Monograph |
Corresponding author: Bing Liu ( jsulb@outlook.com ) Academic editor: Kalina Manoylov
© 2023 Bing Liu.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Liu B (2023) The diatom genus Ulnaria (Bacillariophyta) in China. PhytoKeys 228: 1-118. https://doi.org/10.3897/phytokeys.228.101080
|
This study deals with Ulnaria species found and described from two regions of China with large climate differences during the period of 2014–2022. The first region, located in the Wuling Mountains and Hunan province, has a subtropical climate and the second in Qinghai, a northwest province of China, has a highland continental climate which is characterized by a cold and long winter and warm, short summer. Previously there were nine new Ulnaria species published from the first region. This study describes 14 additional new Ulnaria taxa, nine of which were found in the first region and five of which were found in the second region. A key to the Ulnaria species that have been described from China is provided. The main morphological characteristics for 63 Ulnaria taxa are summarized in Appendices which allow the division of these Ulnaria taxa into three groups: the seven members of group one all possess both uniseriate striae and valve marginal spines, the 42 members of group two all possess uniseriate or mostly uniseriate striae but without the valve marginal spines, and the 14 members of group three all possess mostly biseriate striae and without valve marginal spines. To summarize the morphological characters of the published Ulnaria taxa and 14 taxa described in this study several conclusions for the characterization of Ulnaria are drawn: 1) each cell has two valve-appressed, long plate-like plastids; 2) living cells of many Ulnaria species often lie in girdle view on a slide because they have deep mantles and some copulae associated with either the epivalve or the hypovalve so that the cell depth is often larger than the valve width; 3) the basic structures forming a valve include sternum, virgae, and vimines/viminules; 4) the valvocopula is a closed hoop which has a similar ultrastructure in all Ulnaria taxa but differs from the other copulae in structure; 5) the configuration of girdle bands is a common condition; 6) the life history of Ulnaria can be divided into the four series of successive stages: auxospore, initial cell, pre-normal vegetative cell, and normal vegetative cell, which is very similar to the life history of Hannaea inaequidentata (Lagerstedt) Genkal and Kharitonov; 7) the closed valvocopula is proposed as a definition character for the genus Ulnaria because demonstrating all girdle bands closed is impracticable.
Girdle band structures, life history stages, striation patterns, Ulnaria, valvocopula
Historically, the generic name “Synedra” has been used in two different senses by diatom researchers from around the world. This situation continued until
The history of diatom studies in China from 1848 through 2019 has recently been documented in detail by
No. | Taxon | Reference |
---|---|---|
1 | Synedra ulna var. intermedia Mereschkowsky |
|
2 | Synedra ulna f. curta Skvortzov |
|
3 | Synedra ulna var. mongolica Skvortzov |
|
4 | Synedra affinis var. sinica Skvortzov |
|
5 | Synedra amphicephala var. asiatica Skvortzov |
|
6 | Synedra licenti Skvortzov |
|
7 | Synedra rumpens var. sinica Skvortzov |
|
8 | Synedra tenera var. sinica Skvortzov |
|
9 | Synedra ulna f. constricta Skvortzov |
|
10 | Synedra ulna var. lanceolata f. constricta Skvortzov |
|
11 | Synedra ulna var. tenuirostris Skvortzov |
|
12 | Synedra vaucheriae var. capitata Skvortzov |
|
13 | Synedra ulna var. anhuiensis Yang |
|
14 | Synedra mazamaensis var. changbaiensis Bao and Reimer |
|
15 | Synedra cyclophoroides S.C. Liu |
|
16 | Synedra ulna var. repanda Q.X. Wang & Q.M. You |
|
Except for the above new Synedra taxa described in China, many known Synedra taxa were also reported in China. Fifty-one Synedra taxa which were illustrated and had some descriptions are listed in Table
No. | Taxon | Reference |
---|---|---|
1 | S. acus Kützing |
|
2 | S. acus var. angustissima Grunow |
|
3 | S. acus var. radians (Kützing) Hustedt |
|
4 | S. affinis Kützing |
|
5 | S. amphicephala Kützing |
|
6 | S. amphicephala var. austriaca (Grunow) Hustedt |
|
7 | S. amphicephala var. intermedia Cleve-Euler |
|
8 | S. berolinensis Lemmermann |
|
9 | S. capitata Ehrenberg |
|
10 | S. dorsiventralis Otto Müller |
|
11 | S. familica Kützing |
|
12 | S. fasciculata (C. Agardh) Kützing |
|
13 | S. gaillonii (Bory) Ehrennerg |
|
14 | S. goulardii Brebisson |
|
15 | S. investiens W. Smith |
|
16 | S. mazamaensis Soverergn |
|
17 | S. minuscula Grunow |
|
18 | S. montana Krasske |
|
19 | S. nana Meister |
|
20 | S. parasitica (W. Smith) Hustedt |
|
21 | S. parasitica var. subconstricta (Grunow) Hustedt |
|
22 | S. pulchella (Ralfs ex Kützing) Kützing |
|
23 | S. robusta Ralfs |
|
24 | S. rumpens Kützing |
|
25 | S. rumpens var. familiaris (Kützing) Hustedt |
|
26 | S. rumpens var. meneghiniana Grunow |
|
27 | S. rumpens var. scotica Grunow |
|
28 | S. rumpens var. sinica Skvortzov |
|
29 | S. socia Wallace |
|
30 | S. tabulata (Agardh) Kützing |
|
31 | S. tabulata var. fasciculata (C.A. Agardh) Hustedt |
|
32 | S. tabulata var. parava (Kützing) Hustedt |
|
33 | S. tabulata var. obtusa (Arnott) Hustedt |
|
34 | S. tabulata var. rostrata (Juhlin-Dannfelt) Cleve-Euler |
|
35 | S. tabulata var. genuina Cleve-Euler |
|
36 | S. tenera W. Smith |
|
37 | S. ulna (Nitzsch) Ehrenberg |
|
38 | S. ulna var. aequalis (Kützing) Brun |
|
39 | S. ulna var. amphirhynchus (Ehrenberg) Grunow |
|
40 | S. ulna var. biceps (Kützing) Schönfeldt |
|
41 | S. ulna var. chaseana Thomas |
|
42 | S. ulna var. constracta Østrup |
|
43 | S. ulna var. danica (Kützing) Grunow |
|
44 | S. ulna var. danica f. continua (Kützing) Grunow |
|
45 | S. ulna var. impressa Hustedt |
|
46 | S. ulna var. oxyrhynchus (Kützing) Van Heurck |
|
47 | S. ulna var. oxyrhynchus f. constricta Skvortzow |
|
48 | S. ulna var. ramesi (Héribaud) Hustedt |
|
49 | S. ulna var. spathulifera (Grunow) Van Heurck |
|
50 | S. ulna var. splendcns (Kützing) Grunow |
|
51 | S. vaucheriae (Kütz.) Kützing |
|
From 2017 to 2020, nine new Ulnaria species found in China were published (
No. | Taxon | Reference |
---|---|---|
1 | U. sinensis Bing Liu et D.M. Williams |
|
2 | U. ulnabiseriata D.M. Williams et Bing Liu |
|
3 | U. gaowangjiensis Bing Liu et D.M. Williams |
|
4 | U. rhombus D.M. Williams |
|
5 | U. wulingensis Bing Liu |
|
6 | U. oxybiseriata D.M. Williams & Bing Liu |
|
7 | U. jinbianensis S. Blanco & Bing Liu |
|
8 | U. dongtingensis Bing Liu |
|
9 | U. hunanensis Bing Liu |
|
Although Ulnaria was established in 2001, the use of the name Ulnaria is currently a nomenclatural decision (
Appendices
The diatom samples studied herein were collected from the Wuling Mountains, Dongting Lake and Qinghai province beginning in 2014. The Wuling Mountains, which stretch across Chongqing, Hunan, Hubei, and Guizhou provinces, are one of the ten biodiversity hotspot ecoregions considered as conservation priorities in China (
The collected diatom samples which were not added 70% alcohol were used to observe the living cells. 100 μl diatom samples were transferred into a round chamber (diameter 14 mm, depth 0.35 mm) located in the middle of a custom-made slide by using a pipette, then examined using a Leica DM3000 light microscopy (LM) equipped with a Leica MC190 HD digital camera. The collected diatom samples which were added 70% alcohol were processed (cleaned) for microscopic examination with 10% HCl and 30% H2O2. Permanent slides were prepared using Naphrax mountant and examined using the same light microscopy as above. Slides are deposited in the
Herbarium of Jishou University, Hunan, People’s Republic of China (
Morphological terminology follows
Valvocopula abvalvar edge: the edge that is farther from the valve face.
