Cyst-theca relationship of a new dinoflagellate with a spiny
round brown cyst, Protoperidinium lewisiae sp. nov., and its
comparison to the cyst of Oblea acanthocysta
Kenneth Neil Mertens,
1
* Yoshihito Takano,
2
Haifeng Gu,
3
Aika Yamaguchi,
4
Vera Pospelova,
5
Marianne Ellegaard
6
and
Kazumi Matsuoka
2
1
Research Unit for Palaeontology, Gent University, Gent, Belgium,
2
Institute for East China Sea Research (ECSER),
Nagasaki University, Nagasaki,
4
Research Center for Inland Seas, Kobe University, Kobe, Japan,
3
Third Institute of
Oceanography, SOA, Xiamen, China,
5
School of Earth and Ocean Sciences, University of Victoria, Victoria, British
Columbia, Canada, and
6
Department of Plant and Environmental Sciences, University of Copenhagen, Frederiksberg,
Denmark
SUMMARY
Round spiny brown cysts with apiculocavate processes were
isolated from sediments of Lake Saroma, Japan, Changle
Harbor, East China Sea, China, Jinzhou Harbor, Bohai Sea,
China, and San Pedro Harbor, California, USA. Superficially
similar round spiny brown cysts of the species, Oblea
acanthocysta were, for comparison, restudied through light
microscopy and scanning electron microscopy (SEM) and by
sequencing of small subunit (SSU) and large subunit (LSU)
rDNA obtained through a single cyst from Lake Saroma. These
morphological measurements and SEM observations showed
that the new cysts can be discriminated from O. acanthocysta
by the archeopyle, number of processes, shape of process
bases and its apiculocavate processes. Based on LSU
sequences, the most closely related species was Proto-
peridinium monovelum, for which no cyst stage has been
described so far. However, the thecal morphology of the speci-
mens found in this study differed from P. monovelum in
details of the sulcal plates and shape of apical pore and 2a
plate. We therefore describe Protoperidinium lewisiae sp.
nov., which can be found in estuarine subtropical to temperate
waters of the Pacific Ocean.
Key words: Bohai Sea, Changle Harbor, East China Sea,
Jinzhou Harbor, Lake Saroma, large subunit rDNA, San Pedro
Harbor, small subunit rDNA.
INTRODUCTION
Free-living marine dinoflagellates form a very large and
diverse group of planktonic organisms, currently encompass-
ing 1555 species (Gómez 2005). About 13 to 16% of living
dinoflagellates form so-called resting cysts as part of their
sexual cycle (Head 1996). These cysts can be linked to their
respective motile stage by incubation experiments (Wall &
Dale 1968) and conversely, motile stages may be induced to
form cysts in culture. Molecular phylogenetic analyses based
on either cultured strains or single-cell polymerase chain
reaction (PCR) have helped to elucidate taxonomic relation-
ships within this group (e.g. Bolch 2001; Matsuoka et al.
2006; Takano & Horiguchi 2006; Matsuoka & Head 2013).
Many new cyst species are still being discovered (e.g.
Verleye et al. 2011; Mertens et al. 2014), which is impor-
tant as dinoflagellate cysts are frequently used for
paleoclimate reconstructions (e.g. Mertens et al. 2009;
Price et al. 2013).
Some dinoflagellate cysts have a distinct round, spine-
bearing shape and are brown in color, and are therefore
informally designated as ‘round brown spiny cysts’ (see
review in Radi et al. 2013). The paleontological cyst-based
taxonomy has erected two genera to classify these species.
Cysts that belong to the genus Echinidinium (Zonneveld
1997) open with a theropylic archeopyle (Head et al. 2001),
i.e. an angular slit that follows paraplate boundaries but
without complete release of plates (Matsuoka 1988). The
second genus, Islandinium was erected by Head et al.
(2001) and opens with a saphopylic archeopyle, i.e. having
a free operculum (Matsuoka 1988). At species level, char-
acteristics of processes and wall texture are useful distin-
guishing features of round brown spiny cysts (e.g. Zonneveld
1997; Head et al. 2001; Mertens et al. 2012b; Radi et al.
2013).
Cyst-motile relationships have been established only for a
few species of ‘round brown spiny cysts’: Protoperidinium
monospinum (Paulsen) Zonneveld et Dale (Zonneveld and
Dale 1994), Oblea acanthacysta Kawami, Iwataki et Matsuoka
(Kawami et al. 2006; =cyst of Diplopelta parva (Abé)
Matsuoka sensu Matsuoka 1988), P. tricingulatum Kawami,
van Wezel, Koeman et Matsuoka (Kawami et al. 2009), dif-
ferent types of Archaeperidinium minutum (Kofoid) Jørgensen
– previously called P. minutum (Kofoid) Loeblich III; the
genus Archaeperidinium was reinstated by Yamaguchi et al.
(2011) – and the closely related species A. saanichi Mertens,
*To whom correspondence should be addressed.
Email: kenneth.mertens@ugent.be
Communicating editor: Mona Hoppenrath
Received 6 June 2014; accepted 4 October 2014.
Phycological Research 2015 doi: 10.1111/pre.12083
© 2015 Japanese Society of Phycology