Diatom Research (1996) Volume 11 ( 2 ) ,241-260

CYCLOTELLA STRZATA COMPLEX:

TYPIFICATION AND NEW COMBINATIONS

Hannelore HBkansson

Department of Quaternary Geology, University of Lund,

Tornavagen 13, S-223 63 Lund, Sweden

This morphological investigation deals with taxa in the Cyclotella striata complex. The following
species and varieties were examined: Coscinodiscus striatus, Coscinodiscus minutus, Cyclotella
striata var. baltica, C. striata var. ambigua, C. striata var. mesoleia, C. (striata var.?) subsalina, C.
dallasiana and also Discoplea sinensis. The study also utilized material, drawings and annotations
from the collections of Ehrenberg, Kiitzing, Grunow, Eulenstein and Cleve & Moller; and, where
possible, light and electron microscope studies were made of unmounted original material.
The results show structural differences between species. A lectotype has been selected for
Coscinodiscus striatus Kiitzing (Cyclotella striata (Kiitz.) Grunow) and for Coscinodiscus minutus
Kiitzing, but the name Cyclotella minuta is already occupied and a new name for this species, C.
exigua nom. nov., has had to be chosen. C. striata var. baltica is shown to be a species separate
from C. striata and to be conspecific with C. litoralis Lange et Syvertsen; the correct name for this
species is thus C. litoralis. C. dallasiana W . Smith and Discoplea sinensis Ehrenb., are synonymous
with C. striata, while it is shown that C. ambigua Grunow is a species in its own right.

INTRODUCTION

During recent years the centric diatoms of saline lakes and estuaries have begun to receive
greater attention, especially the genus Cyclotella (Kiitz.) Brtbisson (e.g. Lange & Syvertsen 1989,
Wendker 1991, Prasad et al. 1990, Hikansson et al. 1993, HAkansson & Kling 1994). Detailed
studies of ultrastructure have been published, comparisons between similar species made (e.g.
Takano 1976, Battarbee et al. 1984, HBkansson 1982) and new names established (Lange &
Syvertsen 1989).
The very brief descriptions and poor pictures given by earlier researchers are frequently the
cause of misunderstanding and misidentification (Ehrenberg 1841, 1854, Smith 1856, Grunow in
Cleve & Grunow 1880, Grunow in Van Heurck 1880-1885). Nowadays with light and electron
microscopy we are able to discern the finest detail of the morphological structure of diatoms.
However, there is still very little known about the variability of many taxa during the life cycle.
There are also different opinions about the importance of some morphological characters for
differentiating taxa of centric diatoms, such as number and position of the valve face and mantle
fultoportulae and their satellite pores, the height of the mantle, the structure of the striae, and the
placement and internal orientation of the rimoportula. However, our understanding of these
morphological features in taxa of the genus Stephanodiscus Ehrenberg has increased (HAkansson &
Meyer 1994).
242 H. HAKANSSON

It seems that in Cyclotella some morphological features are more variable than has hitherto
been assumed and so a number of taxa may have to be reduced to synonyms. Meyer & Hikansson
(1996) showed that during the life-cycle taxa may have no stability in the number and position of the
valve face fultoportulae and the internal orientation of the lips of the rimoportula. Furthermore the
two valves of a single frustule can have quite different morphological features.
According to older literature (Grunow in Cleve & Grunow 1880, Grunow in Van Heurck 1880-
1885, Schmidt 1874-1959, Hustedt 1927-1930, Cleve-Euler 1951) the differentiation of the varieties
of C. striatu depends on the density and length of the marginal striae, the number of valve face
fultoportulae, and the presence or absence of granulation on the central area of the valve face. In this
paper an attempt is made to investigate some of the taxa in the Cyclotella striatu complex, to select a
type for C. striata, and, with help of available type material as well as the material newly collected
from Cuxhaven, clarify the taxonomy of this difficult group.

MATERIAL AND METHODS

The following material and slides have been investigated:

