Phycological Research 2004; 52: 419–428
Free-living dinoflagellates in the southern Gulf of Mexico:
Report of data (1979–2002)
Sergio Licea,* María Eugenia Zamudio, Ruth Luna and Jesús Soto
Instituto de Ciencias del Mary Limnología, Universidad Nacional Autónoma de México Apartado postal 70–305 México
04510 D. F., México
SUMMARY
The present study is a report of data of planktonic
dinoflagellates which includes a list of 252 species,
with 10 985 entries in the southern Gulf of Mexico
along with information concerning their occurrence.
Material for the present study consists of water and net
samples obtained during 11 cruises collected at 608
sites between June 1979 and December 2002. Ceratium (47 spp.), Protoperidinium (28 spp.), Dinophysis
(26 spp.), Oxytoxum (19 spp.) and Prorocentrum (15
spp.) were the most diverse genera. The most common
species found are Ceratium breve, Ceratium contortum,
Ceratium furca, Ceratium furca var. eugranum, Ceratium fusus. Ceratium fusus var. seta, Ceratium kofoidii,
Ceratium macroceros, Ceratium massiliense, Ceratium
pentagonum, Ceratium teres, Ceratium trichoceros,
Ceratium tripos, Dinophysis caudata, Ornithocercus
magnificus, Podolampas palmipes, Prorocentrum compressum, Prorocentrum gracile, Prorocentrum micans,
Protoperidinium divergens and Pyrophacus steinii.
Thirteen species are potential toxin producers, among
which Karenia brevis was responsible for fish mass
mortalities. Other toxic species such as Amphidinium
carterae, Dinophysis acuta, Dinophysis caudata, Dinophysis fortii, Dinophysis mitra, Dinophysis rotundata,
Dinophysis tripos, Prorocentrum mexicanum, Prorocentrum micans and Prorocentrum minimum were present
mostly in net samples. The non-toxic species Ceratium
furca, Pyrodinium bahamense var. bahamense, Scrippsiella trochoidea and Gonyaulax polygramma were
found in blooms during the summer. Qualitative data
show that dinoflagellates occurred mostly during July
and August, associated with hydrographic conditions. A
checklist of the species and their occurrence are given.
Key words: database, check list, dinoflagellates, Gulf of
Mexico, toxic species.
Balech (1967a,b), Norris and Berner (1970), Steidinger
and Williams (1970) and Steidinger (1971). Toxic and
harmful species have recently received recognition as a
result of the increase of poisoning events in this
northern area (Steidinger et al. 1967, 1978; Williams
& Ingle 1972; Roberts 1979; Steidinger & Baden
1984; Tester et al. 1991). In the southern region,
earlier plankton investigations were carried out during
the Soviet and Soviet-Cuban expeditions in the 1960s–
1980s (Khromov 1965; Bogdanov et al. 1968; Roujiyaynen et al. 1968; Zernova 1969, 1970, 1974, 1982;
Bessonov et al. 1971; de la Cruz 1971;Krylov 1974;
Vinogradova 1976; Zernova & Zhitina 1985). However,
in most of these studies, dinoflagellates are only mentioned without details concerning species distribution.
Although some investigations on phytoplankton on
the shelf of the southern Gulf of Mexico have been
conducted at the Institute of Marine Science and
Limnology (Santoyo & Signoret 1973, 1975; Licea
1977; Licea & Santoyo 1991), knowledge of dinoflagellates in this region is still scanty. Several harmful
species have recently been observed in great numbers
by Gómez-Aguirre and Licea (1998) and Licea et al.
(2003), but since there has been no consistent assessment and inventory of them, the present paper seeks to
investigate the frequency of species and their distribution based on a taxonomic database obtained by the
authors.
STUDY AREA
The study area, located in the southern Gulf of Mexico
(Fig. 1), has a predominantly cyclonic circulation,
mainly associated with the Yucatan Canal waters
(Nowlin 1972; Merrel & Morrison). Monreal-Gómez and
Salas de León (1985, 1990) confirmed the presence
of cyclonic gyres in a westerly direction that persists in
the whole region from February until March and tends
to vanish in April. In May the gyre disappears, and the
circulation changes from east to west. The cyclonic
INTRODUCTION
There are few records of dinoflagellate taxonomy and
distribution in the southern Gulf of Mexico and contiguous waters. Early investigations were made in the
northern gulf, mainly by Graham (1954), Curl (1959),
*To whom correspondence should be addressed.
Email: licea@mar.icmyl.unam.mx
Communicating editor: K. Matsuoka
Received 1 January 2004; accepted 13 July 2004.
420
S. Licea et al.
Fig. 1. Location of sampling stations in the southern Gulf of Mexico (608 sites). Region-I corresponds to the south-western coast; Region-II
belonging to the Bay of Campeche; and Region-III, is situated on the Yucatan shelf.
gyre sets in August and September throughout the
whole region and lasts until December
Hydrographic conditions in the south-west Gulf of
Mexico are highly influenced by the Loop Current and
the Loop Current rings, as well as by the occurrence of
winter storms between October and February (Hulburt
& Thompson 1980) whose winds contribute to the
cooling and mixing of the surface water column in this
region. In addition, the dominant cyclonic circulation
and riverfronts create quite a dynamic system, which
gives this region unique ecologic conditions.
Some particularly important rivers, such as the
Coatzacoalcos and the Grijalva-Usumacinta RiverSystem, represent approximately one-third of all fluvial
discharges onto the Mexican coast in the southern gulf
(Tamayo 1990). There are two coastal lagoons that
stand out for their estuarine outwelling to the coast:
the Terminos and Tamiahua.
The Yucatan Shelf is highly influenced by the
upwelling that is north of Cape Catoche (Cochrane
1969; Merino 1997). Another significant feature is the
shelf width around the peninsula, which extends
seaward for over 260 km: this makes it the widest
continental shelf in the Caribbean region. In contrast,
in the Caribbean margin, the shelf is very narrow (3 km
wide), which means water upwells onto this shelf and
remains trapped within its euphotic zone for long
periods; this might very well increase the fertilizing
potential in this area (Furnas & Smayda 1987). One
part of the upwelled waters flows toward the west,
leaving the shelf near the Alacranes Reef, while the
other part moves toward the coast, forming a cyclonic
circulation to the north of Cape Catoche. Another
peculiar feature is the absence of rivers along the
Yucatan coast (Merino & Otero 1991); however, ground
water discharge induces negligible salinity variations
(Merino et al. 1990).
