Cembella et al: Cladistic analysis of the toxic dinoflagellate Protogonyaulax 39
Botanica Marina
Vol. 31, pp. 39-51, 1988
Cladistic Analysis of Electrophoretic Variants
within the Toxic Dinoflagellate Genus Protogonyaulax
A. D. Cembella*'**, R J. R. Taylor** and J.-C. Therriault*
* Maurice-Lamontagne Institute, Department of Fisheries and Oceans, 850 route de la M er, P.O. Box 1000,
Mont-Joli, Quebec, Canada G5H 3Z4
** Departments of Oceanography and Botany, University of British Columbia, 6270 University Boulevard,
Vancouver, British Columbia, Canada V6T 1W5
This publication is respectfully and affectionately dedicated to Dr NavalJ. Antia on the occasion of his retirement,
for his outstanding contributions to the field of marine science.
(Accepted 12 October 1987)
Abstract
The systematic position of the Protogonyaulax tamarensisjcatenella 'species complex', a group of morphol-
ogically related toxic marine dinoflagellates, has been the subject of recent debate, at both the infraspecific
and the generic level. Since the conventional taxonomy of thecate dinoflagellates, based almost entirely on
morphological features, has often yielded controversial or inconsistent taxonomic groupings, a biochemical
approach was adopted. Isolates of the genus Protogonyaulax from diverse geographical locations were subjected
to isozyme analysis by polyacrylamide gel electrophöresis (PAGE) of soluble enzymes. The isozyme bands
were coded as independent character data for cladistic reconstruction using a phylogenetic analysis program
to generate Wagner trees. The isozyme character data were also converted to indicate Manhattan distances
between taxa, which were then used to create alternative Wagner distance trees.
The Wagner character analysis produced only a single minimum length mid-point rooted tree, with
tamarensoid and intermediate morphotypes from the southern Strait of Georgia, British Columbia, placed
on the upper major branch, while catenelloid and tamarensoid morphotypes from other locations aligned
along the lower branch. The alternative optimized distance Wagner trees exhibited similar, but not identical,
basic branching patterns, and indicated an closer apparent relationship between the two European tamarensoid
morphotypes than was evident from direct character analysis. Neither cladistic reconstruction supported a
definitive dichotomy between the conventionally recognized major morphotypes within this group.
The extended length of the Wagner trees, as well as the high degree of apomorphism between hypothetical
nodes, indicated substantial genetic divergence, even among contemporaneous geographical populations. This
level of divergence was not reflected in corresponding morphological character differences, and suggests that
sibling speciation may have occurred among these dinoflagellates.
t
. phological criteria employed for taxonomic analysis.
oduction
The systematic
p
OS
ition of these toxin-producing di-
The dinoflagellate genus Protogonyaulax Taylor noflagellates, at both the generic and infraspecific
(1979) comprises photosynthetic toxigenic species levels, remains in dispute (Steidinger 1983, Taylor
which are primarily responsible for paralytic shellfish 1984, 1985, Balech 1985). Attempts to consistently
poisoning (PSP) in coastal marine ecosystems (Taylor discriminate stable species and varieties within this
1984). As for other thecate dinoflagellates, the num- group from biochemical and morphological evidence
ber, shape and orientation of the cellulosic plates have proved to be highly controversial (Loeblich and
forming the cell wall or 'theca' are the principal mor- Loeblich 1979, Taylor 1979, Schmidt et al. 1978,
Botanica Marina / Vol. 31 / 1988 / Fase, l
Copyright © 1988 Walter de jGruyter · Berlin · New York
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