morphology and taxonomy et al. 2004; Pushina 2008). The distribution and of the new taxa were presented earlier in an ab- abundance of diatoms in core PG 1173 are shown stract (Gogorev & Pushina 2006). Valid descrip- in Figure 2. The study revealed 2 thalassiosiroid tions of these taxa are given below. diatoms which could not be assigned to known taxa; these are described here. Thalassiosira longifultoportulata Gogorev In the distribution of these two abundant and & Pushina, sp. nov. Figs 3–17 interesting Thalassiosira species through the long sediment core PG 1173 (1290 cm, water Valvae circulares, subconvexae, margine planae, 32–80 depth 91 m), the fi rst appearance of the species μm in diametro, limbus valvae nanus. Areolae in se- is recorded at 1205 cm depth, where only marine riebus radialibus, fasciculatae, (6)7–8 in 10 μm in faciei valvae et 10–12 in 10 μm in limbui valvae, foraminis conditions were established. The abundance of circularia. Fultoportulae centrales 3–7, raro usque ad these two species was very low in the basal part 9, earum tubuli externi ad 2 μm alt. Fultoportulae mar- of the core and increased only at ca 7000 BP. The ginales in anulo singulari, 6–9(10) in 10 μm, earum tu- highest abundance (31% of total diatom assem- buli externi praecelsi, 2–6 μm alt., per 1 areola inter se blage abundance) of Thalassiosira new species separati. Rimoportula singularis, tubulo magno (quam was determined at 1000–2000 BP (Fig. 2). The fultoportulis grandior) pervio externo, 2.7–3.5 μm alt. 138 POLISH BOTANICAL JOURNAL 55(1). 2010 Figs 3–5. Valves of Thalassiosira longifultoportulata Gogorev & Pushina sp. nov. Figs 4 and 5 are same specimen viewed at different focal planes – on center (4), on valve margin (5). All LM. Scale bars = 10 μm. et 0.7–1.4 μm in diametro, et interno labiato, ad 2.6 μm ECOLOGY. Marine, probably, cold-water neritic alt., anulo fultoportularum marginalium juxtapositus. species. TYPE: ANTARCTICA, Rybiy Khvost Bay, Bunger DISTRIBUTION. Fossil species, recorded only Oasis, in Holocene sediments (1290 cm) at 91 m depth, in the Holocene from the Bunger Oasis. Antarctic 66°15′S, 100°46′E, core PG1173-2, 238–240 cm, 22 endemic. February 1994, leg. M. Melles [HOLOTYPE: Slide no. A1173-2 (238–240), Diatom Collection (LE), Depart- NOTES. In all samples observed, T. longiful- ment of Algology, Komarov Botanical Institute, Russian toportulata was relatively abundant (0.3–31.0% Academy of Sciences, St. Petersburg (Figs 6 & 7); ISO- of the diatom assemblages). From the data avail- TYPE: Slide no. B1173-2 (238–240), Diatom Collection, able, no similar species was noted in Holocene Russian Research Institute for Oceanology and Mineral deposits of other areas (e.g., Cremer et al. 2003) Resources, St. Petersburg]. or in modern plankton of the Antarctic. ETYMOLOGY: The Latin epithet is translated A peculiar morphological feature of the new ‘long marginal strutted processes’. species is its very long and beautiful marginal Valves circular, almost slightly convex, fl at on fultoportulae, 2–6 μm long. The marginal ring of margin, 32–80 μm in diameter, mantle shallow (Fig. the long fultoportulae is diffi cult to see on the 7). Areolae pattern radial, fasciculated (Fig. 6), fo- mantle in LM (Figs 3–5). Extensions of the mar- raminae circular (Figs 13 & 16). Areolae (6–)7–8 ginal fultoportulae can be seen in detailed SEM in 10 μm on valve face and 10–12 on valve mantle. micrographs nearly at any angle if they are not Fultoportulae with 4 satellite pores (Fig. 17). Central broken (Fig. 8). fultoportulae 3–7 or rarely up to 9, external tubes up Interesting data have been obtained on the mor- to 2 μm long (Figs 16 & 17). One ring of densely phology of another Thalassiosira taxon closely re- packed marginal fultoportulae, 6–9(–10) in 10 μm lated to T. australis. Based on some morphological (usually 7–8), with very long external tubes, 2–6 features a new species is proposed. (usually 3.5–5.0) μm long, 1 process for each areola (Figs 6, 7, 11 & 12). Single rimoportula with large Thalassiosira bungerensis Gogorev & Pushina, external tube, 2.7–3.5 μm long and 0.7–1.4 μm in sp. nov. Figs 18–32 diameter, usually longer than those of marginal ful- toportulae, and large internal