MATERIALS AND METHODS
The phytoplankton community in the Gulf of Mexico
has been sampled during long-term oceanographic
surveys (projects: PROGMEX, OGMEX and SGM). For
the present study, 11 cruises were selected. Surveys
were carried out on board the R/V JUSTO SIERRA
between July 1979 and December 2002. The stations
occupied during these cruises are shown in Figure 1
(608 sites were sampled). For practical purposes the
region surveyed was divided into three regions. Water
samples were taken in at least one-third of the region
by a conductivity, temperature and depth recorder
(CTD) Neil Brown with a rosette of Niskin bottles and
Dinoflagellates in the Gulf of Mexico
Table 1.
421
Common species for each region in the southern Gulf of Mexico
Region-I
Occurrence
Region-II
Occurrence
Ceratium breve
Ceratium macroceros
Ceratium tripos
Gyrodinium fusiforme
Oxytoxum gracile
Pyrophacus steinii
196
192
149
126
125
194
Ceratium candelabrum
75
Ceratium carriense
83
Ceratium contortum
134
Ceratium contrarium
103
Ceratium pentagonum var. tenerum 100
Ceratocorys horrida
128
Ornithocercus magnificus
164
Ornithocercus steinii
106
Podolampas bipes
102
Region-III
Occurrence
Amphidinium acutissimum
Ceratium furca
Ceratium horridum
Dinophysis mitra
Lingulodinium polyedrum
Oxytoxum parvum
Prorocentrum aporum
Prorocentrum mexicanum
Prorocentrum micans
Prorocentrum sigmoides
Prorocentrum grande
Protoperidinium mediterraneum
32
154
128
30
44
36
23
94
139
32
104
126
Occurrence corresponds to the number of stations where each species was found.
were fixed in Lugol’s solution for analysis in an inverted
microscope. Cell counts were made following the
Utermöhl method (Hasle 1978). Vertical net tows
within 10 m beneath the surface were made at each
station with a 30 and 45 µm mesh net and were
preserved with 2% buffered formalin. Water mounts by
using Trypan blue (= diamine blue 3B) were examined
to obtain information on plate patterns to aid species
identification. Material has been deposited in the collection MEXU-UNAM at the Institute of Marine Science
and Limnology (ICMYL). In some cases subsamples
rinsed of salt were mounted directly on aluminum
stubs for scanning electron microscopy (SEM) for detail
identification. Electron microscopy was carried out
using a SEM model JSM5410LV. Species frequency
was counted using a database on 11 oceanographic
cruises between 1979 and 2002. Species with their
nomenclatural authorities (Table 2) are arranged alphabetically in each order, according to the classification of
Crétinnot-Dinet et al. (1993). The occurrence data have
also been deposited in a database designed by the
National Commission of Knowledge and Use of Diversity
using Biotica version 4.1 (Fideicomiso Fondo para la
Biodiversidad, CONABIO 2003), which is in active use
for further projects. Although interpretation of the data
is based on or derived from the tables, knowledge of
hitherto unpublished data has also been applied.
RESULTS
Table 1 shows that each region in the southern gulf has
its typical species. As can be seen, region-III had the
highest diversity; in contrast, region-I had fewer characteristic species; although, the composition of these
contrasting species could be related to the hydrographical conditions, or influenced by other factors.
Figures 2–22 illustrate the most common and potentially toxic species.
Table 2 shows a list of 252 free-living dinoflagellates, of which the most common genus was Ceratium
with the species Ceratium breve, Ceratium contortum,
Ceratium furca var. furca, Ceratium furca var. eugranum, Ceratium fusus var. fusus, Ceratium fusus var.
seta, Ceratium kofoidii, Ceratium macroceros, Ceratium massiliense, Ceratium pentagonum, Ceratium
teres, Ceratium trichoceros and C. tripos. Of these,
C. furca and C. fusus had blooms in summer 1987 and
early autumn 2000, respectively, in region-II. Other
frequent species were Dinophysis caudata var. caudata, Ornithocercus magnificus, Podolampas palmipes,
Protoperidinium divergens, Pyrophacus steinii, Prorocentrum gracile, Prorocentrum micans, and Prorocentrum compressum, all being the most widely
distributed species in the area. Because of the use of
preserved material, there are few records of unarmoured species; however, the detection of Karenia
brevis, Amphidinium carterae, among others, is significant since these species are potentially toxic.
The genus Ceratium registered 49 species and 21
infraspecific taxa representing 25.8% of the total
species considered in the database. Its ubiquitous
presence throughout the year makes this genus important not only for its wide distribution but for its relative
abundance. The high frequency of C. furca var. furca,
particularly in summer, is an interesting feature, since
this species was found to be the most common in the
southern gulf.
The presence of K. brevis is particularly significant
since this species is a neurotoxic shellfish poisoning
producer. It was found in 35 localities in late summer
and autumn in regions I and II. Quantitative data show
that the maximum cell concentration of K. brevis
occurred during October 1998 along the shelf of
region-II, which coincided with red tide events, with
fish mass mortalities having densities up to 13.9 × 106
cells/L. The first bloom was observed along the southwestern shelf during the summer of 1979, and was
also associated with fish mortality, according to local
coast guard reports.
