Biochemical Systematics and Ecology,Vol. 21, No. 6/7, pp. 729-735, 1993. 0305-1978/93 $6.00+0.

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Printed in GreatBritain. © 1993PergamonPressLtd.

Biochemical Genetics and Systematics of Millepora

(Coelenterata: Hydrozoa) from the Shore of South Vietnam

GENNADY P. MANCHENKO, ALEXANDER V. MOSCHENKO

and VYATCHESLAV S. ODINTSOV
Institute of Marine Biology, Far East Division of the Russian Academy of Sciences, Vladivostok 690041, Russia

Key Word Index--Hydroid genus Millepora; Hydrozoa; growth forms; isoenzymes; electrophoresis;
biochemical systematics.
Abstract--Throe sympatrically occurring forms of colony growth of hydroid Millepora from the shore of
South Vietnam were electrophoretically compared for the products of 21 loci coding for isoenzymes. "Plate-
like" growth form was found to be genetically isolated from "branch-like" and "comb-like" forms which
presumably represent different morphological variants of a single species.

Introduction
Hydroids of the genus Millepora are widely distributed in warm waters of the World
Ocean. They are characterized by high phenotypic plasticity which is supposed by
some authors to be of intraspecific origin (Hickson, 1898; Estalella, 1982). At the same
time, morphological differences including differences in growth form are usually used
as main taxonomic criteria in their systematics (Wood-Jones, 1907; Boschma, 1948;
Weerdt, 1981, 1984). Based on morphological differences, nine species of Millepora are
described for the Indo-Pacific (Boschma, 1948; Zou, 1978).
It is obvious now that numerous problems existing in coral systematics are caused
by some peculiarities of coral proliferation and spreading. Reproduction by fragmenta-
tion is known to be the predominant mode of proliferation and local dissemination in
many coral species, and is not excluded for Millepora (Highsmith, 1982). In such a
situation numerous colonies originating from one parent colony are genetically
identical. In some coral species, however, sexual reproduction may be predominant. In
these cases the colonies originating from sexually produced meiotic planulae are
genetically heterogenous. Species with different modes of reproduction differ in the
genetic structure of their natural populations. Until recently the immunological test of
histocompatibility was used to define the predominant mode of reproduction of the
local coral populations (Hildemann eta/., 1977; Johnston eta/., 1981; Neigel and Avise,
1983). This approach is based on the supposition that two separate colonies are
histocompatible if they are genetically identical (asexual reproduction) and are not
compatible if genetically different (sexual reproduction). However, direct genotyping of
multiple loci coding for enzymes has shown that this supposition is not absolutely
correct (Heyward and Stoddart, 1985; Willis and Ayre, 1985). Such genotype definition
became possible due to the electrophoretic method of isoenzyme analysis. Using
isoenzymes it was shown, for example, that different growth forms of the scleractinian
coral Pavona cactus are intraspecific variants determined by different genotypes (Willis
and Ayre, 1985). The other successful application of isozymes is the discovery of
ameiotic asexually produced planulae in some scleractinian corals (Stoddart, 1983;
Ayre and Resing, 1986). Isozymes were used as genic markers in detailed studies of the
spatial genetic structure of natural populations in the scleractinian coral Pocillopora
damicornis (Stoddart, 1984a, b). It was found that in spite of a planktonic larval stage in
the life cycle of the coral, separate settlements of R damicomis differ from one another
(Received 13 March 1992)

729
730 G.P. MANCHENKO ETAL.

more drastically than the settlements of other invertebrates with planktonic larvae. The
R damicornis settlements of a few hundred metres in length were found to be a
mixture of a few genetic clones. It is obvious that substantial contribution of genetic
clones in the formation of spatial population structure on the one hand and genetic
determination of intraspecific morphological variants on the other, may be the real
cause of numerous taxonomic problems as well as the coral species problem in
general (Braekel, 1977; Veron, 1981).
Sexual reproduction in Millepora is well defined (Boschma, 1956; Eguchi, 1968).
Though reproduction by fragmentation is also not fully excluded, there are some
reasons to think that sexual reproduction predominates in this hydroid (Highsmith,
1982). The problem of taxonomic status of different growth forms of Millepora is about
three and a half centuries old (Boschma, 1948). Here we describe the results of a
comparative electrophoretic study of enzymes in different growth forms of Millepora
from the shore of South Vietnam in order to clarify their systematic interrrelationships.

