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Polish Botanical Journal 55(1): 49–53, 2010
DIATOM FLORA OF SAN FRANCISCO BAY AND VICINITY. II.
FOGEDIA KRAMMERI SP. NOV.*
ANDRZEJ WITKOWSKI, HORST LANGE-BERTALOT, JOHN PATRICK KOCIOLEK,
MAXIM KULIKOVSKIY, MAŁGORZATA BĄK & MANFRED RUPPEL
Abstract. This is our second paper focusing on new diatom taxa of the San Francisco Bay area. Previously we described 11 spe-
cies of Navicula s.str. as new for science. Here we present another new naviculoid species of the genus Fogedia. This new
species bears the distinguishing features of the genus, including a straight internal raphe slit and short, simple external distal
raphe ends. The species differs from similar Fogedia taxa by being smaller and having coarser stria and lower lineola density.
With this new species of Fogedia, the number of diatoms new for science from San Francisco Bay and vicinity is 12, with more
awaiting description. We discuss the relationships between the newly described and established species of Fogedia, and present
their patterns of biogeographic distribution.
Key words: San Francisco Bay, naviculoid diatoms, diversity, biogeography, new species, Fogedia
Andrzej Witkowski, Horst Lange-Bertalot & Małgorzata Bąk, Palaeoceanology Unit, University of Szczecin, Mickiewicza 18,
70-383 Szczecin, Poland; e-mail: witkowsk@univ.szczecin.pl
John Patrick Kociolek, Museum of Natural History, UCB 218, University of Colorado, Boulder, Co 80309, U.S.A
Maxim Kulikovskiy, Department of Algology, Papanin’s Institute for Biology of Inland Waters, Russian Academy of Sciences,
152742 Yaroslavl, Nekouz, Borok, Russia
Manfred Ruppel, Faculty of Biology, Institute of Ecology, Phylogeny, Diversity, J. W. Goethe-University and Forschungungsinstitut
Senckenberg, Senckenberganlage 31-33, D-60054 Frankfurt am Main, Germany
INTRODUCTION
There have been few studies on the benthic dia-
toms of the San Francisco Bay. Thompson and
Laws (1982) discussed seasonal changes in the
productivity and species composition of the mi-
crophytobenthos, with special attention to seasonal
changes. Later on, Laws (1988) published the re-
sults of his study on diatom assemblages of San
Francisco Bay. Laws published a complete list
of species identifi ed from surface sediments of
the bay.
The material presented in this paper is part of
a larger research program, the purpose of which
is to develop a comprehensive diatom fl ora for
the San Francisco Bay area. Our research was
initiated with samples deposited in the Diatom
Collection of the California Academy of Sciences.
Later, collections were made on the oceanic coast
in the vicinity of San Francisco. Finally, the Bay
area (i.e. Aquatic Park) and the open ocean coast
in San Francisco were sampled. LM and SEM
revealed numerous undescribed taxa to which spe-
cial attention was paid; these were subjected to
detailed ultrastructural examination. In previous
work, 11 new species of the diatom genus Na-
vicula were described (Witkowski et al. 2009);
this paper describes a new species of the genus
Fogedia Witkowski, Lange-Bertalot, Metzeltin
& Bafana.
The genus of Fogedia was established towards
the end of the 1990s based on transfer of several
taxa originally described as Navicula s.l. (Foged
1975; Witkowski et al. 1997, 2000). Most of the
species belonging in Fogedia occur in tropical
waters, with a few identifi ed in temperate and cold
waters. However, the number of species in Fogedia
* Dedicated to Dr. Kurt Krammer on the occasion of his 85th
birthday
50 POLISH BOTANICAL JOURNAL 55(1). 2010
seems to be underestimated. Our current research
shows the occurrence of potential new Fogedia
species in the Pacifi c Ocean.
In Fogedia the most important characters al-
lowing it to be separated from Navicula s.str. in-
clude the internal raphe slit being straight (it is
strongly oblique in Navicula) and the presence of
short, simple apical external raphe endings (they are
strongly hooked on the same side in Navicula s.str.)
(e.g., Cox 1979; Lange-Bertalot 2001; Witkowski
et al. 2009). In addition, the valve face of most Fo-
gedia species bears a lateral area differing from those
observed either in Fallacia A. J. Stickle & D. G.
Mann in F. E. Round et al. or in Lyrella Karayeva
(cf. Round et al. 1990; Witkowski et al. 1997).
MATERIAL AND METHODS
Fogedia krammeri occurred in recently collected samples
from the Aquatic Park (along San Francisco’s northern
shore on San Francisco Bay) and at Ocean Beach, along
the coast, collected by A. Witkowski and J. P. Kociolek
in 2002, which are deposited in the Institute of Marine
Sciences at the University of Szczecin (SZCZ).
