Research Article |
Corresponding author: Weliton José da Silva ( welitondasilva@yahoo.com.br ) Academic editor: Kalina Manoylov
© 2015 Weliton José da Silva, Regine Jahn, Thelma Alvim Veiga Ludwig, Friedel Hinz, Mariângela Menezes.
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:
da Silva WJ, Jahn R, Ludwig TAV, Hinz F, Menezes M (2015) Typification and taxonomic status re-evaluation of 15 taxon names within the species complex Cymbella affinis/tumidula/turgidula (Cymbellaceae, Bacillariophyta). PhytoKeys 53: 1-25. https://doi.org/10.3897/phytokeys.53.4782
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Specimens belonging to the C. affinis / C. tumidula / C. turgidula species complex have many taxonomic problems, due to their high morphological variability and lack of type designations. Fifteen taxon names of this complex, distributed in five species, were re-evaluated concerning their taxonomic status, and lectotypified based on original material. In addition to light microscopy, some material was analyzed by electron microscopy. Four new combinations are proposed in order to reposition infraspecific taxa.
Diatoms, typification, Cymbellales , Cymbella tropica , Cymbella subturgidula , Cymbella rheophyla , Cymbella uenoi
The history of the genus Cymbella C.Agardh is replete with taxonomic complexities. Within these complexes many species are similar in valve morphology. Much of the confusion in these complexes was caused by poor species descriptions including specimen images and the lack of designation of types, which has been required by the International Code of Nomenclature for algae, fungi, and plants (ICN) only since 1958 (
In the most recent revision of the genus Cymbella,
Krammer in
The material of C. uenoi Skvortsov was also revisited by
Despite the changes, the revision of
The aim of the present study was to elucidate the current taxonomic status of C. tumidula, C. excisa and C. subturgidula, as well as of C. uenoi and C. uenoi f. nipponica [≡ C. turgidula var. nipponica; ≡ Cymbella rheophila]; to revisit the infraspecific taxa encompassed in this complex of species; and to lectotypify the ambiguous taxa.
We analyzed the protologues and morphological features from materials of C. affinis var. affinis, C. affinis var. procera, C. excisa var. excisa, C. excisa var. procera, C. excisa var. angusta, C. excisa var. subcapitata, C. tumidula var. tumidula, C. salinarum, C. turgida, C. tropica, C. subturgidula, C. turgidula var. nipponica, C. reophila, C. uenoi f. uenoi, and C. uenoi f. nipponica (Table
Metric features of the material of the complex C. affinis/tumidula/turgidula analyzed in this study (n= 29–34 individuals).
Taxon (published name) | Current name | Material | Length (µm) | Breadth (µm) | L/B ratio | Striae in middle part (in 10 µm) | Striae close to the ends (in 10 µm) | Puncta (in 10 µm) | Stigmata |
---|---|---|---|---|---|---|---|---|---|
C. affinis Kütz. sensu |
C. tumidula var. tumidula | Preparation 1198F IOK in BRM, from France, Normandy, Falaise, coll. Brébisson, in BRM (Figs |
25.4–29.8 | 7.8–8.7 | 3.4–3.9 | 10–15 | 12–19 | 26–33 | 2–3 |
C.
affinis
|
C. affinis var. affinis | Lectotype designated by |
22.5–26.7 | 7.0–8.5 | 2.9–3.4 | 9–12 | 11–18 | 19–27 | 1 |
C.
affinis
var.
procera
|
C. tumidula var. procera (Krammer) W.Silva, comb. nov. | Holotype, preparation 714 IOK, from Serbia, Rogatica, 1976, in BRM (Figs |
27.0–40.3 | 9.0–10.2 | 2.9–4.2 | 9–13 | 13–18 | 25–31 | 2–5 |
C.
excisa
var.
procera
|
C. affinis var. neoprocera W.Silva, comb. nov. et nom. nov. | Holotype, preparation 212A IOK, from West Germany, Eifel, Totenmaar, 22.2.1974, in BRM (Figs |
21.5–41.0 | 8.0–11.0 | 2.7–4.0 | 9–13 | 11–14 | 22–27 | 1 |
C.