Other toxic species observed were Dinophysis acuta
var. acuta, Dinophysis caudata, Dinophysis fortii, Dinophysis mitra, Dinophysis rapa, Dinophysis rotundata
422
S. Licea et al.
Figs 2–22. 2. Ceratium furca var. eugrammun, dorsal view. 3. Ceratium fusus var. fusus, dorsal view. 4. Ceratium trichoceros, ventral
view. 5. Ceratium teres, dorsal view. 6. Ceratium pentagonum, ventral view. 7. Ceratium kofoidii, dorsal view. 8. Ceratium furca var.
furca, ventral view. 9. Ceratium breve var. breve, ventral view. 10. Ceratium massiliense, dorsal view. 11. Ornithocercus magnificus,
right lateral view. 12. Ceratium fusus var. seta, ventral view. 13. Dinophysis caudata var. caudata, right lateral view. 14. Ceratium
macroceros, ventral view. 15. Protoperidinium divergens, ventral view. 16. Podolampas palmipes, ventral view. 17. Pyrophacus steinii,
antapical view. 18. Prorocentrum gracile. 19. Ceratium contortum var. contortum, vental view. 20. Prorocentrum micans. 21. Prorocentrum
compressum. 22. Ceratium tripos var. tripos, ventral view. Scales: Figs 13,15,22,23 = 5 µm; Figs 4,8,7,17,18,20 = 10 µm;
Figs 5,6–10,11,14,19 = 20 µm; Fig. 21 = 40 µm; Figs 16,24 = 50 µm; Fig. 12 = 100 µm.
Dinoflagellates in the Gulf of Mexico
Table 2.
Taxa
423
Table 2.
List of taxa and their occurrence†
Occurrence†
Actiniscales Sournia, 1984
Actiniscaceae Kützing, 1844
Actiniscus pentasterias Ehrenberg, 1854
Dinophysiales Lindemann, 1928
Citharistaceae Kofoid et Skogsberg, 1928
Citharistes apsteinii Schütt, 1895
Citharistes regius Stein, 1883
Dinophysiaceae Stein, 1883
Amphisolenia asymmetrica Kofoid, 1907
Amphisolenia bidentata Schröder, 1900
Amphisolenia bifurcata Murray et Whitting, 1899
Amphisolenia cf. elongata Kofoid et Skogsberg, 1928
Amphisolenia globifera Stein, 1883
Amphisolenia schroederi Kofoid, 1907
Dinophysis acuta Ehrenberg, 1839
Dinophysis argus (Stein) Abé, 1967
Dinophysis caudata var. caudata Saville Kent, 1881
Dinophysis caudata var. abbreviata Jörgensen, 1923
Dinophysis caudata var. pedunculata (Schmidt)
Schröder, 1906
Dinophysis circumsutum (Karsten) Balech, l967
Dinophysis cuneus (Schütt) Abé, 1967
Dinophysis diegensis Kofoid, 1907
Dinophysis doryphora (Stein) Abé, 1967
Dinophysis exigua Kofoid et Skogsberg, 1928
Dinohysis fortii Pavillard, 1923
Dinophysis hastata Stein, 1883
Dinophysis hindmarchii (Murray et Whitting)
Balech, 1967
Dinophysis laevis Cleparède et Lachmann, 1859
Dinophysis mitra (Schütt) Abé, 1967
Dinophysis operculoides (Schütt) Balech, 1967
Dinophysis ovata Cleparède et Lachmann, 1859
Dinophysis parvula (Schütt) Balech, 1967
Dinophysis pusilla Jörgensen, 1923
Dinophysis rapa (Stein) Abé, 1967
Dinophysis rotundata Cleparède et Lachmann,
1859
Dinophysis schroederi Pavillard, 1909
Dinophysis schuettii Murray et Whitting, 1899
Dinophysis sphaerica Stein
Dinophysis tripos Gourret, 1883
Dinophysis whittingae Balech, 1967
Heteroschisma inaequale Kofoid et Skogsberg, 1928
Heteroschisma cf. subantartica Balech, 1971
Histioneis crateriformis Stein, 1883
Histioneis hippoperoides Kofoid et Michener, 1911
Histioneis inclinata Kofoid et Michener, 1911
Histioneis jorgensenii Schiller, 1928
Histioneis pulchra Kofoid, 1907
Histioneis rotundata Kofoid et Michener, 1911
Histioneis striata Kofoid et Michener, 1911
Metaphalacroma skogsbergi Tai, 1934
Ornithocercus heteroporus Kofoid, 1907
Ornithocercus magnificus Stein, 1883
Ornithocercus quadratus Schütt, 1900
Ornithocercus splendidus Schütt, 1893
Ornithocercus steinii Schütt, 1900
Ornithocercus thumii (Schmidt) Kofoid et
Skogsberg, 1928
Triposolenia depressa Kofoid, 1906
Triposolenia intermedia Kofoid et Skogsberg, 1928
Oxyphysaceae Sournia, 1984
Oxyphysis oxytoxoides Kofoid, 1926
Gymnodiniales Lemmermann, 1910
Akashiwo sanguinea (Hirasaka)
G. Hansen et Moestrup, 2000
18
1
3
2
124
38
4
18
1
11
2
185
27
99
2
22
7
8
2
2
76
6
4
33
2
2
6
7
44
11
5
60
2
3
1
1
1
2
18
3
6
1
3
28
11
18
184
86
30
107
96
1
3
4
5
Taxa
List of taxa and their occurrence† (continued )
Occurrence†
Gymnodiniaceae Lankester, 1885
Amphidinium acutissimum Schiller, 1937
Amphidinium acutum Lohmann, 1920
Amphidinium carterae Hulburt, 1957
Amphidinium extensum Wulff, 1916
Amphidinium globosum Schröder, 1911
Amphidinium operculatum Cleparède et Lechmann,
1858–1859
Amphidinium schroderi Schiller, 1928
Amphidinium sphenoides Wulff, 1916
Balechina coerulea (Dogiel) Taylor, 1976
Gymnodinium grammaticum (Pouchet) Kofoid et
Swezy, 1921
Gymnodinium marinum Seville-Kent, 1880–1882
Gyrodinium falcatum sensu Kofoid et Swezy, 1921
Gyrodinium fusiforme Kofoid et Swezy, 1921
Karenia brevis (Davis) G. Hansen et Moestrup, 2000
Noctilucales Haeckel, 1894
Kofoidiniaceae Taylor 1976
