Marine Benthic Algae of the Russian Coasts of the Bering Sea
(from Ozernoi Gulf to Dezhnev Bay, including Karaginskii Island)
O.N. Selivanova
Kamchatka Branch of Pacific Institute of Geography,
Far Eastern Division, Russian Academy of Sciences,
683000, Petropavlovsk-Kamchatsky, Russia
ABSTRACT
A
list of the marine benthic algae of the Russian coasts of the Bering Sea that
belong to the administrative region Kamchatka Oblast is
given in the paper.
Since the marine
benthic flora of this region is insufficiently studied and the
literature is minimal, the information presented in the article is
based mostly on the analysis of the author's own materials.
This
inventory of the flora of marine algae of the western coasts of
the Bering Sea reveals 170 species of macrophytes, including 33 species
of green, 39 species of brown, and 98 species of red algae.
Twenty species are newly recorded from the area, and four are new to the
flora of the Far Eastern seas of Russia.
A new species of Phycodrys (Delesseriaceae) was discovered.
The information is updated
in accordance with new data on taxonomy and nomenclature of the
species.
INTRODUCTION
The marine benthic flora of the western coasts of
the Bering Sea is poorly investigated.
It has been studied by Russian
phycologists for many years, but these studies were episodic and
uncoordinated. Remoteness and inaccessibility, severe climate and ice
conditions, and a short navigation season make this area very
inconvenient for natural studies.
Practically no seasonal field
observations have been conducted there, no marine biological stations
have ever existed, and scientific expeditions have been infrequent
and sporadic.
Therefore information on the structure and composition
of benthic vegetation is limited.
Special publications on this
subject are rare (e.g.,
Vinogradova 1973a; 1978;
Perestenko 1988;
Zhigadlova and Selivanova 2003 (in press),
although information on
the marine algae of this area may be found in some general taxonomic,
floristic and detailed hydrobiological studies (Kongisser 1933;
Petrov 1972;
Vinogradova 1973b, 1973c; 1974; 1979;
Kussakin and Ivanova 1978;
Vinogradova et al. 1978;
Klochkova and Demeshkina 1985;
Gusarova and Semkin 1986;
Perestenko 1994;
Klochkova and Berezovskaya 1997;
Klochkova 1998).
Short publications (conference
abstracts) by T. Klochkova (1999) and Zhigadlova (2000) concerned
algae of only the Karaginskii Gulf.
Data from some of these
publications are cited in
Table 1.
The
laboratory of Hydrobiology of the Kamchatka Branch of Pacific
Institute of Geography (KBPIG) conducted studies on flora and fauna
of the Russian continental coasts of the Bering Sea (from Ozernoi Gulf to
Dezhnev Bay) during a 7-month-long expedition in 1988. However, processing of
phycological material collected during this expedition was delayed.
The species were difficult to identify
because many of them deviate morphologically from typical forms.
This may be due to geographical factors such as the phenomenon of
gigantism in northern races of plants and animals. For example, some
samples of
Porphyra miniata (C.Agardh) C.Agardh from the bays
north of Olyutorskii Gulf reach 50–55 cm in length.
It
should be noted that taxonomic viewpoints of Russian and western
phycologists differ quite often, and that is the case with the above
mentioned Porphyra.
Lindstrom and Cole (1992a, 1992b) treated
species resembling our specimens as
P. cuneiformis (Setch. & Hus) Krishnamurthy
but their viewpoint is not recognized in
Russian literature.
Phycologists in our country prefer to follow
Perestenko's (1982, 1983, 1994) position, so we identified our
samples in accordance with the concept of the latter author as
P. miniata.
Systematics of the other species within the genus
Porphyra is also controversial.
For instance, Porphyra purpurea (Roth) C.Agardh
was revised by Lindstrom and Cole (1992a, 1992b)
who showed by biochemical and genetic analyses that true
P. purpurea was distributed only in the Atlantic Ocean,
while its
Pacific vicariant was described by these authors as a new species
P. kurogii S.C.Lindstr.
(Lindstrom and Cole 1992b).