excisa
|
C. affinis var. excisa (Kütz.) Grunow | Isolectotype, preparation 1131G IOK, Hauck No. 72 from Italy, 26.3.1837, in BRM (Figs |
23.1–28.5 | 7.0–9.5 | 2.7–3.9 | 9–12 | 11–17 | 24–26 | 1 |
C.
excisa
var.
angusta
|
C. affinis var. angusta (Krammer) W.Silva, comb. nov. | Holotype, preparation 752 IOK, from Croatia, Lake Gavanovac, 18.5.1976, in BRM (Figs |
20.5–35.0 | 6.0–8.5 | 3.2–4.6 | 9–15 | 11–18 | 19–29 | 1 |
C.
excisa
|
C. affinis var. excisa (Kütz.) Grunow | Lectotype (designated here), preparation BM 18543, from Hauk No. 72 from Italy, 26.3.1837, in BM (Figs |
21.5–26.5 | 7.0–8.7 | 2.6–3.5 | 9–13 | 11–16 | 21–25 | 1 |
C.
excisa
var.
subcapitata
|
C. affinis var. subcapitata (Krammer) W.Silva, comb. nov. | Holotype, preparation 717A IOK, from Hungary, Balaton, 6.77, in BRM (Figs |
21.0–31.7 | 7.4–9.0 | 2.7–4.0 | 8–12 | 9–15 | 26–31 | 1 |
C.
subturgidula
|
C. subturgidula Krammer | Lectotype, preparation 1046E IOK, from Korea, Ulchin County, Kyungsang Pukdo, Kwangchun River, in BRM (Figs |
30.3–37.4 | 9.0–12.8 | 2.6–3.5 | 9–11 | 12–15 | 21–24 | 2 |
C.
tropica
|
C. tropica Krammer | Holotype, preparation 1015D IOK, from Venezuela, Rio Manizanes, coll. Rumrich 4.4.1990, in BRM (Figs |
34.5–42.7 | 10.0–12.0 | 3.1–4.0 | 9–12 | 11–12 | 21–24 | 1 |
C. tumidula Grunow in |
C. tumidula var. tumidula | Lectotype (designated here); Epitype (designated here), preparation BM 18543, from Hauck No. 72, from Italy, 26.3.1837, in BM (Figs |
26.8–34.7 | 7.8–8.7 | 3.4–4.0 | 10–15 | 12–19 | 26–33 | 2–4 |
C. salinarum Grunow in |
C. tumidula var. salinarum (Grunow) Cleve | Holotype illustration provided |
34.0 | 10.7 | 3.2 | 14 | 15 | – | 1 |
C. turgidula Grunow in |
C. turgidula Grunow | Epitype designated by |
36.5–45.0 | 10.8–13.3 | 3.2–3.4 | 9–12 | 11–17 | 21–24 | 1–3 |
C. turgidula var. nipponica Skvortsov (1936) [≡ C. rheophila Ohtsuka in |
C. subturgidula Krammer | Isolectotype, preparation R 214.928, from sample 0983 from Japan, Lake Biwa, coll. Tamiji Kawamura, 03.11.1915, in R (Figs |
28.1–39.0 | 10.0–13.2 | 2.8–3.0 | 9–12 | 12–15 | 21–26 | 1–2 |
Preparations R 214.929 and R 214.930, from sample 1062, from Japan, Yamanoshita Bay in Lake Biwa, Otsu City, Shiga Prefecture, coll. Yasuko Iwao in 23.01.1993, in R (Fig. |
27.7–39.2 | 10.3–13.1 | 2.5–3.6 | 9–12 | 12–15 | 22–26 | 1–2 | ||
Preparations R 214.931 and R214.932, from from sample 1093, from Japan, cobble in Lake Biwa, at Uchidehama, Otsu City, Shiga Prefecture, coll.Yasuko Iwao, 03.03.1993, in R (Figs |
26.5–40.1 | 11.1–13.5 | 2.3–3.4 | 9–13 | 12–14 | 21–24 | 2–3 | ||
C. uenoi Skvortsov ex |
Epitype and isotype illustrations provided |
26.3–41.0 | 9.0–12.7 | 2.9–3.2 | 9–11 | 12-15 | 21–24 | 2 |
Materials of C. affinis (BM18530), C. excisa (BM 18543), C. tumidula (BM 18543), C. turgidula var. nipponica (R 214.928, R 214.929, R 214.930, R 214.931 and R 214.932) were analyzed in the Laboratório de Ficologia, Museu Nacional, Rio de Janeiro, Brazil, using an Olympus BX 51 microscope (Olympus, Tokyo, Japan) fitted with an Olympus Q-Color digital camera. Images were processed with Q capture Pro QImaging© software.