Valvocopula advalvar edge: the edge that is closer to the valve face.
Closing plate: a plate that does not completely occlude the outside opening of areola. This plate is usually solid (sometimes with several perforations) and has a few struts attaching it to each areolar wall.
Configuration of girdle bands: the ratio between the number of girdle bands associated with the epivalve and the number associated with the hypovalve in a diatom cell.
Ghost striae: the faint “striae” composed of unperforated grooves in the central area of the valve interior (visible under SEM). These grooves look like “striae” under LM but are not true striae.
Mixed stria: the striae are usually divided into three types: uniseriate, biseriate and multiseriate striae. A mixed stria is a stria composed of at least two of the previous three stria types.
Pre-normal vegetative frustule: the pre-normal vegetative period is the time between immediately after the initial cell’s first division and the presence of the first new normal vegetative cells. The frustule occurring during this period is termed pre-normal vegetative frustule.
Vimines (s. vimen): the cross-connecting tiny ribs between two adjacent virgae defining areolae in uniseriate striae.
Viminules (s. viminule): the interconnecting tiny ribs between two adjacent virgae which define areolae in the biseriate or multiseriate striae.
Virga (pl. virgae): the transverse silica rib between two adjacent striae.
(Figs
The life history of Ulnaria can be divided into the following four series of successive stages: auxospore, initial cell, pre-normal vegetative cell, and normal vegetative cell. Unfortunately, we did not find an auxospore of Ulnaria, but the initial cell, pre-normal vegetative cell, and normal vegetative cell of U. ulnabiseriata were all documented using LM and SEM. The life history, from the initial cell, via pre-normal vegetative cells, to normal vegetative cell, is a process from chaos to order, i.e., the shapes of initial cell and pre-normal vegetative cells gradually become more and more regular and symmetrical from irregular and asymmetrical (see Figs
Life history of Ulnaria ulnabiseriata, from initial cell, via pre-normal cells, finally to normal cell, ×400, LM A initial frustule showing its cylindrical and twisted outline, central area flanked by striae on one side B–N pre-normal frustules, note twisted frustules with laterally located sterna (B–E), one apex twisted and with almost centrally located sterna (F, G), almost centrally located sterna and complete central areas (H–N). O Normal valve.
Life history of Ulnaria ulnabiseriata, from initial cell, via pre-normal cells, finally to normal cell, external view, SEM A initial frustule, note its cylindrical and twisted frustule (for details see Fig.
Initial cells: Two initial cells were found and measured. One is 240 μm long (Fig.
Initial frustule of Ulnaria ulnabiseriata, external view, SEM A Initial frustule, note its arcuate, cylindrical, and twisted outline B, C details of middle part from A, note small central area flanked by striae on one side (C, arrow), mixed striae, and underdeveloped sternum (i.e., striae continue across valve surface) D apical detail from A, note mantle not well differentiated from valve face (double-headed arrow), and virgae and vimines/viminules occurring nearly on the same plane E, F other apical details from A, note laterally located sternum present only near one apex (F, double-headed arrow), ocellulimbus flush with and extending onto valve surface, and its partial pervalvar rows of porelli not perpendicular to the valve plane (F, arrow). Scale bars: 20 μm (A); 2 μm (B–F).
Pre-normal vegetative cells: The valve length range of pre-normal vegetative cells is 196–250 μm (n = 41). Early pre-normal vegetative frustules may be twisted and have laterally located sterna (Fig.
Pre-normal frustule of Ulnaria ulnabiseriata, external view, SEM A pre-normal frustule, note its cylindrical and twisted outline B, C middle part details from A, note small central area flanked by striae on one side (B, two arrows), girdle bands lacking D, E apical details from A, note laterally located sternum (E, double-headed arrow) and flush ocellulimbus F other half from A, note laterally located sternum (double-headed arrow). Scale bars: 20 μm (A); 4 μm (B–F).
Pre-normal frustule of Ulnaria ulnabiseriata, external view, SEM A pre-normal valve B middle part detail from A, note its constricted outline in the middle C, D proximal parts of valve from A, note closing plates underdeveloped E, F apical details from A, note slightly inset ocellulimbus. Scale bars: 20 μm (A); 3 μm (B–F).
Details of late pre-normal frustule of Ulnaria ulnabiseriata from Fig.
Details of late pre-normal valve of Ulnaria ulnabiseriata from Fig.
Normal vegetative cells: The valves of U. ulnabiseriata (Figs
Normal valve of Ulnaria ulnabiseriata, external view, SEM A complete valve B middle detail, note the rectangular central area C, D two apices, note centrally located sternum and rostrate apices E, F apical details, note inset ocellulimbus and some outgrowths protruding over it (F, three arrows). Scale bars: 20 μm (A); 3 μm (B, C, E); 1 μm (D, F).
The morphological features that change during the life circle of U. ulnabiseriata are summarized in Table
Features of initial cell, pre-normal and normal vegetative frustule/valve in U. ulnabiseriata.