We followed this
viewpoint in our publications dealing with the flora of the Commander
Islands and
replaced
P. purpurea in our list of algae with
P. kurogii
(Selivanova and Zhigadlova 1997a, 1997c).
The morphological description of P. kurogii however was
considerably different from that of P. purpurea given by
Perestenko (1982; 1994) who insisted on wide Atlantic-Pacific
distribution of P. purpurea.
There were specimens of Porphyra
in our material from Olyutorskii Gulf that morphologically
represented P. purpurea in Perestenko's interpretation.
There were also specimens that fit into P. kurogii. Additional
taxonomic studies are necessary to resolve the problem. Since we
have no opportunity to carry out biochemical or genetic analyses, we
prefer to keep the morphological approach and to treat P. kurogii
and P. purpurea as separate species. They are both presented
in
Table 1.
According to personal information of Gayle Hansen
both species also occur in Alaska.
The
interpretation of families within the orders has also been disputed.
For instance, Perestenko (1975) described the family Crossocarpaceae
(Cryptonemiales, Rhodophyta) on the basis of vegetative structure and
the morphology of the post-fertilization fusion cell. Independence of
this family was supported by Russian phycologists but rejected by
Hansen and Lindstrom (1984), who considered the Crossocarpaceae a
subgroup within the Kallymeniaceae sensu lato. The circumscription of the
family Dumontiaceae is also treated differently by Russian and
western phycologists: thus Neoabbotiella araneosa described
by
Perestenko (1975)
is still included by this author (Perestenko 1994)
in the Dumontiaceae
in spite of the proposal of
Lindstrom (1985)
to transfer this species from the Dumontiaceae into the
Cryptonemiaceae (=Halymeniaceae).
Another example, the family Arthrothamnaceae
was segregated by
Yu. Petrov (1974)
from the Laminariaceae on the
basis of distinctive features of lamina formation. The
Arthrothamnaceae is widely recognized in Russian taxonomic literature
but overlooked by other phycologists.
In
addition, there are many other unsolved systematic and nomenclature
problems. For example,
Kraft and Robins (1985)
synonymized the
order Cryptonemiales with the Gigartinales within the division
Rhodophyta. This viewpoint was supported by some phycologists,
including leading specialists (e.g.,
Scagel et al. 1986, 1993;
Perestenko 1994). Others (e.g.,
Silva et al. 1996;
Silva 2002)
retained the Cryptonemiales as a separate order.
Silva (2002)
considered the proposal of Kraft and Robins to be
retrogressive because the order Cryptonemiales was sufficiently
heterogeneous and several new orders had already been segregated from
it, including: Hildenbrandiales Pueschel & Cole (1982) and
Corallinales Silva & Johansen (1986).
Later the Halymeniales was
proposed by
Saunders and Kraft (1996)
to replace the remaining
Cryptonemiales on the basis of molecular, histochemical and
ultrastructural features. This proposal was disputed by
Masuda et al. (1999), who chose the older name, based on priority and the
personal recommendation of P. Silva.
These
examples show that even higher taxa (orders and families) are still
controversial. Systematics of lower taxa (genera and species) is also
in many cases unresolved. For example, the number of genera within
the order Acrochaetiales has been disputed. Woelkerling (1971, 1973)
applied the common generic name Audouinella to the species
representing the previously described genera Acrochaetium,
Rhodochorton, Colaconema, Meiodiscus etc. Several subsequent
authors shared this concept (e.g.,
Garbary, Hansen, and Scagel 1982)
but the others considered transfer of these species to Audouinella
unreasonable
(Perestenko 1994;
Silva et al. 1996).
Harper and Saunders (2002)
have recently published a new classification of
acrochaetioid algae which includes the establishment of the new order
Colaconematales. Taxonomy of the group is clearly still in flux.
The
main purpose of our work was to make an inventory of the flora of the Russian
coasts of the Bering Sea using our own material, taking into
consideration current data on taxonomy and nomenclature of the
species. We have also tried to include species from earlier studies
dealing with benthic algae of the Bering Sea. After revision these data
were included in the list of the marine benthic algae of the coasts
of the Bering Sea belonging to Kamchatka Oblast in its former boundaries
(including Koryak Autonomous Disrtict): from Ozernoi to Olyutorskii
Gulf and its northwestern coasts up to Dezhnev Bay.