Materials and Methods

Three sympatric growth forms of hydroid M/I/epora--"branch-like" (B), "comb-like" (C) and "plate-like" (P)--
were collected from the shore of South Vietnam (Kondor Island: B--five colonies, P--seven colonies; Tkhotyu
Island: P--18 colonies; Sharlott Bank: B--eight colonies, P--11 colonies; Nha Trang Bay: B--24 colonies, C--13
colonies, P--13 colonies) during the eighth Soviet-Vietnamese marine expedition on the scientific ship
"Academic Alexander Nesmejanov" in January-March 1986.
Starch gel electrophoresis was carried out using enzymes derived from the tissues of colony branch tips free
of such macrosymbionts as sponges, crustaceans and polychaetes. Branch tips were homogenized in 2 vol of
distilled water. Individual homogenates were absorbed on to small squares of Whatman 3 MM chromato-
graphic paper and inserted into a 14% starch gel block (5×120x200 mm). Two continuous buffer systems were
used: Tris-EDTA-boric acid, pH 8.5 (TEB) and Tris-citric acid, pH 7.0 (TC). During electrophoresis the gel blocks
were cooled with ice. After completion of electrophoresis a gel block was sliced horizontally into 1.5 mm slices.
The gel slices were then histochemically stained for enzyme activity as previously described (Harris and
Hopkinson, 1976; Serov et al., 1977). The enzymes studied, their abbreviations, presumptive gene loci coding
for isozyme and electrophoretic conditions are listed in Table 1.
Genetic interpretations were inferred from comparisons of electrophoretic patterns of individual colonies
with those described for the same enzymes in other animals where a genetic basis of enzyme polymorphisms
was well established. Electrophoretic patterns of the majority of studied enzymes were similar to those of other
diploid sexually reproducing organisms. We did not observe any serious complications in isozyme patterns
caused by symbiotic zooxanthellae. The comparison of isozyme patterns obtained after electrophoresis of
homogenates prepared from upper layers of branch tips enriched by zooxanthellae with deeper layers revealed
no substantial differences, at least in sets of major bands. After prolonged gel staining, additional bands
were observed on GOT, GPI, MDH and TPEP zymograms. These bands perhaps originated from symbiotic
zooxanthellae. They were not taken into account in calculations of genetic identity and genetic distance. The

TABLE 1. ENZYMES STUDIED, GENIC LOCI CODING FOR ISOENZYMES AND ELECTROPHORETIC
CONDITIONS

Electrophoretic
Enzymes Loci* bufferst

Hexokinase (HK) Hk TEB

Glutamateoxaloacetatetransaminase(GOT) Got4,2,3 TEB
Glutamatepyruvatetransaminase (GPT) Gpt-1,2 TEB
Glucosephosphateisomerase(GPI) Gpi TEB
Dipeptidase Gly-Leu (DPEP) Dpep-1,2,3 TEB
Isocitrate dehydrogenase (IDH) Idh TC
Malate dehydrogenase (MDH) Mdh-1,2 TC
Methylumbelliferylacetate esterase (MUEST) Muest TEB
Superoxide dismutase (SOD) Sod-1,2 TEB
Tripeptidase Leu-Gly-Gly (TPEP) Tpep-1,2,3,4 TEB
Phosphoglucomutase (PGM) Pgm TEB

*Multiple loci are numbered according to the position of corresponding isoenzymes from anode to
cathode.
l-The composition of buffer systems is given in the text.
GENETICSAND SYSTEMATICSOF MILLEPORA 731

estimates of Nei's indices were calculated using allelic frequencies at polymorphic and monomorphic loci (Nei,
1972).