Feature | Initial frustule/valve | Pre-normal vegetative frustule/valve | Normal vegetative frustule/valve |
---|---|---|---|
Girdle bands | Not found | Not found to a few present | 4 copulae associated with epivalve |
Valve outline | Cylinder-like, twisted | Irregular and asymmetrical | Linear-lanceolate |
Valve apex | Rounded | Rounded, rostrate, or sub-capitate | Rostrate to sub-capitate |
Sternum | Non-existent except lateral sternum present only at apex | Lateral to nearly central sternum | Central sternum, i.e., normal, situated on the midline of valve |
Central area | Present, small, with short striae at one side | From small to rectangular | Rectangular |
Virga/viminule | Virgae and vimines/viminules almost flush with each other | Vimines/viminules slightly lower than virgae | Virgae raised, viminules sunken |
Stria | Mixted striae in the middle, uniseriate striae near apex | Gradually become mostly biseriate striae (striae near apex become biseriate too) | Mostly biseriate striae |
Closing plate | Not found | Not found to present | Well developed |
Rimoportula number per valve | Two | Two | Two |
Ocellulimbus | Extending on valve face, pervalvar row of porelli not perpendicular to the valve plane | Pervalvar rows of porelli gradually becoming perpendicular to the valve plane | Pervalvar rows of porelli all perpendicular to the valve plane |
1 | Biseriate striae | 2 |
– | Uniseriate striae | 9 |
2 | Panduriform valve outline | 1. U. pandurata-biseriata |
– | Not panduriform valve outline | 3 |
3 | Apiculate apices | 2. U. oxybiseriata |
– | Not apiculate apices | 4 |
4 | Linear valve outline | 3. U. wuling-biseriata |
– | Linear-lanceolate or lanceolate valve outline | 5 |
5 | Linear-lanceolate valve outline | 6 |
– | Lanceolate valve outline | 7 |
6 | Constricted central valve margins with rostrate apex | 4. U. constricta-biseriata |
– | Parallel central valve margins with rostrate apices | 5. U. gaowangjiensis |
7 | Slightly constricted central valve margins with capitate to sub-capitate apices | 6. U. sangzhi-biseriata |
– | Parallel central valve margins | 8 |
8 | Central area apically rectangular | 7. U. ulnabiseriata |
– | Central area very variable | 8. U. jishou-biseriata |
9 | Interlocking spines produced on valve margins | 9. U. sinensis |
– | Interlocking spines lacking | 10 |
10 | Undulate valve outline | 10. U. repanda |
– | Not undulate valve outline | 11 |
11 | Panduriform valve outline | 11. U. pandurata-uniseriata |
– | Not panduriform valve outline | 12 |
12 | Parallel distal regions of valve present | 13 |
– | Parallel distal regions of valve lacking | 15 |
13 | Rhombic valve outline | 12. U. rhombus |
– | Not rhombic valve outline | 14 |
14 | Parallel distal region of valve more than 20 μm long | 13. U. wulingensis |
– | Parallel distal region of valve less than 20 μm long | 14. U. hunanensis |
15 | Central area completely lacking | 16 |
– | Central area present | 18 |
16 | Valve width less than 5 μm | 15. U. qinghainensis |
– | Valve width more than 5 μm | 17 |
17 | Lanceolate valve outline | 16. U. hupingensis |
– | Linear-lanceolate valve outline | 17. U. xieriverensis |
18 | Central area complete | 19 |
– | Central area very variable | 21 |
19 | Linear-lanceolate valve outline | 18. U. jinbianensis |
– | Lanceolate valve outline | 20 |
20 | Length of parallel central margins equal to the length of the central area | 19. U. dongtingensis |
– | Length of parallel central margins much larger than the length of the central area | 20. U. fanjingensis |
21 | Capitate apices | 21. U. neobiceps |
– | Not capitate apices | 22 |
22 | Linear-lanceolate valve outline | 22. U. chengduoensis |
– | Lanceolate valve outline | 23 |
23 | Rostrate apices | 23. U. menyuanensis |
– | Sub-capitate apices | 24. U. blancoi |
Slide JIUDIA202301, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103807
China. Guizhou province: Fanjing Mountain, Temple Longquan, Heiwan River (27°51'36"N, 108°45'51"E, 532 m a.s.l.), collected by Bing Liu, December 31, 2015.
LM
(Fig.
SEM
(Figs
Ulnaria constricta-biseriata sp. nov., girdle view, SEM A a frustule with a collapsed hypovalve B–D details from A, note three copulae (labeled B1 to B3) associated with the epivalves and two rows of poroids present on B2 (wavy arrows) and B3 (arrows) E, F the apices with three copulae visible (labeled B1 to B3) associated with the epivalves and two rows of poroids produced on B2 (wavy arrow) and B3 (arrow). Scale bars: 20 μm (A); 3 μm (B–F).
Ulnaria constricta-biseriata sp. nov., internal view, SEM A a complete valve with a valvocopula B middle detail from A, note serrated projections at the advalvar edge (two arrows) C, D two apical details from A, note the closed nature of valvocopula and serrated projections at the advalvar edge (two arrows respectively). Scale bars: 20 μm (A); 4 μm (B–D).
Ulnaria constricta-biseriata sp. nov., external view, SEM A a complete valve B middle part from A showing the constricted middle margins and central area forming a rectangular hyaline region C, D two apical details from A, note a few serrated apical projections protruding over the ocellulimbus (two black arrows) E another middle part, note two constricted middle margins F another apex, note a few serrated apical projections protruding over the ocellulimbus (arrow) and the closing plates (wavy arrow). Scale bars: 20 μm (A); 2 μm (B–F).
Ulnaria constricta-biseriata sp. nov., internal view, SEM A a complete valve B middle part from A, note two constricted middle margins and some ghost striae C, D two apical details from A, note the striae mostly biseriate and two helictoglossae E another complete valve F middle detail from E, note two constricted middle margins and some ghost striae. Scale bars: 20 μm (A, E); 2 μm (B–D, F).
The specific epithet is formed from two terms: constrict and biseriate, reflecting the constricted valve central margins and the mostly biseriate striae of the valve.
The sampling site is close to the headwaters of the Heiwan River, which originates in the Fanjing Mountain National Nature Reserve. The diatom samples were scraped off of the stone surfaces. The following environmental parameters were measured in the field. Conductivity was 49.7 ± 0.2 μS∙cm-1, pH was 7.7 ± 0.1 and water temperature was 9.4 ± 0.1 °C. So far, its distribution is known only from the type locality. To sum up, U. constricta-biseriata lives on the stone surfaces of the headwaters of a mountainous river.
Ulnaria constricta-biseriata is characterized by its linear-lanceolate valve outline, constricted valve central margins, and mostly biseriate striae. Ulnaria contracta (Østrup) E.A. Morales & M.L. Vis has also constricted valve central margins, but it differs from U. constricta-biseriata by its lanceolate valve outlines and mostly uniseriate striae (see
Slide JIUDIA202302, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103808.
China. Hunan province: Jishou City, Lianaiqiao, Donghe River (28°18'51.3"N, 109°43'41.6"E, 200 m a.s.l.), collected by Bing Liu, December 9, 2016.
LM
(Figs
Ulnaria jishou-biseriata sp. nov., ×400, LM A A cell in girdle view, note two long valve-appressed chloroplasts per cell B a dividing cell in girdle view, note the chloroplasts are distributed along the pervalvar axis C, D two cells in valve view, note the long plate of chloroplast. Scale bar: 20 μm.
SEM
(Figs
Ulnaria jishou-biseriata sp. nov., girdle view, SEM A a dividing cell in girdle view B middle detail from A, note the mantles are hyaline in the central area C, D two apical details from A, showing both epivalve and hypovalve with associated three girdle bands (labelled B1 to B3) E, F two other apical details also show both epivalve and hypovalve with associated three girdle bands in a dividing cell (labelled B1 to B3). Scale bars: 20 μm (A); 2 μm (B–F).
Ulnaria jishou-biseriata sp. nov., internal view, SEM A a valve with valvocopula B, C middle details from A, note serrated projections over each virga (C, arrows) D, E two apical details from A, note unornamented valvocopula at both apices (arrow respectively). Scale bars: 20 μm (A); 2 μm (B, D, E); 1 μm (C).
Ulnaria jishou-biseriata sp. nov., external view, SEM A a frustule in valve view B middle detail from A, showing central area flanked by a few marginal striae only on one side C, D two apical details from A, note a few serrated projections protruding over the ocellulimbus (C, three arrows) E a middle part showing the central area flanked by shortened striae on both sides F a middle part showing a rectangular central area. Scale bars: 20 μm (A); 2 μm (B, E, F); 1 μm (C, D).
Ulnaria jishou-biseriata sp. nov., internal view, SEM A a complete valve B middle detail from A, note the rectangular central area C apical detail from A, note two rimoportulae present D other apical detail from A, note only one rimoportula present E, F two middle parts, note the variable central areas. Scale bars: 20 μm (A); 2 μm (B–F).
The epithet jishou-biseriata is formed from the city name Jishou and the term biseriate to reflect its type locality (Jishou) and its mostly biseriate striae.
The sampling site is close to Jishou City and many anthropogenic influences affect the environment and hence the diatoms. The diatom samples were scraped off of the stone surfaces. The following environmental parameters were measured in the field: Conductivity was 202.3 ± 1.2 μS∙cm-1, pH was 8.5 ± 0.1, and water temperature was 13.2 ± 0.3 °C. So far, its distribution is known only from the type locality. To sum up, U. jishou-biseriata lives on the stone surfaces of a mountainous river running through a small city.
