Stauroneis lacusvulcani sp. nov. (Bacillariophyceae), a new diatom from volcanic lakes in...

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Accepted by David M. Williams: 31 Oct. 2013; published: 28 Nov. 2013 47 PHYTOTAXA ISSN 1179-3155 (print edition) ISSN 1179-3163 (online edition) Copyright © 2013 Magnolia Press Phytotaxa 148 (1): 4756 (2013) www.mapress.com/ phytotaxa/ Article http://dx.doi.org/10.11646/phytotaxa.148.1.3 Stauroneis lacusvulcani sp. nov. (Bacillariophyceae), a new diatom from volcanic lakes in northeastern China PATRICK RIOUAL 1* , QIANG GAO 1 , YUMEI PENG 2 & GUOQIANG CHU 1 1 Key Laboratory of Cenozoic Geology and Environment, Institute of Geology and Geophysics, Chinese Academy of Sciences, #19 Beitucheng Xilu, P.O. box 9825, Beijing 100029, China * E-mail: [email protected] (Corresponding author). 2 State Key Laboratory of Loess and Quaternary Geology, Institute of Earth Environment, Chinese Academy of Sciences, Xi’an 710075, China *Corresponding author: E-mail: [email protected] Abstract Stauroneis lacusvulcani sp. nov. is described from two small volcanic lakes in northeastern China. The morphology of S. lacusvulcani is illustrated with light and scanning electron micrographs and discussed in comparison with several species of the Stauroneis gracilior group. S. lacusvulcani can be distinguished by the size of the valves and its long-protracted and strongly capitate apices. Diatom analysis of the sedimentary record from Lake Xiaolongwan (Jilin Province) showed that this new species was most abundant during the early Holocene (~8950 to 10,640 yrs BP). Key words: China, morphometric analysis, new species, palaeolimnology, Stauroneis Introduction In recent years, our understanding of the taxonomy and species diversity in the genus Stauroneis Ehrenberg (1843: 45) has considerably improved thanks to the work of Lange-Bertalot et al. (2003), Van de Vijver et al. (2004) and Bahls (2010). In their overview for China, Li & Qi (2010) listed 43 species and varieties of Stauroneis. In the past few years, sediment cores were retrieved from numerous volcanic lakes in northeastern China. During this research, a new species of Stauroneis was discovered and is described herein using light (LM) and scanning electron microscopy (SEM). Material and methods Samples collection and preparation In the context of an ongoing project that aims at investigating the past and present diversity and distribution of diatoms in Northeastern China (Fig. 1), surface-sediment and sediment core samples have been investigated from a suite of lakes. The samples were retrieved from the deepest point of the lakes using either an Uwitec® gravity corer or a modified piston corer (Chu et al. 2009). In parallel with the collection of sediment samples, surface water samples (~0.3 m depth) were also collected and analysed for a wide range of chemical and physical variables. The methods used to measure these various environmental variables are given in Rioual et al. (2013). The species described herein was found in 2 localities: first in the surface-sediment of Lake Xiaolongwan (October 2005) and then in the surface sediment of Sifangshan Tianchi (April 2012). We also investigated the distribution of this new species in a sediment core retrieved in 2006 from Lake Xialongwan. The age model TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Transcript of Stauroneis lacusvulcani sp. nov. (Bacillariophyceae), a new diatom from volcanic lakes in...

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PHYTOTAXA

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2013 Magnolia Press

Phytotaxa 148 (1): 47–56 (2013)

www.mapress.com/phytotaxa/Article

http://dx.doi.org/10.11646/phytotaxa.148.1.3

Stauroneis lacusvulcani sp. nov. (Bacillariophyceae), a new diatom from volcanic

lakes in northeastern China

PATRICK RIOUAL1*, QIANG GAO1, YUMEI PENG2 & GUOQIANG CHU1

1Key Laboratory of Cenozoic Geology and Environment, Institute of Geology and Geophysics, Chinese Academy of Sciences, #19

Beitucheng Xilu, P.O. box 9825, Beijing 100029, China * E-mail: [email protected] (Corresponding author).2State Key Laboratory of Loess and Quaternary Geology, Institute of Earth Environment, Chinese Academy of Sciences, Xi’an 710075,