Results and Discussion

The electrophoretic examination of 11 enzyme systems (Table 1) allowed the resolution
of the gene products of 21 presumptive loci in the branch-like and comb-like growth
forms and 20 loci in the plate-like form of milleporine hydroids, According to the
modern Mi//epora classification based on species specificity of growth form (Boschma,
1948), these forms were identified as M. intricata, M. dichotoma and M. platyphylla,
respectively (Fig. 1).
AIIozymic variation with one-banded homozygotes and two-banded heterozygotes
indicating monomeric subunit structure of enzyme molecules was revealed in HK and
PGM. One-banded homozygotes and three-banded heterozygotes were observed on
zymograms of dimeric enzymes GPI and GOT. Subunit structures of other polymorphic
enzymes were not inferred from their electrophoretic patterns because different
allozymes of these enzymes were developed on zymograms as broad diffuse bands
which were not clearly resolved in heterozygotes.
The allele frequencies, observed and expected heterozygosity (H) values for poly-
morphic and monomorphic loci in three Millepora growth forms are presented in Table
2. Genetic variability estimates are summarized in Table 3. These estimates are
approximately the same as in other coelenterates including actinian (Nevo et al., 1984)
and scleractinian (Manchenko and Latypov, unpublished observations) species.
No fixed differences in allele frequencies are observed between branch-like and
comb-like growth forms (Table 2). Such differences, however, are found between the
plate-like growth form on the one hand and the branch-like and comb-like form on the

FIG. 1. DIFFERENT GROWTH FORMS OF MILLEPORA. (A) "Plate-like" (M. plab/phylla); (B) "comb-like" (M. dichotoma); (C)
"branch-like" (114.intricata).
732 G.P. MANCHENKO ETAL

TABLE 2. ALLELE FREQUENCIESAND OBSERVED (Hob) AND EXPECTED (He~) HETEROZYGOSITIESOF GENE LOCI STUDIED IN
THREE GROWTH FORMS OF MILLEPORA: "PLATE-LIKE" (P), "COMB-LIKE" (C) AND "BRANCH-LIKE" (B)

Alleles
Growth
Loci form (N)* 1 2 3 4 5 6 Sst Hob H~×

1 2 3 4 5 6 7 8 9 10 11

Dpep-1 P (19) 1.000 0.000 0.000

C (13) 1.000 ? 0.000 0.000
B (20) 0.050 0.725 0.225 0.450 0.421
Dpep-2 P (19) 1.000 0.000 0.000
C (13) 1.000 -- 0.000 0.000
B (27) 1.000 0.000 0.000
Dpep-3 P (19) 1.000 0.000 0.000
C (13) 1.000 ? 0.000 0.000
B (29) 0.966 0.034 0.069 0.066
Got-1 P (37) 1.000 0.000 0.009
C (13) 1.000 D 0.000 O.000
B (31) 0.032 0.968 0.064 0.062
Got-2 P (29) 0.897 0.103 0.138 0.185
C (13) 1.000 ? 0.000 0.000
B (11) 0.818 0.182 0.364 0.298
Got-3 P (37) 0.027 0.960 0.013 0.081 0.077
C (13) 1.000 ? 0.000 0.000
B (31) 1.000 0.000 0.000
Gpi P (41) 0.707 0.110 0.183 0.537 0.455
C (13) 0.192 0.769 0.038 D 0.308 0.370
B (35) 0.643 0~314 0.043 0.400 0.486
Gpt-1 P (34) 1.000 0.000 0.000
C (8) 1.000 ? 0.000 0.000
B (23) 0.087 0.348 0.565 0174 0.552
Gpt-2 P (29) 0.034 0.793 0.172 0.138 0.340
C (8) 1.000 ? 0.000 0.000
B (14) 0.893 0.071 0.036 0214 0.196
Hk P (34) 0.029 0.897 0.073 0.088 0.189
C (13) 1.000 M 0.000 0.000
B (30) 1.000 0.000 0.000
Idh P (5) 0.300 0.700 0.200 0.420
C (3) 0.667 0.333 ? 0.000 0.444
B (28) 0.054 0.769 0.143 0.034 0.429 0.384
Mdh-1 P (40) 0.975 0.025 0.050 0.049
C (13) 0.038 0.808 0.154 ? 0.385 0.322
B (15) 0.033 0.834 0.133 0.200 0.286
Mdh-2 P (40) 1.000 0.000 0.000
C (13) 1000 ? 0.000 0.000
B (19) 1.000 0.000 0.000
Muest P (32) 0.094 0.906 0 156 0.170
C (8) 0.562 0.438 ? 0125 0.492
B (10) 0.650 0.350 0.700 0.455
Pgm P (23) 0.065 0.674 0.261 0.435 0.473
C (9) 0.056 0.tll 0.666 0.167 M 0.556 0.513
B (13) 0.077 0.115 0.808 0.231 0.328
Sod-1 P (11) 1.000 0.000 0.000
C (5) 1.000 - 0.000 0.000
B (10) 1.000 0.000 0.000
Sod-2 P (11 ) 1.000 0.000 0.000
C (5) 1.000 0.000 0.000
B (10) 1.000 0.000 0.000
GENETICS AND SYSTEMATICSOF M/LLEPORA 733