Materials of C. affinis sensu Krammer (1198F IOK BRM) and C. affinis var. procera (714 IOK BRM) were examined in the laboratory of the Botanischer Garten und Botanisches Museum (BGBM), Berlin, Germany, using a Zeiss Axio Imager 4.2 microscope (Carl Zeiss MicroImaging GmbH, Berlin), and the images were captured through an MRc/MRm system (Carl Zeiss MicroImaging) and the software AxioVision Rel. 4.8 (Carl Zeiss MicroImaging).
We also analyzed materials C. excisa sensu Krammer (1131G IOK BRM), C. excisa var. procera (212A IOK BRM), C. excisa var. angusta (752 IOK BRM), C. excisa var. subcapitata (717A IOK BRM), C. tropica (1015D IOK BRM) and C. subturgidula (1046E IOK BRM) with a Zeiss Axioplan microscope (Carl Zeiss, Jena, Göttingen, Germany) with an Olympus XC50 capture system (Olympus, Tokyo, Japan) and the software analySIS Image Processing (Soft Imaging System, Münster, Germany), at the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
Scanning Electron Microscopy (SEM) analyses were carried out only for C. turgidula var. nipponica. Samples were deposited on cover slips and attached to aluminum stubs using LeitSilver® (Sigma-Aldrich, Berlin, Germany). The material was coated with 150–200 Å of gold in an Emitech K 550 sputter coater (Quorum Technologies Ltd., Kent, UK). The material was analyzed in a Jeol JSM-6390 scanning electron microscope (Jeol, USA), operated at 6–8 kV, spot size 10–30, in the electron microscopy laboratory in the Museu Nacional, Rio de Janeiro, Brazil.
The term “degree of dorsiventrality” is used here to define how dissimilar the sides of the valvae are on the apical axis. The symbols “≡”, “=” and “–” preceding specific and infraspecific names are used to represent homotypic or nomenclatural, heterotypic or taxonomic, and concept synonyms, respectively, as used in the ICN (
Cymbella affinis Kütz., Bacill., 80, Pl. 6 Fig. 15, 1844.
3rd figure from the left in
“Falaise (France): De Brébisson! Herb. Binder., Schleswig: Herb. Binder.!”
An individual on preparation BM18530, from Kützing packet 333, in the Natural History Museum (BM), designated by
Falaise, France, coll. De Brébisson.
Valvae lanceolate, dorsiventral, dorsal and ventral margins convex; ends barely protracted, rounded, to slightly subrostrate or subcapitate; length 22.5–26.5 µm, breadth 7.0–8.5 µm, L/B ratio 2.9–3.4; axial area narrow, linear-arched, indistinct central area; striae 9–12 in 10 µm, becoming 11–18 towards ends, one isolated pore at end of central striae on ventral side; 19–27 punctae in 10 µm.
Although
The lectotype valve of C. affinis is similar to the lectotype valve of C. excisa, except for the excision in the middle part of the valve present in the majority of specimens (Figs
Cymbella affinis species complex 1–7C. affinis var. affinis8–14C. affinis var. excisa (Kütz.) Grunow, specimens from Trieste in preparation BM 18543 11 Lectotype, designated here 15–21C. affinis var. excisa [– C. excisa sensu Krammer], specimens from Trieste in preparation 1131 IOK 16 Isolectotype, designated here 22–28C. affinis var. neoprocera W.Silva, specimens from Germany in preparation 212A IOK, holotype 22 Lectotype, designated here. Scale bar: 10 µm.
The two taxa were both proposed by
Cymbella affinis valves sensu
C. excisa var. procera Krammer (Figs
Cymbella affinis var. excisa (Kütz.) Grunow, Beitr. Paläont. Österreich.-Ungarns Orients, 2: 142, Pl. 19(1), Fig. 26, 1882.
