C Nova Hedwigia Vol.

107 (2018) Issue 1–2, 141–165

published online December 19, 2017; published in print August 2018
Article

A review on the synonymy of the dinoflagellate genera

Oxytoxum and Corythodinium (Oxytoxaceae, Dinophyceae)

Fernando Gómez
Carmen Campos Panisse 3, E-11500 Puerto de Santa María, Spain
fernando.gomez@fitoplancton.com

With 7 figures and 1 table

Abstract: The thecate dinoflagellates of the family Oxytoxaceae are reviewed. The synonymy of the
species of the genera Oxytoxum and Corythodinium (=Pyrgidium) is analysed in order to facilitate
the identifications. Seventy-one species have been placed in the genus Oxytoxum. Only twenty-
nine species are accepted after a comparison of the original descriptions and recent observations
from Mediterranean, Atlantic and Pacific Oceans. Most of the remaining species are considered
synonyms of other species of Oxytoxum or placed in Corythodinium or other genera. Species such
as O. gladiolus or O. schauinslandii are resucitated as senior synonyms of species more commonly
reported in the literature. From the sixteen species placed in Corythodinium, only C. carinatum and
C. michaelsarsii are considered synonyms of other congeneric species. Six taxa initially described
as species of Oxytoxum are transferred into Corythodinium. The new combinations are C. brunellii
comb. nov., C. milneri comb. nov., C. mucronatum comb. nov., C. radiosum comb. nov., C. robustum
comb. nov. and C. strophalatum comb. nov. A new species is proposed, C. hasleae sp. nov., for a
taxon misidentified as O. milneri.
Keywords: armored Dinophyta, new combinations, thecate Dinoflagellata.

Historical account of the species descriptions

Stein (1883) described the earlier species of the genera Oxytoxum and Pyrgidium
based on preserved material collected from stomachs of salps. The samples were
collected from several oceans, and Stein (1883) did not report details of the type
locality (with the exception of P. mitra). He reported turned upside down illustrations
of the new taxa Oxytoxum cribrosum, O. diploconus, O. gladiolus, O. scolopax,
O. sphaeroideum, and Pyrgidium constrictum, P. mitra, P. reticulatum, P. sceptrum
and P. tesselatum. Elongated cells with pointed apex and antapex were placed in
the genus Oxytoxum, while Pyrgidium encompasses broader cells with brunt apices
lacking spines. However, the appearances of O. diploconus and P. sceptrum are closer
to Pyrgidium and Oxytoxum, respectively. The delimitation between the two genera
was unclear, and the species of Pyrgidium were transferred into Oxytoxum by the later

© 2017 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.de

Germany. DOI: 10.1127/nova_hedwigia/2017/0460 0029-5035/2017/0460 $ 6.25
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authors. Bütschli (1885) transferred P. constrictum, Schütt (1895, 1896) transferred
P. tesselatum and P. reticulatum, Schröder (1900) and Cleve (1901) transferred
P. sceptrum, and Schiller (1937) transferred P. mitra. Pyrgidium was considered a
subgenus of Oxytoxum (Lemmermann 1905, Paulsen 1908). The dinoflagellate name
Pyrgidium was not further used, with the exception of Haeckel (1899) that proposed
P. pyriforme based on a single illustration.
Murray & Whitting (1899) described Oxytoxum milneri and Lemmermann (1905)
described O. schauinslandii from the warm Atlantic and Pacific Oceans, respectively.
From samples collected in the open tropical Pacific, Kofoid (1907b) described
O. challengeroides, O. compressum, O. cristatum, O. turbo and O. subulatum. He also
described Prorocentrum curvatum that he later transferred into Oxytoxum (Kofoid &
Michener 1911). Kofoid (1907a, b) described Amphidoma biconica and Murrayella
globosa that were later placed in the genera Oxytoxum and Corythodinium, respectively
(Gaarder 1954, Taylor 1976, Dodge & Saunders 1985). Kofoid (1907b) described
O. gigas that was further considered a synonym of Schuettiella mitra and unrelated to the
Oxytoxaceae (Balech 1988). Kofoid & Michener (1911) described without illustrations
O. breve, O. recurvum and O. robustum. Meunier (1910) described O. belgicae
and Pavillard (1916) described O. elegans from the Arctic and Mediterranean Seas,
respectively. From the Greek coasts, Athanassopoulos (1931) described O. phalericum,
but the species is unrecognizable because the description and illustration are insufficient.
Schiller (1937) published a monograph with the species known at that time, with the
exceptions of P. pyriforme, O. phalericum and O. schauinslandii. Schiller (1937)
proposed numerous new species from the coastal Mediterranean Sea: O. adriaticum,
O. caudatum, O. coronatum, O. crassum, O. depressum, O. globosum, O. gracile, O. laticeps,
O. longum, O. mediterraneum, O. obliquum, O. ovale, O. parvum, O. variabile and
O. viride. Rampi (1939, 1941, 1943, 1951, 1969a, b) described O. aceratum, O. areolatum,
O. brunellii, O. frenguellii, O. minutum, O. obesum, O. punctulatum, O. radiosum,
O. tenuistriatum and O. spinosum from the Ligurian Sea, NW Mediterranean Sea. Rampi
(1951, 1969b) described two distinct species using the name O. ligusticum. Sournia (1973)
proposed O. rampii for the posterior homonym O. ligusticum in Rampi (1969b). Rampi
(1969a) also described O. tonollii and O. margalefii that were considered synonyms of
Amphidoma caudata and currently placed within the genus Azadinium (Halldal 1953,
Nezán et al. 2012). From Norwegian waters, Halldal (1953) and Hope (1954) described
O. nanum and O. mucronatum, respectively. Other Norwegian researcher, Gaarder (1954)
described O. carinatum, O. latum and O. michaelsarsii from the open North Atlantic
Ocean. Wood (1963) described O. elongatum from the Australian coasts, and Balech
described O. criophilum from the Antarctic Ocean (Balech & El-Sayed 1965).
Loeblich & Loeblich (1966) noted that the dinoflagellate generic name Pyrgidium
F.Stein 1883 was pre-occupied by the lichen Pyrgidium Nylander 1867. They proposed
the new name Corythodinium based on exclusively on the rules of priority, without
making new taxonomical observations. There was no a need to replace Stein’s Pyrgidium
because that name was not in use and considered a synonym of Oxytoxum. Loeblich
& Loeblich (1966) only transferred P. tesselatum as Corythodinium tesselatum, and
they proposed this species and O. scolopax as type species for their respective genera.
Taylor (1976) described the species O. semicollatum and O. lativelatum from the open

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Indian Ocean. Taylor (1976) re-opened the discussion on the synonymy of Oxytoxum
and Pyrgidium (now Corythodinium) and he supported the generic split. Taylor (1976)
amended the diagnosis of Corythodinium for "relatively robust species, rounded or
broadly elongated, lacking 'affixed' apical species (although the epitheca may rise to
a pointed cone) but usually possessing an antapical spine..." (Taylor 1976, p. 123).
All members of Oxytoxum have an anteriorly situated cingulum with a relatively
small epitheca, whereas the epitheca is relatively larger in Corythodinium (broader
across its base) and the cingulum may be either median or anterior in position (Taylor
1976). Taylor transferred P. constrictum and P. reticulatum, while he maintained
P. mitra and P. sceptrum within the genus Oxytoxum. Taylor (1976) also transferred
into Corythodinium: O. belgicae, O. carinatum, O. compressum, O. cristatum,
O. curvicaudatum, O. diploconus, O. elegans, O. frenguellii, O. latum, O. michaelsarsii
and O. recurvum. He also transferred Amphidoma biconica (=Murrayella kofoidii) and
Murrayella globosa into Corythodinium.
Dodge & Saunders (1985) published a partial revision of Oxytoxum and Corythodinium
based on scanning electron microscopy observations. They merged Corythodinium into
Oxytoxum based on the similar plate pattern despite the considerable range of variation
of the shape and plate ornamentation. Dodge & Saunders (1985) proposed the new
species O. pyramidale and O. strophalatum, and the new combination O. biconicum
for Amphidoma biconica.