1) The slides BM 19362 and BM 19363 (marked ‘Type’) labelled Coscinodiscus rninutus and
material from packet 825 in the Kutzing Collection at the Natural History Museum, London (BM)
and in the Eulenstein Collection at the Royal Zoological Society in Antwerp (AWH).
2) Cyclotella striutu (Kutz.) Grunow, slide 905, “Hafenschlamm von Cuxhaven”, in the Grunow
Collection, Naturhistorisches Museum, Wien (W).
3) Rabenhorst exsiccatum 1697 (Rabenhorst 1864): ‘Coscinodiscus striutus Ehrbg. Ktz. Bacill. pag.
131. T.l. fig. VIII. Flor. Eur. p. 34. Auf Holzwerk welches den Sommer uber unter Wasser gelegen
hat. Oct. 1863 im Kieler Hafen. ges. von Joh. Luders’. Collection at the Botanical Museum, Lund,
Sweden (LD).
4) C. striutu var. baltica Grunow (Grunow Collection 2241, W) with material from the Baltic.
According to Grunow’s notebook (kept in the Grunow Collection at W, in which he listed all the
diatoms found on a slide, with his personal comments): ‘Im Kieler Hafen auf Holzwerk, welches im
Sommer unter Wasser (Rabenhorst 1864, Alg. Eur. no. 1697); Cyclotella striatu var. baltica m, viel’.
5 ) C. striata var. arnbigua Grunow (Grunow Collection 2030a and b, W) with material from Jenissey.
According to Grunow‘s notebook: ‘Jenissey Korepovskoj 1876’.
6) C. striata var. rnesoleia Grunow (Grunow Collection 2230, W) with material from Delaware.
According to Grunow’s notebook: ‘C. striata var. genuina, Dallasiana, baltica, etc.’ .
7) C. (striata var.?) subsalina Grunow (Grunow Collection 676, W) with material (according to
Grunow’s notebook) from ‘Fossile Vaucheria, Greenwich Pier bei London, Jahr 1861’.
8) Cyclotella dallasiana W. Smith, BM 23192 “March 22, 1854, Medway EWD’.
9) Slide 178 from the Cleve & Moller Slide Collection (1879). Collection at the Swedish Museum of
Natural History, Stockholm, Sweden (S).
10) Discoplea sinensis Ehrenberg from the Ehrenberg Collection, Museum fur Naturkunde der
Humboldt-Universitat zu Berlin, Germany (BHU) “Blumenerde von Canton, China”.
 CYCLOTELLA STRIATA COMPLEX 243

11) Recent material from the North Sea (Cuxhaven), Germany. Collected by S. HAkansson near the
harbour. Slide Ld 443 in the H&ansson Collection, Department of Quaternary Geology, Lund,
Sweden (LD).
Unfortunately very little raw material from the Kutzing Collections (and none from the Grunow
Collection) was available. Only measurements and light micrographs could be made from the slides.
Material from Rabenhorst exsiccatum 1697 and packet 825 (from Kutzing’s and Van Heurcks
Collections), Ehrenberg’s sample of Discoplea sinensis and the recent material were prepared for
light microscopical (LM) and scanning electron microscopical (SEM) investigation as described by
HAkansson (1984).

OBSERVATIONS

Some of the following descriptions are less than ideal, because of missing EM detail, but even
so it is possible to solve some of the nomenclatural and taxonomical problems.

Coscinodiscus striatus Kutzing and C. minutus Kutzing (see Figs 1-24)

No sample in Kutzing’s herbarium in the Natural History Museum, London (BM) was labelled
by him as containing either Coscinodiscus striatus or C. minutus. However, his sample no. 825,
which was in his possession in 1844, is labelled in his handwriting ‘Verschiedene Arten von
Coscinodiscus. Cuxhaven’ and also ‘minutus Kg.’ in Eulenstein’s handwriting. The slides BM 19362
and BM 19363 were prepared from this sample, and there is also part of it in the Van Heurck Diatom
Collection, AWH. Whether the slide 905 in Grunow’scollection in W, came from the same gathering
by Sonder or from another is uncertain.
However, the most important question arising is: which species had Kutzing seen, when
describing Coscinodiscus minutus and C. striatus from the same locality “Cuxhaven” (Kutzing 1844,
p. 131, Taf. I, figs VIII and XIV; reproduced as Figs 8 and 1 respectively). The main difference
seems to be the diameter, C. minutus being smaller, with a diameter of 18 pm, compared to C.
striatus with 56 pm. This might indicate that they are based on specimens from opposite ends of the
size range of the same species or that two different species are present.
Amongst other centric diatoms (Coscinodiscus, Actinocyclus, Paralia) on slide BM 19363, I
found a few Cyclotella species with a morphology typical of the genus - a differently structured
central area compared to the striated marginal area - exactly as Kutzing showed (op. cit.). I assume
that some of them (Figs 2-7) are what Kutzing (1844, p. 131, Taf. I, fig. XIV; Fig. 1) had seen when
he prepared his very brief description of Coscinodiscus minutus. In his description, however, Kutzing
(op. cit.) gives only one diameter, 18 pm, whereas I found a few specimens on slide 19363 with
diameters between 12.5 and 18 pm. They all have the same appearance: the central area is
tangentially undulate with a strongly colliculate structure, a valve face fultoportula is present, even if
difficult to find with certainty (Fig. 2 arrowed a), and 8-9 striae in 10 pm of even length. It was also
difficult to see the exact position of the mantle fultoportulae. I detected at least one rimoportula (Fig.
2, arrowed b). A tilted specimen (Figs 5 , 6), even if smaller (only c. 13.5 pm) shows the short mantle
and that the costae are straight and all of the same length (no recessed ones).
The name Cyclotella minuta, however, cannot be used for this species because of the existence
of Cyclotella minuta (Skvortzov) Antipova (Antipova 1956). Since no later name can be found that
can be applied with certainty to this form, the name Cyclotella exigua is proposed at the end of this
paper.
244 H. HAKANSSON

Figs 1-7. Cyclotella exigua. Fig. 1 . Reproduction of Kutzing’s (1844, Taf. 1 , fix. XIV) drawing at twice the
original size; i.e. 840/1. Figs 2 4 . BM 19363; lectotype, different foci. Figs 5,6. BM 19363; another specimen on
the same slide as the lectotype, different foci. Fig. 7. Scanning electron micrograph on a whole frustule (packet
825). Photo taken at Natural History Museum, London. Scale bars represent 10 p(Figs 2-6), 5 pm (Fig. 7).