Data on algae of the
Commander Islands are excluded from the present list as they were
published earlier in a series of papers
(Selivanova and Zhigadlova 1993; 1997a; 1997b; 1997c; 1997d; 1999; 2000; 2002a).
Information on the algae
of the northern part of the Bering Sea located on the territory of
Chukchi Autonomous District (Anadyrskii Gulf, north of Dezhnev Bay to
Bering Strait) is not included in the present paper, because the
author did not have opportunity to work in this area. Data on the
algae of this area may be found in the papers of Vinogradova (1973a,
b, c), Tolstikova (1974), Kussakin and Ivanova (1978); Perestenko
(1988; 1994), but the inventory of marine algae of this region is far
from complete, and additional floristic and taxonomic studies are
necessary.
MATERIALS AND METHODS
The list of algae presented in the paper is based on
phycological material collected by the author during expedition of
the Laboratory of Hydrobiology of KBPIG in the Bering Sea in 1988. The
material was collected from August through October on the littoral
fringe during low tides, with the help of a long hook called “kanza”
from the depths of 1 to 3 m, and with usage of SCUBA technique from
the depths of 1 to 30 m and with a dredge from deeper waters (up to
85 m). Algae cast ashore were also picked up.
Material was sectioned freehand with razor blades, placed in a drop
of fresh water on the slides and examined using the light microscope.
The sections were studied uncoloured or stained with Lugol's
solution or aniline dyes.
The processing of
collections was conducted at Kamchatka Branch of the Pacific
Institute of Geography (Petropavlovsk-Kamchatsky, Russia).
I have examined a total of 808 herbarium sheets. All of them are stored in
the herbarium of KBPIG (Petropavlovsk-Kamchatsky).
RESULTS AND DISCUSSION
The list of benthic
algae of the Ozernoi Gulf to Dezhnev Bay, including Karaginskii
Island
(Fig.1),
totals to 170 species of macroalgae (records from the literature included):
33 species of Chlorophyta, 39 Phaeophyta and 98 Rhodophyta. The
author's own collections constitute 90 of the species.
The material is presented in the form of a table
(Table 1)
for convenience of information retrieval and with the purposes of
shortening printed space. Higher taxa (orders within divisions) are
arranged in the table according to their systematic position, while
families within the orders, genera within the familes and species
within the genera are given in alphabetic order. A dash in the column
“Depths” means the absence of data. Comments to the
species names that may arouse questions are given in the footnotes
below the table.
In quoting the data of the other authors I do not always
share their opinion on taxonomic status of the species and their
nomenclature because quite often the data cited are outdated, not
taking into account changes in the nomenclature and systematics of
algae which have occurred recently. Nevertheless, they are included
in the table with corrections, where it is possible, in conformity
with modern taxonomic data. The original information of the other
authors in the case I do not accept their interpretation of the
species is given in the column ‘Outdated, misapplied or
incorrect species name’
(Table 1).
The
20 species new to the flora are marked by an asterisk (*) in the
table. The species marked with a double asterisk (**) in the table are
recorded from the Far Eastern seas of Russia for the first time. Some
of them (Palmaria mollis (Setch. & N.L.Gardner) Van der Meer et
Bird, Opuntiella californica (Farlow) Kylin,
and Membranoptera setchellii N.L.Gardner) are
recorded in our recent study (Zhigadlova and Selivanova 2002) of
the marine algae of Karaginskii Gulf. However this work is still in
press, so I repeat them here as new to the flora of the Far Eastern
seas. Two of the species found in the study area are proposed for
inclusion in the Red Data Book of Kamchatka: Membranoptera serrata
(Postels & Rupr.) Zinova.
(Fig. 2)
and Pantoneura juergensii
(J.Agardh) Kylin
(Fig. 3).
This book is a regional edition analogous to
the Red Data Book of the Russian Federation, including information on
rare and endangered species of flora and fauna of Kamchatka. I
inscribed the above mentioned species of algae in this list because
of their rarity in our water areas. The Red Data Book of Kamchatka is
planned to be published in the near future.