Current status

Molecular data of the type species of Oxytoxum and Corythodinium support the generic
split (Gómez et al. 2016). These genera of the family Oxytoxaceae are unrelated
to any other known dinoflagellate, including former members of the Oxytoxaceae
[Amphidiniopsis, Pseudadenoides (formerly Adenoides), Roscoffia, Sabulodinium and
Thecadinium]. Some species possess intermediate characteristics between Oxytoxum
and Corythodinium (i.e., O. challengeroides) and other species such as O. sphaeroideum
and allied species do not fully fit with the characteristics of the type species of Oxytoxum
or Corythodinium. The species of the Oxytoxaceae do not usually reach high densities,
they are uncultivable with the available methods, and they usually inhabit in the open
warm ocean, far of the specialized laboratories on molecular phylogeny. These oceanic
species are of no interest for public health when compared to the harmful bloom-
forming coastal species. We cannot expect more advances in terms of molecular data
in the next years. The synonymy of the species of Oxytoxum and Corythodinium needs
to be revised because there is a species overestimation. Dodge & Saunders (1985),
Balech (1988) and Gómez et al. (2008) misidentified several species, and there is a
need to review the Oxytoxaceae to facilitate the identifications.

Problems on the species descriptions

Numerous species of the Oxytoxaceae were described from the observation of a

single individual, thus, ignoring the intraspecific variability. The original illustrations

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were often restricted to a line drawing, and sometimes the cell ventral view was not
reported. The cells show a different degree of cell compression, and the length-width
ratio vary according to the angle of view. Consequently, cells may be more robust
or slender than in the original descriptions. The intraspecific variability is evaluated
based on the co-occurrence of cells with different morphologies in the same sample,
and the morphology of the recently divided daughter cells. Gómez et al. (2016)
illustrated dividing cells of Oxytoxum and Corythodinium. After the cytokinesis two
daughter cells share a part of the parent theca and each daughter will regenerate the
missing theca. One of the daughter cells has an incomplete epitheca with a rounder
apex, lacking the spines and thecal ornamentation. This suggests that some species
descriptions, especially cells with round epitheca and lacking spines, may correspond
to immature cells of other known species. More stable diagnostic characters seem to
be the shape of the hypotheca, omitting the relative length of the spines, and the width
of the cingulum and the basis of the epitheca.
The present study is based on the long-term observations carried out in the
Mediterranean Sea, Caribbean Sea, North, Equatorial and South Pacific Oceans, and
South Atlantic Ocean. Sampling and observation methods were described in Gómez et
al. (2004, 2011, 2016). I have never observed cells of the Oxytoxaceae in my samplings
from the north English Channel or sporadic surveys in the Arctic Sea. The present
study provides the first micrographs of numerous species. The original descriptions,
posterior line drawings by other authors, and light micrographs are combined here for
comparative purposes, and to provide a more realistic species concepts and synonymy
of the Oxytoxaceae, and to facilitate the species identification.

Oxytoxum scolopax and allied species

Stein (1883) described Oxytoxum scolopax for cells with an elongated hypotheca, a
narrow cingulum and a minute spherical epitheca. The anterior hypotheca showed
straight shoulders (forming an almost straight angle with the lateral contour) (Figs
1A–C). Stein provided three illustrations of O. scolopax with different degree of
development of the apical and antapical spines (Figs 1A–C). This is coherent taken
into account that one of the daughter cells is devoid of spines after the cell division
(Gómez et al. 2016). Dodge (1982) illustrated O. scolopax with the typical morphology
(Fig. 1D), while Dodge & Saunders (1985) reported a cell with a conical epitheca,
wider cingulum and a broad apical spine (Fig. 1E) that is closer to O. challengeroides.
Wood (1963) illustrated O. scolopax with an epitheca that differed from the usual
morphology (Fig. 1F). Stein (1883) illustrated the variability in the spines of O. scolopax
with three cells of similar length (excluding the spines). It is common to observe the
co-occurrence in the field samples of cells of O. scolopax with variable cell length
(Figs 1G–O). Wood (1963) described these longer cells as O. elongatum (Fig. 1U).
The presence of intermediate forms between O. elongatum and the typical O. scolopax
makes difficult to establish the limits between these two species (Figs 1P–T). Dodge
& Saunders (1985) reported O. elongatum for a cell more robust than O. scolopax
(Fig. 1X). The figure 1Y illustrated a cell close to Dodge & Saunder’s O. elongatum.
Schiller (1937) proposed the new species O. longum for a cell with a long epitheca

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Fig. 1. Line drawings and light micrographs of Oxytoxum scolopax and allied species. A–C)
O. scolopax in Stein (1883). D) O. scolopax sensu Dodge (1982). E) O. scolopax sensu Dodge &
Saunders (1985). F) O. scolopax sensu Wood (1963). G–O) O. scolopax. P–T) Intermediate cells
between O. scolopax and O. elongatum. P–S) Epifluorescence pictures. Q) Note the chloroplast in red.
S) Note the thecal plates. U) O. elongatum in Wood (1963). V–W) O. elongatum. X) O. elongatum
sensu Dodge & Saunders (1985). Y) Oxytoxum sp. Z) O. longum in Schiller (1937). AA) O. longum
sensu Hasle (1960). AB) O. caudatum. Scale bar = 20 μm.

as in O. elongatum, but the cingulum was broader (Fig. 1Z). The apical spine and the
antapical swelling that characterized O. scolopax and O. elongatum were missing in
O. longum (Fig. 1Z). The lack of an apical spine in O. longum may be due to a
description based on an immature individual, but the cingulum is wider when compared

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to O. elongatum. Hasle (1960) illustrated O. longum from the Pacific Ocean (Fig. 1AA),
but her cells showed a narrow cingulum when compared to Schiller’s description, and it
may correspond to an elongated cell of O. caudatum (Fig. 1AB). I have observed cells
that agree with O. elongatum (Figs 1V–W), and Hasle’s O. longum (Fig. 1AA), but I
have not observed cells with the morphology of O. longum. Despite the problems to
delimitate O. scolopax and O. elongatum, it is coherent to classify the more elongated
individuals as O. elongatum. Oxytoxum longum needs further research before it might
be considered as an earlier description of O. elongatum.

