Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea
According to the rule of de Candolle-Walles a species diversity decreases from low latitudes regions to high latitudes regions. However in the Weddell Sea the biodiversity of the fauna of some groups are approximately the same (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) or more...
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Національний антарктичний науковий центр МОН України
2012
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| Cite this: | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea / B.I. Sirenko, I.S. Smirnov // Український антарктичний журнал. — 2011-2012. — № 10-11. — С. 186-200. — Бібліогр.: 71 назв. — англ. |
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| author | Sirenko, B.I. Smirnov, I.S. |
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| citation_txt | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea / B.I. Sirenko, I.S. Smirnov // Український антарктичний журнал. — 2011-2012. — № 10-11. — С. 186-200. — Бібліогр.: 71 назв. — англ. |
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| description | According to the rule of de Candolle-Walles a species diversity decreases from low latitudes regions to high latitudes regions. However in the Weddell Sea the biodiversity of the fauna of some groups are approximately the same (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) or more than this in the Magellan region (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea, Ascidiacea). In spite of the rather high species diversity of bivalves in the Weddell Sea, communities where bivalves dominated (like in Arctic seas) practically are absent and average size of gastropods and bivalves is much less than in the Arctic Ocean. Moreover Reptantia from Decapoda crustacean is absent in the Weddell Sea. We will try to explain the reasons of the breach of the Walles rule of the fauna and will try to explain some other peculiarities of the Weddell Sea. For this purpose we will compare the fauna of some groups in the Magellan region and in the Weddell Sea (Table. Number of species of different genera the Magellan Region and in the Weddell Sea).
Согласно правилу меридионального изменения разнообразия Декандоля–Уоллеса, разнообразие видов уменьшается от низких широт к высоким. Сравнение фауны моря Уэдделла с фауной соседнего с ним Магелланова района обнаруживает нарушение этого правила для некоторых групп морских донных животных. У актиний, мизид, кумовых раков, брюхоногих моллюсков, мшанок и морских ежей (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) видовое разнообразие в море Уэдделла почти не уступает таковому в Магеллановом р-не, а у губок, морских пауков, голотурий, морских звезд, офиур и асцидий (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea, Ascidiacea) оно даже выше, вопреки правилу Уоллеса. Несмотря на довольно высокое разнообразие видов двустворчатых моллюсков в море Уэдделла, сообщества, где двустворки доминируют (как в арктических морях), фактически отсутствуют, а средний размер гастропод и двустворчатых моллюсков – намного меньше, чем в Северном Ледовитом океане. Кроме того, ракообразные из группы Reptantia (Decapoda) в море Уэдделла отсутствуют. Мы попытаемся объяснить причины нарушения правила Декандоля–Уоллеса и некоторые другие особенности фауны моря Уэдделла. С этой целью мы сравним фауну некоторых групп беспозвоночных Магелланова района и моря Уэдделла.
Відповідно до правила меридіональної зміни різноманіття Декандоля–Уоллеса, різноманіття видів зменшується від низьких широт до високих. Порівняння фауни моря Уедделла з фауною сусіднього з ним Магелланового району виявляє порушення цього правила для деяких груп морських донних тварин. У актиній, мізид, кумових раків, черевоногих молюсків, мшанок та морських їжаків (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) видове різноманіття в морі Уедделла майже не поступається такому в Магеллановому районі, а у губок, морських павуків, голотурій, морських зірок, офіур і асцидій (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea, Ascidiacea) воно навіть вище, всупереч правилу Уоллеса. Попри досить велике різноманіття видів двостворчастих молюсків у морі Уедделла, угруповання, де двостворки домінують (як в арктичних морях), фактично відсутні, а середній розмір гастропод і двостворчастих молюсків набагато менший, аніж у Північному Льодовитому океані. Окрім того, ракоподібні з групи Reptantia (Decapoda) в морі Уедделла відсутні. Ми спробуємо пояснити причини порушення правила Декандоля–Уоллеса та деякі інші особливості фауни моря Уедделла. З цією метою порівняємо фауну деяких груп безхребетних Магелланового району й моря Уедделла.