Cymbella excisa Kütz., Bacill., 80, Pl. 6, Fig. 17, 1844.
(designated here). An individual marked with blue ring on preparation BM 18543, from Hauck No. 72, 26.4.1837, in the Natural History Museum (BM), London, United Kingdom, represented by Fig.
(designated here). An individual on preparation 1131G IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
“Unter Oscillatorien in Bächen bei Triest”, Italy, 26.4.1837.
Valvae dorsiventral, dorsal margin broadly convex, ventral margin straight, usually with an excision in middle portion; ends subrostrate to rostrate; length 21.5–28.5 µm, breadth 7.0–9.5 µm, L/B ratio 2.6–3.9; axial area narrow, linear-arched, central area indistinct; striae 9–13 in 10 µm, becoming 11–17 toward ends, one isolated pore at end of central striae on ventral side; 21–26 punctae in 10 µm.
Similarly to C. affinis, the lectotypification of Cymbella excisa designated by
The main difference between C. affinis var. excisa and the nominate variety is the presence of an excision in the middle portion of the ventral side of the valve, a characteristic common to populations of this taxon around the world (
Cymbella excisa var. procera Krammer, Diatoms Europe 3: 159, Figs 9:1–7, 2002 (Figs 22–28).
Preparation 212A IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
(designated here). An individual on preparation 212A IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany; represented by Fig.
Germany, Eifel, Totenmaar, 22.2.1974.
Valvae dorsiventral, dorsal margin broadly convex, ventral margin straight; ends not protracted, rounded, or subrostrate to rostrate; length 21.5–41.0 µm, breadth 8.0–11.0 µm, L/B ratio 2.7–4.0; axial area narrow, linear-arched, indistinct central area; striae 9–13 in 10 µm, becoming 11–14 toward ends, one isolated pore at end of central striae on ventral side; 22–27 punctae in 10 µm.
The combination of C. excisa var. procera with Cymbella affinis would be illegitimate unless it was given a new name, because it would be a later homonym of C. affinis var. procera Krammer. Specimens designated by
A broader range of metric characteristics were highlighted in this study compared to the characterization of the type population (
Cymbella excisa var. angusta Krammer, Diatoms Europe 3: 159, Figs 9: 8–13, 2002.
Preparation 752 IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
(designated here). An individual on preparation 752 IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
Cymbella affinis species complex 29–35C. affinis var. angusta (Krammer) W.Silva, specimens from Croatia in preparation 752 IOK, holotype 29 Lectotype, designated here 36–42C. affinis var. subcapitata (Krammer) W.Silva, specimens from Hungary in preparation 717 IOK, holotype 39 Lectotype, designated here. Scale bar: 10 µm.
Croatia, watercourse near Lake Gavanovac, Plitvice.
Valves dorsiventral, dorsal margin broadly convex, ventral margin straight, usually with excision in middle portion; ends not protracted to slightly protracted, rounded, subrostrate or subcapitate; length 20.5–35.0 µm, breadth 6.0–8.5 µm, L/B ratio 3.2–4.6; axial area linear-arched, indistinct central area; striae 9–15 in 10 µm, becoming 11–18 towards ends, one isolated pore at end of central striae of ventral side; 19–29 punctae in 10 µm.
The material recorded here presented slight differences in the valve metrics compared to
Similar to C. affinis var. neoprocera, specimens designated by
Cymbella excisa var. subcapitata Krammer, 2002, Diatoms of Europe 3: 159, Figs 10: 14–18.
Preparation 717A IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
(designated here). An individual on preparation 717A IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
Hungary, Balaton.
Valves dorsiventral, dorsal margin broadly convex, ventral margin straight, usually with excision in middle portion; ends barely protracted, rounded, to broadly protracted, subcapitate; length 21.0–31.7 µm, breadth 7.4–9.0 µm, L/B ratio 2.7–4.0; axial area linear-arched, indistinct central area; striae 8–12 in 10 µm, becoming 9–15 towards ends, one isolated pore at end of central striae of ventral side; 26–31 punctae in 10 µm.
Specimens designated by
Cymbella tumidula var. tumidula, in Schmidt, A. Schmidt’s Atlas Diatom.-Kunde, Pl. 9, Fig. 33, 1875.