Oxytoxum gladiolus and allied species

Stein (1883) described O. gladiolus for cells with an elongated hypotheca (almost
straight shoulders), a narrow cingulum and a minute spherical epitheca (Figs 2B–C).
Oxytoxum gladiolus resembled a robust form of Stein’s O. scolopax devoid of spines
(Fig. 2A). While O. scolopax is a distinctive species and the most common generic
species in the literature, O. gladiolus has almost disappeared. More than a form of
O. scolopax, it seems that the records of O. gladiolus have been further reported under
other names. In field samples, these cells showed a variable curvature of the hypotheca
(Figs 2D–I). Schiller (1937) described O. variabile with two cells with different degree
of cell elongation and rounder shoulders (Figs 2J–K) and O. gracile with straight
shoulders and a pointed apex (Fig. 2L). Intermediate cells between O. variabile and
O. gracile can be found (Fig. 1M). Hope (1954) illustrated O. nanum with two cells
of variable elongation (Figs 2N–O). Sournia (1973, p. 50) corrected the epithet as
'nanus'. The genus Oxytoxum is neuter, and this correction seems to be unsupported.
Taylor (1976) illustrated O. nanum (Fig. 2P). Rampi (1969b) described O. rampii
(as O. ligusticum) (Fig. 2Q) that is similar to O. nanum (Figs 2N–O). Schiller (1937)
described more elongated cells as O. coronatum (Fig. 2R) and O. caudatum (Fig. 2S).
Hasle (1960) also illustrated the variability in the elongation of the hypotheca of cells
identified as O. variabile and O. caudatum (Fig. 2T). Coinciding with Hasle (1960),
I have observed cells with variable degree of elongation (Figs 2U–V). These cells
with a longitudinally straight hypotheca and more or less straight shoulders should be
considered conspecific. Schiller’s O. gracile and O. variabile, Hope’s O. nanum, and
Rampi’s O. rampii should be considered junior synonyms of O. gladiolus. Cells with a
more elongated hypotheca corresponded to the morphology of Schiller’s O. caudatum
and O. coronatum. Schiller’s illustrations did remark the longitudinal curvature of the
hypotheca (Figs 2R–S). This is a feature that depends on the angle of view, and difficult
to note in the descriptions based on a single line drawing. Oxytoxum caudatum and
O. coronatum should be considered synonyms. Kofoid (1907a) described Prorocentrum
curvatum as a cell with a curved hypotheca (Figs 2W–X). Hasle (1960) reported
with more detail the morphology of O. curvatum (Fig. 2Y). Wood (1963) under the
name O. curvatum reported a cell of O. caudatum (Fig. 2Z). Certainly, O. curvatum
is a distinct and interesting taxon without a constriction at the cingulum level and an
atypical epitheca (Figs 2AA–AB). That species deserves detailed studies and it could
not belong to Oxytoxum.

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Fig. 2. Micrographs and line drawings of Oxytoxum gladiolus and allied species. A) O. scolopax devoid
of spines in Stein (1883). B–C) O. gladiolus in Stein (1883). D–I) O. gladiolus. J–K) O. variabile in
Schiller (1933). L) O. gracile in Schiller (1933). M) O. gladiolus. N–O) O. nanum in Halldal (1953).
P) O. nanum sensu Taylor (1976). Q) O. ligusticum (=O. rampii) in Rampi (1969b). R) O. coronatum
in Schiller (1937). S) O. caudatum in Schiller (1937). T) O. caudatum sensu Hasle (1960). U–V)
O. caudatum. W–X) Prorocentrum curvatum in Kofoid (1907a). Y) O. curvatum sensu Hasle (1960).
Z) O. curvatum sensu Wood (1963). AA–AB) O. curvatum. AC) O. turbo in Kofoid (1907b). AD)
O. turbo sensu Schiller (1937). AE) O. obliquum in Schiller (1937). AF) O. nanum sensu Taylor
(1976). AG) O. turbo var. in Wood (1963). AH–AM) O. turbo. AN) Robust cell close to O. turbo.
AO–AP) Pyrgidium mitra in Stein (1883). AQ) O. mitra. AR) O. laticeps in Schiller (1937). AS)
O. mitra sensu Dodge & Saunders (1985). AT) O. pyramidale in Dodge & Saunders (1985). AU)
O. laticeps sensu Wood (1963). AV) C. reticulatum sensu Dodge (1982). AW–AY) Three unidentified
species. AX) O. parvum sensu Dodge & Saunders (1985). BA) Oxytoxum sp. Scale bar = 20 μm

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Other group of species is characterized by a spherical epitheca with a very short apical
spine and a marked constriction at the cingulum level. This morphology fit well with
O. turbo as described by Kofoid (1907b) (Fig. 2AC). Schiller (1937) also reported
two illustrations of O. turbo, but he did not report the origin of his observations (Fig.
2AD). Schiller (1937) described O. obliquum (Fig. 2AE) that is a junior synonym of
O. turbo. Taylor (1976) under the name O. nanum illustrated a cell that resembled
O. turbo (Fig. 2AF). Wood (1963) illustrated a species as “O. turbo var.” (Fig. 2AG).
Oxytoxum turbo is a distinctive species (Figs 2AH–AM), and some cells showed a
more robust hypotheca (Fig. 2AN).
Stein (1883) described Pyrgidium mitra with two illustrations of robust cells with
a broad cingulum and epitheca (Figs 2AO–AP). The general appearance of the
hypotheca and the cingulum is closer to Corythodinium, while the spherical epitheca
is closer to Oxytoxum (Fig. 2AQ). Taylor (1976) doubted on the transfer of P. mitra
into Corythodinium. Oxytoxum mitra is a rare but distinctive species that needs further
research because it possesses intermediate characteristics between the two genera.
Schiller (1937) described O. laticeps with a hemispherical epitheca, lacking the short
apical spine (Fig. 2AR). Oxytoxum laticeps resembled O. mitra. Dodge & Saunders
(1985) illustrated O. mitra with a more elongated hypotheca than Stein’s illustration
(Fig. 2AS). Dodge & Saunders (1985) described O. pyramidale with similar shape
than O. mitra, but about one half smaller in size (Fig. 2AT). Oxytoxum pyramidale is
here considered a valid species, although we cannot discard based on the morphology
of the epitheca that it was described from an immature cell.
There are other cells with a very anterior cingulum such as O. laticeps in Wood (1963)
(Fig. 2AU) or O. reticulatum in Dodge (1982) (Fig. 2V). There are undescribed
species with these characteristics (Figs 2AW–AY). Dodge & Saunders (1985) mistook
O. parvum (Fig. 2AX) for a species that resembled an elongated cell of O. pyramidale
(Fig. 2AT). Cells with this morphology have been also recorded in this study (Fig. 2BA).

Oxytoxum sceptrum and allied species

Stein (1883) described O. cribrosum and P. sceptrum for spindle-shaped cells with
apical and antapical spines. Oxytoxum cribrosum showed a relatively wide cingulum
and epitheca, and the contour of the epitheca was almost straight (Fig. 3A), while the
cingulum of O. sceptrum is narrow and the contour of the hypotheca is convex (Fig.
3D). The records of O. cribrosum are rare (Fig. 3B; Rampi & Bernhard 1980). Figure
3C showed a cell that may correspond to O. cribrosum. Stein (1883) exaggerated the
length of the antapical spine of O. sceptrum (Fig. 3D). More than a long antapical spine,
O. sceptrum possesses a pointed antapex similar to the illustration of O. parvum.
Oxytoxum sceptrum is a distinctive species (Fig. 3E–I). Schiller (1937) described
O. parvum (Fig. 3J–K) with a more robust morphology, wider cingulum and broader
epitheca than O. sceptrum. Dodge & Saunders (1985) under the name O. parvum
illustrated a cell closer to O. pyramidale (Fig. 3L). Later, Rampi (1941, 1951)
described two close species, O. aceratum with a narrower cingulum (Fig. 3M) and
O. tenuistriatum with a wide cingulum (Fig. 3N). Both Rampi’s species are here