Samples were processed by standard methods, in-
cluding 10% HCl treatment in order to remove calcium
carbonate, followed by rinsing with distilled water. Sam-
ples were boiled in hydrogen peroxide (37%) and again
washed with distilled water. Naphrax® was used as the
mounting medium. Diatoms were examined by light
and electron microscopy. LM observations were made
with a Leica DMLB with a Plan-apochromatic 100×
(1.4 n.a.) oil immersion objective and Zeiss Axioscop
with a Plan-apochromatic 100× (1.4 n.a.) oil immersion
objective. For SEM, cleaned material was air-dried on
fi lter paper. The fi lter paper strips were mounted on stubs
and sputter-coated with gold. Observations were made
with a Hitachi S-4500 microscope. Terminology related
to valve ultrastructure follows Round et al. (1990).
RESULTS
Fogedia krammeri Witkowski, Lange-Bertalot,
Kociolek & Kulikovskiy, sp. nov. Figs 1 & 2
Valvae variabiles ad instar enim late elliptico-lance-
olatae ad lineari-ellipticas apicibus ex cuneo curte et
anguste subrostratis. Longitudo 20–30 μm, latitudo 9.5–
12.0 μm. Raphe parum lateralis extremis centralibus
terminalibusque punctiformibus. Area axialis anguste
linearis. Area centralis in media parte latitudinis fas-
ciam distinctam formans circiter rectangulata ad instar
delimitata areis lateralibus et duabus striis abbreviatis
utrimque. Striae transapicales subparallelae ad parum
radiantes in media parte sed distinctius radiantes in
partibus ad margines et sub apices, 10–12 in 10 μm.
Lineolae striarum comparate bene discernendae, etiam
microscopio photonico, 26–29 in 10 μm. Aspectus ul-
tramicroscopicus externus internusque omnes structurae
generaliter fi ssurae raphis solum usque prope poros
centrales rectae apparentes ibi aliquid curvatae et in
poros fortiter dilatatos guttiformes infl uentes. Fissurae
terminales absunt quia extrema terminalia aliquid di-
latata curtissime ad latus declinata. Fissurae raphis
internae – ut communiter in genere – rectae omnino
nec distortae ut in Navicula s.str. Fogedia fi nmarchica
et F. acuta cum dimensionibus similibus differunt aliquid
ad instar valvarum sed proprie densitate lineolarum
circiter 35–40 nec 26–29 in 10 μm.
TYPE: U.S.A., Brackish water of San Francisco Bay,
Aquatic Park macrophytes, May 2002, leg. A. Witkowski
& J. P. Kociolek [HOLOTYPE: Praep. No. 5051 (1) in
Coll. Witkowski, Institute of Marine Sciences, Uni-
versity of Szczecin (SZCZ); ISOTYPE: Praep. No. CAS
accession number 627380, slide number 223002 in
Diatom Collection, California Academy of Sciences,
San Francisco].
ETYMOLOGY: The species name honors Dr. Kurt
Krammer on the occasion of his 85th birthday.
LM. Valves moderately variable in outline
from broad-elliptic or broadly elliptic-lanceolate
to linear-elliptic, ends cuneate and at the terminus
shortly-protracted subrostrate. Length 20–30 μm,
breadth 9.5–12.0 μm. Raphe slightly lateral with
central and terminal ends appearing punctiform by
LM. Axial area narrow, linear throughout. Central
area forming a shortened, almost rectangular fascia
connected to the typical lateral areae (of Fogedia);
in the marginal part on either side the transapical
striae are interrupted. Striae subparallel to slightly
radiate proximally, becoming more strongly radiate
distally up to the ends, where they arevery slightly
denser, 10–12 in 10 μm. Lineolae of the striae
comparatively coarse, easily discernible by LM,
26–29 in 10 μm.
SEM. External and internal view (Fig. 1: 8–11,
2: 1–3): the general pattern of structures as in the
few other known taxa of Fogedia. Valve surface
A. WITKOWSKI ET AL.: DIATOM FLORA OF SAN FRANCISCO BAY AND VICINITY. II 51
fl at, slightly depressed in the form of lateral areae.
External central raphe endings largely expanded
and very shortly declined. External apical raphe
endings very unlike the terminal fi ssures of Na-
vicula s.str., short and slightly bent on one side.
Raphe slit internally quite straight from the central
nodule to the helictoglossa; it is not twisted, which
is the condition observed in Navicula s.str.
Fig. 1. Fogedia krammeri Witkowski, Lange-Bertalot, Kociolek & Kulikovskiy, sp. nov. 1–7 – valve view of specimens from
type habitat (6 & 7a represent holotype specimen); 8 & 9 – external valve view of whole specimens; note the slightly undulating
raphe, expanded external central raphe endings and short, slightly bent terminal raphe endings; 10 & 11 – closeup of valve; note
depression of lateral area. 1–7 – LM: 1–5 – brightfi eld optics, 6 & 7 – DIC optics; 8–11 – SEM. Scale bar for 1–7 = 10 μm.