Cymbella affinis Kütz. sensu Krammer, Diatoms Europe 3: 41, Figs 21:2–21, 22:1–20, 23:1–18, 2002. (Figs
Cymbella tumidula species complex 43 Original illustration provided by Grunow in
(designated here). Plate 9, Fig. 33 from Schmidt (1875a) (Fig.
Italy, Trieste, 26.4.1837.
(designated here). An individual marked with a red ring on preparation BM 18543 from Hauck No. 72, 26.4.1837, in the Natural History Museum (BM), London, United Kingdom, represented by Fig.
(designated here). An individual on preparation 1131G IOK, from Hauck No. 72, 26.4.1837, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
Italy, Trieste, 26.4.1837.
Valves lanceolate, dorsiventral, dorsal and ventral margins convex; subcapitate ends; length 25.4–34.7 µm, breadth 7.8–8.7 µm, L/B ratio 3.4–4.0; axial area linear-lanceolate, straight to arched, central area irregular to rounded; striae 10–15 in 10 µm, becoming 12–19 toward ends, 2–4 isolated pores at end of central striae on ventral side; 26–33 punctae in 10 µm.
The lectotypification of C. affinis allowed us to consider C. affinis and C. tumidula [– C. affinis sensu Krammer], lectotypified and epitypified here, as independent species. C. tumidula has a more lanceolate outline, subcapitate ends, and a lower degree of dorsiventrality than C. affinis. The striae in the middle part of the valve are shorter and unevenly distributed in C. tumidula, forming a distinct central area (Figs
Specimens from the population of the holotype material of C. affinis var. procera were very similar in outline but larger and wider than C. tumidula var. tumidula, resulting in higher maximum length/breadth ratios. However, all metric characteristics of C. affinis var. procera intergraded with C. tumidula var. tumidula, and therefore this taxon was transferred to C. tumidula var. procera.
The characteristics of C. affinis sensu
Cymbella affinis var. procera Krammer, Diatoms Europe 3: 161, Figs 22: 8–13, 2002.
Preparation 714 IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
(designated here). An individual on preparation 714 IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
Serbia, Rogatica (abundant in chalk-rich spring), 1976.
Valves lanceolate, dorsiventral, dorsal and ventral margins convex; ends subrostrate or subcapitate; length 27.0–40.3 µm, breadth 9.0–10.2 µm, L/B ratio 2.9–4.2; axial area linear-lanceolate, slightly arched, central area rounded; striae 9–13 in 10 µm, becoming 13–18 toward ends, 2–5 isolated pores; 25–31 punctae in 10 µm.
According to
Specimen designated by
Cymbella tumidula var. salinarum (Grunow) Cleve, Kongl. Svenska Vetensk.-Akad. Handl., ser. 4, 26(2): 171, 1894.
Cymbella salinarum Grunow in Schmidt, A. Schmidt’s Atlas Diatom.-Kunde, Pl. 9, fig. 28, 1875.
Preparation 1603 in the Grunow Collection in the Naturhistorisches Museum Wien (W).
(designated here). An individual on preparation 1603, in the Grunow Collection in the Naturhistorisches Museum Wien (W), represented by the illustration in
Salinen von Zaule (Trieste, Italy).
Valves lanceolate, dorsiventral, dorsal and ventral margins convex; ends subcapitate; length 34.0 µm, breadth 10.7 µm, L/B ratio 3.2; axial area linear-lanceolate, slightly arched, central area rounded; striae 14 in 10 µm, becoming 15 toward ends, 1 isolated pore.
This taxon presents morphometric characteristics similar to C. tumidula var. tumidula, except it has wider valves.
Cymbella turgidula Grunow, in Schmidt, A. Schmidt’s Atlas Diatom.-Kunde, Pl. 9, Figs 23–26, 1875.
Plate 9, Fig. 23 in
Puerto Rico and Kahyenmathay.
An individual on slide 1504 in the Grunow Collection in the Naturhistorisches Museum Wien (W), designated by
Puerto Rico.