Ulnaria jishou-biseriata is characterized by its lanceolate valve outline, mostly biseriate striae, and very variable central areas. With respect to the valve outline, it is similar in some ways to U. ulnabiseriata, but the former has smaller and very variable central areas whereas the latter’s central areas are always a rectangular fascia (see
Slide JIUDIA202303, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103809
China. Hunan province: Zhangjiajie National Forestry Park, Jinbian stream, at Shuirao Simen (29°20'36"N, 110°28'13"E, 467 m a.s.l.), collected by Bing Liu, December 29, 2015.
LM
(Fig.
Ulnaria pandurata-biseriata sp. nov., external view, SEM A a frustule in valve view B middle part detail from A, showing the central area with a few shortened marginal striae on one side C, D two apical details from A, note some partially triseriate striae (arrows) E another middle part showing the fascia-shaped central area F another apex, note a few serrated projections protruding over ocellulimbus. Scale bars: 10 μm (A); 2 μm (B–F).
Ulnaria pandurata-biseriata sp. nov., internal view, SEM A a valve with a valvocopula B middle detail from A, note the unornamented middle portion of valvocopula (arrow) C, D apical details from A, note serrated projections over each virga (arrows) and valvocopula without ornamentation at each apex. Scale bars: 10 μm (A); 2 μm (B–D).
SEM
(Figs
The epithet pandurata-biseriata is a combination of the terms pandurate and biseriate to reflect the valvar panduriform outline of the valve and its mostly biseriate striae.
The sampling site is situated close to the headwaters of Jinbian stream, which originates in the Zhangjiajie National Forestry Park. The diatom samples were scraped off of stone surfaces. The following environmental parameters were measured in the field: Conductivity was 102.7 ± 0.8 μS∙cm-1, pH was 8.5 ± 0.7 and water temperature was 8.7 ± 0.2 °C. So far, its distribution is known only from the type locality. To sum up, U. pandurata-biseriata lives on the stone surfaces of the headwaters of a mountainous stream.
Ulnaria pandurata-biseriata is characterized by its panduriform valve outline, mostly biseriate striae, variable central areas, and smaller valves. Similar taxa include U. goulardii D.M. Williams, Potapova & C.E. Wetzel (see
Slide JIUDIA202304, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103810.
China. Hunan province: Sangzhi County, Wudaoshui Town, Jinlong power station, Li River (29°43′7.1″N, 109°54′50.9″E, 398 m a.s.l.), collected by Bing Liu, September 30, 2015.
LM
(Fig.
SEM
(Figs
Ulnaria sangzhi-biseriata sp. nov., girdle view, SEM A a partly collapsed frustule B middle part from A, note variable central mantles and serrated projections of valvocopula (three arrowheads) C, D two apical details from A, showing four girdle bands associating the epivalve (labelled B1–B4). Scale bars: 10 μm (A); 2 μm (B–D).
Ulnaria sangzhi-biseriata sp. nov., internal view, SEM A a valve with a valvocopula B middle part from A, note the serrated projections (two arrowheads) C, D two apical details from A, note the closed nature of valvocopula and serrated projections (arrowheads) E another valve with a valvocopula F apical detail from E, note the serrated projections (two arrowheads). Scale bars: 10 μm (A, E); 2 μm (B–D, F).
Ulnaria sangzhi-biseriata sp. nov., external view, SEM A a complete valve B middle detail from A, note rectangular central area C, D two apical details from A, note a few serrated projections protruding over the ocellulimbus (arrows) E another middle part showing rectangular central area with ghost striae F another apical detail, note a few serrated projections (two arrows) and the closing plates (two curved arrows). Scale bars: 10 μm (A); 2 μm (B, E); 1 μm (C, D, F).
The epithet sangzhi-biseriata is a combination of Sangzi and the term biseriate to reflect its type locality (Sangzi) and its mostly biseriate striae.
The sampling site is near Wudaoshui Town, and some human activities may have affected the environment and hence the diatoms. The diatom samples were scraped off of the stone surfaces. The following environmental parameters were measured in the field: Conductivity was 219.3 ± 1.2 μS∙cm-1, pH was 8.5 ± 0.2 and water temperature was 17.9 ± 0.3 °C. To sum up, U. sangzhi-biseriata lives on the stone surfaces of a mountainous river flowing away from the Town. So far, its distribution is known only from the type locality.
Ulnaria sangzhi-biseriata is characterized by its linear-lanceolate valve outline, mostly biseriate striae, rectangular or square central area, and capitate to sub-capitate apices. It is similar to U. goulardii which has a more constricted valve central margin. Moreover, the former has capitate to sub-capitate apices whereas U. goulardii has rostrate apices (see
Slide JIUDIA202305, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103811
China. Hunan province: Zhangjiajie National Forestry Park, Jinbian stream, at Shuirao Simen (29°20'36"N, 110°28'13"E, 467 m a.s.l.), collected by Bing Liu, December 29, 2015.
LM
(Fig.
Ulnaria wuling-biseriata, sp. nov., external view, SEM A a complete valve B middle detail from A, showing the rectangular central area C, D two apical details of A, note some uniseriate striae at the apices E another complete valve F middle detail from E, showing a trapezoidal central area. Scale bars: 20 μm (A, E); 2 μm (B–D, F).
Ulnaria wuling-biseriata, sp. nov., internal view, SEM A a complete valve B middle detail from A, showing the rectangular central area C, D two apical details from A. E Another complete valve F middle detail from E showing the trapezoidal central area. Scale bars: 20 μm (A, E); 2 μm (B–D, F).
SEM
(Figs
Ulnaria wuling-biseriata, sp. nov., internal view, SEM A a valve with a valvocopula B middle detail from A. C, D two apical details of A, note the valvocopula unornamented at each apex E, F details of the valvocopula, note its closed nature and the shelf-like projection (F, arrow). Scale bars: 20 μm (A); 2 μm (B–F).
The epithet "wuling-biseriata" is a combination of Wuling and the term biseriate to reflect its type locality (Wuling Mountains) and its mostly biseriate striae.
Ulnaria wuling-biseriata was commonly found in the surface sediment collected in Jinbian stream with U. pandurata-biseriata and U. ulnabiseriata. Thus, U. wuling-biseriata lives on the stone surfaces of the headwaters of a mountainous stream. So far, its distribution is known only from the type locality.
Ulnaria wuling-biseriata is characterized by its linear valve outline, mostly biseriate striae, rectangular or trapezoid central area, and rostrate to sub-capitate apices. The apices of Ulnaria ulna var. spathulifera Aboal sometimes have an inflation before tapering to form subrostrate to rostrate ends whereas U. wuling-biseriata does not have this inflation, i.e., its apex is not spatulate. Moreover, the valves of U. ulna var. spathulifera are wider than the ones of U. wuling-biseriata (8–9 μm vs 6.5–8 μm) and the former has lower stria density than the latter (9–10 in 10 μm vs 10–11 in 10 μm) (see
Slide JIUDIA202306, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103812.
China. Qinghai province: Menyuan County, an unnamed river, at a sampling location named Kengtan (37°27'28"N, 101°23'15"E, 2940 m a.s.l.), collected by Bing Liu, July 18, 2019.
LM
(Fig.
Ulnaria blancoi sp. nov., external view, SEM A, C two complete valves, note central area present or absent B middle part from A, note central area completely absent D middle part from C, note small central area flanked by marginal short striae E, F two apical details, note two horn-like projections protruding over the ocellulimbus (E, two arrows). Scale bars: 20 μm (A, C); 2 μm (B, D, F); 1 μm (E).
SEM
(Figs
Named after the Spanish diatomist, Dr. Saúl Blanco.
The sampling site is located in the plateau which belongs to the highland continental climate zone. The diatom samples were scraped off of the stone surfaces. The following environmental parameters were measured in the field: Conductivity was 448.3 ± 0.5 μS∙cm-1, pH was 8.3 ± 0.1 and water temperature was 11.9 ± 0.5 °C. To sum up, U. blancoi lives on the stone surfaces of a plateau river. So far, its distribution is known only from the type locality.