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B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
186
UDK 591.9+99
SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF
THE FAUNA IN THE WEDDELL SEA
B.I. Sirenko, I.S. Smirnov
Zoological Institute, St. Petersburg, RU, marine@zin.ru
Abstract. According to the rule of de Candolle-Walles a species diversity decreases from low latitudes
regions to high latitudes regions. However in the Weddell Sea the biodiversity of the fauna of some groups
are approximately the same (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) or more
than this in the Magellan region (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea,
Ascidiacea). In spite of the rather high species diversity of bivalves in the Weddell Sea, communities where
bivalves dominated (like in Arctic seas) practically are absent and average size of gastropods and bivalves is
much less than in the Arctic Ocean. Moreover Reptantia from Decapoda crustacean is absent in the Weddell
Sea. We will try to explain the reasons of the breach of the Walles rule of the fauna and will try to explain
some other peculiarities of the Weddell Sea. For this purpose we will compare the fauna of some groups in
the Magellan region and in the Weddell Sea (Table. Number of species of different genera the Magellan
Region and in the Weddell Sea).
Key words: Antarctic fauna, Walles’ rule, Weddell Sea, Magellan region.
Нарушение правила Уоллеса и другие особенности фауны моря Уэдделла. Б.И. Сиренко,
И.С. Смирнов.
Реферат. Согласно правилу меридионального изменения разнообразия Декандоля–Уоллеса,
разнообразие видов уменьшается от низких широт к высоким. Сравнение фауны моря Уэдделла с
фауной соседнего с ним Магелланова района обнаруживает нарушение этого правила для некоторых
групп морских донных животных. У актиний, мизид, кумовых раков, брюхоногих моллюсков, мшанок
и морских ежей (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) видовое
разнообразие в море Уэдделла почти не уступает таковому в Магеллановом р-не, а у губок, морских
пауков, голотурий, морских звезд, офиур и асцидий (Spongia, Pycnogonida, Holothuroidea, Asteroidea,
Ohpiuroidea, Ascidiacea) оно даже выше, вопреки правилу Уоллеса. Несмотря на довольно высокое
разнообразие видов двустворчатых моллюсков в море Уэдделла, сообщества, где двустворки
доминируют (как в арктических морях), фактически отсутствуют, а средний размер гастропод и
двустворчатых моллюсков – намного меньше, чем в Северном Ледовитом океане. Кроме того,
ракообразные из группы Reptantia (Decapoda) в море Уэдделла отсутствуют.
Мы попытаемся объяснить причины нарушения правила Декандоля–Уоллеса и некоторые другие
особенности фауны моря Уэдделла. С этой целью мы сравним фауну некоторых групп
беспозвоночных Магелланова района и моря Уэдделла.
Ключевые слова: антарктическая фауна, правило Уоллеса, море Уэдделла, Магелланов район.
Порушення правила Уоллеса та інші особливості фауни моря Уедделла. Б.І. Сіренко, І.С. Смирнов.
Реферат. Відповідно до правила меридіональної зміни різноманіття Декандоля–Уоллеса, різноманіття
видів зменшується від низьких широт до високих. Порівняння фауни моря Уедделла з фауною
сусіднього з ним Магелланового району виявляє порушення цього правила для деяких груп морських
донних тварин. У актиній, мізид, кумових раків, черевоногих молюсків, мшанок та морських їжаків
(Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) видове різноманіття в морі Уедделла
майже не поступається такому в Магеллановому районі, а у губок, морських павуків, голотурій,
морських зірок, офіур і асцидій (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea,
Ascidiacea) воно навіть вище, всупереч правилу Уоллеса. Попри досить велике різноманіття видів
УКРАЇНСЬКИЙ АНТАРКТИЧНИЙ
ЖУРНАЛ
УАЖ, № 10-11, 186-200 (2011/2012)
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
187
двостворчастих молюсків у морі Уедделла, угруповання, де двостворки домінують (як в арктичних
морях), фактично відсутні, а середній розмір гастропод і двостворчастих молюсків набагато менший,
аніж у Північному Льодовитому океані. Окрім того, ракоподібні з групи Reptantia (Decapoda) в морі
Уедделла відсутні.