Valves lanceolate, dorsiventral, dorsal margins broadly convex and ventral margin convex; ends barely protracted, narrowly rounded, or subcapitate; length 36.5–45.0 µm, breadth 10.8–13.3 µm, L/B ratio 3.2–3.4; axial area linear, arched, central area indistinct to rounded; striae 9–12 in 10 µm, becoming 11–17 toward ends, 1–3 isolated pores; 21–24 punctae in 10 µm.
Cymbella tropica Krammer, Diatoms Europe 3: 61, Figs 44:1–10, 49: 12, 13, 2002.
Preparation 1015D IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
(designated here). An individual on preparation 1015D IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
Venezuela, Rio Manizanes, coll. Rumrich, 4.4.1990.
Valves lanceolate, dorsiventral, dorsal and ventral margins convex; ends barely protracted, rounded, or subcapitate, slightly deflected to ventral margin; length 34.5–42.7 µm, breadth 10.0–12.0 µm, L/B ratio 3.1–4.0; axial area linear, arched, central area indistinct to slightly rounded; striae 9–12 in 10 µm, becoming 11–12 toward ends, 1 isolated pore; 21–24 punctae in 10 µm.
Cymbella subturgidula Krammer, Diatoms Europe 3: 166, Figs 44: 19–21, 2002.
= Cymbella turgidula var. nipponica Skvortzov, Philipp. J. Sci. 61: 283, Figs 2:8, 4:4, 1936. (Figs
– Cymbella uenoi Skvortsov in Skvortsov & Noda, Sci. Rep. Niigata Univ., ser. D (Biol.) 8: 19, Pl. 3, Fig. 3, 1971. (nom. inval.)
= Cymbella rheophila Ohtsuka in Ohtsuka & Tuji, Phycol. Res. 50: 245, Figs 7, 8, 2002.
= Cymbella uenoi Skvortsov ex Tuji, Diatom 23: 50, Figs 20–25, 2007.
= Cymbella uenoi f. nipponica (Skvortsov) Tuji, Diatom 23: 54, Fig. 26, 2007.
Preparation 1046E IOK [not “1046c IOK”], in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany.
(designated here). An individual on preparation 1046e IOK, in the Alfred-Wegener-Institut für Polar- und Meeresforschung (BRM), Bremerhaven, Germany, represented by Fig.
Cymbella subturgidula Krammer 71–77 Specimens from Korea in preparation 1046E IOK, holotype 73 Lectotype, designated here 78–80 Specimens from Japan in preparation R 214.928 78Isolectotype, designated here, of C. turgidula var. nipponica Skvortsov [≡ C. rheophila Ohtsuka], heterotypic synonym of Cymbella subturgidula 81 Specimen from Japan in preparation R 214.930 82–84 Specimens from Japan in preparation R 214.932. Scale bar: 10 µm.
Korea, Ulchin County, Kyungsang Pukdo, Kwangchun River.
Valvae slightly lanceolate to lanceolate, dorsiventral, dorsal margin broadly convex and ventral margin straight to convex; ends barely protracted, subrostrate to broadly subcapitate; length 26.3–41.0 µm, breadth 9.0–13.5 µm, L/B ratio 2.3–3.6; axial area linear to linear-lanceolate, arched, central area indistinct to slightly rounded; striae 9–13 in 10 µm, becoming 12–15 toward ends, 1–3 isolated pores; 21–26 punctae in 10 µm. In SEM, the striae showed lineolate punctae externally and internally, the striae are composed by an alveolus internally, surrounded by thick costae; the isolated pores are rounded externally; internally, the alveoli of the isolated pores are irregularly obovate and connected to intercostae, the margins with tooth-like structures (brocca); one apical pore field (APF) not divided by the external terminal fissure of the raphe can be observed on each pole of the valvae; the terminal nodule extends to the dorsal side, under the APF and has a short branch that penetrates the APF apically; the helictoglossae lie under the terminal nodule and are deflected to the dorsal side.
Cymbella subturgidula and C. turgidula are closely related species. However, C. turgidula is more lanceolate and has a higher degree of dorsiventrality than C. subturgidula. Moreover, C. turgidula is slightly broader than C. subturgidula, with a more prominent ventral side of the valve. The ends in the two species are different, being subrostrate-rounded in C. turgidula and slightly subrostrate-truncate in C. subturgidula. The central area is more distinct in C. turgidula than C. subturgidula. Although the number of punctae in 10 µm is the same in both species, the striae in C. subturgidula seem to be more coarsely punctuated than in C. turgidula.