Kofoidinium pavillardii J. Cachon et M. Cachon, 1967
Kofoidinium velleloides Pavillard, 1928
Noctilucaceae Kent, 1881
Pronoctiluca pelagica Fabre-Domergue, 1889
Oxyrrhinales Sournia, 1984
Oxyrrhinaceae Sournia, 1984
Oxyrrhis marina Dujardin, 1841
Peridiniales Haeckel, 1894
Ceratiaceae Kofoid, 1907
Ceratium arietinum var. arietinum Cleve, 1900
Ceratium arietinum var. gracilentum (Jörgensen)
Sournia, 1966
Ceratium azoricum Cleve, 1900
Ceratium belone Cleve, 1900
Ceratium bigelowi Kofoid, 1907
Ceratium breve var. breve (Ostenfeld et Schmidt)
Schröder, 1906
Ceratium breve var. parallelum (Schmidt) Jörgensen,
1911
Ceratium candelabrum var. candelabrum
(Ehrenberg) Stein, 1883
Ceratium candelabrum var. depressum (Pouchet)
Jörgensen, 1920
Ceratium carriense var. carriense Gourret, 1883
Ceratium carriense var. volans (Cleve) Jörgensen,
1911
Ceratium cephalotum (Lemmermann) Jörgensen,
1911
Ceratium concilians Jörgensen, 1920
Ceratium contortum var. contortum (Gourret)
Cleve, 1900
Ceratium contortum var. karsteinii (Pavillard)
Sournia, 1966
Ceratium contortum var. robustum (Karsten)
Sournia, 1966
Ceratium contrarium (Gourret) Pavillard, 1905
Ceratium declinatum (Karsten) Jörgensen, 1911
Ceratium deflexum (Kofoid) Jörgensen, 1911
Ceratium digitatum Schütt, 1895
Ceratium euarcuatum Jörgensen, 1920
Ceratium extensum (Gourret) Cleve, 1900
Ceratium falcatiforme Jörgensen, 1920
Ceratium falcatum (Kofoid) Jörgensen, 1920
Ceratium furca var. furca (Ehrenberg) Cleparède
et Lechmann 1859
Ceratium furca var. eugrammum (Ehrenberg)
Schiller, 1937
32
33
17
7
11
9
7
19
38
1
1
36
126
35
13
9
37
22
75
28
20
27
7
231
80
77
31
89
29
9
8
149
122
47
122
26
44
13
34
129
38
49
160
326
424
Table 2.
Taxa
S. Licea et al.
List of taxa and their occurrence† (continued)
Table 2.
Occurrence†
Ceratium fusus var. fusus (Ehrenberg) Dujardin,
1841
Ceratium fusus var. seta (Ehrenberg) Sournia, 1966
Ceratium geniculatum (Lemmermann) Cleve, 1901
Ceratium gibberum var. gibberum Gourret, 1883
Ceratium gibberum var. dispar (Pouchet) Sournia,
1966
Ceratium gravidum Gourret, 1883
Ceratium hexacanthum f. hexacanthum Gourret, 1883
Ceratium hexacanthum f. spirale (Kofoid) Schiller,
1937
Ceratium hircus var. horridum Schröder, 1909
Ceratium horridum (Cleve) Gran, 1902 var. horridum
Ceratium horridum var. molle (Kofoid) Jörgensen,
1911
Ceratium incisum (Karsten) Jörgensen, 1911
Ceratium inflatum (Kofoid) Jörgensen, 1911
Ceratium kofoidii Jörgensen, 1911
Ceratium limulus Gourret, 1883
Ceratium lineatum (Ehrenberg) Cleve, 1899
Ceratium longirostrum Gourret, 1883
Ceratium longissimum (Schröder) Kofoid, 1907
Ceratium lunula (Schimper) Jörgensen, 1911
Ceratium macroceros var. macroceros (Ehrenberg)
Vanhöffen, 1897
Ceratium macroceros var. gallicum (Kofoid) Sournia,
1966
Ceratium massiliense (Gourret) Jörgensen, 1911
Ceratium minutum Jörgensen, 1920
Ceratium paradoxides Cleve, 1900
Ceratium pentagonum var. pentagonum Gourret,
1883
Ceratium pentagonum var. tenerum Jörgensen, 1920
Ceratium platycorne Daday, 1888
Ceratium praelongum (Lemmermann) Kofoid, 1907
Ceratium ranipes Cleve, 1900
Ceratium setaceum Jörgensen, 1911
Ceratium symmetricum var. symmetricum Pavillard,
1905
Ceratium symmetricum var. coarctatum (Pavillard)
Graham et Bronikowsky, 1944
Ceratium tenue var. tenue Ostenfeld et Schmidt,
1901
Ceratium tenue var. buceros Balech, 1988
Ceratium tenue ver. tenuissimum (Kofoid) Graham
et Bronikovsky, 1944
Ceratium teres Kofoid, 1907
Ceratium trichoceros (Ehrenberg) Kofoid, 1908
Ceratium tripos var. tripos (O. F. Müller) Nitzsch,
1817
Ceratium tripos var. atlanticus (Ostenfeld) Paulsen,
1907
Ceratium tripos var. neglectum (Ostenfeld) Paulsen,
1907
Ceratium tripos var. pulchellum (Schröder) López,
1955
Ceratium vultur f. vultur Cleve, 1900
Ceratium vultur f. japonicum (Schröder) Wood, 1954
Ceratium vultur f. recurvum (Jörgensen) Schiller,
1937
Ceratium vultur f. sumatranum (Karsten) Sournia,
1966
Ceratocorythaceae Lindemann, 1928
Ceratocorys armata (Schütt) Kofoid, 1910
Ceratocorys horrida Stein, 1883
255
254
4
12
11
9
104
9
87
147
6
3
16
142
17
81
44
13
27
229
97
223
14
3
218
100
1
8
20
4
83
51
95
55
20
240
245
186
12
40
19
61
17
3
9
24
140
Taxa
List of taxa and their occurrence† (continued )
Occurrence†
Cladopyxidaceae Stein emend. Balech, 1913
Cladopyxis brachiolata Stein, 1883
Cladopyxis hemibrachiata Balech, 1964
Paleophalacroma unicinctum Schiller, 1928
Goniodoma sphaericum Murray et Whitting, 1899
Triadinium polyedricum (Pouchet) Dodge, 1981
Pyrodinium bahamense Plate, 1906 var. bahamense
Gonyaulacaceae Lindemann, 1928
Gonyaulax birostris Stein, 1883
Gonyaulax ceratocoroides (Murray et Whitting)
Kofoid, 1910
Gonyaulax fusiformis Graham, 1942