labiate tube, up to 2.6 Thalassiosira australis sensu McMinn & Hodgson 1993, Fig. 9. μm long, located very nearly in ring of marginal fultoportulae (Figs 12 & 13). Slit of rimoportula Frustulae tympaniforme, plus minusve aequale in altitu- nearly radial (Figs 14 & 15). dini et diametro. Valvae convexae, 26–47 μm in diametro, R. M. GOGOREV ET AL.: NEW THALASSIOSIRA SPECIES FROM EAST ANTARCTICA 139 Figs 6–11. Valve of Thalassiosira longifultoportulata Gogorev & Pushina sp. nov. 6–9 – exterior (8 & 9 – well visible central fultoportulae), 10 – interior, 11 – marginal fultoportulae. All SEM. Scale bars: 6–10 = 10 μm; 11 = 1 μm. limbus valvae altus. Areolae in seriebus radialibus, 12–13 in 10 μm, earum tubuli externi breves, minus 1 μm alt. in 10 μm in faciei valvae et 11–20 in 10 μm in limbui Rimoportula singularis, in anulo processibus occlusis valvae, foraminis circularia in faciei valvae, elongata positus, rima tubuli interni radiatum versa. Processus elliptica prope centrum valvae. Fultoportulae subcen- occlusi aut desunt aut in anulo singulari, 2–3 in 10 μm, trales numerosae, in 4–8 gregibus, 9–20(28) in gregi earum tubuli externi praecelsi, 4–15 μm alt., angustati quovis. Fultoportulae marginales in anulo singulari, 3–4 et curvati ad centrum valvae. 140 POLISH BOTANICAL JOURNAL 55(1). 2010 Figs 12–17. Valvar structure of Thalassiosira longifultoportulata Gogorev & Pushina sp. nov. 12–15 – marginal fultoportulae and rimoportula (12 & 13 – exterior, 14 & 15 – interior), 16 & 17 – central fultoportulae (16 – exterior, 17 – interior, showing satellite pores). All SEM. Scale bars: 6–10 = 10 μm; 11 = 1 μm. TYPE: ANTARCTICA, Rybiy Khvost Bay, Bunger ISOTYPE: Slide no. B1173-2 (238–240), Diatom Col- Oasis, in Holocene sediments (1290 cm) at 91 m depth, lection, Russian Research Institute for Oceanology and 66°15′S, 100°46′E, core PG1173-2, 238–240 cm, 22 Mineral Resources, St. Petersburg]. February 1994, leg. M. Melles [HOLOTYPE: Slide no. Frustule drum-shaped in girdle view, with A1173-2 (238–240), Diatom Collection (LE), Depart- pervalvar axis length less than or equal to diam- ment of Algology, Komarov Botanical Institute, Russian Academy of Sciences, St. Petersburg (Figs 21 & 23); eter (Figs 18 & 21). Valves convex, 26–47 μm in R. M. GOGOREV ET AL.: NEW THALASSIOSIRA SPECIES FROM EAST ANTARCTICA 141 Figs 18–20. Thalassiosira bungerensis Gogorev & Pushina sp. nov. 18 – mantle view of frustule, 19 – valve face, 20 – frustule with putative endogenous resting spore. All LM. Scale bars: 18 & 20 = 10 μm; 19 = 5 μm. diameter, with high mantle (Fig. 21). Areolae in 1985). Some species were described in fossil ma- radial rows, 12–13 in 10 μm on valve face, 11–20 terials or surface sediments. in 10 μm on mantle, foraminae circular and elon- Comparisons are drawn between Thalassio- gated elliptical at valvar center (Figs 22 & 25). sira longifultoportulata and species having several 4–8 clusters of numerous subcentral fultoportulae, central fultoportulae (CF), one ring of marginal in cluster 9–20(–28) fultoportulae (Figs 23, 27, fulotoportulae (MF) and one rimoportula (Table 1). 30 & 31). Single ring of marginal fultoportulae, Similar characteristics are seen in 22 species; 6 of 3–4 in 10 μm, with short external extensions less them are chosen for comparison here. than 1 μm (Fig. 29). Fultoportulae with 4 satellite The new species is externally similar and seems pores. One rimortula with radially located slit of to be closely related to Thalassiosira densannula. labiate tube outside (Fig. 32). Occluded processes Many morphological features of the two species, 2–3 in 10 μm when present, arranged in single including MF density (one per areola), are very ring that includes the rimoportula, external tubes similar. The main distinctions are in the arrangement of processes very long (4–15 μm), curved to valve of areolae on the valve (linear in T. densannula and center and narrowed on their distal part (Figs 21, radial in T. longifultoportulata) and MF length. 