Ми спробуємо пояснити причини порушення правила Декандоля–Уоллеса та деякі інші особливості
фауни моря Уедделла. З цією метою порівняємо фауну деяких груп безхребетних Магелланового
району й моря Уедделла.
Ключові слова: антарктична фауна, правило Уоллеса, море Уедделла, Магелланів район.
1. Introduction
According to the rule of de Candolle-Walles a species diversity decreases from low latitudes
regions to high latitudes regions. However in the Weddell Sea the biodiversity of the fauna of
some groups are approximately the same (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa,
Echinoidea) or more than this in the Magellan region (Spongia, Pycnogonida, Holothuroidea,
Asteroidea, Ohpiuroidea, Ascidiacea). In spite of the rather high species diversity of bivalves in
the Weddell Sea, communities where bivalves dominated (like in Arctic seas) practically are
absent and average size of gastropods and bivalves is much less than in the Arctic Ocean.
Moreover Reptantia from Decapoda crustacean is absent in the Weddell Sea.
We will try to explain the reasons of the breach of the Walles rule of the fauna and will try to
explain some other peculiarities of the Weddell Sea. For this purpose we will compare the fauna of
some groups in the Magellan region and in the Weddell Sea.
2. Results
We will compare some general abiotic and biotic peculiarities in the Weddell Sea and the
Magellan region as well.
2.1. Temperature, salinity, ice conditions and light regime
Weddell Sea
Water temperatures of the shelf are normally low and stable (-1.8±0.2°C) with the exemption
of irregularly occurring intrusion of Warm Deep Water into the shelf which increase the
temperature (Bathmann et al., 1991; Gerdes and Montiel, 1999). The salinity shows little variation
with a normal range between 34.6 to 34.9‰S (Gerdes and Montiel, 1999). There are ice coverage
and clearly defined seasonality in light regime.
Magellan region
Temperatures in the Magellan Straits are comparably high and vary seasonally between 6.5 to
9.0°C (Artegiani and Pachini, 1991). The salinity is usually lower than in the Weddell Sea and
vary from 34.0 to 31.0‰ and less in the vicinity of the glaciers (Antezana, 1999). Ice coverage is
absent and seasonality in light regime is less defined than in the Weddell Sea.
2.2. Currents
Weddell Sea
The Weddell Sea is included in circulation system of the Weddell Gyre (Fig. 1). Northern
boundary of the Weddell Gyre is the northern boundary of the Weddell Sea current (Fig. 2). On the
east part of the Gyre approximately in 20-400E from south edge of the Antarctic Circumpolar
Current relatively warm waters go to south and then westward along the Antarctic coast coming
back to the Weddell Sea.
Magellan region
The waters of West Wind Drift from Pacific Ocean penetrate in the Magellan Strait and are
transformed there.
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
188
Fig. 1. Circulation of surface waters of the Fig. 2. Antarctic and Subantarctic Convergence
Southern Ocean (Ledenev, 1969). аnd Northern boundary the Weddell Sea
current (Deacon, 1977).
2.3. Bottom sediments
Weddell Sea
The usual sediments of the Weddell Sea shelf are muddy sand with gravel and boulders of
various sizes (Gerdes et al., 1992). The main specific character of bottom in Antarctic shelf is very
poor sorting deposits (Pasternak, Gusev, 1960). There are a lot of various sizes rough stones on the
surface of the bottom. They were transported by icebergs from Antarctic coast (Ushakov, 1962).