Gonyaulax fragilis (Schütt) Kofoid, 1911
Gonyaulax pacifica Kofoid, 1907
Gonyaulax polygramma Stein, 1883
Gonyaulax spinifera (Cleparède et Lachmann)
Diesing, 1866
Gonyaulax striata Manguin, 1922
Lingulodinium polyedrum (Stein) Dodge, 1989
Peridiniella danica (Paulsen) Okolodkov et Dodge,
1995
Peridiniella sphaeroidea Kofoid et Michener, 1911
Protoceratium spinulosum (Murray et Whitting)
Schiller, 1937
Heterodiniaceae Lindemann, 1928
Heterodinium agassizii Kofoid, 1907
Heterodinium dispar Kofoid et Adams, 1933
Heterodinium scrippsii Kofoid, 1906
Heterodinium whittingae Kofoid, 1906
Oxytoxaceae Lindemann, 1928
Corythodinium constrictum (Stein) Taylor, 1976
Corythodinium tesselatum (Stein) Loeblich et
Loeblich III, 1966
Oxytoxum adriaticum Schiller, 1937
Oxytoxum caudatum Schiller, 1937
Oxytoxum crassum Schiller, 1937
Oxytoxum elegans Pavillard, 1916
Oxytoxum elongatum Wood, 1962
Oxytoxum globosum Schiller, 1937
Oxytoxum gracile Schiller, 1937
Oxytoxum laticeps Schiller, 1937
Oxytoxum longiceps Schiller, 1937
Oxytoxum mediterraneum Schiller, 1937
Oxytoxum milneri Murray et Whitting, 1899
Oxytoxum ovale Schiller, 1937
Oxytoxum pachyderme Schiller, 1937
Oxytoxum parvum Schiller, 1937
Oxytoxum sceptrum (Stein) Schröder, 1900
Oxytoxum scolopax Stein, 1883
Oxytoxum sphaeroideum Stein, 1883
Oxytoxum turbo Kofoid, 1907
Oxytoxum variabile Schiller, 1937
Schuettiella mitra (Schütt) Balech, 1988
Peridiniaceae Ehrenberg, 1828
Diplopsalopsis globula Abé, 1941
Ensiculifera angulata Balech, 1988
Ensiculifera mexicana Balech, 1967
Protoperidinium abei (Paulsen) Balech, 1974
Protoperidinium brochii (Kofoid et Swezy) Balech,
1974
Protoperidinium cassum (Balech) Balech, 1974
Protoperidinium conicum (Gran) Balech, 1974
Protoperidinium crassipes (Kofoid) Balech, 1974
Protoperidinium depresum (Bailey) Balech, 1974
Protoperidinium divergens (Ehrenberg) Balech, 1974
Protoperidinium elegans (Cleve) Balech, 1974
26
18
8
3
6
14
39
5
33
2
35
132
6
6
45
8
1
8
1
1
3
1
19
16
4
11
9
46
1
20
125
67
7
20
10
25
1
36
28
103
1
28
5
13
5
1
4
1
1
26
64
9
14
202
48
Dinoflagellates in the Gulf of Mexico
Table 2.
425
List of taxa and their occurrence† (continued)
Taxa
Occurrence†
Protoperidinium fartum Balech, 1979
Protoperidinium grande (Kofoid) Balech, 1974
Protoperidinium jorgenseni Balech, 1971
Protoperidinium longipes (Karsten) Balech, 1974
Protoperidinium mediterraneum (Kofoid) Balech,
1974
Protoperidinium nipponicum (Abé) Balech, 1974
Protoperidinium oblongum (Aurivillus) Park et
Dodge, 1976
Protoperidinium oceanicum (Vanhöffen) Balech, 1974
Protoperidinium oviforme (Dangeard) Balech, 1974
Protoperidinium ovum (Schiller) Balech, 1974
Protoperidinium pallidum var. daedalum Balech,
1978
Protoperidinium pellucidum var. stellatum Balech,
1978
Protoperidinium pentagonum (Gran) Balech, 1974
Protoperidinium poucheti (Kofoid et Michener)
Taylor et Balech, 1988
Protoperidinium punctulatum (Paulsen) Balech, 1974
Protoperidinium cf. rectum (Kofoid) Balech, 1974
Protoperidinium subsphaericum (Balech) Balech,
1974
Protoperidinium tuba (Schiller) Balech, 1974
Protoperidinium venustum (Matzenauer) Balech, 1974
Protoperidinium vulgare Balech, 1978
Scrippsiella trochoidea (Stein) Fine et Loeblich III,
1976
Podolampadaceae Lindemann, 1928
Blepharocysta splendor-maris (Ehrenberg) Ehrenberg,
1873
Podolampas bipes Stein, 1883
Podolampas elegans Schütt, 1895
Podolampas palmipes Stein, 1883
Podolampas reticulata Kofoid, 1907
Podolampas spinifera Okamura, 1912
Pyrophacus horologium Stein, 1383
Pyrophacus steinii (Stein) Wall et Dale, 1971
Pyrophacus vancanpoae (Rossignol) Wall et Dale,
1972
Peridiniales incertae sedis
Spiraulax kofoidii Grabam, 1942
Prorocentrales Lemmermann, 1910
Prorocentraceae Stein, 1883
Prorocentrum aporum (Schiller) Dodge, 1975
Prorocentrum balticum (Lohmann) Loeblich, 1970
Prorocentrum compressum (Bailey), Abé 1967
Prorocentrum cordatum (Ostenfeld), Dodge 1976
Prorocentrum dentatum, Stein 1883
Prorocentrum gibbosum (Schiller) Schiller, 1933
Prorocentrum gracile Schütt, 1895
Prorocentrum cf. lima (Ehrenberg) Dodge, 1975
Prorocentrum mexicanum Osorio-Tafall, 1942
Prorocentrum micans Ehrenberg, 1833
Prorocentrum minimum (Pavillard) Schiller, 1933
Prorocentrum rostratum Stein, 1883
Prorocentrum sigmoides Böhm, 1933
Prorocentrum triestinum Schiller, 1918
Pyrocystales Apstein, 1909
Pyrocystaceae (Schütt) Lemmermann, 1899
Dissodinium elegans (Pavillard) Matzenauer, 1933
Pyrocystis fusiformis Wyviile-Thomson, 1885
Pyrocystis humulus Cleve, 1900
Pyrocystis lunula (Schütt) Schütt, 1896
Pyrocystis obtusa Pavillard, 1931
Pyrocystis robusta Kofoid, 1907
†
Number of sites where species were found.