Cymbella turgidula var. nipponica was described by
Skvortsov and Noda (1971) described C. uenoi Skvortsov, but did not indicate any type. According to Article 40.1 of the ICN, names of new genera or taxa of lower ranks published after 1958 are valid only when the type is indicated (
The re-analysis of the type material of C. subturgidula allows us to broaden the metric data of this species compared to its original description. We observed a wider range of values for length (30.3–37.4 vs. 36–37 µm), breadth (9.0–12.8 vs. 10.0–11.0 µm), striae (9–11 vs. 10–11 in 10 µm) and punctae (21–24 vs. 24 in 10 µm) compared to
Few studies have discussed the criteria to delimit infraspecific ranks in diatoms.
However, taxonomic analyses of some species complexes in diatoms using molecular data have demonstrated that
The criteria adopted by
Because of the lack of taxonomic studies on the Cymbella affinis/tumidula/turgidula species complex using molecular data, we opted to use Krammer’s concept in order to attempt to organize this confusing group. This criterion is usual as a reference to circumscribe groups with similar morphologies, even if individuals with these morphologies sometimes overlap. Some authors prefer to define similar groups as “morphodemes”, which have no nomenclatural status. However, in some cases, this definition sounds more similar to the old taxonomy when any taxonomic unit could be denominated by different names, but scientifically by a long sentence that is more similar to the current “diagnosis”. Although the concept of varieties used here implies the publication of nomenclatural novelties, none of them are new proposals, but rather are simply adjustments of already existing names that are in confusing combinations, because of a misinterpretation by
The criteria of delimitation of taxa and the weight of characters in diatoms are variable from group to group (
In this context, several characters must be considered in their characterization and identification of taxa. The degree of dorsiventrality of the valve, for example, is slightly higher in C. affinis than in C. tumidula, and even higher in C. schilleri
Ultrastrutural characterizations by SEM are also important in diatoms, but not always possible using the original material. Records from the literature have demonstrated that representatives of the C. affinis/tumidula/turgidula species complex present similar internal ultrastructure of the isolated pores (i.e., aperture covered by small teeth) and the intercostae. This is the case of C. affinis (=C. excisa sensu
The economic and ecological uses of diatoms require a refined taxonomy, which is more detailed than simply species complexes. This is especially true for bioassessments using diatoms (
Naturally, beside the knowledge of types, supplementary studies are necessary to record phenotypic plasticity resulting from different ecological conditions or by life cycles. Such studies should be carried out in natural or cultivated samples in order to observe a more realistic concept of the species (
The process of lectotypification can markedly influence the identity of some taxa and can sometimes substantially change the relation to other taxa. The designation of a type for C. affinis resulted in a profound restructuring of C. affinis, C. excisa and C. tumidula. C. excisa has been shown to be the same taxon at the species level as C. affinis, but because of its specific morphology is treated herein at a different rank. The epithet affinis has priority over the epithet excisa, as defined by the criterion of the first effective publication. Thus, four infraspecific taxa of C. excisa were transferred to C. affinis.
The lectotypification of C. tumidula Grunow and comparisons with the lectotype of C. affinis allowed us to conclude that the two species are independent. C. affinis var. procera was treated as a new species, which is closer to C. tumidula than C. affinis because of morphological similarities. Infraspecific taxa described by
The analysis of the type and the history of taxa such as C. subturgidula Krammer, C. rheophila Ohtsuka, and C. uenoi Skvortsov ex Tuji allowed us to conclude that these taxa are conspecific, and to determine that the epithet subturgidula has priority.
The authors thank Jovita Yesilyurt, from the Natural History Museum, London, and Taisuke Ohtsuka, from the Lake Biwa Museum, Kusatsu, Shiga Prefecture, who provided the material for analysis; and the Laboratório de Microscopia Eletrônica do Museu Nacional for the SEM analyses. The first author is grateful to CAPES and DAAD/CAPES for the award of a doctoral scholarship and a “sandwich” scholarship for study at the BGBM, Berlin, Germany.