Magellan region
Bottom sediments in the Magellan region are very various. There are rocks, stones, gravels,
sand, mud, clay and different their combination in the Magellan region.
2.4. River outflow
Weddell Sea
River outflow in the Weddell Sea is absent. There is transport of terrestrial mineral particles
with icebergs.
Magellan region
River outflow is moderate.
2.5. Diversity of facies
Weddell Sea
Monotony of sediments in the Weddell Sea shelf, domination of flat landscapes which are
interrupted by rare depressions and hills and absence of a shoals and islands are responsible for
low diversity of facies.
Magellan region
Contrary of the Weddell Sea, Magellan region possesses high diversity of sediments, different
depths from littoral to bathyal zones, and a lot of small and large islands and straits. Taken
together these characters create big diversity of facies.
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
189
2.6. Nutrients and primary production
Weddell Sea
Concentrations of Phosphate and Nitrogen compounds are very high in an Antarctic waters
and even in summer period during bloom of phytoplankton they are more than in temperate waters
of North Hemisphere during winter maximum (Bogojavlensky, 1958). The reason of richness of
nutrients in Antarctic are in constant renew of their stocks owing to circulations of deepwaters
where regeneration of these salts are occured. Concentration of silicic acid in Antarctic water also
high and the highest in the region of the Weddell Sea and Weddell circulation. Gorshkov et all.
(1993) note that the interesting peculiarity of distribution of silicic acid in deep water is increase
its concentration along stream from the Weddell Sea to Dreak strait (Fig. 3). Authors suppose that
the reason of this increase is in accumulation of silicic acid as silicic remains of organism
skeletons from deep and nearbottom dissolve.
Fig. 3. Distribution of solved silicic acid in surface waters (microgramms-atom/litter)
(Gorshkov et al., 1993).
In whole South Ocean including Subantarctic regions the concentrations nutrients do not limit
the development of phytoplankton. The main reason which limits quantitative development of
phytoplankton is density of water structure that is the intensive vegetation of phytoplankton begins
after a development of summer pycnocline (Voronina, 1984). On the assessment of different
scientists primary production of the Antarctic waters vary from 36 to 182 g C/m2, often about 46 g
C/m2 (Koblents-Mishke, 1977). However Voronina (1984) considers that this figures of the
production are lower than real ones. According to Knox and Lowry (1977) the Antarctic waters are
about 400% more productive than the rest of the oceans.
Magellan region
In subantarctic waters nutrients concentrations are much less then in Antarctic waters. There
are seasonal alterations of concentrations of these salt and their reduction towards north, like in
Antarctic waters (Fedorov, 1970)
About phytoplankton vide supra.
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
190
2.7. Zooplankton
Weddell Sea
The most abundant species of zooplankton in Antarctic waters are three species of copepods
(Rincalanus gigas, Calanoides acutus, Calanus propinquus) and one species of krill (Euphausia
superba). The average biomasses of zooplankton in Antarctic waters are 72-75 mg/m3 (Voronina,
1984). The biomass of krill in rich Antarctic regions is in the average 30 g/m2 (Marr, 1962).
Magellan region
The most abundant species of zooplankton in Subantarctic region are three species of
copepods (Rincalanus gigas, Calanus simillimus and Calanus tonsus). Share of macroplankton is
small (3% from whole zooplankton)(Voronina, 1984). The average annual biomasses of
zooplankton of Antarctic and Subantarctic region are measured in the same order (Foxton, 1956).
2.8. Composition of benthic dominant groups and structure of benthic settlements
Weddell Sea
There is clear domination of sessile groups and some slow moving benthic organisms
(Porifera, Bryozoa, Ascidiacea, Pterobranchia, Holothuroidea and Crinoidea) in the most regions
of the Weddel Sea shelf (Sirenko et al., 2001). In the same sea sponges and holothurians were in
dominant group even in the samples taken with multibox corer (Gerdes et al., 1992) which usually
did not show real dominants owing to its too small boxes. The same main group of sessile fauna
(Porifera, Bryozoa, Ascidiacea) were marked for the depths 100-500 m in the eastern part of the
Weddell Gyre and eastward up to 900E (Pasternak, Gusev, 1960).