Unfortunately Coscinodiscus minufus could not be found in material from the packet 825 (from
both Kutzing’s and Eulenstein’s Collection) prepared for SEM, In SEM pictures kindly sent by Miss
Patricia Sims (BM) the diatoms are covered with debris and it is therefore very difficult to be certain
about any detail (Fig. 7).
On the same slide (BM 19363) there is a larger specimen with a diameter of c. 35 pm, (Figs 9,
lo), which agrees with the brief description and figure of Coscinodiscus striatus given by Kutzing
(1844, p.131, Taf. I, fig. VIII; reproduced as Fig. 8) except for the diameter which should be 56 pm.
Kutzing also gave only one measurement for this species. This species also clearly belongs to the
genus Cyclotella. It has a tangentially undulate, colliculate central area and a striated marginal zone,
but the marginal striation differs from that of C. exigua (compare Figs 2-4 with Figs 9, 10). The
interstriae (or costae) are pronounced, and each has an elongate “hollow” (alveolus) in the middle of
its length (Figs 9, 10). In Fig. 9 (arrowed a) the mantle fultoportulae are visible at every second or
third (seldom at every fourth) costa. The arrow (b) on Fig. 9 and on Fig. 10 (with a questionmark)
possibly shows the position of the rimoportula. There are several artefacts on the valve face (debris or
mountant?) which make it impossible to be certain whether valve face fultoportulae are present. I
could not find C. striatu in material from Kiitzing’s packet 825 either in London or Antwerp.
 CYCLOTELLA STRIATA COMPLEX 245

Figs 8-12. Cyclotella striata. Fig. 8. Reproduction of Kiitzing’s (1844, Taf. 1, fig. VIII) drawing at twice the
original size; i.e. 840/1. Figs 9, 10. BM 19363, lectotype, different foci. Figs 11, 12. Grunow slide 905
(material from Cuxhaven); different foci. Scale bars represent 10 pm.

Coscinodiscus striatus is the species Grunow (in Cleve & Grunow 1880) transferred to
Cyclotella as C. striata. He also made a clear differentiation between C. striata and his newly
described C.ambigua (Figs 40-44). According to Grunow, C. striata has markings on the valve face
(now known to be fultoportulae) organized in a semicircle. By studying a great number of samples
from different localities, however, he found that the pattern of the semicircular “crown” of puncta
and the development of spinulae at the valve facehalve mantle junction are variable. Grunow
246 H. HAKANSSON
 CYCLOTELLA STRIATA COMPLEX 247

thought therefore that it would be better to make C. ambigua a variety of C. striatu (Grunow in Cleve
& Grunow 1880, p. 119), and, although he did not make the appropriate combination there, he did so
later (Grunow in Van Heurck, 1882: pl. 92. fig. 12).
According to Grunow’s notebook, slide 905 is from Cuxhaven, as is Kutzing’s material.
Grunow’s notebook drawing is reproduced as Fig. 13, and his drawing in Van Heurck 1882 (pl. 92,
fig. 6) as Fig. 14. Slide 905 contained several different CycZoteZZu species, some of which have valve
face fultoportulae while others apparently do not (Figs 15-17). Valves with fultoportulae nearly
forming a semicircle on the valve face are present, but these have a different structure in the
marginal, striated area (Fig. 16). There are, however, some specimens with the characteristic
morphological structure that I found on Kutzing’s slide BM 19363 (compare Figs 9, 10 with Figs 11,
12), with a diameter between 25 and 49 pm (one example had a diameter of 67 pm). It is possible
that Grunow had confused C. striutu with what he later (in Van Heurck 1882 called C. striutu var.
bulticu and this will be discussed later.
For SEM investigation, I collected new material from Cuxhaven (Figs 18-24). This material
contained the same characteristic CycZoteZZa species I had found on Kutzing’s and Grunow’s slides:
the central area of the valve face is tangentially undulate and colliculate, while the marginal striated
area shows the typical dark or light (depending on focus) elongate alveoli (compare Figs. 18, 19 with
Figs 9, 10). The diameter is between 28 pm and 43 pm with 8-10 striae in 10 pm.
The external view of the valve in SEM shows the typical central colliculate structure, while the
marginal striation consists of elevated interstriae (costae) and striae in the depressions (Fig. 20). Short
spinulae are present at the valve facehalve mantle junction (Fig. 21). Both the interstriae and the
striae continue onto the relatively short mantle (Fig. 21). On the mantle just below the spinulae are
the openings of the mantle fultoportulae, on every second, third or fourth interstria (Figs 20-21).
Between two of these is the rimoportula (Fig. 21 arrowed), with a slightly larger opening than those
of the fultoportulae. All external openings have a thickening around them. Valve face fultoportulae
are found, although very difficult to discern due to debris. The internal view of the SEM shows that
the central area is smooth and, apart from fultoportulae, unstructured. The striation at the margin has
both equal and more or less distinctly recessed costae (Figs 22-24). All the fultoportulae have three
satellite pores (Fig. 24). The rimoportula has an unusual shape (Fig. 23): a thickened base bears a
broad lip orientated perpendicular to the costa.