Ulnaria blancoi is characterized by its lanceolate valve outline, uniseriate striae, very variable central areas, and rostrate to sub-capitate apices. It differs from U. vitrea (Kützing) E. Reichardt by its much longer valves (104–236 μm vs 90–120 μm) and its often-present central area whereas U. vitrea often lacks central area (see
Slide JIUDIA202307, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103813
China. Qinghai province: Menyuan County, an unnamed river, at a sampling location named Kengtan (37°27'28"N, 101°23'15"E, 2940 m a.s.l.), collected by Bing Liu, July 18, 2019.
LM
(Fig.
Ulnaria menyuanensis sp. nov., external view, SEM A a complete valve B middle part detail from A, note the shallow grooves and short striae at the central region C, D two apical details from A. E, F Other middle parts, note the shallow grooves and short striae. Scale bars: 10 μm (A); 2 μm (B–F).
SEM
(Figs
Ulnaria menyuanensis sp. nov., internal view, SEM A a complete valve B middle part from A, note shallow grooves and short striae C, D two apical details from A, note the bilabiate rimoportulae and apical pore fields E, F other middle parts, note shallow grooves and short striae. Scale bars: 10 μm (A); 2 μm (B–F).
Ulnaria menyuanensis sp. nov., internal view, SEM A a valve with associated valvocopula B middle part detail from A, note serrated projections (three arrows) C, D apical details from A, note serrated projections (C, three arrows) and closing plates (D, two arrows) E, F details from other apex from A, note serrated projections (E, three arrows) and closing plates (F, two arrows). Scale bars: 10 μm (A); 5 μm (B, C, E); 2 μm (D, F).
Named after Menyuan County, where the species was found.
Ulnaria menyuanensis was commonly found in the surface sediment collected in Kengtan with U. blancoi. Thus, U. menyuanensis lives on the stone surfaces of a plateau river. So far, its distribution is known only from the type locality.
Ulnaria menyuanensis is characterized by its fusiform valve outline, the presence of distinct ghost striae, and cuneate to rostrate apices. It differs from U. ramesii (Héribaud) T. Ohtsuka by the latter’s linear-lanceolate valve outline (see
Slide JIUDIA202308, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103814.
China. Qinghai province: Menyuan County, an unnamed river, at a sampling location named Kengtan (37°27'28"N, 101°23'15"E, 2940 m a.s.l.), collected by Bing Liu, July 18, 2019.
LM
(Fig.
Ulnaria neobiceps sp. nov., internal view, SEM A a complete valve B middle part detail from A, note the very small central area (arrow) C, E two poles from A, note that valve margins do not constrict near the capitate apices (two arrows, respectively) D, F two apical details, note the capitate apices and apical pore field. Scale bars: 20 μm (A); 2 μm (B–F).
SEM
(Figs
Ulnaria neobiceps sp. nov., valvocopula, SEM A a complete valvocopula B–D details from A, showing row of poroids interrupted in the middle and shelf-like projection at each apex (C, D, two arrows respectively) E, F other two apical details of valvocopula have the same apical structure. Scale bars: 20 μm (A); 3 μm (B); 2 μm (C–F).
The epithet biceps has been occupied by Ulnaria biceps (Kützing) Compère, so here the epithet neobiceps is used, which is a combination of neo (new) and biceps (two capitate) reflecting its two distinctly capitate poles.
Ulnaria neobiceps, U. blancoi, and U. menyuanensis were commonly found in the same sampling site of Kengtan. Thus, U. neobiceps lives on the stone surfaces of a plateau river. So far, its distribution is known only from the type locality.
Ulnaria neobiceps is characterized by its linear-lanceolate valve outline, variable central area, distinctly capitate apices, and long and slender valve. Both U. neobiceps and U. capitata (Ehrenberg) Compère have capitate apices, but the apices of U. capitata are rhomboid-capitate (see
Slide JIUDIA202309, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103815.
China. Qinghai province: Chengduo County, Baima River, at a sampling location (33°22'21"N, 97°0'18"E, 3690 m a.s.l.), collected by Bing Liu, July 22, 2019.
LM
(Fig.
SEM
(Figs
Ulnaria chengduoensis sp. nov., external view, SEM A a broken valve B middle part detail from A, note the central area flanked by marginal short striae C, D two apical details from A, note the rostrate apices E a complete valve F middle part detail from E, note central area absent. Scale bars: 10 μm (A, E); 2 μm (B–D, F).
Named after Chengduo County, where this species was found.
Epilithic in a plateau river. The following environmental parameters were measured in the field: Conductivity was 422.7 ± 1.3 μS∙cm-1, pH was 8.2 ± 0.1 and water temperature was 12.2 ± 0.5 °C. So far, its distribution is known only from the type locality.
Ulnaria chengduoensis is characterized by its linear valve outline, variable central area, narrow rostrate valve apices. It differs from U. ramerii and U. verhaegeniana by its variable central area, i.e., it may completely lack a central area whereas U. ramerii always possesses an ellipsoid to rectangular central area (see
Slide JIUDIA202310, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103816.
China. Qinghai province: Chengduo County, Baima River, at a sampling location (33°22'21"N, 97°0'18"E, 3690 m a.s.l.), collected by Bing Liu, July 22, 2019.
LM
(Fig.
Ulnaria qinghainensis sp. nov., external view, SEM A a valve with valvocopula B detail of middle part from A, note absent central area C–F two poles from A, note two horn-like projections (D, two arrows) and row of poroids terminating before each apex of valvocopula (E, F, wavy arrow respectively). Scale bars: 20 μm (A); 4 μm (B–F).
SEM
(Figs
Named after Qinghai province, where this species was found.
Epilithic in a plateau river. Ulnaria qinghainensis and U. chengduoensis were commonly found in the same sampling site. The environmental parameters were measured in the field and the results are given in the description of the former taxon above. So far, its distribution is known only from the type locality.
Ulnaria qinghainensis is characterized by its linear-lanceolate valve outline, lacking central area, sub-capitate apices, and slender valves. It differs from U. obtusa (W. Smith) E. Reichardt by its narrower valves (3.1–5.0 μm vs 5–8 μm) and its much higher stria density (9–11 in 10 μm vs 3–4 in 10 μm) (see
Slide JIUDIA202311, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103817.
China. Guizhou province: Fanjing Mountain National Nature Reserve, Heiwang river, near Mile Daochang (27°49'10"N, 108°46'18"E, 494 m a.s.l.), collected by Bing Liu, December 31, 2015.
LM
(Figs
SEM
(Figs
Ulnaria fanjingensis sp. nov., girdle view, SEM. A a frustule in girdle view B–D details from A, note the 3:2 configuration of girdle bands E, F apical details from D, showing the closing plates (two arrows respectively) and two horn-like projections (F, two arrowheads). Scale bars: 20 μm (A); 2 μm (B–D); 1 μm (E, F).
Ulnaria fanjingensis sp. nov., external view, SEM A a complete valve B middle detail from A, showing the clear central area C middle detail of another valve showing the central area D a pole from A, note the slightly radiated striae and sub-capitate apex E, F two apices from A, note two largest horn-like projections and ocellulimbus. Scale bars: 20 μm (A); 2 μm (B–D); 1 μm (E, F).
Ulnaria fanjingensis sp. nov., internal view, SEM A a complete valve B middle part detail from A, note the clear central area C detail of another valve center showing the clear central area D a pole from A, note the tapering valve and rostrate apex E, F two apical details from A, note bilabiate rimoportulae and rostrate apices. Scale bars: 20 μm (A); 5 μm (B–D); 1 μm (E, F).