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Fig. 3. Micrographs and line drawings of Oxytoxum sceptrum and allied species. A) O. cribrosum in
Stein (1883). B) O. cribrosum sensu Rampi & Bernhard (1980). C) O. cf. cribrosum. D) Pyrgidium
sceptrum in Stein (1883). E–I) O. sceptrum. J–K) O. parvum in Schiller (1937). L) O. parvum sensu
Dodge & Saunders (1985). M) O. aceratum in Rampi (1951). N) O. tenuistriatum in Rampi (1941).
O–Q) O. parvum. R–S) O. longiceps in Schiller (1937). T–U) O. longiceps. V) O. schauinslandii in
Lemmermann (1905). W) O. sceptrum sensu Rampi & Bernhard (1980). X–Y) O. schauinslandii.
Scale bar = 20 μm.

considered synonyms of O. parvum. It is difficult to establish the limit between

O. sceptrum and O. parvum, but it is coherent to assign the more robust cells to
O. parvum (Figs 3O–Q). Schiller (1937) described O. longiceps for cells larger than
O. parvum (Fig. 3R–S). It is difficult to discern whether these larger cells (Figs 3T–U)
should be considered as an independent species. Lemmermann (1905) described
O. schauinslandii for very elongated cells (Fig. 3V). That description, omitted in
Schiller (1937), could go unnoticed for further authors. Rampi & Bernhard (1980)

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illustrated under the name O. sceptrum a cell with an appearance of O. schauinslandii
(Fig. 3W). Oxytoxum schauinslandii is a distinctive species (Figs 3X–Y) which records
have been misidentified as O. longiceps.

Oxytoxum sphaeroideum and allied species

The species of this group are small, with round bodies and lacking long spines. These
species are inefficiently collected in the net samplings, and the information is more
limited. This makes difficult the identifications (Gómez et al. 2008). Stein (1883)
described O. sphaeroideum with six illustrations (Figs 4A–E, AU). Stein’s figure 8
showed a smaller cell with round apex and antapex that he considered an immature
cell (Fig. 4A). The figure 9 showed a cell with round apex and a pointed antapex (Fig.
4B). The figure 10 showed a cell with pointed antapex and round antapex (Fig. 4C).
It was later named as O. sphaeroideum var. conicum (Lemmermann 1905). The figure
11 showed a slightly larger cell, with pointed apex and antapex and a wider cingulum
and epitheca (Fig. 4D). It was later named as O. sphaeroideum var. steinii (Ostenfeld &
Paulsen 1904). The figure 12 corresponded to a cell with a partially dissected hypotheca
(Fig. 4E). The figure 13 showed a cell with a broad cingulum that is unrelated to that
species (Fig. 4AU). Schiller (1937) proposed O. boehmii based on Stein’s figure 13.
Dodge (1982) provided an illustration of O. sphaeroideum that does not correspond to
that species (Fig. 4F). Haeckel (1899) described Pyrgidium pyriforme that is close to
O. sphaeroideum var. steinii (Stein’s figure 11), although the epitheca is anomalously
ornamented (Fig. 4G). The description of Pyrgidium pyriforme cannot be considered
an illustration with analysis, and the species description is invalid (I.C.N., article
38.1, 38.8, 38.10). Oxytoxum sphaeroideum is a distinctive species with a very short
antapical spine that was not illustrated by Stein (1883). I did not observe any cells of
O. sphaeroideum with pointed epithecae. The cell robustness varied with the angles
of view (Figs 4H–L). Rampi (1969b) described O. obesum, an invalid description
due to the lack of Latin diagnosis (Fig. 4M). It is here considered a synonym of
O. sphaeroideum. Dodge & Saunders (1985) under the name O. ovale illustrated a cell
than is close to O. mediterraneum (Fig. 4N). Taylor (1976) described O. lativelatum
(Figs 4O–P) and O. semicollatum (Fig. 4Q) that are here considered synonyms of
O. sphaeroideum. Wood (1963) under the name O. sphaeroideum var. steinii reported
a cell with straight shoulders (Fig. 4R). Dodge & Saunders (1985) reported the same
species as O. variabile (Fig. 4S). Schiller (1937) described O. mediterraneum with
more or less round shoulders (Fig. 4T), while Rampi & Bernhard (1980) illustrated that
species with straight shoulders and a very short antapical spine (Fig. 4U). It should be
noted that Schiller (1937) illustrated other species such as O. caudatum or O. variabile
with rounder shoulders than the real morphology. Rampi (1941) also described O.

Fig. 4. Micrographs and line drawings of Oxytoxum sphaeroideum and allied species. A–E, AU)
O. sphaeroideum in Stein (1883). C) O. sphaeroideum var. conicum. D–E) O. sphaeroideum var.
steinii. F) O. sphaeroideum sensu Dodge (1982). G) Pyrgidium pyriforme in Haeckel (1889). H–L)

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O. sphaeroideum. M) O. obesum in Rampi. N) O. ovale sensu Dodge & Saunders. O–P) O. lativelatum
in Taylor (1976). Q) O. semicollatum in Taylor (1976). R) O. sphaeroideum var. steinii sensu Wood
(1963). S) O. variabile sensu Dodge & Saunders (1985). T) O. mediterraneum in Schiller (1937).
U) O. mediterraneum sensu Rampi & Bernhard (1980). V) O. punctulatum in Rampi. W–X)
O. mediterraneum sensu Balech 1988. Y–AE) O. mediterraneum. AF) O. laticeps in Schiller (1937).
AG) O. cf. laticeps. AH) O. laticeps sensu Taylor (1976). AI) O. cf. laticeps. AJ) O. depressum in
Schiller (1937). AK) O. cf. depresum. AL) Oxytoxum sp. AM). O. crassum sensu Dodge & Saunders
(1985). AN) O. laticeps sensu Dodge & Saunders (1985). AO. O. ovale in Schiller (1937). AP)
O. cf. ovale. AQ) O. adriaticum in Schiller (1937). AR–AT) O. adriaticum. AU) O. spheroideum
(=O. boehmii) in Stein (1883). AV–AW) O. boehmii. AX–AY) O. ovale in Gaarder (1954). AZ–BA)
O. ovale. BB) O. minutum in Rampi (1941). BC) O. minutum. BD) O. viride in Schiller (1937). BE)
O. crassum in Schiller (1937). BF) O. pachyderme in Schiller (1937). BG) O. globosum in Schiller
(1937). BH) O. spinosum in Rampi (1941). BI–BJ) O. cf. viride. BK–BM) O. spinosum. Scale bar
= 20 μm.

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punctulatum that is a junior synonym of O. mediterraneum. Oxytoxum mediterraneum
is distinctive species and it has the priority (Figs 4Y–AE). Schiller (1937) described O.
laticeps (Fig. 4AF) that is close to O. mediterraneum (Fig. 4AG, AI). Schiller (1937)
described O. depressum with a scarce constriction at the cingulum level (Fig. 4AJ).
Other difference between these species is that O. depressum possessed a short apical
spine (Fig. 4AK) that is missing in the original description of O. laticeps (Fig. 4AF),
and present in other illustrations as in Taylor (1976, Fig. 4AH) and in Hasle (1960, Fig.
6H). Probably, this short spine was unnoticed in the original description of O. laticeps.
The cells of the figures 4AK–AL can be assigned to O. depressum. The illustration of
O. crassum (Fig. 4AM) and O. laticeps (Fig. 4AN) in Dodge & Saunders (1985) were
misidentified. Schiller (1937) described O. ovale as an elongated cell with round antapex
and a flat epitheca (Fig. 4AO). This species could not belong to the Oxytoxaceae (Fig.
4AP). The illustration of O. ovale (Fig. 4N) in Dodge & Saunders (1985) is closer to O.
sphaeroideum. Oxytoxum adriaticum is a distinct species (Figs 4AQ–AT). Oxytoxum
boehmii showed a conical epitheca (Figs 4AU–AW). Gaarder (1954) described O. ovum
with scarce detail (Figs 4AX–AY), and it could be a junior synonym of Rampi’s (1941)
O. minutum (Fig. 4BB–BC). Schiller (1937) described four species with almost spherical
hypotheca: O. viride (Fig. 4BD), O. crassum (Fig. 4BE), O. pachyderme (Fig. 4BF) and
O. globosum (Fig. 4BG), and Rampi (1941) described O. spinosum (Fig. 4BH). The
identification of O. breve is difficult because it was described without illustration (Kofoid
& Michener 1911). Further studies are necessary before to establish the synonymy among
these species. The cells observed in the present study fit with the cell shape of O. viride
(Figs 4BI–BJ) and O. spinosum (Figs 4BI–BM).