Cyclotella striata var. baltica Grunow (Figs 25-39)

Grunow never published a description for C. striutu var. bulticu; only pictures (in Van Heurck
1882, pl. 92, figs 13-15; reproduced as Fig. 25). In a copy of Van Heurck’s Atlas (op. cit.) at W, the
slide number to those figures is given as 2241. According to Grunow’s annotation to this slide in his
notebook, this is the same material as Rabenhorst’s exsiccatum 1697 (compare Figs 26-31 with Figs
32, 33). Rabenhorst called this species Coscinodiscus striutus Ehrbg. Ktz. (‘Ehrbg.’ because
Ehrenberg had described the genus Coscinodiscus); Grunow wrote in his annotations ‘CycloteZlu
striutu var. bultica m, vie]’.

Figs 13-19. Cyclotella striatu. Fig. 13. Reproduction of Grunow’s drawing (“Bilder-Sammlung” No. 9304).
Fig. 14. Reproduction of figure in the Van Heurck (1882, pl. 92, fig. 6) as C. striata (Kiitz.) Grunow; original
size. Figs 15-17. Different Cyclotella species on Grunow’s slide 905. Figs 18, 19. Newly collected material
from Cuxhaven (type locality). Scale bars represent 10 pn.
248 H. HAKANSSON

Figs 20-24. Cyclofellu striufu. Scanning electron micrographs of material from Cuxhaven, type locality. Fig.
20. Exterior view of a whole frustule with spinulae at the valve facehalve mantle juntion. Fig. 21. Detail with
the openings of the mantle fultoportulae and the opening of the rimoportula (arrowed). Fig. 22. Interior view of
a valve; note the recessed costae bearing the mantle fultoportulae. Figs 23, 24. Detailed interior view with the
rimoportula (Fig. 23) and the mantle fultoportulae with three satellite pores on recessed costae (Fig. 24). Scale
bars represent 5 km (Figs 20-23), 1 Frn (Fig. 24).
 CYCLOTELLA STRIATA COMPLEX 249

This is a diatom with a relatively wide size range: 11-45 pm, and 9-12 striae in 10 pm. It has a
pronounced tangential undulation of the central area with a colliculate structure (Figs 27, 29, 31, 32).
Part of the marginal striation is included in the undulation (Figs 34-36). On the elevation of the valve
face are 2-7(8) fultoportulae organized in a semicircle (Figs 26, 27, 32, 34-36). The striated marginal
area consists of striae and interstriae continuing onto the mantle. Granulae more or less covering the
valve can be observed. On every second to third interstria (or costa) near the edge of the valve are the
openings of the mantle fultoportulae (Figs 35, 36). A single rimoportula is located slightly above the
ring of mantle fultoportulae (Fig. 36, arrowed). Externally this has a simple opening; internally it is
like an elongated slit protruding towards the centre (Figs 37, 39, arrowed). The valve face
fultoportulae have three satellite pores and the mantle fultoportulae two, orientated radially (Figs 37-
39).
This diatom is similar to Cyclotella litoralis Lange & Syvertsen. Lange & Syvertsen (1989)
made a very detailed investigation and compared their new species with both C. striata and C.
stylorum. The size range they recorded for C. litoralis is wider (10-60 pm) than that found in C.
striata var. baltica, with 2-20 valve face fultoportulae and 9-14 striae in 10 pm. There is also a
minor difference in the arrangement of the mantle fultoportulae. In C. litoralis they occur on every
costa or every second one; in C. striata var. baltica, however, they are located every second or third
costa.
C. striata and C. striata var. baltica differ in the marginal structure, the arrangement of the
valve face fultoportulae, and the position and number of the satellite pores belonging to the mantle
fultoportulae. These two taxa should not, therefore, be combined within one species. The differences
between C. striata var. baltica and C. litoralis described above are minor and may depend on the size
range and on environmental effects (e.g. of salinity), and so I consider C. striata var. baltica and C.
litoralis to be conspecific. Thus according to the Botanical (Tokyo) Code (Art. 11:2; Greuter et al.
1994), which states that a name has no priority outside its rank, the name C. litoralis has to be used.
It seems that this diatom is often misidentified as Cyclotella striata because of Grunow’s
statement (in Cleve & Grunow 1880, p. 119) that the central area of the valve face in C. striata
should contain fultoportulae arranged in a semicircle. Hustedt (1927-30), Cleve-Euler (1951),
Helmcke & Krieger (1954), Helmcke et al. (1974), Takano (1976) and Prasad et al. (1990) all
described diatoms as Cyclotella striata, that in reality were C. litoralis (= C. striata var. baltica).