China

*Corresponding author: E-mail: [email protected]

Abstract

Stauroneis lacusvulcani sp. nov. is described from two small volcanic lakes in northeastern China. The morphology of S.

lacusvulcani is illustrated with light and scanning electron micrographs and discussed in comparison with several species

of the Stauroneis gracilior group. S. lacusvulcani can be distinguished by the size of the valves and its long-protracted

and strongly capitate apices. Diatom analysis of the sedimentary record from Lake Xiaolongwan (Jilin Province) showed

that this new species was most abundant during the early Holocene (~8950 to 10,640 yrs BP).

Key words: China, morphometric analysis, new species, palaeolimnology, Stauroneis

Introduction

In recent years, our understanding of the taxonomy and species diversity in the genus Stauroneis Ehrenberg

(1843: 45) has considerably improved thanks to the work of Lange-Bertalot et al. (2003), Van de Vijver et al.

(2004) and Bahls (2010). In their overview for China, Li & Qi (2010) listed 43 species and varieties of

Stauroneis. In the past few years, sediment cores were retrieved from numerous volcanic lakes in northeastern

China. During this research, a new species of Stauroneis was discovered and is described herein using light

(LM) and scanning electron microscopy (SEM).

Material and methods

Samples collection and preparation

In the context of an ongoing project that aims at investigating the past and present diversity and

distribution of diatoms in Northeastern China (Fig. 1), surface-sediment and sediment core samples have been

investigated from a suite of lakes. The samples were retrieved from the deepest point of the lakes using either

an Uwitec® gravity corer or a modified piston corer (Chu et al. 2009). In parallel with the collection of

sediment samples, surface water samples (~0.3 m depth) were also collected and analysed for a wide range of

chemical and physical variables. The methods used to measure these various environmental variables are

given in Rioual et al. (2013).

The species described herein was found in 2 localities: first in the surface-sediment of Lake Xiaolongwan

(October 2005) and then in the surface sediment of Sifangshan Tianchi (April 2012). We also investigated the

distribution of this new species in a sediment core retrieved in 2006 from Lake Xialongwan. The age model

Accepted by David M. Williams: 31 Oct. 2013; published: 28 Nov. 2013 47

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established for this core combines the count of annual laminations (= varves) and 14C dating (Chu et al. 2009).

For diatom analysis, at least 500 diatom valves were enumerated to establish abundances values expressed as

relative percentages. Diatom concentration was estimated using the microsphere method of Battarbee &

Kneen (1982) and expressed as number of valves per gram dry matter (valves g dm-1). Collection data and

details on the localities of the samples investigated are presented in Table 1.

TABLE 1: Physical and chemical parameters recorded at the localities of the samples studied.

The sediment samples were prepared using the water-bath method (Renberg 1990). Sub-samples of the

cleaned, homogenized suspension were diluted with distilled water and left to settle onto glass coverslips and

dry overnight in ambient conditions. Coverslips were mounted on glass slides with Naphrax® (a commonly

used mountant with a refraction index of 1.73). To establish abundance values expressed as relative

percentages at least 500 diatom valves were enumerated.

Counting was performed with a Leica DM LB2 light microscope using bright-field and phase contrast oil

immersion optics at 1000× magnification.

Morphometric analysis

For LM analysis, measurements of valve length, width and striae density were made on digitized images

calibrated against a slide micrometer using the public domain ImageJ software (http://imagej.nih.gov/ij/). In

total, 85 valves were investigated under the LM.

For SEM analyses, aliquots of cleaned samples were air-dried onto glass coverslips of 12 mm diameter.

These were mounted onto aluminum stubs using Ted Pella® double-coated carbon conductive adhesive tape

and coated with gold using a Polaron SC7640 sputter coater for viewing on a LEO 1530 VP at the State Key

Laboratory of Paleobiology and Stratigraphy (Chinese Academy of Sciences, Nanjing). For areolae density,

the length of 5 areolae was measured on 5 striae located immediately near the stauros. These values were

converted to a number of areolae per 10 μm (Cortese & Gersonde 2007) and averaged. Only 7 valves could be

investigated under the SEM.