TABLE 2--CONTINUED

Alleles
Growth
Loci form (/V) 1 2 3 4 5 6 Sat Hob H,~

1 2 3 4 5 6 7 8 9 10 11

Tpep-1 P (36) 0.153 0.014 0.819 0.014 0.250 0.305

C (13) 0.923 0.077 ? 0.154 0.142
B (30) 0,733 0.267 0.200 0.391
Tpep-2 P (36)-~ 0.153 0.014 0.819 0.014 -- --
C (13) 0.692 0.231 0,077 ? 0.308 0.462
B (33) 1.000 0.000 0.000
Tpep-3 P (12) 0.958 0.042 0,083 0,080
C (11) 0.045 0.910 0.045 ? 0.182 0.168
B (11) 0.955 0.045 0.091 0.086
Tpep-4 P (37) 0,040 0.947 0.013 0,108 0.101
C (13) 0.731 0.269 ? 0.231 0.393
B (30) 0.233 0.767 0.333 0.357

*N = number of colonies studied;

tS s = Subunit structure; M = monomer; D = dimer; ? = subunit structure was not inferred from electrophoretic pattern in
heterozygotes because of poor resolution of separate allozymea.
Tpep-2 locus is absent in ~P" form. Therefore, allele frequencies of the Tpep-1 locus are presented here (assuming
homology between Tpepl and Tpep-2loci) to involve the Tpep-2locus in calculation of Nei's indices.

TABLE 3. ESTIMATESOF GENETICVARIABILITYIN THREE GROWTH FORMS OF MILLEPORA

Mean heterozygosity (±S.E.)

Growth form Observed Expected Polymorphic loci (%)*

B 0.187+0.043 0.208+0.043 52
C 0.107±0,036 0.157:1:0.045 43
P 0.113±0.039 0.165±0,040 45
B, C (pooled) 0.147±0.028 0.183±0.031 48

Calculations are based on the gene products of 21 presumptive loci in "branch-like" (B) and "comb-like"
(C) forms and 20 loci in "plate-like" (P) form.
*The locus is assumed to be polymorphic if the frequency of more common alleles does not exceed
0.95.