Cyclotella striata var. ambigua (Grunow) Grunow (Figs 4 0 4 4 )

The material on slides 2030 a and b is from Jenissey, which Grunow in Cleve & Grunow (1880,
p. 119) gave as the locality for C. ambigua. He described C. ambigua in this publication, compared
his species with C. meneghiniana and C. striata, and said at the end of his article that it would
possibly be better if C. ambigua were to be considered as a variety of C. striata. As mentioned
before, he did not make the appropriate combination until later (Grunow in Van Heurck 1882, pl. 92,
fig. 12). In his notebook he called this taxon ‘C. striata var. ambigua m’. In an annotated copy of
Van Heurcks Atlas (1880-1885) at the Naturhistorisches Museum, Wien, the slide 2863 is cited for
plate 92, fig. 12, but this must be wrong: according to Grunow’s notebook this is material from
Nimrod Sound and no Cyclotella species is mentioned. On drawing 9308 (reproduced as Fig. 41) in
the Grunow Collection (W) he himself indicated that this is the taxon he described and illustrated in
Cleve & Grunow (1880, fig. 133; my Fig. 40). He also gives the co-ordinates (both on the drawing to
slide 2030 b and on slide 2030 a), to be sure that this species could be found. Unfortunately the
millimetre paper Grunow so carefully put around the coverslips on slides 2030a and b has been lost. I
250 H. HkCANSSON
 CYCLOTELLA STRIATA COMPLEX 25 1

investigated the slides and found them to contain freshwater diatoms, including a Cyclotella species
which agrees with Grunow’s picture (Figs 42-44). My measurements (18-34 pm diameter; 9-10
striae in 10 pm) are only slightly different from Grunow’s (22-31 pm and 9 striae in 10 pm). I also
found that there were 1 or 2 valve face fultoportulae on the specimens (Fig. 43). It was sometimes
difficult to see these, however, because of the bad condition of the slides. Opposite the valve face
fultoportulae the rimoportula is visible as a slight thickening of one costa (Figs 42 and 46, arrowed).
The hyaline “ring” on the striation visible in the three figures (Figs 42-44) is possibly caused by the
smooth inside siliceous layer covering the chambers. It is very difficult to see the position of the
mantle fultoportulae. There was no material available to make a detailed SEM investigation.
In the literature (Grunow in Van Heurck 188&1885, Fricke in Schmidt 1874-1959, Hustedt
1927-1930, Cleve-Euler 1951), this taxon was given as a variety of C. striata. However, as shown in
this investigation, C. ambigua has no similarity to C. striata and should be kept as a species in its
own right: C. ambigua Grunow.

Cyclotellu striata var. mesoleia Grunow, C. striata formae minores and C. striata var. intermedia
Grunow (Figs 45-53)

Slide 2230 in the Grunow Collection, representing material from the Delaware river mouth
contains several taxa of the striata complex. According to the legend of the figures in Van Heurck
(1882, pl. 92, figs 7-9; reproduced as Figs 45, 46) these should include C. striuta var. mesoleia and
C. striata formae minores (possibly meaning “small” forms of C. striata). In his notes to the slides
themselves, however, Grunow mentioned ‘C. striata var. genuina, Dallasiana, baltica etc.’. The slide
is thickly covered with material, several specimens lying over each other, which makes it very
difficult to know which of the taxa are which (Figs 48-53). Nearly all the valves found had valve
face fultoportulae, but there were often differences in the morphology of the marginal striation.
Comparing Grunow’s figures (in Van Heurck 1882, pl. 92, figs 7-9; my Figs 45, 46) with the
figures given here (Figs 48-53) it could very well be that these are the diatoms Grunow had seen and
called C. striata var. mesoleia and formae minores of C. striata.
The figures given by Grunow (in Van Heurck, op. cit.) all show the valve face fultoportulae
arranged in a semicircle. According to Grunow (in Cleve & Grunow 1880) this is a typical character
of Cyclotella striata, but none are visible on the figure of C. striata forma major Grunow (probably
meaning “large” forms of C. striata) (in Van Heurck 1882, pl. 92, fig. 6; my Figs 13, 14).
Grunow validly published C. striata var. intermedia in Van Heurck’s Atlas (1882, pl. 92, fig. 10;
reproduced as Fig. 47). In an annotated copy of this Atlas (at W), however, no slide number is given,
only “Moller’sTypenplatte”. Nothing more could be found in Grunow’sCollection. Unfortunately the
original drawing (in Van Heurck 1882, pl. 92, fig. 10; my Fig. 47) is insufficient to serve as a basis
for a distinction between C. striata var. intermedia and similar species in the C. striata complex. C.
striata var. intermedia must therefore be treated as “species inquirenda” (i.e. needing further study).