52 POLISH BOTANICAL JOURNAL 55(1). 2010
DIFFERENTIAL DIAGNOSIS. The only spe-
cies that has a similar valve outline is F. acuta
Witkowski, Lange-Bertalot & Metzeltin, known
from salt marshes and found also in Netarts Bay,
Oregon, U.S.A. (Riznyk 1973). Fogedia krammeri
is distinguished easily from F. acuta by its smaller
cell dimensions, coarser striae and lower lineola
density. Valves of Fogedia fi nmarchica differ from
those of F. krammeri by their lanceolate outlines
and having 35–40 lineolae per 10 μm, which are
very diffi cult to resolve by LM.
DISTRIBUTION. Fogedia krammeri is known only
from San Francisco Bay. Specimens fi gured and iden-
tifi ed as Navicula fi nmarchica (Cleve & Grunow)
Cleve (= Fogedia fi nmarchica Witkowski, Metzeltin
& Lange-Bertalot in Witkowski) by Riznyk (1973)
from Yaquina Bay in Oregon, appear more similar
to Fogedia krammeri.
DISCUSSION
Our light and electron microscopy-based re-
search on samples from the San Francisco Bay
area revealed the presence of a new species of
Fogedia. In this paper we also provided data on
the ultrastructure of Fogedia as a genus. Based on
our observations, the gross morphology of Fogedia
is consistent, exhibiting little variation within the
genus (e.g., Witkowski et al. 1997, 2000), unlike
in Navicula s.str. (Lange-Bertalot 2001; Witkowski
et al. 2009).
In its generic characters, Fogedia krammeri
conforms with the generitype F. fi nmarchica. Fo-
gedia species are characterized by linear-lanceo-
late, rarely elliptic valve shape. With the exception
of F. heterovalvata (Simonsen) Witkowski, Lange-
Bertalot & Metzeltin (Witkowski et al. 2000),
Fogedia species possess a lateral area. At the ul-
trastructural level, on the exterior of Fogedia are
simple distal raphe ends slightly bent to the same
side, while internally there is a straight and simple
slit-like raphe (Witkowski et al. 2000). It is inter-
esting to note that taxa inhabiting temperate wa-
ters have robust, heavily silicifi ed valves, whereas
those from tropical waters are fragile (Witkowski
et al. 1997).
Fig. 2. Fogedia krammeri Witkowski, Lange-Bertalot, Kociolek & Kulikovskiy, sp. nov. 1 & 2 – internal valve view, note internal
raphe slit running straight; 3 – closeup of internal view of central part of valve; note structureless internal view of lateral area,
simple internal central raphe endings and domed areolae occlusions. All SEM.
A. WITKOWSKI ET AL.: DIATOM FLORA OF SAN FRANCISCO BAY AND VICINITY. II 53
Some variation in the distribution of Fogedia
species was observed. In general we have few
species, including Fogedia giffeniana (Foged)
Witkowski, Lange-Bertalot & Metzeltin, that
are confi ned to warm waters. They occur in the
western Indian Ocean (Oman, Yemen, Kenya)
and in the Mediterranean Sea (Witkowski et al.
2000; A. Witkowski, unpubl.). Taxa similar to Fo-
gedia fi nmarchica have a very broad geographic
distribution, though apparently absent from the
Southern Ocean. Fogedia fi nmarchica and related
taxa represent the epipelon and inhabit marine lit-
torals including tidal fl ats of the North Atlantic
and neighboring seas (Salah 1953, 1955; Hendey
1964; Witkowski et al. 2000). They also were re-
ported from the United States Pacifi c coast, Oregon
(Riznyk 1973). Here we report Fogedia krammeri,
which is known so far from the San Francisco
Bay area, a region infl uenced by the cold-water
California Current.
With respect to valve morphology, Fogedia
may be a rather close relative of Hippodonta
Lange-Bertalot, Witkowski & Metzeltin. This is
suggested by the very similar, slit-like, apically
elongate striae in most of Hippodonta and in
all Fogedia species (Lange-Bertalot et al. 1996;
Witkowski et al. 1997; Lange-Bertalot 2001). Fo-
gedia and Hippodonta also share the same type of
plastid, that is, two girdle-apprised lobes, the same
as in Navicula s.str. (Cox 1979; Lange-Bertalot
2001; J. Park et al., unpubl.). The two genera also
have signifi cant differences. The valve face of Hip-
podonta species, for example, is always arched,
whereas in Fogedia taxa it is fl at. Finally, Hippo-
donta bears apical striae, which are missing in Fo-
gedia (e.g., Lange-Bertalot et al. 1996; Witkowski
et al. 1997; Witkowski et al. 2000; Lange-Bertalot
2001). Further work is required to elucidate the
phylogenetic relationships of this branch of the
diatom tree of life.
ACKNOWLEDGEMENTS. The research was supported by
the California Academy of Sciences, San Francisco,
and by grant no. N306 468538 from the Polish Ministry
of Science and Higher Education. We are grateful to
the anonymous reviewer for valuable comments and
suggestions.
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Received 11 February 2010