other at Sod-1 and Sod-2 loci. Not fixed but significant differences in allele sets and
frequencies of common alleles are observed at many of the loci compared, but are
more pronounced at the Tpep-1 locus when these two groups of growth forms are
compared. Moreover, branch-like and comb-like growth forms express an additional
TPEP locus (Tpep-2) which is absent in the plate-like form. Because the Tpep-1 locus of
the plate-like form and the Tpep-2 locus of the branch-like and comb-like forms have
one allele in common (Table 2), we assumed that the Tpep-2 locus resulted from
duplication of the Tpep-1 locus. However, an opposite situation with the loss of gene
from a pre-existing two-locus TPEP system is also not excluded. The appearance of a
new gene, or the loss of a gene in a pre-existing multilocus enzyme system may serve
as a synapomorphy in a particular lineage and thus be of considerable systematic
value (Buth, 1984).
Obtained results indicate with certainty that the plate-like growth form is genetically
isolated from the branch-like and comb-like forms. The three growth forms of
Millepora fall into two genetically distinct groups (Table 4). The branch-like and comb-
like forms are genetically very similar (•=0.95), whereas the plate-like form is
considerably different from the branch-like ( / = 0.74) and comb-like ( / = 0.77) forms. In
terms of genetic distance and similarity, two groups (one represented by plate-like and
734 G.E MANCHENKO ETAL.

TABLE 4. NEI'S GENETIC IDENTITY (/) AND GENETIC DISTANCE

(D) INDICES BETWEEN DIFFERENT GROWTH FORMS OF
MILLEPORA

Compared growth
forms* / D

P-C 0.767 0.265

P-B 0.744 0.296
C-B 0.948 0.053

*Abbreviations are the same as in Tables 2 and 3.

another by branch-like and comb-like forms) exhibit considerably greater similarity

than was found for morphologically distinct congeneric species of scleractinian corals
of the family Acroporidae (Manchenko and Latypov, unpublished observations). This
similarity is also greater than that observed for sibling species pairs in nemerteans
(Manchenko and Kulikova, 1988) bryozoans (Thorpe et al., 1978a,b), crustaceans
(Salmon et al., 1979), freshwater molluscs (Chambers, 1978) and Drosophila (Ayala et
al., 1974; Ayala 1975), but approximately the same value as between sibling species of
mosquitoes (Bullini, 1982) and somewhat lower than between hymenopteran sibling
species (Metcalf et al., 1984). So it is clear that the level of genetic divergence of
morphologically close species differs in phylogenetically distinct taxa. This means
that in some cases morphological similarity of species reflects real phylogenetic
proximity and a short period of divergent evolution, while in other cases similarity is
the result of conservation of morphology along with substantial divergence at the
molecular level during a very long period of independent evolution. As to compared
growth forms of Millepora, we believe that we are dealing just with the former case.
The situation as regards interrelationship between branch-like and comb-like growth
forms is not absolutely clear. We assumed that these forms are intraspeciflc variants.
Some differences observed between these forms in allele frequencies and in the
number of alleles at homologous loci may be explained in terms of spatial genetic
differentiation of conspecific populations (or local coral settlements) and the small
sample size examined. As was shown, significant spatial genetic differentiation is
characteristic for some coral species (Stoddart, 1984a, b) and perhaps is caused by
contribution of asexual reproduction in formation of local settlements.
Indirect evidence for conspecificity of branch-like and comb-like forms follows from
additional weakly stained monomorphic bands observed on GOT, MDH and TPEP
zymograms. These bands, perhaps derived from endosymbionts, were not taken into
account during genetic comparisons of different growth forms. It is interesting,
however, that no differences in these bands were observed between branch-like and
comb-like forms, whereas both these forms differ considerably in this respect from the
plate-like form. These differences are even higher than those revealed in enzymes
presumably originating from zooids. This phenomenon may reflect species specificity
of endosymbionts in Millepora and be of great systematic value. Of course, the
possibility that some fixed genetic differences between branch-like and comb-like
forms will be found after examination of a more representative set of isoenzymes is
not excluded. However, the results of this investigation do not give reason to think that
sympatrically occurring branch-like and comb-like growth forms of Millepora are
genetically isolated species, while the same results strongly suggest that the plate-like
growth form is such a species.

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