Figs 25-30. Grunow‘s slide 2241 “Cyclotella striata var. baltica”. Fig. 25. Reproduction of figures in Van
Heurck (1882, pl. 92, figs 13-15) at twice the original size. Figs 26-30. Different specimens on Grunow’s slide
2241 (light micrographs). Figs 3 1-33. Light micrographs on material from Rabenhorst’s exsiccatum 1697. Figs
34 and 35. Scanning electron micrographs of Rabenhorst’s exsiccatum 1697. Fig. 34. Exterior of a whole
frustule. Fig. 35. Detail of Fig. 34 with the valve face fultoportulae positioned in a semicircle (arrowed). Scale
bars represent 10 pm (Figs 26-34), 5 pm (Fig. 35).
252 H. HAKANSSON

Figs 36-39. Scanning electron micrographs of Rabenhorst's exsiccatum 1697. Fig. 36. External view of a whole
frustule with the external opening of the rimoportula (arrowed). Fig. 37. Internal view with the labium of the
rimoportula (arrowed). Fig. 38. Tilted specimen showing the position of the mantle fultoportulae with two
satellite pores. Fig. 39. Detail of a valve with the rimoportula arrowed. Scale bars represent 5 pm (Fig. 36),
1 pm (Figs 37-39).

As can be seen so far in this investigation several of the taxa in the C. striata complex have
valve face fultoportulae. So have members of the C. meneghiniana complex, but these do not have a
colliculate structure in the central area of the valve face. But the most important differentiating
characters in both complexes lies in the marginal area of the valve face: the type of striation, the
position of the mantle fultoportulae, the position and number of the fultoportula satellite pores, and
the position and structure of the rimoportula. Only these morphological features can give us certain
identification. Therefore more detailed investigation is needed for the diatoms which Grunow called
C. striata var. mesoleia and C. striata formae minores.
 CYCLOTELLA STRIATA COMPLEX 253

Figs 40-44. Cyclotella ambigua. Fig. 40. Reproduction of Grunow’sfigure 133 in Cleve & Grunow (1880). Fig.
4 1. Reproduction of Grunow’s drawing (“Bilder-Sammlung” 9309). Figs 42-44. Light micrographs of C.
arnbigua from Grunow’s slides 2030 a and b. Note the two valve face fultoportulae on Fig. 43 (mowed) and on
Fig. 44 the position of the rimoportula. Scale bars represent 10 pm.

Cyclotellu (striata var.?) subsalina Grunow (Figs 54-56)

A taxon found on Grunow’s slide 676, containing material from ‘Greenwich Pier bei London,
Jahr 1861’, shows very great similarity with the figure given in Van Heurck (1882, pl. 92, fig. 11;
reproduced as Fig. 54), and also with Grunow’s drawings, where he indicated that this taxon has 8-9
striae in 10 pm.
The valves (Figs 5 5 , 56) are nearly flat and do not exhibit colliculate structure in the central
area of the valve face, unlike the previous taxa. The marginal striae are short. Very few specimens
were present on the slide, which was, however, crowded with other diatoms. I could not find any
valve face fultoportulae. The diameter is c. 11-15 pm with 8-10 striae in 10 pm.
This taxon also requires more detailed investigation using SEM before its identity can be
established.
254 H. HAKANSSON

Fig. 45. Reproduction of figures (C. striuta formae minores) in Van Heurck (1882, pl. 92, figs 7, 8): original
size. Fig. 46. Reproduction of figure (C. striutu var. mesoleia) in Van Heurck (1882, pl. 92, fig. 9); original
size. Fig. 47. Reproduction of figure (C. striatu var. intermedia) in Van Heurck (1882, pl. 92, fig. 10); original
size. Figs 48-53. Light micrographs of Grunow's slide with material from the Delaware river mouth (different
species in the C. striatu complex). Note the valve face fultoportulae in a semicircular ring in Figs 48-50. Scale
bars represent 10 pm.

Cyclotella dallasiuna W. Smith (Figs 57,58)

It has sometimes happened that diatom taxa have been described from just one specimen. This
was the case when W. Smith (1856) described Cyclotella dallasiana (Figs 57 and 58 -kindly taken
 CYCLOTELLA STRIATA COMPLEX 255

Fig. 54.Reproduction of figure (C. striata (var. ?) subsalina Grunow in Van Heurck (1882, pl. 92, fig. 1 I);
twice original size. Figs 55, 56. Light micrographs of Cyclotella subsalina, Grunow slide 676. Figs 57, 58. BM
23192, different focus on Cyclotella dallasiana W. Smith (Photo: Mr. P. York, Natural History Museum,
London). Fig. 59. Slide 178 from the Cleve & Moller slide Collection (1879). Figs 60, 61. Scanning electron
micrographs of Ehrenberg's material Discoplea sinensis (= Cyclotellu sinensis (Ehrenb.) Ralfs). Note the
recessed costae and the unusual shape of the rimoportulae (arrowed). Scale bars represent 10 pm (Figs 55-59),
5 pm (Fig. 61), 3 pm (Fig. 60).
256 H. HAKANSSON

by Mr Peter York are of this specimen on slide BM 23192 at the BM). Smith gives a diameter of
0.0022” with a costa-length of 0.0002“ (= 56 pm and 5.1 pm). This species has the same
characteristic morphology, the colliculate central area and the typical marginal striation as C. striata,
so I think (as did Grunow in Van Heurck 1882) that C. dallasiana is conspecific with C. striata
(compare Figs 57,58 with Figs 9, 10).