Named after Fanjing Mountain, where the species was found.
The sampling site is close to the headwaters of Heiwan River, which originates in the Fanjing Mountain National Nature Reserve. The diatom samples were scraped off of the stone surfaces. The following environmental parameters were measured in the field. Electric conductivity was 54.9 ± 1.4 μS∙cm-1, pH was 7.6 ± 0.1, and water temperature was10.4 ± 0.1 °C. Since the diatom sample was scraped off of the surfaces of stones and the conductivity is below 100 μS∙cm-1, U. fanjingensis can be considered an epilithic diatom characteristic of poor electrolyte content fresh water. So far, its distribution is known only from the type locality.
Ulnaria fanjingensis is characterized by its lanceolate valve outline, apically rectangular central area, rostrate to sub-capitate apices, and long valves. It is similar to U. dongtingensis Bing Liu, but they have different valve outlines: U. dongtingensis has narrow-lanceolate valves with parallel central margins the length of the central area whereas U. fanjingensis has parallel central margins extending beyond the central area (see
Slide JIUDIA202312, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103818.
China. Hunan province: Huping Mountain National Nature Reserve, Xie River, a sampling location (30°1'9"N, 110°37'46"E, 400 m a.s.l.), collected by Bing Liu, March 14, 2021.
LM
(Figs
SEM
(Figs
Ulnaria hupingensis sp. nov., internal view, SEM A a valve with valvocopula B middle detail from A, note the row of poroids interrupted in the middle (two arrows) C, E two poles from A. D Apical detail from C, note two rimoportulae at one apex (two arrows) F apical detail from E, note one rimoportula at one apex. Scale bars: 20 μm (A); 2 μm (B–F).
Ulnaria hupingensis sp. nov., external view, SEM A a complete valve B middle part detail from A, note central area complete absent C, D two apical details from A, showing two horn-like projections protruding over the oceullulimbus (two arrows, respectively) E, F two other apical details showing two horn-like projections protruding over the ocellulimbus (two arrows, respectively). Scale bars: 20 μm (A); 1 μm (B–F).
Named after Huping Mountain National Nature Reserve, where the species was found.
Epilithic in a mountain stream with oligotrophic waters. The following environmental parameters were measured in the field: Conductivity was 263 ± 1 μS∙cm-1, pH was 8.4 ± 0.1 and water temperature was 12.4 ± 0.4 °C. So far, its distribution is known only from the type locality.
Ulnaria hupingensis is characterized by its lanceolate valve outline, lacking central area, and rostrate to sub-capitate apices. It differs from U. qinghainensis by its wider valves (5.4–7.0 μm vs 3.1–5.0 μm) and from U. obtusa by its much higher stria density (9.5–11.5 in 10 μm vs 3–4 in 10 μm) (see
Slide JIUDIA202313, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103819.
China. Hunan province: Huping Mountain National Nature Reserve, Xie River, a sampling location (30°1'9"N, 110°37'46"E, 400 m a.s.l.), collected by Bing Liu, March 14, 2021.
LM
(Figs
Ulnaria xieriverensis sp. nov., ×1000, LM A–H eight valves exhibiting a size diminution series, note the almost parallel but slightly undulate valve margins in larger specimens (A–C) and parallel valve margins in smaller specimens (D–H). A Micrograph of holotype specimen. Scale bar: 10 μm.
SEM
(Figs
Ulnaria xieriverensis sp. nov., external view, SEM A a complete valve B middle part detail from A, note central area is completely absent C, D two apical details from A. E, F Two other apical details, note two horn-like projections protruding over the ocellulimbus (two arrows respectively). Scale bars: 10 μm (A); 1 μm (B–F).
Ulnaria xieriverensis sp. nov., internal view, SEM A a complete valve B middle part detail from A, note central area is completely absent C, D two apical details from A, note one rimoportula located against the mantle (D, arrow) E, F two other apical details. Scale bars: 10 μm (A); 1 μm (B–F).
Ulnaria xieriverensis sp. nov., internal view, SEM A a complete valve with valvocopula B, C two complete valves with valvocopula, note undulate valve margins D middle detail from A, note the central area is completely absent and a row of serrated projections over virgae (three arrows) E, F two apical details from A, note serrated projections over virgae (E, three arrows). Scale bars: 10 μm (A–C); 1 μm (D–F).
Named after Xie River, where the species was found.
Epilithic in a mountain stream with oligotrophic waters. Ulnaria hupingensis was found in the same sampling site with U. xieriverensis and the environmental parameters see above. So far, its distribution is known only from the type locality.
Ulnaria xieriverensis is characterized by its linear valve outline, absence of central area, and rostrate apices. It differs from U. hupingensis by its linear valve outline.
Slide JIUDIA202314, specimen circled on slide, illustrated as Fig.
PhycoBank http://phycobank.org/103820.
China. Hunan province: Lanshan County, Shun River, a sampling location (25°11'29"N, 112°7'47"E, 490 m a.s.l.), collected by Bing Liu, October 5, 2021.
LM
(Figs
Ulnaria pandurata-uniseriata sp. nov., external view, SEM A, B two complete valves, note panduriform valve outline C, D two middle part details from A, B, respectively, note rectangular central areas E, F two apical details from B, note two horn-like projections protruding over ocellulimbus. Scale bars: 10 μm (A, B); 1 μm (C, D); 2 μm (E, F).
SEM
(Figs
The epithet pandurata>-uniseriata is a combination of the terms pandurate and uniseriate reflecting the panduriform outline of the valve and its mostly uniseriate striae.
Epilithic in a mountain stream with oligotrophic waters. The following environmental parameters were measured in the field: Conductivity was 53.9 ± 0.6 μS∙cm-1, pH was 8.4 ± 0.3 and water temperature was 21.9 ± 0.1 °C. So far, its distribution is known only from the type locality.
Ulnaria pandurata-uniseriata is characterized by its panduriform valve outline, rectangular central area, mostly uniseriate striae, and its rostrate apices. It differs from U. jumlensis (Jüttner) D.M. Williams & Karthick by its narrower valve (6.2–8.2 μm vs 7.5–11.2 μm) and lower stria density (9–11 in 10 μm vs 13–14 in 10 μm) (see
Valvocopula ultrastructure: In the above “Taxonomic treatment”, we illustrate a whole valvocopula that is separate from the valve in U. neobiceps sp. nov. (Fig.
Configuration of girdle bands: The number of girdle bands associated with the hypovalve may vary, as can the number of girdle bands associated with the epivalve in the same species. Evidence is found in U. oxybiseriata D.M. Williams & Bing Liu and U. jishou-biseriata sp. nov. as the number of girdle bands associated with the epivalve can be either 3 (see
Ultrastructure and configurations of girdle bands in 11 Ulnaria species.
Species | Mantle depth (μm) | Poroids in girdle bands | Configuration of girdle bands (dividing cells) | Configuration of girdle bands (non-dividing cells) |
---|---|---|---|---|
U. constricta-biseriata sp. nov. | 2–4 | Two rows of poroids produced in both 2nd and 3rd band, but the 2nd row of poroids discontinuous | 3:3 | 3:2 |
U. dongtingensis | 2–3 | One row of poroids in all girdle bands | No data (one ratio should be 3:3) | 3:2 |
U. fanjingensis sp. nov. | 2–3 | One row of poroids in girdle bands except third copula which has second interrupted row of poroids | No data (one ratio should be 3:3) | 3:2 |
U. gaowangjiensis | 3–4 | Second interrupted row of poroids in 4th copula | 4:4 | No data |
U. hupingensis sp. nov. | 2–3.5 | Second interrupted row of poroids in 2nd copula | No data | Three girdle bands associated with the epivalve |
U. jinbianensis sp. nov. | 2–3 | One row of poroids in all girdle bands | No data (one ratio should be 4:4) | 4:2 at beginning |
U. jishou-biseriata sp. nov. | 2–3 | Second interrupted row of poroids in 3rd copula | No data | 3:3 or 4:3 |
U. oxybiseriata | 2–3 | Second interrupted row of poroids in 3rd copula | 3:3 | 4:3 |
U. sinensis | 2–4 | One row of poroids in all girdle bands | No data (one ratio should be 4:4) | 4:2 |
U. sangzhi-biseriata sp. nov. | 2–3 | Second interrupted row of poroids in 3rd and 4th copulae | No data (one ratio should be 4:4) | 4:2 |
U. ulnabiseriata | 2–3 | One row of poroids in all girdle bands | 4:4 | 4:2 |
In Ulnaria constricta-biseriata sp. nov., the epivalve is associated with a valvocopula and two copulae, so it has an overall configuration of 3:3 in dividing cells or 3:2 in non-dividing cells (Fig.