24, 27 & 30). The new species is closely related to Thalassio- sira gerloffi i but has less dense areolae on the valve ETYMOLOGY: The new species is named after face (7–8 instead of 13–18 in 10 μm) and a more its type locality. variable areolae pattern. In general, Thalassiosira ECOLOGY. Marine, probably, cold-water neritic longifultoportulata seems similar to several taxa species. of the genus and differs in having longer strutted processes extending. DISTRIBUTION. The new species is recorded in Occluded processes are known for 13 species Holocene sediments from the Bunger Oasis and of Thalassiosira. Morphological data for some in modern phytoplankton from the Vestfold Hills. taxa are summarized in Table 2. Antarctic endemic. Thalassiosira bungerensis, like T. australis, NOTES. Thalassiosira bungerensis seems to be has a combination of characters such as clusters a spore-forming taxon. We observed frequently of subcentral fultoportulae and occluded processes resting spores inside frustules only in LM (Fig. 20), which are not recorded for any other Thalassiosira but we could not study them in SEM. species. Thalassiosira bungerensis can be distinguished DISCUSSION from T. australis by the following: (i) there are more numerous clusters of subcentral fultoportulae The genus Thalassiosira is represented at least by (4–8 rather than 2–4); (ii) the location of these 19 species in Antarctic waters (Johansen & Fryxell clusters is closer to the valve margin than the valve 142 POLISH BOTANICAL JOURNAL 55(1). 2010 Figs 21–26. Thalassiosira bungerensis Gogorev & Pushina sp. nov. 21 – valve mantle with occluded processes, 22, 23 – valve exterior, 24 – adjacent valves, 25, 26 – valve center (25 – outside, 26 – interior with velum occluding the areolae). All SEM. Scale bars: 21–24 = 10 μm; 25 & 26 = 1 μm. center, and so should be deemed submarginal; (iii) of occluded processes; (v) the distance between the number of fultoportulae per cluster is smaller occluded processes; and (vi) the probable presence (9–28 rather than 11–46); (iv) the shape and size of resting spores (Table 2). R. M. GOGOREV ET AL.: NEW THALASSIOSIRA SPECIES FROM EAST ANTARCTICA 143 Figs 27–32. Valve structure of Thalassiosira bungerensis Gogorev & Pushina sp. nov. 27 & 30 – occluded processes and cluster of ‘submarginal’ fultoportulae, 28 & 29 – marginal fultoportula(e) (28 – exterior, 29 – interior with satellite pores), 31 – cluster of ‘submarginal’ fultoportulae outside, 32 – adjacent valves showing cluster of ‘submarginal’ fultoportulae (white arrow) and rimoportula (black arrow) inside. All SEM. Scale bars = 1 μm. Thalassiosira australis has been observed 1985), southern Argentina and South Orkney in benthic habitats in the Antarctic sector of Islands (Fryxell 1977), rarely in plankton of the southwestern Atlantic (Johansen & Fryxell eastern Antarctic waters (Gogorev 2009), so 144 POLISH BOTANICAL JOURNAL 55(1). 2010 5; 0 5; 0 5; 0 5; 0 7 3; 198199 198199 198199 9; 198199 MF) and one rimoportula. e DistributionReference Bunger Oasis (East This paperAntarctica) Indian OceanHasle & Fryxell 197 SW Atlantic, Indian Fryxell & Hasle 198& SW Pacifi c sec-Johansen & Fryxell tors of AntarcticaFryxell & Johansen SW Atlantic sector Johansen & Fryxell of AntarcticaFryxell & Johansen NE Pacifi c, Indian & Pacifi c sectors of Makarova 1988Antarctica Coastal waters of Johansen & Fryxell Chile & PeruFryxell & Johansen Fryxell & Hasle 197North Atlantic, Johansen & Fryxell AntarcticFryxell & Johansen ulae ( atellitpores 4 3–4 3 ginal fultoport MF length, Sμm 2–6 1.2–1.5* 1.5 inside fl ared skirt easily visible in LM ring of mar MF in 10 μm 6–9(10) 7–8 3–7 6–9 (5–11) 2.5–3 4–6 1–2 CF), one Number of CF 3–7(9), < 2 mk 5–6 1–6 2–9 3–5 1–8 1–8 (2–5) ( e ultoportula Areolae in 10 μm (6)7–8 (10–12) 6–7 ca 30 13–18 6–7, 8–10 10–15 11–16 f eral central Areola pattern radial, asciculated linear radial asciculated adial-linear asciculated asciculated v f f r f f e cies with s Diameter, μm 32–80 27–52 15–22 16–77 21.5–53 17–40.5 14–38 e Table 1. Comparison of Thalassiosira sp Taxon Thalassiosira longifultoportulata Gog-orev & Pushina sp. nov. Thalassiosira densannula Hasle & Fryxell Thalassiosira dichotomica (Kozl.) Fryxell & Hasle Thalassiosira gerloffi i Rivera Thalassiosira latimarginata Makarova Thalassiosira poro-irregulata Hasle & Heimdal Thalassiosira poroseriata (Ramsf.) Hasle * – data obtained from illustrations only.

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