Cyclotella “dallasiana” and C. sinensis (Ehrenb.) Ralfs in Pritchard (Figs 59-6 1)

The taxon found on slide 178 in the Cleve & Moller slide series (1879: material from Elephant
Point, Bengal; Fig. 59), is said to be Cyclotella dallasiana Smith, but looks different in LM (compare
Figs 59 with Figs 57, 58). It is more similar to a diatom found in Ehrenberg’s material ‘Blumenerde
von Canton, China’ under the name Discoplea sinensis (the basionym of Cyclotella sinensis
(Ehrenberg) Ralfs in Pritchard 1861; Fig. 60), which Hikansson (1986) transferred to C. striata. As
can be seen from Fig. 61, the inside view of this taxon shows the same morphological features in the
marginal area of the valve face as C. striata (Figs 22, 23). Both taxa have recessed costae, similar
mantle fultoportulae and the same form and orientation of the rimoportula. It cannot be stated with
certainty that C. “dallasiana“found on Cleve & Moller slide 178 is conspecific with C. sinensis, but
C. sinensis is certainly conspecific with C. striata, as Hustedt (1927-1930) and Hikansson (1986)
have already stated.

DISCUSSION

Our knowledge of morphological variation in Cyclotella has increased during recent years
(Battarbee et al. 1984, Hikansson 1989, 1990, Meyer & Hikansson 1996). We are able to distinguish
much finer morphological features and the differences in these between taxa; it has also been found
that a single morphological character is generally insufficient to differentiate between taxa, and
several characters in combination are needed. Many characters can only be seen using SEM.
As Cox (1995) has pointed out, we must remember, when interpreting the literature of the last
century, that the early workers did not have the optics we have in our present microscopes and that
they were not able to interpret accurately and consistently the morphological characters now known
to be important for differentiating taxa. It is therefore not at all surprising that since the first
descriptions of many taxa, very different opinions have been expressed in the literature about their
circumscription (e.g. Hustedt 1927-30, Cleve-Euler 1951, Helmcke & Krieger 1954, Gerloff &
Helmcke in Helmcke et al. 1974, Takano 1976, Lange & Syvertsen 1989). C. striata has been
compared with C. meneghiniana (Grunow in Cleve & Grunow 1880, Helmcke & Krieger 1954, plate
118, Hustedt 1957, Hasle 1962, Hikansson 1982). The identification of these centric taxa is often a
problem, especially in lakes of high conductivity, inland salt lakes, or in estuarine sediment studies (
where there may be a mixture of freshwater and marine taxa) and ecology has often guided the
choice of name.
The diatoms investigated from the Cyclotella striata complex clearly do belong to the genus
Cyclotella as defined by Round (1970), Round et al. (1990) and Round & Hikansson (1992). In valve
view the valve face is clearly divided into a marginal striated area and a central area which is
variously ornamented.
Some members of the C. striata complex exhibit a fine structure that has not been described
before. All have a similar morphology of the central area of the valve face, which is strongly
tangentially undulate and has a colliculate structure, visible even in immature or corroded specimens
 CYCLOTELLA STRIATA COMPLEX 257

(see Figs 34-38). Most of them have valve face fultoportulae, although these may be difficult to
discern; and it seems that C. subsalina has none (Figs 53, 54), although this needs to be confirmed
because only a few specimens have been found and no SEM investigation was possible. It is also
unclear whether the taxon found on Cleve & Moller slide 178 under the name C. dallasiana (Figs 55,
56) has any. C. striata, C. exigua H&ansson (see below) and also C. ambigua Grunow have at least
one valve face fultoportula. In C. litoralis Lange & Syvertsen (Lange & Syvertsen, 1989) the valve
face fultoportulae are organized in a semicircular ring.
The most important morphological criteria are the colliculate structure of the central area (that
can, however, sometimes be “missing”), the presence or absence of valve face fultoportulae, the
position of the costae (recessed or not), the number and position of the mantle fultoportulae, the
number and position of the satellite pores, and the position of the rimoportula (some of these are only
visible in SEM investigations). However, none of these characters are diagnostic on their own. It is
the combination of characters which distinguishes taxa. This has been stated already by Geissler
(1970) and is ever more evident in the literature.