Based on our published data (
There are rarely reports of the whole life history in the araphid diatoms, but a few papers mention the transverse perizonal bands of the initial cell. For example, Williams noted “no sign of transverse perizonal bands at all” in Fragilariforma virescens (Ralfs) D.M. Williams & Round (
We observed living cells of five Ulnaria species (U. fanjingensis sp. nov., Fig.
Ross et al. defined the basal siliceous layer as “the layer that forms the basic structure of the various components of the frustule” (
The striae are usually divided into three types: uniseriate, biseriate and multiseriate striae (
Closing plates occlude the external openings of areolae in Ulnaria (sensu
At first, the ocellulimbus was defined as a special type of pore field, a “plate set into the polar valve mantle” (
The rimoportulae in all Ulnaria have the same structure (more or less) – they are simple holes externally and bilabiate internally – with the number present and some aspects of the shape and position varying. One rimoportula is present at each apex in most Ulnaria species. However, in three Ulnaria species, U. colcae Van de Vijver & Cocquyt, U. macilenta E. Morales, C.E. Wetzel & S.F. Rivera, and U. titicacaensis E. Morales, Ector & P.B. Hamilton, only one rimoportula is present at one apex per valve (
In the 14 Ulnaria taxa described in this study, only in U. chengduoensis sp. nov. was the valvocopula not observed. We also document the whole valvocopula that is separate from the valve in U. neobiceps sp. nov. (Fig.
In a diatom cell, the configuration of girdle bands is defined as the ratio between the number of girdle bands associated with the epivalve and those associated with the hypovalve (sensu
The life history of Ulnaria was revealed in this study for the first time. A distinct pre-normal vegetative cell stage exists during the process from the initial cells to the normal vegetative cells.
The configuration of girdle bands of 11 species and the valvocopula ultrastructure of 13 species are observed, illustrated, and described. A conclusion can be drawn at this moment that the configuration of girdle bands in Ulnaria cells is a common condition based on this study and our previous studies. The configuration of girdle bands changes during the cell cycle and the number of girdle bands in a cell will reach its maximum at the late stage of cell dividing. Thus, to prove all the girdle bands are closed, you must observe the cell at the late stage of cell division, which is difficult to achieve in practice. Herein, I recommend the closed valvocopula as a defining character for the genus Ulnaria because it is more feasible in practice and an equally effective alternative to “closed girdle bands”.
Most Ulnaria species possess uniseriate striae. Three types of mixed striae in Ulnaria are observed in this study. If we hypothesize that the first type of mixed striae is a transitional type from the uniseriate striae to mostly biseriate (the second type of mixed striae), and the third type of mixed striae is a transitional type from the mostly biseriate to the mostly triseriate striae, then we predict that the existence of Ulnaria species, which possesses mostly triseriate striae, is waiting to be found in nature.
I thank three master’s degree candidates, Wen-Hui Mo, Ji-Yan Long and Li Yuan, for their assistance in the field and the laboratory. I am also very grateful to two reviewers and the editors for their careful revisions and helpful comments.
The author has declared that no competing interests exist.
No ethical statement was reported.
This work was supported by the National Natural Science Foundation of China (No. 31760051) and the Natural Science Foundation of Hunan Province [No. 2018JJ2311].
Writing – review and editing: BL.
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Seven Ulnaria taxa possessing both uniseriate striae and marginal spines.
No. | Taxon | Valve outline | Valve dimensions (μm): length (L), width (W) | Central area | Striae density (in 10 μm) | Nature of girdle bands | References |
---|---|---|---|---|---|---|---|
1 | U. cochabambina | Linear-lanceolate with subrostrate or sub-capitate apices | L: 88–296; W: 8–9 | Square or rectangle fascia | 8–10 | Valvocopula closed |
|
2 | U. fragilariaeformis | Linear-lanceolate with rostrate or sub-capitate apices | L: 105–120; W: 3–5 | Absent | No data | Closed |
|
3 | U. monodii | Linear with with sub-capitate apices | L: 140–400; W: 7.5–8.5 | Absent | 8–9 | Closed |
|
4 | U. pseudogaillonii | Linear with attenuate and rounded apices | L: 220–410; W: 8–10 | Absent | 7–9 | Unknown |
|
5 | U. sinensis | Linear with rostrate apices | L: 296–512; W: 6–8 | Absent | 8–9 | Closed |
|
6 | U. undulata | Linear with rostrate to sub-capitate apices | No data | Absent | No data | Closed |
|
7 | U. ungeriana | Linear-lanceolate with cuneately rostrate apices | L: 85–230; W: 8–10 | Very variable | (7) 8–10 | Unknown |
|
No. | Taxon | Valve outline | Valve dimensions (μm): length (L), width (W) | Central area | Striae density (in 10 μm) | Nature of girdle bands | References |
---|---|---|---|---|---|---|---|
1 | U. acuscypriacus | Linear-lanceolate with sub-capitate apices | L: 140–300; W: 6–6.8 | Rectangular or square as incomplete fascia | 8.5–10 | At least two copulae closed |
|
2 | U. constricta-biseriata sp. nov. | Linear-lanceolate, constricted at centre, with rostrate to sub-capitate apices | L: 66–166; W: 5.5–8 | Rectangular or square fascia | 10–12 | Closed | This paper |
3 | U. gaowangjiensis | Linear with rostrate apices | L: 61–108; W: 6.5–8.5 | Square fascia | 9–11 | Closed |
|
4 | U. goulardii | Lanceolate with a distinctive constriction at the centre, apex cuneately rostrate | L: 55–90; W: 8–10 | Rectangular or square fascia | 9–11 | Unknown |
|
5 | U. gusliakovii | Linear with rostrate apices | L: 110–256; W: 4.5–6 | Absent | 10 | Closed |
|
6 | U. jishou-biseriata sp. nov. | Lanceolate with rostrate apices | L: 139–200; W: 6–8 | Variable | 10–12 | Closed | This paper |
7 | U. lanceolata | Linear with cuneately rostrate apices | L: 55–75; W: 8–10 | Rectangular or square fascia | 8–10 | Unknown |
|
8 | U. nyansae | Linear with sub-capitate to capitate apices | L: 55–175; W: 8–15 | Half fascia | (12) 13–15 | Unknown |
|
9 | U. oxybiseriata | Linear-lanceolate to lanceolate with apiculate apices | L: 56–78; W: 6–9 | Rectangular or trapezoid fascia | 10–12 | Valvocopula closed |
|
10 | U. pandurata-biseriata | Panduriform with rostrate apices | L: 37–60; W: 7–9.5 | Variable | 9–11 | Valvocopula closed | This paper |
11 | U. sangzhi-biseriata sp. nov. | Linear-lanceolate with capitate to sub-capitate apices | L: 49–91; W: 6.5–8.2 | Rectangular or square fascia | 10–12 | Closed | This paper |
12 | U. ulnabiseriata | Linear-lanceolate with rostrate to sub-capitate apices | L: 105–229; W: 6–8 | Rectangular fascia | 9–11 | Closed |
|
13 | U. ulna var. spathulifera | Linear with spatulate apices | L: 134–190; W: 8–9 | Circular to rectangular, sometimes flanked by short striae | 9–10 | Unknown |
|
14 | U. wuling-biseriata sp. nov. | Linear with rostrate to sub-capitate apices | L: 160–200; W: 6.5–8 | Rectangular or trapezoid fascia | 10–11 | Valvocopula closed | This paper |
Forty-two Ulnaria taxa possessing uniseriate striae but no marginal spines.