1
120
1
35
125
6
37
97
1
30
3
1
78
1
2
4
1
68
5
3
10
13
102
65
192
14
92
81
133
18
and Dinophysis tripos, all of which widely distributed,
although in small numbers. A. carterae has been noted
to produce toxic substances (Steidinger 1993). Prorocentrum cf. lima was found in 32 localities in the
region-II (Bay of Campeche). A. carterae, D. caudata,
D. fortii, D. tripos, D. mitra and D. rotundata were
present mostly in net samples.
Based on the analysis of water samples, Amphidinium carterae were found to have a brief blooming
period in the region-III (2 × 104 cells l–1). P. micans
and D. caudata had fluctuations between >103 and
<104 cells/L in March 2000. Pyrodinium bahamense
var. bahamense had values of <105 cells/L, mostly at
inlets of some coastal lagoons.
Occurrence data show that Prorocentrum mexicanum and D. tripos are species widely distributed in
small numbers. Prorocentrum minimum, D. acuta and
D. mitra were occasionally found only in net samples.
P. micans showed its maximum frequency during
summer and autumn.
A red tide of Scrippsiella trochoidea was found in
concentrations of about 300 000 cells/L north-east of
Cape Catoche on the Yucatan Shelf. This bloom caused
a brown-greenish water discoloration in early summer
(July–August), representing 99.6% of the total phytoplankton population (Licea unpubl. data 1998).
The potentially harmful species C. furca var. furca,
P. bahamense var. bahamense, S. trochoidea and Gonyaulax polygramma were found forming blooms during
summer. In all cases every species was widely distributed, with the exception of S. trochoidea, which
bloomed at only 7 stations located east of region-III,
where a regular upwelling occurs.
DISCUSSION
6
23
29
168
1
18
1
160
32
100
143
10
2
34
4
34
39
4
20
47
83
Our results agree with the considerable constancy in
the dinoflagellate species composition found in the
Caribbean Sea (Balech 1967a; Wood 1968, 1969;
Zernova 1969, 1970, 1974) and the Straits of Florida
(Steidinger & Williams 1970). The latter author
explains this similarity on the basis of the fact that a
great part of the waters of Florida are derived from the
Caribbean Sea through the Yucatan Channel, with only
a small contribution from the Gulf of Mexico. The small
differences found in species composition are probably
a result of regional hydrographic conditions that occur,
particularly in region-II. D. acuta, Heteroschisma cf.
subantartica, Amphidinium acutissimum, A. globosum
and Prorocentrum cf. lima, which were found in that
region, are reported for the first time for the Gulf of
Mexico.
Data suggest that species distribution is mainly
associated with local hydrographic conditions and
species requirements. According to the distribution
patterns of Ceratium arietinum, Ceratium azoricum,
C. contortum, Ceratium euarcuatum, Ceratium tenue,
426
Ornithocercus quadratus, Oxytoxum sceptrum and
Podolampas spinifera, these species were found close
to the limit of a hyaline front located in the northern
coast of the Tamiahua lagoon (Region-I), where several
rivers discharge and nutrients were relatively high, as
compared with the adjacent area. In contrast, the same
species were absent at salinities below 32 PSU and
because of oligotrophic conditions (Estradas-Romero
2004). In addition, this region was characterized by
the presence of anticyclonic gyres that drift in from the
Loop Current (Vidal et al. 1992), which indicates a
lack of nutrients. The same species were present at
the western part of region-II near the shelf where the
discharge of rivers was evident. These facts, combined
with littoral circulation associated with cyclonic gyres
drifting from a branch of the Loop Current (Monreal &
Salas de León 1990), created contrasting hydrographical conditions that might or might not support the
survival of species, therefore affecting their distribution. The same group of species occurred simultaneously along the shelf of Yucatan, associated with
the upwelled water described by Merino (1997) at the
eastern region-III. In contrast, the eastern part of
region-II has no rivers, but it does have a wide shelf
and high temperature and salinity (Alatorre et al. 1987),
which are advantageous conditions for the settlement
of blooms of Oxyrrhis marina and P. bahamense.
Therefore, it can be concluded that region-II is a
transition between region-I and region-III.
In the early summer at the eastern region-III (Fig. 1),
a water discoloration by S. trochoidea was observed,
apparently associated with the upwelling that occurs in
the north of Cape Catoche, suggesting that nutrients
contribute to the bloom settlement of this species.
However, the presence of many cysts (not counted)
during this time indicates the beginning of the development of the first two of four blooming stages of algae,
described by Steidinger (1983). Therefore, the blooming
in question manifests itself more prominently, followed
by the third and fourth stages of the development of this
species, when water discoloration is visible; then it
moves and is scattered over the Yucatan Shelf. The
dominance of dinoflagellates during summer shows that
it is precisely at this time of the year when the most
appropriate conditions exist in the Gulf of Mexico.
ACKNOWLEDGMENTS
Funds for this research were provided by the Institute
of Marine Science and Limnology (ICMYL-UNAM). We
are grateful to National Commission for the Knowledge
and Use of Diversity for financial support for 2 years
(grants: FB683/S088 and A0012/2002). We extend
our deep appreciation to the crew members of the
RV JUSTO SIERRA and the participating scientists
in the PROGMEX, OGMEX and SGM cruises for their
invaluable support, to Yuri Okolodkov for some species
S. Licea et al.
identification, to Joseph Doshner for reading and
commenting on the English and to Jorge Sepulveda for
scanning electron microscopy technical assistance.
REFERENCES
Alatorre, M. A., Ruiz, F. and Salas de León, D. 1987. Efecto
Del Paso de Frentes Fríos Atmosféricos Sobre la Bahía de
Campeche. In Gonzàlez, J. F., Medina, M. and Martínez,
M. (Eds) Memoria Reunión Anual 1987. Unión Geofísica
Mexicana, México, pp. 186–93.