Lake

Parameter Sifangshan Xiaolongwan

Sample collector G.Chu P.Rioual

No water samples and dates 1; Apr–2012 8; from Jul–05 to Oct–12

Elevation (masl) 869 657

Latitude (N) 49º 22'33'' 42º 17'59''

Longitude (E) 123º 27'51' 126º 21'35''

Lake area (ha) 0.1 7.7

Max depth (m) 0.5 15

pH 5.99 6.54–6.93

Elec. Conductivity (µS/cm) 146 52–76

Alkalinity (µeq/L) 1177 364–498

TP (µg/L) 330 4–26

TN (µg/L) 5186 784–1818

DOC (mg/L) 35.7 2.4–12.3

Cl (mg/L) 5.1 1.0–2.5

NO3 (mg/L) 1.6 0.0–4.2

SO4 (mg/L) 1.9 3.1–5.3

Ca (mg/L) 22.1 5.7–9.4

K (mg/L) 5.6 1.3–2.9

Mg (mg/L) 5.4 1.5–2.2

Na (mg/L) 2.4 0.4–1.4

Dissolved Si (mg/L) 38.3 0.1–1.3

% Stauroneis lacusvulcani in surface sediment 0.7 0.2

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FIGURE 1: Geographical location of the study sites. Lake Xiaolongwan is marked by an X, Sifangshan Tianchi by a S. Shaded areas

represent mountainous regions with an elevation >1000 m.

Statistical analyses

To check if the populations of Stauroneis found at the two localities show significant morphometric

differences, single characters analyses (boxplots) and bivariate analyses (scatter plots) were performed. In

particular, the relationship between valve length and width is traditionally used to distinguish diatom taxa

(Mann et al. 2004). For single characters analyses, non-overlapping notches on boxplots were used to infer

statistically significant dissimilarity at the 95% confidence interval (Chambers et al. 1983, Paull et al. 2008).

Boxplots and scatterplots were drawn using R (http://www.r-project.org/).

Abbreviations

Light microscopy (= LM); scanning electron microscopy (= SEM). Author abbreviations follow the

International Plant Name Index (IPNI; http://www.ipni.org/); herbarium acronyms follow Index

Herbariorum (http://sciweb.nybg.org/science2/IndexHerbariorum.asp) with IGGDC = Herbarium of

Bacilliarophyceae, Institute of Geology and Geophysics, Chinese Academy of Sciences, Beijing, People's

Republic of China

New species description

Stauroneis lacusvulcani Rioual sp. nov. (Figs 2–21)

Type:—CHINA: Inner Mongolia, Great Xing’An Mountains, Sifangshan Tianchi, 869 m, 49º 22'33''N , 123º 27'51''E,

April-2008, surface sediment collected by Dr Guoqiang Chu, IGG-CAS, Beijing. (holotype slide IGGDC-DB-SIFA-

1204!, holotype specimen illustrated in Fig 7, located using England Finder K45/1); BM 101 674 (isotype slide).

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FIGURES 2–15. Light micro-photographs of Stauroneis lacusvulcani. Scale bar = 10 µm. Figs 3,4,7,8. Specimen from Sifangshan

Tianchi, surface-sediment. Figs 2, 5, 6. Specimens from Xialongwan surface-sediment. Figs 9–15. Specimens from Xiaolongwan core

sample.

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LM (Figs 2–15): Valves lanceolate with long protracted capitate apices (Figs 2–15); length 62–83 μm, width

9.2–12.0 μ m. Raphe moderately lateral, becoming filiform at distal ends, with strongly unilaterally deflected

(hooked) distal ends. Axial area narrow, linear. Shape of central area varies from narrow rectangular (Figs 2–

3, 5–6, 10–13) to asymmetrical stauros expanding towards valve margins (Figs 1, 4, 7–9, 14–15). Shortened

striae may be present at valve margins (Figs 2, 5–8, 9–12) or not (Figs 3–4, 13–15). Striae parallel along

border of stauros becoming radiate to more steeply radiate near valve apices, 21–28 in 10 µm.