Oxytoxum tesselatum and allied species

Stein (1883) described Pyrgidium constrictum with a distinctive transversal constriction

in the hypotheca (Fig. 5A). The degree of constriction is variable, and in some cells
is hardly visible (Figs 5B–J). When the constriction is less evident, P. constrictum
can be mistaken for P. reticulatum (Fig. 5K). Pyrgidium tesselatum is a distinctive
species due to the transversal ridges (Fig. 5O). Burns & Mitchell (1982) mistook
O. robustum for C. tesselatum. Oxytoxum elegans is larger (Fig. 5Y) and showed elongated
apices, especially a pointed cone in the epitheca when compared to P. tesselatum.
Oxytoxum elegans is distinct species (Figs 5Z–AB). Dodge & Saunders (1985) mistook
O. tesselatum for C. elegans (Fig. 4X). There are no doubts on the identity of
P. constrictum (Figs 5B–J), P. tesselatum (Figs 5P–S, V–W) and O. elegans (Figs
5Z–AB). A cell of C. tesselatum devoid of tessellation or C. constrictum devoid of the
transversal constriction (as in immature cells) could be also mistaken for P. reticulatum.
It is uncertain whether the observations of C. reticulatum corresponded to immature
cells of C. constrictum or C. tesselatum. The illustrations of O. reticulatum in Dodge
(1982) (Fig. 5L) and Dodge & Saunders (1985) (Fig. 5M) corresponded to two different
species, and they differed from Stein’s Pyrgidium reticulatum (Fig. 5K). The illustration
of C. reticulatum in Balech (1988) did not correspond to that species (Fig. 5N).
Amphidoma biconica was described with a median cingulum and a conical epitheca
and hypotheca (Kofoid 1907a) (Fig. 5AC). Gaarder (1954) reported this species (Fig.

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Fig. 5. Micrographs and line drawings of Corythodinium constrictum and allied species. A) Pyrgidium
constrictum. B–J) C. constrictum. K) P. reticulatum in Stein (1883). L) O. reticulatum sensu Dodge
(1982). M) O. reticulatum sensu Dodge & Saunders (1985). N) C. reticulatum sensu Balech (1988).
O) P. tesselatum in Stein (1883). P–S) C. tesselatum. T) O. michaelsarsii in Gaarder (1954). U) O.
areolatum in Rampi (1941). V–W) C. tesselatum. X) O. elegans sensu Dodge & Saunders (1985).
Y) O. elegans in Pavillard (1916). Z–AB) C. elegans. AC) Amphidoma biconica in Kofoid (1907b).
AD) Murrayella kofoidii in Gaarder (1954). AE) O. biconicum sensu Dodge & Saunders (1985).
AF–AH) C. biconicum. AI) Murrayella kofoidii f. elongata Gaarder (1954). AJ–AN) C. biconicum.
AO) O. diploconus in Stein (1883). AP) C. diploconus. AQ) O. milneri sensu Hasle (1960). AR–AV)
C. hasleae sp. nov. AW) O. milneri in Murray & Whitting (1889). AX–BE) C. milneri (G.Murray &
Whitting) F.Gómez, comb. nov. BF) O. subulatum in Kofoid (1907b). BG) O. milneri sensu Dodge
(1982). BH) O. milneri sensu Dodge & Saunders (1985). BI) C. diploconus sensu Balech (1988).
BJ) 'Corythodinium sp. cf. milneri' sensu Balech (1988). Scale bar = 20 μm.

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5AD). Dodge & Saunders (1985) provided a line drawing of O. biconicum that reported
a cell in ventral view with conical epitheca and scarce pointed apex, but missing the
spines (Fig. 5AE). Balech (1988) provided a line drawing of C. diploconus (Fig. 5BI)
that is closer to C. biconicum (Fig. 5AC). The figures 5AF–AH illustrated C. biconicum.
Gaarder (1954) described an elongated cell as Murrayella kofoidii var. elongata (Fig.
5AI). This taxon could correspond to an independent species (Figs 5AJ–AN). Stein
(1883) described O. diploconus (Fig. 5AO) as a bi-conical cell with a more anterior
cingulum than A. biconica (Fig. 5AC). Corythodinium diploconus is rare species (Fig.
5AP). Hasle (1960) under the name O. milneri (Fig. 5AQ) reported a cell that is closer
to C. diploconus. Corythodinium diploconus and O. milneri possess pointed apices,
while Hasle’s O. milneri possesses long apical and antapical spines. Oxytoxum milneri
sensu Hasle (1960) possesses a conical epitheca, while the epitheca is flat in O. milneri
(Fig. 5BH). I have observed cells in the open Mediterranean Sea and Pacific Ocean
that fit with Hasle’s O. milneri. This suggests that this is an undescribed species. It is
proposed here as the new species Corythodinium hasleae sp. nov. (Figs 5AR–AV).
Murray & Whitting (1899) described O. milneri with a detailed illustration (Fig. 5AW),
although the cells tended to show a more flat epitheca than in the original illustration
(Figs 5AX–BD). In some angles of view, the pointed cone of the apex is eccentric
(Fig. 5BE), and Kofoid (1907b) described this morphology as O. subulatum (Fig.
5BF). Oxytoxum milneri is a high distinctive species, but there are misidentifications
in the literature. Dodge (1982) under the name O. milneri reported a cell similar to
O. challengeroides (Fig. 5BG). Later, Dodge & Saunders (1985) reported O. milneri
with its typical morphology (i.e., broad and flat epitheca, Fig. 5BH). Balech (1988)
reported the name 'Corythodinium sp. cf. milneri', but there is no a formal proposal
of that new combination. Balech’s illustration of 'Corythodinium sp. cf. milneri' (Fig.
5BJ) is unrelated to that species, and it is closer to Amphidoma biconica (Fig. 5AC)
or his illustration of C. diploconus (Fig. 5BI). With no doubt, O. milneri should be
placed into Corythodinium.