Balech, E. 1967a. Dinoflagellates and tintinnids in the northeastern Gulf of Mexico. Bull. Mart. Sci. 17: 280–98.
Balech, E. 1967b. Dinoflagelados nuevos o interesantes del
Golfo de México y Caribe. Rev. Mus. Argent. Cienc. Nat.
(Hidrobiol.) 2: 77–126.
Bessonov, N., Gonzalez, O. and Elizarov, A. 1971. Resultados
de Las Investigaciones Cubano-Soviéticas En El Banco de
Campeche. In UNESCO (Eds) Coloquio sobre investigaciones y recursos del mar Caribe y regiones adyacentes,
Curaçao, Antillas Holandesas, November 1968. UNESCO,
París, pp. 317–23.
Bogdanov, D. V., Sokolov, V. A. and Khromov, N. S. 1968.
Regions of high biological and commercial productivity in
the Gulf of Mexico and the Caribbean Sea. Oceanology
(USSR) 8: 466–78. (In Russian with English summary.)
Cochrane, J. D. 1969. Water and circulation on Bank of
Campeche in May. Bull. Jpn. Soc. Fish. Oceanogr. Spec.
no. (Prof. Uda’s Commemorative Papers), pp. 123–8.
Crétinnot-Dinet, M. J., Sournia, A., Ricard, M. and Billard, C.
1993. A classification of the marine phytoplankton of the
world from class to genus. J. Phycol. 32: 159–79.
de la Cruz, A. 1971. Estudios del plancton en el Banco de
Campeche. In UNESCO (Ed.) Coloquio sobre investigaciones y recursos del Mar. Caribe y regiones adyacentes,
Curacao, Antillas Holandesas, Nov. 1968. UNESCO,
Paris, pp. 375–83.
Curl, H. Jr. 1959. The phytoplankton of Apalache Bay and the
northeastern Gulf of Mexico. Publ. Inst. Mar. Sci. Univ.
Texas 6: 277–320.
Estradas-Romero, A. 2004. Phytoplankton abundance and
distribution in two transects off the Coatzacoalcos river
and Grijalva-Usumasinta river systems (March 2000).
Ms Thesis, Universidad Nacional Autónoma de México,
Mexico, 70 pp. (In Spanish with English summary.)
Fideicomiso Fondo para la Biodiversidad, CONABIO. 2003.
Sistema de Información Biótica. Versión 4.1. México, D.F,
704 pp.
Furnas, M. J. and Smayda, T. J. 1987. Inputs of subthermocline waters and nitrate onto the Campeche Bank. Continental Shelf Res. 7: 161–75.
Gómez-Aguirre, S. and Licea, S. 1998. Blooms of Pyrodinium
bahamense (Dinophyceae) in coastal lagoons of the southern Gulf of Mexico and Mexican Caribbean. In Reguera,
B., Fernández, M. L. and Wyatt, T. (Eds) Harmful Algae.
Xunta de Galicia and International Oceanogr. Comm.
UNESCO, Paris, pp. 61–2.
Dinoflagellates in the Gulf of Mexico
Graham, H. W. 1954. Dinoflagellates of the Gulf of Mexico.
In Galtsoff, P. (Ed.) Gulf of Mexico-its origin, waters, and
marine life. US Fish and Wildlife Serv., Fishery Bulletin
89, Vol. 55, Washington, 223–6.
Hasle, G. R. 1978. The inverted microscope method In
Sournia, A. (Ed.) Phytoplankton Manual, UNESCO, Paris,
pp. 136–42.
Hulburt, H. E. and Thompson, J. D. 1980. A numerical study
of Loop Current intrusions and eddy shedding. Journal of
Physical Oceanography. 10: 1611–51.
Khromov, N. S. 1965. Distribution of plankton in the Gulf of
Mexico and some aspect of its seasonal dynamics. In.
Bogdanov, D. V. (Ed.) Soviet Cuban Fishery Research.
(VNIRO-CIP), Moscow, pp. 47–69.
Krylov, V. V. 1974. Distribución del fitoplancton y de las
biocenosis planctónicas en el Banco de Campeche. Rev.
Invest. Inst Nal. Pesca 1: 75–9.
Licea, S. 1977. Variación estacional del fitoplancton de la
Bahía de Campeche, México (1971–72). FAO Fish. Report
200: 253–73.
Licea, S. and Santoyo, H. 1991. Algunos aspectos ecológicos
del fitoplancton de la región central de la Bahía de
Campeche. Ann. Int. Cienc. Mar. Limnol. University Nal.
Autón. México 18: 157–67.
Licea, S., Zamudio, M. E., Luna, R., Okolodkov, Y. B. and
Gómez-Aguirre, S. 2003. Toxic and harmful dinoflagellates in the southern Gulf of Mexico In Steidinger, K. A.,
Landsberg, J. H., Tomas, C. and Vargo, G. A. (Eds)
Harmful Algae 2002. Proceedings of the Xth International
Conference on Harmful Algae. Florida Fish. Wild. Conserv.
Comm. Inter. Oceanogr. Comm. UNESCO, St. Petersburg,
Florida, (in press).
Merino, M. 1997. Upwelling on the Yucatan Shelf: hydrographic evidence. J. Mar. Syst. 13: 101–21.
Merino, M., Czitrom, S., Jordan, E., Martín, E., Thomé, P. and
Moreno, O. 1990. Hydrology and rain flushing of the
Nichupté Lagoon System, Cancum, Mexico. Estuarine,
Coastal Shelf Sci. 30: 223–37.
Merino, M. and Otero, L. 1991. Atlas Ambiental Costero,
Puerto Morelos-Quintana Roo. Centro Investigaciones
Quintana Roo. (Eds). Chetumal, México, 80 pp.
Merrel, W. J. Jr and Morrison, J. M. 1981. On the circulation
of the western Gulf of Mexico with observations from
1978., J. Geophy. Res. 86: 4181–5.
Monreal-Gómez, M. A. and Salas de León, D. A. 1985.