SEM (Figs 16–21): Areolae round to transapically oblong, irregularly spaced, 18–28 in 10 µm. Weakly

developed hyaline area present, thus striae next to stauros appear discontinuous (Figs 17, 20). Raphe bordered

by wide sternum on each side. Proximal raphe ends straight and slightly expanded (Fig. 17), apical external

raphe endings hooked with hooks opening toward secondary side (Fig. 18). Internally, raphe-sternum bears

longitudinal ribs and well-developed helictoglossae (Fig. 21).

FIGURES 16–21. SEM of Stauroneis lacusvulcani from Sifangshan. 16. Exterior view of whole valve. 17. Close-up of valve centre.

18. Close-up of distal end. 19. Interior view of whole valve. 20. Interior view of centre showing ridge bearing raphe fissure. 21.

Interior view of distal end showing helictoglossa.

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Etymology:—Based on its occurrence in volcanic lakes.

Distribution and ecology:—Stauroneis lacusvulcani was found in the surface sediment of two volcanic

lakes of NE China, Sifangshan Tianchi and Lake Xiaolongwan. Lake Xiaolongwan located in the Long Gang

volcanic field in Jilin Province, is a small maar lake (i.e. formed by a phreatomagmatic eruption) with a

maximum water depth of 15 m (Chu et al. 2009). Sifangshan Tianchi, located in the Great Xing’An Mountains

(Inner Mongolia) is a much smaller water body as only a small pond, less than 1 m deep, persists at the center of

the crater. In both sites, the water is slightly acidic with low electrical conductivity and relatively high dissolved

organic carbon concentration (Table 1). For Sifangshan, however, the values for total phosphorus (TP), total

nitrogen (TN) and dissolved silica were measured on a water sample that was contaminated with sediment.

In both surface-sediment samples, S. lacusvulcani is rare and represents less than 1% of the assemblages.

In the sediment core from Lake Xiaolongwan, S. lacusvulcani is most abundant (14.4%) in the sample taken at

400 cm core depth from the early part of the Holocene.

Discussion

Stauroneis lacusvulcani can be usefully compared with the group of species that include Stauroneis gracilior

Reichardt (1995: 34) and Stauroneis neohyalina Lange-Bertalot & Krammer (in Lange-Bertalot & Metzeltin

1996: 104) (Table 2), the most similar being Stauroneis gracilior “Zackenberg” (a manuscript name for a taxon

described, but not named, by Van de Vijver et al. 2004: 37) from East Greenland (Zackenberg Area) and

Stauroneis pikuni Bahls (2010: 115) described from Montana (USA). Bahls (2011) suggested that these two taxa

are synonyms. S. lacusvulcani differs from both by its larger dimension (valve length is 62–83 instead of 42–62

µm for S. gracilior ‘Zackenberg’, 53–64 µm for S. pikuni), its lanceolate shape (instead of rhombic-lanceolate),

the long-protracted and strongly capitate apices, the presence of a weakly developed hyaline zone and, in most

specimens, the presence of shortened striae in the stauros. S. gracilior sensu stricto, originally described from

French Guyana, is smaller and has a more conspicuous hyaline area around the stauros. S. neohyalina is much

smaller, has more linear valves and has a higher striae density. Stauroneis sofia Van De Vijver & Le Cohu (in

Van de Vijver, Gremmen, Beyens, & Le Cohu 2004: 104), described from the sub-Antarctic region, has valves

of similar dimension to S. lacusvulcani but its apices are less protracted and is characterized by a large hyaline

area. These differences justify the description of S.lacusvulcani as a new species.

The boxplots for valve length (Fig. 22) and width (Fig. 23) indicate that the population from Sifangshan

has larger valve dimension than that from Xiaolongwan. However, there was no significant difference in striae

density (Fig. 24). Moreover, the ‘length vs width’ scatter plot shows that the two populations overlap and that

they do not occupy different size trajectories (Fig. 25). There is therefore no reason to suspect that these two

populations belong to different taxa. The difference in the distribution of lengths and widths between the two

populations may be due to a difference in the number of generations included in the samples (Tropper 1975).