Corythodinium belgicae and allied species

In this group are pooled the other species with the characteristics of Corythodinium.
The delimitation of the species is difficult due to the smaller sizes and the absence of
distinctive features such as the distinctive ornamentation (constrictions, tessellation
or spines). Hope (1954) described O. mucronatum from the Norwegian waters
(Figs 6A–B). He did not report the ventral view, and he illustrated a high number of
longitudinal ridges (Fig. 6A). Balech & El-Sayed (1965) described O. criophilum
from the Weddell Sea (Figs 5C–D). Balech (1976) reported other illustration (Fig.
6E). Oxytoxum mucronatum and O. criophilum are similar in shape, with a low
conical epitheca, and a curved antapex. Balech’s illustration of O. criophilum showed
the ventral view and a lower number of ridges. Hope (1954) was not precise in the
description of the ridges because they anomalously continued along the cingulum
and the epitheca (Fig. 6A). Although Balech’s illustrations were more complete,
there are reasons to consider that both species are synonyms with priority for Hope’s
species. Dodge & Saunders (1985) described O. strophalatum with a more reduced

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Fig. 6. Micrographs and line drawings of Corythodinium belgicae and allied species. A–B) Oxytoxum
mucronatum in Hope (1954). C–D) O. criophilum in Balech and El-Sayed (1965). E) O. criophilum
sensu Balech (1976). F) O. strophalatum in Dodge & Saunders (1985). G) O. laticeps in Schiller
(1937). H) O. laticeps sensu Hasle (1960). I) O. laticeps sensu Dodge & Saunders (1985). J–Q)
C. mucronatum (B.Hope) F.Gómez, comb. nov. R) O. latum in Gaarder (1954). S) C. latum. T)
O. belgicae in Meunier (1910). U) O. belgicae sensu Balech (1988). V) O. crassum sensu Dodge
& Saunders (1985). W) O. brunellii in Rampi (1939). X) O. frenguellii in Rampi (1941). Y–AD)
C. brunellii (Rampi) F.Gómez, comb. nov. AE) Murrayella globosa in Kofoid (1907b). AF–AG)
C. globosum. AH) O. radiosum in Rampi (1941). AI) O. ligusticum in Rampi (1941). AJ–AK)
C. radiosum (Rampi) F.Gómez, comb. nov. Scale bar = 20 μm.

and flat epitheca and less longitudinal ridges (Fig. 6F) than O. mucronatum. These
species need to be placed into Corythodinium (Figs 6J–Q). Schiller (1937) described
O. laticeps with a hemispherical shape of the epitheca that resembled O. mitra (Fig.
6G). In contrast, Hasle (1960) illustrated O. laticeps with a more pointed apex with
short spine (Fig. 6H). Dodge & Saunders (1985) illustrated O. laticeps with a very
anterior and flat epitheca (Fig. 6I). The identity of O. laticeps is unclear. Gaarder
(1954) described O. latum as a robust species with conical epitheca and hypotheca (Fig.
6R). The illustration named as C. reticulatum in Balech (1988) is close to C. latum
(Fig. 5N). Meunier (1910) described O. belgicae as a robust species (Fig. 6T) that
was later illustrated by Balech (1988) (Fig. 6U). Dodge & Saunders (1985) reported
O. crassum (Fig. 6V) with more similarity to O. belgicae. Rampi (1939) described
O. brunellii without a ventral view of the cell (Fig. 6W), while Rampi (1941) described
O. frenguellii with an illustration of the ventral view (Fig. 6X). The general shape
of the species is similar, and they could be considered synonyms with priority for

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O. brunellii. Taylor transferred O. frenguellii, and he commented that O. brunellii
should be placed into Corythodinium (Taylor 1976, p. 124). If these taxa are considered
synonyms, O. brunellii should be transferred into Corythodinium and accepted as a
senior synonym of C. frenguellii. If these taxa are not considered synonyms, O. brunellii
also needs to be placed into Corythodinium. Kofoid (1907b) described Murrayella
globosa (Fig. 6AE) that Taylor (1976) transferred into Corythodinium (Figs AF–AG).
Rampi (1941) described two species, O. radiosum (Fig. 6AH) and O. ligusticum (Fig.
6AI), with almost hemispherical hypotheca, and round antapex and a low conical
epithecae. Rampi (1941) did not provide the ventral views. Oxytoxum radiosum and
O. ligusticum should be considered synonyms. My observations fit with O. radiosum.
This is a distinctive species and fit with the characteristics of Corythodinium (Figs
6AJ–AK).

Corythodinium cristatum and allied species

This group of species is characterized by relative large sizes, robustness, some

degree of compression and asymmetrical hypothecae. Kofoid (1907b) described
O. cristatum as one of the largest species of the Oxytoxaceae, with a distinctive crest,
cell compression and a curved antapical spine (Fig. 7A). The cell shape, size and
compression is quite different when compared to the type species of Corythodinium,
C. tesselatum. However, the molecular data showed that C. cristatum and C. tesselatum
are closely related (Gómez et al. 2016). Kofoid & Michener (1911) described
O. robustum without illustration as a cell similar to O. cristatum with a rounder apex.
Gaarder (1954) described O. carinatum (Fig. 7D). Wood (1963) reported an illustration
under the name O. robustum (Fig. 7E) that is similar to O. carinatum. Despite the
absence of illustration in the original description, O. robustum should be considered an
earlier description of O. carinatum. Corythodinium cristatum is a rare species because
it is found in deep waters where the sampling effort is lower (Figs 7B–C) as well as
O. robustum (Fig. 7F–G). When it divides, one of the recently formed daughter cell will
probably show a round epitheca instead of a crest. Consequently, we cannot discard
that O. robustum could be an immature cell of O. cristatum. Kofoid (1907b) described
O. compressum (Fig. 7I) that is smaller, with a low and slightly pointed epitheca and
long antapical spine when compared to O. cristatum or O. robustum. The figure 7J
illustrated C. compressum. Wood (1963) under the name O. compressum illustrated
a cell with elongated hypotheca that does not correspond to that species (Fig. 7H).
Kofoid & Michener (1911) described O. recurvum without illustration. According
to the description, the cells showed a low conical epitheca and a curved and pointed
antapex (Figs 7K–O). The SEM pictures of cells identified as O. compressum in
Burns & Mitchell (1982) corresponded to O. recurvum. Kofoid (1907b) described
O. curvicaudatum as a cell with asymmetric hypotheca with longitudinal ridges curved
to an eccentric antapex (Fig. 7P). This is a distinctive species with a rare occurrence
(Figs 7Q–R).
The synonymy of all the species placed in the genera Oxytoxum and Corythodinium
is reported in the Table 1.

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Fig. 7. Micrographs and line drawings of Corythodinium cristatum and allied species. A)
Oxytoxum cristatum in Kofoid (1907b). B–C) C. cristatum. D) O. carinatum in Gaarder (1954). E)
O. robustum sensu Wood (1963). F–G) C. robustum (Kof. & J.R.Michener) F.Gómez, comb. nov. H)
O. compressum sensu Wood (1963). I) O. compressum in Kofoid (1907b). J) C. compressum. K–O)
C. recurvum. P) O. curvicaudatum in Kofoid (1907b). Q–R) C. curvicaudatum. Scale bar = 20 μm.

Taxonomical innovations

Several taxa described initially under the genus Oxytoxum need to be transferred into
Corythodinium.
Corythodinium brunellii (Rampi) F.Gómez, comb. nov.
Basionym: Oxytoxum brunellii Rampi (1939, Nuovo Giorn. Bot. Ital. ser. 2, 46: 466, fig. 24).
Corythodinium milneri (G.Murray & Whitting 1899) F.Gómez, comb. nov.
Basionym: Oxytoxum milneri G.Murray & Whitting (1899, Trans. Linn. Soc. London, Bot. 5: 328,
pl. 27, fig. 6)
Corythodinium mucronatum (B.Hope) F.Gómez, comb. nov.
Basionym: Oxytoxum mucronatum B.Hope (1954, Nytt Mag. Bot. 2, 153, fig. 1c–e)

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 Table 1. List of species names placed within the genera Oxytoxum and Corythodinium. The names reported in bold in the first column are the accepted
names.