Barotropic and baroclinic modes in the Gulf of Mexico.
In Grieken, V. and Wollast, R. (Eds). Proceedings of
the Progress in Belgian Oceanogr. Res., Antwerpen
University Press, Brussels, pp. 81–91.
Monreal-Gómez, M. A. and Salas de León, D. 1990. A.
Simulación de la circulación en la Bahía de Campeche.
Geofisica Int. 29: 101–11.
Norris, D. R. and Berner, L. D. Jr. 1970. Thecal morphology
of selected species of Dinophysis (Dinoflagellata) from the
Gulf of Mexico. Contrib. Mar. Sci. 15: 145–92.
Nowlin, W. D. Jr. 1972. Winter circulation patterns and
property distributions. In Capurro, L. R. A. and Reid, J. L.
427
(Eds) Contributions on the Physical Oceanography of the
Gulf of Mexico. Gulf Publishers Co, Houston, pp. 3–52.
Roberts, B. S. 1979. Occurrence of Gymnodinium breve red
tides along the west and east coasts of Florida during
1976 and 1977. In Taylorand, D. L. and Selioger, H. H.
(Eds) Toxic Dinoflagellate Blooms. Elsevier/North Holland,
New York, pp. 199–202.
Roujiyaynen, M. I., Georgieva, L. V. and Senichkina, L. G.
1968. Composition. quantitative development and distribution of phytoplankton in the Central-American seas.
Studies on the Central-American seas, 2, Kiev, pp. 14–39.
(In Russian with English and Spanish summaries.)
Santoyo, H. and Signoret, M. 1973. Hidrología y fitoplancton
en un transecto en la plataforma continental de la Bahía
de Campeche, México (agosto 1972). Rev. Lat.-Am.
Microbiol. 15: 207–15.
Santoyo, H. and Signoret, M. 1975. Variación nictemeral del
fitoplancton en la Bahía de Campeche, México. Rev. Lat.Am. Microbiol. 17: 567–96.
Steidinger, K. 1971. Gonyaulax balechi sp. Nov (Dinophyceae)
with a discussion of the genera Gonyaulax and Heteraulacus. Phycologia. 10: 183–7.
Steidinger, K. A. and Baden, D. G. 1984. Toxic marine
dinoflagellates. In Spector, D. L. (Ed.) Dinoflagellates.
Academic Press, Orlando, pp. 201–61.
Steidinger, K. A., Davis, J. T. and Williams, J. 1967. A key to
the marine dinoflagellate genera of the west coast of
Florida. Technical Series, no. 52, Florida Department Nat.
Res. Mar. Res. Laboratory, St. Petersburg, Florida, 45 pp.
Steidinger, K. A., Truby, E. W. and Dawes, C. 1978. J:
Ultrastructure of the red tide dinoflagellate Gymnodinium
breve. I. General description. J. Phycol. 14: 72–9.
Steidinger, K. A. and Williams, J. 1970. Dinoflagellates,
Memoirs of the Hourglass Cruises, Vol. II. Florida Department Nat. Res. Mar. Res. Laboratory, St. Petersburg,
Florida, 251 pp.
Steidinger, K. A. 1983. A re-evaluation of toxic dinoflagellate
biology and ecology. In Round, F. E. and Chapman, D. J.
(Eds) Progress in Phycological Research. Elsevier. Science
Publishing, New York, pp. 147–88.
Steidinger, K. A. 1993. Some taxonomic and biologic aspects
of toxic dinoflagellates. In Falconer, I. R. (Ed.) Algal toxins
in sea food and drinking water. Academic Press, London,
pp. 1–28.
Tamayo, J. L. 1990. Geografía Moderna de México. Trillas
(Eds.) México.
Tester, P. A., Stumpf, R. P., Vukovich, F. M., Fowler, P. K.
and Rurner, J. T. 1991. An expatriate red tide bloomTransport, distribution, and persistence. Limnol. Oceanogr. 36: 1053–61.
Vidal, V. M., Vidal, F. V. and Pérez-Moreno, J. M. 1992.
Collision of a Loop Current anticyclonic ring against the
continental shelf slope of the western Gulf of Mexico. J.
Geophys. Res. 97: 2155–72.
Vinogradova, L. A. 1976. Influence of upwelling on the development of phytoplankton in the area of the Bank of Campeche.
Proceedings AtlantNIRO 63: 81–9. (In Russian.)
428
Williams, J. and Ingle, R. M. 1972. Ecological notes on
Gonyaulax monilata (Dinophyceae) blooms along the west
coast of Florida. Leaflet Series, 1, Pt. No. 5. Florida
Department Nat. Res. Mar. Res. Laboratory, St. Petersburg, Florida, pp. 1–12.
Wood, E. J. F. 1968. Dinoflagellates of the Caribbean Region
and Adjacent Waters. Miami University Press, Miami,
Florida, 144 pp.
Wood, E. J. 1969. F: Relations of phytoplankton to marine
habitat in the Straits of Florida and adjacent areas. I. The
Dinoflagellates. Nova Hedwigia 28: 645–763.
Zernova, V. V. 1969. The horizontal distribution of phytoplankton in the Gulf of Mexico. Oceanology (USSR) 9:
695–706. (In Russian with English summary.)
Zernova, V. V. 1970. About planktonic algae of the Gulf of
Mexico and the Caribbean Sea. Oceanol. Studies (USSR)
S. Licea et al.
20: 69–104. (In Russian with English and Spanish summaries.)
Zernova, V. V. 1974. Species structure of the phytocene in
the southern Gulf of Mexico. Soviet-Cuban Fishery Res.
(VNIRO-CIP) Moscow 4: 117–31. (In Russian with
Spanish summary.)
Zernova, V. V. 1982. On the dependence of the quantitative
development of phytoplankton on the abiotic factors in the
Gulf of Mexico. Inst Oceanol. USSR Acad. Sci. 114:
60–72. (In Russian with English summary.)
Zernova, V. V. and Zhitina, L. S. 1985. About the phytoplankton community of the American Mediterranean
Sea. Studies on the Oceanic Phytoplankton. Russian Institute of Marine Fisheries and Oceanography, Moscow,
pp. 27–36. (In Russian.)