Descriptions and typification

New and more detailed investigations are necessary for several taxa in the C. striata complex:
these include C. striata var. mesoleia, C. striata var. intermedia and C. subsalina. C. striata var.
intermedia is treated as “species inquirenda” because of missing material. For the other taxa involved
the following descriptions can be given:

Cyclotellu exiguu HIkansson nom. nov.

Synonym: Coscinodiscus minutus Kutzing 1844, p. 131, Taf. 1, fig. XIV
Type locality: Cuxhaven, Germany
Lectotype: Slide BM 19363, Kutzing Collection, BM, designated by H. Hbkansson.
Valve circular, central area tangentially undulate and colliculate, the marginal area striate.
Diameter c. 12.5-18(24) pm with c. 8 striae in 10 pm and a relatively short mantle. One valve face
fultoportula with a single rimoportula close to the opposite margin. The rimoportula is positioned
slightly more towards the valve face than the ring of mantle fultoportulae, which seem to be placed
on every second or third costa.

CycZoteZZu striutu (Kutz.) Grunow in Cleve & Grunow 1880, p.119.

Basionym: Coscinodiscus striatus Kutzing 1844, p. 131, Taf. 1, fig. VIII.
Synonyms: Discoplea sinensis Ehrenberg 1848, p. 484; 1854, pl. 3913 (2), fig. 30; Cyclotella sinensis
(Ehrenb.) Ralfs in Pritchard 1861, p. 812, Cyclotella dullasiana W. Smith 1856, p. 87.
Type locality: Cuxhaven, Germany
Lectotype: Slide BM 19363, Kutzing Collection, BM ,London, designated by H. Hikansson.
Cells drum-shaped to shortly cylindrical. Valve circular, strongly tangentially undulate,
especially in the central area. Valve face sharply divided into a colliculate central area and a striated
marginal area. Diameter (20)25-67 pm with 8-10 striae in 10 pm. Striation (c. 5.5-6 pm long)
consisting of striae and interstriae. Mantle relatively short. At the valve facehalve mantle junction a
short thick spine (the smaller dots mentioned before) is visible on every interstriae; beneath the
spines, on every third or fourth interstria are the openings of the mantle fultoportulae. Between two
of the mantle fultoportulae lies the slightly bigger opening of the rimoportula. The striae consist of
one row of small areolae which are surrounded by larger ones on either side. Internal views show
258 H. HAKANSSON

short chambers, which are seen in LM as “elongated hollows”. It is difficult to discern the number of
satellite pores, but there are three. A single rimoportula is present with a short tubulus which opens
into a very broad “mouth”.

Cyclotella litoralis Lange & Syvertsen 1989, p. 3431344, figs 1-30.

Synonyms: Coscinodiscus striatus sensu Rabenhorst 1864, exsiccatum no. 1697; Cyclotella striata
var. baltica Grunow in Van Heurck 1880-1885, pl. 92, figs 13-15
Type locality: South Atlantic, 34” 21‘ S, 52” 16‘W.
Holotype: BM 81403, BM, London.
Isotype: IMBB (Dep. Biology, Mar. Bot. Univ. Oslo).
Cells drum-shaped. Valve circular, strongly tangentially undulate, especially in the central area.
Valve face sharply divided into a colliculate central area and a striated marginal area. Diameter 10-
60 pm with 9-14 striae in 10 pm. Each stria is alveolate with an outer layer of rows of areolae and an
inner smooth layer of silica covering the alveoli except for a small oblong opening. The raised part of
the central area bears a semicircle of 2-20 fultoportulae with 3 satellite pores. Valve mantle with a
ring of fultoportulae located on every, every second or every third costa, each having two satellite
pores in a radial position. A single rimoportula is present, only slightly below the ring of mantle
fultoportulae.

Cyclotella ambigua Grunow in Cleve & Grunow 1880, p. 119, fig. 133.
Synonym: Cyclotella striata var. ambigua (Grunow) Grunow in Van Heurck 1880-1885, pl. 92, fig.
12
Type locality: Jenissey, Korepovskoj.
Lectotype: Slide no. 2030a in the Grunow Collection, W, designated by H. Hikansson.
Valve circular, strongly tangentially undulate, especially in the central area of the valve face.
The valve face divided into a colliculate central area and a striated marginal area. Diameter 18-34 pm
with 9-10 striae in 10 pm. One to two valve face fultoportulae, with a single rimoportula opposite.
Mantle fultoportulae present.

ACKNOWLEDGEMENTS

I am very grateful for discussions and help in various ways from Dr K. Sabbe, Dr B. Mayer, Dr
R. Jahn, Dr F.E. Round. Dr D. Mann, Mr P. York, Mr P. Lassen and two anonymous reviewers. I
want to thank Dr H. Riedl. Dr. U Passauer, Miss Patricia Sims, Dr W. Krutzsch and Dr Stridh for
loan of slides and for copies of annotations from the collections. I would like to thank especially Mr
R. Ross who looked at the difficult nomenclature and corrected it where necessary, and for his
historical information concerning the Kutzing Collection. Mr R. Ross and Dr D. Mann corrected the
language and made suggestions to improve the text.

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