Taxa Basionym Accepted name

Amphidoma biconica Kof. - C. biconicum (Kof.) F.J.R. aylor
Corythodinium belgicae (Meunier) F.J.R.Taylor O. belgicae Meunier C. belgicae (Meunier) F.J.R.Taylor
Corythodinium biconicum (Kof.) F.J.R.Taylor Amphidoma biconica Kof. C. biconicum (Kof.) F.J.R.Taylor
Corythodinium brunellii (Rampi) F.Gómez, comb. nov. O. brunellii Rampi C. brunellii (Rampi) F.Gómez, comb. nov.
C. robustum (Kof. & J.R.Michener) F.Gómez,
Corythodinium carinatum (Gaarder) F.J.R.Taylor O. carinatum Gaarder
comb. nov.
Corythodinium compressum (Kof.) F.J.R.Taylor O. compressum Kof. C. compressum (Kof.) F.J.R.Taylor
Corythodinium constrictum (F. Stein) F.J.R.Taylor P. constrictum F.Stein C. constrictum (F.Stein) F.J.R.Taylor
Corythodinium cristatum (Kof.) F.J.R.Taylor O. cristatum C. cristatum (Kof.) F.J.R.Taylor
O. curvicaudatum (Kof.)
Corythodinium curvicaudatum (Kof.) F.J.R.Taylor C. curvicaudatum (Kof.) F.J.R.Taylor
F.J.R.Taylor

eschweizerbart_xxx
Corythodinium diploconus (F.Stein) F.J.R.Taylor O. diploconus F.Stein C. diploconus (F. Stein) F.J.R.Taylor

158
Corythodinium elegans (Pavill.) F.J.R.Taylor O. elegans Pavill. C. elegans (Pavill.) F.J.R.Taylor
Corythodinium frenguellii (Rampi) F.J.R.Taylor O. frenguellii Rampi C. frenguellii (Rampi) F.J.R.Taylor
Corythodinium globosum (Kof.) F.J.R.Taylor M. globosa Kof. C. globosum (Kof.) F.J.R.Taylor
Corythodinium hasleae F.Gómez, sp. nov. O. milneri sensu Hasle (1960) C. hasleae F.Gómez, sp. nov.
Corythodinium latum (Gaarder) F.J.R.Taylor O. latum Gaarder C. latum (Gaarder) F.J.R.Taylor
Corythodinium michaelsarsii (Gaarder) F.J.R.Taylor O. michaelsarsii Gaarder C. tesselatum (F.Stein) Loebl. & A.R.Loebl.
Corythodinium milneri (G.Murray & Whitting) F.Gómez, comb. C. milneri (G.Murray & Whitting) F.Gómez,
O. milneri G.Murray & Whitting
nov. comb. nov.
Corythodinium mucronatum (B.Hope) F.Gómez, comb. nov. O. mucronatum B.Hope C. mucronatum (B.Hope) F.Gómez, comb. nov.
Corythodinium radiosum (Rampi) F.Gómez, comb. nov. O. radiosum Rampi C. radiosum (Rampi) F.Gómez, comb. nov.
Corythodinium recurvum (Kof. & J.R.Michener) F.J.R.Taylor O. recurvum Kof. & J.R.Michener C. recurvum (Kof. & J.R.Michener) F.J.R.Taylor
Corythodinium reticulatum (F.Stein) F.J.R.Taylor O. reticulatum F.Stein C. reticulatum (F.Stein) F.J.R.Taylor
Corythodinium strophalatum (J.D.Dodge & R.D.Saunders) O. strophalatum J.D.Dodge & C. strophalatum (J.D.Dodge &
F.Gómez, comb. nov. R.D.Saunders R.D.Saunders) F.Gómez, comb. nov.
Corythodinium robustum (Kof. & J.R.Michener) F.Gómez, C. robustum (Kof. & J.R.Michener) F.Gómez,
O. robustum Kof. & J.R. Michener
comb. nov. comb. nov.
Corythodinium tesselatum (F.Stein) Loebl. & A.R.Loebl. P. tesselatum F.Stein C. tesselatum (F.Stein) Loebl. & A.R.Loebl.
 Murrayella globosa Kof.
Murrayella kofoidii Gaarder
Murrayella kofoidii f. elongata Gaarder
Oxytoxum aceratum Rampi
Oxytoxum adriaticum J.Schiller
Oxytoxum areolatum Rampi
Oxytoxum belgicae Meunier
Oxytoxum biconicum (Kof.) J.D.Dodge & R.D.Saunders
Oxytoxum boehmii J.Schiller
Oxytoxum breve Kof. & J.R.Michener
Oxytoxum brunellii Rampi
Oxytoxum carinatum Gaarder
Oxytoxum caudatum J.Schiller
Oxytoxum challengeroides Kof.
Oxytoxum compressum Kof.
159
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Oxytoxum constrictum (F.Stein) Buetschli
Oxytoxum coronatum J.Schiller
Oxytoxum crassum J.Schiller
Oxytoxum cribrosum F.Stein
Oxytoxum criophilum Balech in Balech & El-Sayed
Oxytoxum cristatum Kof.
Oxytoxum curvatum (Kof.) Kof. & J.R.Michener
Oxytoxum curvicaudatum Kof.
Oxytoxum depressum J.Schiller
Oxytoxum diploconus F.Stein
Oxytoxum elegans Pavill.
Oxytoxum elongatum E.J.F. Wood
Oxytoxum frenguellii Rampi
Oxytoxum gigas Kof.
Oxytoxum gladiolus F.Stein
= C. globosum (Kof.) F.J.R.Taylor
A. biconica Kof. C. biconicum (Kof.) F.J.R.Taylor
= C. biconicum (Kof.) F.J.R.Taylor
= O. sceptrum (F.Stein) Schröd.
= O. adriaticum J.Schiller
= C. tesselatum (F.Stein) Loebl. & A.R.Loebl.
= C. belgicae (Meunier) F.J.R.Taylor
A. biconica Kof. C. biconicum (Kof.) F.J.R.Taylor
O. sphaeroideum F.Stein O. boehmii J.Schiller
= O. breve Kof. & J.R.Michener
= C. brunellii (Rampi) F.Gómez, comb. nov.
C. robustum (Kof. & J.R.Michener) F.Gómez,
=
comb. nov.
= O. caudatum J.Schiller
= O. challengeroides Kof.
= C. compressum (Kof.) F.J.R.Taylor
P. constrictum F.Stein C. constrictum (F.Stein) F.J.R.Taylor
= O. caudatum J.Schiller
= O. crassum J.Schiller
= O. cribrosum F.Stein
= C. mucronatum (B.Hope) F.Gómez, comp. nov.
= C. cristatum (Kof.) F.J.R.Taylor
Prorocentrum curvatum Kof. O. curvatum (Kof.) Kof. & J.R.Michener
= C. curvicaudatum (Kof.) F.J.R.Taylor
= O. depressum J.Schiller
= C. diploconus (F.Stein) F.J.R.Taylor
= C. elegans (Pavill.) F.J.R.Taylor
= O. elongatum E.J.F.Wood
= C. frenguellii (Rampi) F.J.R.Taylor
= Schuettiella mitra (F.Schütt) Balech
= O. gladiolus F.Stein
 Taxa
Oxytoxum globosum J.Schiller
Oxytoxum gracile J.Schiller
Oxytoxum laticeps J.Schiller
Oxytoxum lativelatum F.J.R.Taylor
Oxytoxum latum Gaarder
Oxytoxum ligusticum Rampi 1941
Oxytoxum ligusticum Rampi 1969b
Oxytoxum longiceps J.Schiller
Oxytoxum longum J.Schiller
Oxytoxum margalefii Rampi
Oxytoxum mediterraneum J.Schiller
Oxytoxum michaelsarsii Gaarder
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Oxytoxum milneri G.Murray & Whitting
160
Oxytoxum milneri sensu Hasle (1960)
Oxytoxum minutum Rampi
Oxytoxum mitra (F.Stein) J.Schiller
Oxytoxum mucronatum B.Hope
Oxytoxum nanum Halldal
Oxytoxum obesum Rampi
Oxytoxum obliquum J.Schiller
Oxytoxum ovale J.Schiller
Oxytoxum ovum Gaarder
Oxytoxum pachyderme J.Schiller ex F.J.R.Taylor
Oxytoxum parvum J. Schiller
Oxytoxum punctulatum Rampi
Oxytoxum pyramidale J.D.Dodge & R.D.Saunders
Oxytoxum radiosum Rampi
Oxytoxum rampii Sournia
Oxytoxum recurvum Kof. & J.R.Michener
Basionym Accepted name
= O. globosum J.Schiller
= O. gladiolus F.Stein
= O. laticeps J.Schiller
O. sphaeroideum F.Stein
= C. latum (Gaarder) F.J.R.Taylor
= C. radiosum (Rampi) F.Gómez, comp. nov.
= O. gladiolus F.Stein
= O. longiceps J.Schiller
= O. longum J.Schiller
Azadinium caudatum (Halldal) Nézan &
=
Chomérat
= O. mediterraneum J.Schiller
= C. tesselatum (F.Stein) Loebl. & A.R.Loebl.
C. milneri (G.Murray & Whitting) F.Gómez,
=
comb. nov.
= C. hasleae F.Gómez, sp. nov.
= O. minutum Rampi
P. mitra F. Stein O. mitra (F.Stein) J.Schiller
= C. mucronatum (B.Hope) F.Gómez, comb. nov.
= O. gladiolus F.Stein
= O. sphaeroideum F.Stein
= O. turbo Kof.
= O. ovale J.Schiller
= O. ovum Gaarder
O. pachyderme J.Schiller O. pachyderme J.Schiller ex F.J.R.Taylor
= O. parvum J.Schiller
= O. mediterraneum J.Schiller
= O. pyramidale J.D.Dodge & R.D.Saunders
= C. radiosum (Rampi) F.Gómez, comp. nov.
O. ligusticum Rampi 1969b O. gladiolus F.Stein
= C. recurvum (Kof. & J.R.Michener) F.J.R.Taylor
 Oxytoxum reticulatum (F.Stein) F.Schütt = C. reticulatum (F.Stein) F.J.R.Taylor
C. robustum (Kof. & J.R.Michener) F.Gómez,
Oxytoxum robustum Kof. & J.R.Michener =
comb. nov.
Oxytoxum sceptrum (F.Stein) Schröd. P. sceptrum F. Stein O. sceptrum (F.Stein) Schröd.
Oxytoxum schauinslandii Lemmerm. = O. schauinslandii Lemmerm.
Oxytoxum scolopax F.Stein = O. scolopax F.Stein
Oxytoxum semicollatum F.J.R.Taylor = O. sphaeroideum F.Stein
Oxytoxum sphaeroideum F.Stein = O. sphaeroideum F.Stein
Oxytoxum sphaeroideum var. conicum Lemmerm. O. sphaeroideum F.Stein O. sphaeroideum F.Stein
Oxytoxum sphaeroideum var. steinii Ostenf. & Paulsen O. sphaeroideum F.Stein O. sphaeroideum F.Stein
Oxytoxum spinosum Rampi = O. spinosum Rampi
C. strophalatum (J.D.Dodge & R.D.Saunders)
Oxytoxum strophalatum J.D.Dodge & R.D.Saunders =
F.Gómez, comb. nov.
C. milneri (G.Murray & Whitting) F.Gómez,
Oxytoxum subulatum Kof. =
comb. nov.
Oxytoxum tenuistriatum Rampi = O. sceptrum F.Stein
Oxytoxum tesselatum (F.Stein) F.Schütt P. tesselatum C. tesselatum (F.Stein) Loebl. & A.R.Loebl.