We
have also discovered several samples of a new species of the genus
Phycodrys (Delesseriaceae, Rhodophyta)
(marked by
the triple asterisk *** in the table). Sterile and tetrasporic
samples representing the new species were found in two bays of the Bering
Sea (Lavrov and Glubokaya Bays). We have named the new species
in honour of the well-known Russian phycologist Valentina
F. Przhemenetskaya (Makienko) who has been studying marine algae of
the Far Eastern seas of Russia for about 30 years. The new species
differs from other species of the genus Phycodrys by having
distromatic lamina and localization of generative proliferations at
the base of the lamina. Its description in Russian with Latin
diagnosis is given in our paper (Selivanova and Zhigadlova 2002b)
that is in press. I include here illustrations and a
description in English:
Phycodrys sp. nov.
(Figs. 4 a-e)
Plants
up to 25 cm long, with marginal prolifications. Laminae
membranaceous, pink-reddish to brownish, up to 8 cm wide and 30–70 µm
thick in cross section, nearly always
distromatic, very rarely monostromatic. Lobes liguliform or
cuneiform. Midribs and veins dark and prominent in older parts of the
lamina and light and flattened in younger parts. Old parts of the
lamina eroded up to the midribs. Glandular cells absent.
Tetrasporangia up to 90 µm in diam. (70 µm without cell
wall), located on prolifications at the basal part of the lamina.
Cystocarps not found.
Type:
Russia, Bering Sea, Glubokaya Bay, 5–7 m depth, 26 VIII 1988,
collected by O.N. Selivanova ( 1872). Kept in Kamchatka Branch
of the Pacific Institute of Geography, Far Eastern Division Russian
Academy of Sciences.
Growing in subtidal zone, on rocks and stones, at the
depths of 5–7 m.
The new species differs from the other species of the genus
Phycodrys by having a mostly distromatic lamina and localized
generative proliferations at the base of the lamina.
Specimens examined:
1872 (type): Glubokaya Bay, 5–7 m depth, rocks,
tetrasporic, 26 VIII, 1988, coll. A.G. Bazhin; 1860,
1868, 1869; 8470; 1871-1873, Glubokaya Bay, 5–7 m
depth, rocks and stones, tetrasporic, 26 VIII 1988, coll. A.G.
Bazhin; 1861 Olyutorskii Gulf, Lavrov Bay, 6–7 m depth,
rocks, sterile, 22 VIII 1988, coll. O.N. Selivanova.
According
to Vinogradova and Perestenko (1978) the marine benthic flora of the
western part of the Bering Sea differs considerably from that of the
southeastern Kamchatka and Commander Islands. Comparison of the
species composition of the macrophytes from these areas in our
collection confirmed this statement. There are principal differences
between floristic complexes of the areas possibly due to the system
of marine currents. Cold arctic currents in the Bering Sea cause invasion
of elements of flora from the Arctic Ocean, in its turn, the flora of
the Commander Islands is enriched by the species from the American
coasts. We also noted an interesting phenomenon: there were no
samples of Yendonia crassifolia (Rupr.) Kylin in our
collections from the Commander Islands, while we have many of them
from the Bering Sea, including Karaginskii Island; on the contrary, its
close relative Mikamiella ruprechtiana (A. Zin.) Wynne was
abundant in the Commander Islands and absent in our collections from
the Bering Sea. It should be noted that these two species are recorded by
other authors from both water areas. In addition, Pantoneura
juergensii is relatively abundant on Karaginskii Island whereas
it is very rare in other areas and has been proposed for
inclusion as a rare species in the Red Data Book of Kamchatka.
However comparison of floristic complexes of the three water areas of
the Russian Pacific (eastern Kamchatka, the Bering Sea, and Commander
Islands) is not the task of the present publication: we plan to
prepare a separate paper concerning this problem.
ACKNOWLEDGEMENTS
I am thankful to all colleagues from Hydrobiology Lab
(KB PIG) who participated in collecting of algae, especially to A.G.
Bazhin and D.D. Danilin. I would also like to thank my colleague
Galina G. Zhigadlova for her participation in the identification of
algal species.
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