The suspension feeding communities consisted of epibenthic animals are the most common in
the shelf of the Weddell Sea. They are replaced by infauna communities in the not numerous shelf
depressions.
It is interesting to consider a structure of epibenthic settlements. There are a lot of high
animals in the epibenthic communities. Large sponges and ascidians, small sponges and ascidians
arranged on high shaft, high gorgonarians and hydroids form highest level of the settlement.
Bryozoans, tube worms, sea anemones, pterobranchs, corals and other small sessile groups of
invertebrates including small individuals of sponges, ascidians, gorgonarians and hydroids form
one or more lower levels of settlements. Small sessile animals (bryozoans, sponges, hydroids,
byssus bivalves etc.) attach to high animals as high as possible. Different representatives of vagile
fauna (ophiuroids, holothurians, crinoids, crustaceans etc.) go up on sessile organisms. Most of
animals try to occupy place as high as possible in order to be closer to seston – their main food.
Therefore they form settlement with several levels. Each level of such settlement has own special
habitat and own feeding conditions. As a result of this multilevel settlement a number of niches
increase.
Biomass of benthos in Antarctic suspension feeding communities at the depth 100-500 m in
Indian Ocean Sector (Ocean grab samples, square 0,25 m2) is 319.46-1244.82 g/m2 (Pasternak,
Gusev, 1960) and 450-500 g/m2 (Belyaev, 1964), off Sabrina Coast at the depth 200-300 m (Ocean
grab samples, square 0.25 m2) is 183-1363 g/m2 (Ushakov, 1962), in the Weddell Sea at the depth
170-2037 m (Multibox corer samples, 9x0.0225 m2) is from 0.12 to 1673.0 g/m2 (Gerdes at al.,
1992). Thus the average biomass of benthos on the Antarctic shelf is more than 400-500 g/m2 that
is rather high.
Magellan region
There are many different communities in the Magellan region owing to high diversity of
faces. Main dominants in the Magellan region are bivalves, polychaetes, echinoids, holothurians
and crustaceans (Gerdes, Montiel, 1999). The average biomass in the Magellan Straits at the depth
8-459 m is 96.8 g/m2 in the Beagle Channel at the depth 38-348 m is 301.6 g/m2 (Gerdes, Montiel,
1999).
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
191
2.9. Pelagic – benthic coupling
Weddell Sea
The main source of food in the Weddell Sea is phytoplankton of the open waters and diatoms
of criopelagic communities. Zooplankton feed on algae and than both including dead and alive and
their faces are the food for zoobenthos.
The Weddell Gyre seemingly influences on productivity of this region and can explain the
abundance of both plankton and benthos. The Gyre is large-scale cyclonic circulating system
disposed southwards from Polar Frontal Zone, from Antarctic Peninsula to 20-400E (Klepikov,
1963; Deacon, 1979; Danilov and Guretsky, 1993). Cold waters from the south-western Weddell
Sea extend first to north then to east. The cold waters which go out from the Weddell Sea near
north part of Antarctic Peninsula are rich with nutrients and krill. According to Marr’s counts
(Marr, 1962) the most abundant krill concentrations are in the Atlantic Sector of South Ocean
exactly in the Antarctic Circumpolar Current and the Weddell Gyre Boundary (Fig. 4).
Approximately in 20-400E from south edge of the Antarctic Circumpolar Current relatively warm
waters with abundant phyto- and zooplankton (including krill) go to south and then mainly
westward along the Antarctic coast coming back in to the Weddell Sea. There the waters rush over
the bottom communities dominated by sessile suspension feeders and feed them on abundant dead
and alive phyto- and zooplankton and faeces.
Fig. 4. Distribution of krill in the Southern Ocean (Marr, 1962).