161
Azadinium caudatum (Halldal) Nézan &

eschweizerbart_xxx
Oxytoxum tonollii Rampi =
Chomérat
Oxytoxum turbo Kof. = O. turbo Kof.
Oxytoxum variabile J.Schiller = O. gladiolus F.Stein
Oxytoxum viride J.Schiller = O. viride J.Schiller
Pyrgidium constrictum F.Stein = C. constrictum (F.Stein) F.J.R. Taylor
Pyrgidium mitra F.Stein = O. mitra (F.Stein) J.Schiller
Pyrgidium pyriforme Haeckel = O. sphaeroideum F.Stein
Pyrgidium reticulatum F.Stein = C. reticulatum (F.Stein) F.J.R.Taylor
Pyrgidium sceptrum F.Stein = O. sceptrum (F.Stein) Schröd.
Pyrgidium tesselatum F.Stein = C. tesselatum (F.Stein) Loebl. & A.R.Loebl.
Corythodinium radiosum (Rampi) F.Gómez, comb. nov.
Basionym: Oxytoxum radiosum Rampi (1941, Ann. Mus. Civ. St. Nat. (Genova) 61: 65, pl. 1, fig. 12)

Corythodinium robustum (Kof. & J.R.Michener) F.Gómez, comb. nov.

Basionym: Oxytoxum robustum Kof. & J.R.Michener (1911, Bull. Mus. Comp. Zool. Harvard. Coll.
54: 288, no fig.)

Corythodinium strophalatum (J.D.Dodge & R.D.Saunders) F.Gómez, comb. nov.

Basionym: Oxytoxum strophalatum J.D.Dodge & R.D.Saunders (1985, Bot. Mar. 28: 108, figs 26,
27, 76I)

Corythodinium hasleae F.Gómez, sp. nov.

Basionym: Oxytoxum milneri in Hasle (1960, Skrifter utgitt av Det Norske Videnkaps-Akademi i
Oslo. I. Matematisk-Naturvidenskapelig Klasse, no. 2, p. 37, fig. 32a).
Iconotype: Fig. 32a in Hasle (1960) and Fig. 5AQ in the present study.
Isotype: Fig. 5AR.

Type locality: Pacific Ocean (Hasle 1960), and this study (Fig. 5AR). Other records
from the Mediterranean Sea (Figs 5AS–AV).
Etymology: In honour of Grethe R.Hasle who first described this species as Oxytoxum
milneri.
The cells were bi-conical with a slightly anterior epitheca with apical and antapical
spines. Total size of about 80 µm long and the width at the anterior hypotheca was 20–
35 µm wide. The apical spine was slightly ventrally eccentric. The thecal surface showed
longitudinal ridges and the contour of the hypotheca is serrated in the mature cells.
This new species differed from C. biconicum in the presence of apical and antapical
spines. This new species showed an elongated conical epitheca, while the epitheca of
C. milneri is flat with a pointed cone.

Acknowledgements

I was supported by the contract JCI-2010-08492 of the Ministerio Español de Ciencia y Tecnología,
and the Brazilian contract BJT 370646/2013-14 by Conselho Nacional de Desenvolvimento Científico
e Tecnológico. This is a contribution to the French ANR Biodiversity program (ANR BDIV 07 004-
02 'Aquaparadox').

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Manuscript submitted September 18, 2017; accepted October 25, 2017.

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