No. | Taxon | Valve outline | Valve dimensions (μm): length (L), width (W) | Central area | Striae density (in 10 μm) | Nature of girdle bands | References |
---|---|---|---|---|---|---|---|
1 | U. acus | Lanceolate | L: 90–100; W: 4–6 | Variable | 12–15 | Closed |
|
2 | U. aequalis | Linear | L: 50–200; W: 5–10 | Variable | 4–6 | Closed |
|
3 | U. amphirhynchus | Linear with rostrate apices | No data | No data | No data | Unknown |
|
4 | U. biceps | Lanceolate with capitate apices | No data | No data | No data | Unknown |
|
5 | U. blancoi sp. nov. | Lanceolate with rostrate to sub-capitate apices | L: 104–236: W: 4.6–6.8 | Very variable | 10–13 | Valvocopula closed | This paper |
6 | U. capitata | Linear with capitate, rhomboid apices | L: 255–406; W: 8 | No clear central area | 10–11 | Unknown |
|
7 | U. chengduoensis sp. nov. | Linear with rostrate apices | L: 42–66; W: 6–8 | Not clearly visible due to presence of ghost striae or absent | 12–15 | Unknown | This paper |
8 | U. colcae | Linear to linear-lanceolate with subrostrate apices | L: 18–46; W: 3.9–5.2 | Asymmetrical, rectangular to circular fascia | 16–17 | Closed |
|
9 | U. contracta | Constricted valve outline with rostrate to subrostrate apices | L: 84–140; W: 5–6 | Rectangular of variable length | 11–14 | Unknown |
|
10 | U. danica | Lanceolate with rostrate to sub-capitate apices | No data | No data | No data | Unknown |
|
11 | U. delicatissima | Needle-shaped with sub-capitate apices | L: 142–330; W: 4.2–5 | Variable | 8.5–10 | Unknown |
|
12 | U. dongtingensis | Narrow-lanceolate with rostrate to capitate apices | L: 106–260; W: 5–7 | Square fascia | 10–12 | Closed |
|
13 | U. fanjingensis sp. nov. | Lanceolate with rostrate to sub-capitate apices | L: 165–291; W: 4.8–6.3 | Apically rectangular fascia | 9–12 | Closed | This paper |
14 | U. ferefusiformis | Linear-lanceolate with sub-capitate apices 76–152, 4–4.8 | L: 76–152; W: 4.0–4.8 | Indistinctly defined | 12–14 | Unknown |
|
15 | U. grunowii | Narrowly fusiform with sub-capitate apices | L: 100–380; W: 2.0–4.0 | Not or not distinctly separated | 11–15.5 | Unknown |
|
16 | U. hunanensis | Linear-lanceolate with sub-capitate to rostrate apices | L: 62–166; W: 3.3–5.2 | Variable | 12–14 | Unknown |
|
17 | U. hupingensis sp. nov. | Lanceolate with rostrate to sub-capitate | L: 81–200; W: 5.4–7.0 | Completely absent | 9.5–11.5 | Closed | This paper |
18 | U. japonica | Needle-shaped | No data | Present | 11–14 | Closed |
|
19 | U. jinbianensis | Linear-lanceolate with rostrate to cuneate apices | L: 42–91; W: 5.5–7.5 | Rectangular to trapezoid fascia | 10–11 | Closed |
|
20 | U. jumlensis | Slightly panduriform with apiculate apices | L: 37–48; W: 6.5–9.5 at centre | Present | 13–14 | Closed |
|
21 | U. macilenta | Linear-lanceolate with subrostrate to broadly rounded apices | L: 18.5–95; W: 2.5–3.5 | Square to rectangular fascia | 18–19 | Unknown |
|
22 | U. menyuanensis sp. nov. | Lanceolate with cuneate to rostrate apices | L: 60–104; W: 5–7 | Not clearly visible due to presence of ghost striae | 12–14 | Valvocopula closed | This paper |
23 | U. neobiceps sp. nov. | Linear-lanceolate with distinct capitate apices | L: 202–307; W: 4.5–6.7 | Very variable | 9–11 | Valvocopula closed | This paper |
24 | U. obtusa | Linear with rounded apices | L: 150–200; W: 5–8 | Absent | 3–4 | Closed |
|
25 | U. oxyrhynchus | Linear with acute, conical to subrostrate apices | L: 43–64; W: 6–7 | Present, flanked by short striae | 13–14 | Unknown |
|
26 | U. pandurata-uniseriata sp. nov. | Panduriform with rostrate apices | L: 70–116; W: 6.2–8.2 | Distinct, fascia-shaped | 9–11 | Valvocopula closed | This paper |
27 | U. pilum | Needle-shaped with sub-capitate apices | L: 218–295; W: 5.6–6.3 | Rectangular | 10–11.5 | Unknown |
|
28 | U. qinghainensis sp. nov. | Linear-lanceolate with sub-capitate apices | L: 88–223; W: 3.1–5.0 | Completely absent | 9–11 | Valvocopula closed | This paper |
29 | U. ramesii | Linear-lanceolate with rostrate apices | L: 50–92; W: 8–9 | Ellipsoid to rectangular | 11–14 | Closed |
|
30 | U. repanda | Undulate with broadly rounded apices | L: 380–425; W: 5–9 | Absent | 7–12 | Unknown |
|
31 | U. rhombus | Rhombic with sub-capitate apices | L: 90–184; W: 4–5.5 | Rhomboid, circumscribed by shortened striae | 13–14 | Unknown |
|
32 | U. rostrata | Linear-lanceolate with rostrate apices | No data | Rectangular | No data | Closed |
|
33 | U. schroeteri | Filiform | L: 300–450; W: 2.5–4 | Rectangular | 15–18 | Closed |
|
34 | U. splendens | Linear with rostrate apices | L: 60–300; W: 5–10 | Round to oval hyaline portion | 6–10 | Closed |
|
35 | U. titicacaensis | Lanceolate with subrostrate apices | L: 15.5–98.5; W: 4–6.5 | Oval to square fascia, asymmetrical | 16–18 | Unknown |
|
36 | U. tooelensis | Linear to linear-elliptical with sub-capitate apices | L: 14–97.5; W: 3.5–5.5 | Rectangular central area on one side | 11–15 | Closed |
|
37 | U. tortuos | Arc | L: 120–450; W: 10–12 | Present | 5–10 | Closed |
|
38 | U. ulna | Needle-shaped with sub-capitate apices | L: 230–320; W: 6.3–7.7 | Absent | 9.2–10.4 | Closed |
|
39 | U. verhaegeniana | Linear with acute apices | L: 55–90; W: 5–6 | Apically rectangular | 10–12 | Unknown |
|
40 | U. vitrea | Lanceolate | L: 90–120; W: 4.5–6 | Present, variable | No data | Closed |
|
41 | U. wulingensis | Narrowly linear-lanceolate | L: 153–233; W: 2.7–4.8 | Rectangular fascia | 12–14 | Valvocopula closed |
|
42 | U. xieriverensis sp. nov. | Linear with rostrate apices | L: 64–120; W: 6.0–8.6 | Absent | 10.5–12 | Valvocopula closed | This paper |