The down-core sample from Xiaolongwan represents a much longer time span than the surface sediment

collected in Sifangshan and therefore may include a population than underwent more divisions and is hence

skewed towards the smaller sizes.

Most species of Stauroneis species are motile, epipelic diatoms (Round et al. 1990, Bahls 2011) and this is

probably also true for S. lacusvulcani. Currently in Lake Xiaolongwan, the development of the epipelon is

limited by a dense belt of Phragmites in the littoral zone and by rapid light attenuation under the deeper water.

Consequently, the assemblage found in the uppermost section of Lake Xiaolongwan sedimentary record is

dominated by planktonic (e.g. Discostella, Fragilaria) and epiphytic (e.g. Achnanthidium, Brachysira,

Encyonopsis, Tabellaria flocculosa) diatoms (Panizzo et al. 2013). The reduced extent of the lake area

available for the development of the epipelic community may explain why S. lacusvulcani is so rare in the

surface sediment sample we analyzed.

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FIGURES 22–24. Box plots showing quantile distributions of valves characters for populations of Stauroneis lacusvulcani observed

in the 2 localities: Sifangshan (n=35) and Xialongwan (n=7 for surface sediment and n=50 for core sample). 22. Valve length. 23.

Valve width. 24. Striae density (= number of striae in 10 µm). The 25–75 percent quartiles (excluding outliers) are drawn using a box.

The median is shown with a horizontal line inside the box. The whiskers represent the upper and lower “inner fence”, i.e. are drawn

from the edge of the box up to the largest/lowest data point less than 1.5 times the box height. Outliers, i.e. values outside the inner

fences, are shown as circles if they lie further from the edge of the box than 3 times the box height.

The diatom analysis of the sedimentary record from Lake Xiaolongwan indicates that S. lacusvulcani has

been present in that lake since at least the Late glacial with four peaks in relative abundances at ~11,450

(7.2%), between ~10,650 and ~8950 (14.4%), at ~2350 (7.4%) and ~600 (4.6%) calibrated years BP (Fig. 26).

The palaeolimnological investigation of this sedimentary record is still on-going and at this stage of our study

we cannot propose a definitive interpretation of these peaks of Stauroneis. However, it is interesting to note

that high values of S. lacusvulcani coincide with low values for diatom concentration. The peaks in

S.lacusvulcani may therefore correspond with intervals when the productivity of planktonic and epiphytic

diatoms communities decreased and when conditions became more favorable to the epipelon. The highest peak

of S. lacusvulcani occurred during an interval when the deposition of laminated sediment was interrupted by a

slump deposit (between the core depths 388 and 414 cm) clearly identified by sharp changes in water content

and dry bulk density (Fig. 26). This disturbance in sedimentation was most likely related to the volcanic

eruption of regional origin that occurred at ~10,460 cal. yrs BP (Stebich et al. 2009).

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FIGURE 25. Length vs width scatter plot for Stauroneis lacusvulcani. The valve width is significantly related to valve length (Width

= 3.916 + 0.092 × Length, r2=0.52, F1,90

=101.3, p<0.001, n=92).

FIGURE 26. Sedimentary record of Lake Xiaolongwan plotted versus core depth (primary y-axis) and calibrated years BP (secondary

y-axis). A) Simple stratigraphy showing laminated sediments interrupted by a slump deposit. B) Position of samples analyzed for

diatoms in this study. C) Water content of the sediment. D) Dry bulk density. E) Diatom concentration. F) Variations of the relative

percentages of Stauroneis lacusvulcani.

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So far, S. lacusvulcani has been found in only two lakes. However, its distribution is not restricted to a

small geographical zone as the two lakes are located more than 800 km apart and the type of lake in which it

was found is rather common in northeastern China.

Acknowledgements

Dr Yan Fang (State Key Laboratory of Paleobiology and Stratigraphy, Nanjing) is thanked for assistance with

the SEM. This research was supported by the "Strategic Priority Research Program" of the Chinese Academy

of Sciences, Climate Change: Carbon Budget and Relevant Issues (grant # XDA05120201-2) and the National

Science Foundation of China (grant # 40502018).

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