Magellan region
There are several sources of food in the Magellan region. Main from them are phytoplankton
and benthic algae. As it was mentioned krill are not abundant in the Subantarctic waters (3% from
whole zooplankton) in contrast to the abundance of krill in Antarctic waters. There is a belt of
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
192
kelps including huge Macrocystis pyrifera in the shallow waters of the Magellan region. Both
benthic algae and the their detritus serve as a main or additional food for different bottom animals.
2.10. Species diversity
The comparison of species diversity in the Weddell Sea and Magellan region (Table 1) shows
that first group of invertebrates in the Weddell Sea have less diversity than in the Magellan
Region. They are Hydrozoa, Polychaeta, Sipuncula, Gammaridea, Isopoda, Polyplacophora,
Bivalvia, Scaphopoda,. Second group of invertebrates have approximately the same species
diversity (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea). Third group of
animals have more rich fauna in the Weddell Sea than in the Magellan Region (Porifera,
Pycnogonida, Holothuroidea, Asteroidea, Ophiuroidea and Ascidiacea). First group of
invertebrates follows the Walles’ rule, but second and especially third groups of animals do not
follow this rule.
3. Discussion
Comparison of abiotic and biotic peculiarities in the Weddell Sea and the Magellan Region
allows us to select several general ones. Among them: temperature, currents, ice coverage, light
regime, bottom sediments, diversity of facies, composition of benthic dominant groups and
structure of their settlement.
On the one hand warmer temperature, absence of ice coverage, longer light time, diversity of
bottom sediments and facies in the Magellan Region favor to development of more rich fauna in
this region as compare with the Weddell Sea. On the other hand a composition of benthic
dominant groups in the Weddell Sea and a structure of their settlements and as well a peculiar
disposition of the Weddell Sea in the system of currents allow the fauna of this sea support the
high species diversity at least in several dominant groups of invertebrates and high productivity of
ecosystem of Weddell Gyre.
Indeed the Weddell Sea is included in the Weddell Gyre, which is giant circulating system
and occupies about 25% of the Southern Ocean. Northern boundary of the Weddell Gyre dispose
to south from the Antarctic Circumpolar Current where the most abundant krill concentrations are
reported (Marr, 1962). Alive and dead phyto- and zooplankton (including krill) and its faeces
seemingly came back in the Weddell Sea as a result of westward moving of waters along Antarctic
coast. The waters of the Weddell Gyre pass whole way of Gyre during about 3 year. According to
Voronina (1984) to the end of first year larvae of krill finish their development, during second year
they feed on and grow up and during third year they mature and spawn. The life span of Euphausia
superba is 3-4 years (Siegel, 1987; Spiridonov, 1987) but the experimental investigations show a
possibility of the longevity of krill up to 8-9 years (Ikeda, 1984). Be it as it way the longevity of
krill is sufficiently for they to pass whole way in the Weddell Gyre one time at least. During this
way krill (alive, dead and its facies) along with other components of plankton serve as a main food
for sessile suspension feeders. Sokolova (1993) reported that krill was main food for abyssal brittle
stars in the Weddell and Scotia seas. Moreover krill was in 100% studied specimens and it was
main component of food (94.2% in the Weddell Sea and 89.5% in the Scotia Sea). Dearborn
(1977) registers several species of stars and brittle stars that feed on euphausiids and supports the
idea that kril is of considerable direct importance to benthic invertebrates. According to data of
Tseitlin and Voronina (1996): “The average flux of Corg from the layer 0-100 m in the areas rich in
krill is nearly the same as in the rest part of the Antarctic, but in the places of extremely high
concentrations of krill it may be several times more than the average value. In such regions the
flux of dead krill is much more intensive than that of faeces, and thus, it serves as the main source
of food for the bottom fauna”. Makarov and Spiridonov (1993) consider that the stable and large-
scale caring out of krill from the Weddell Sea and from Antarctic Peninsula region is a cause of
high biomass of E. superba in the Scotia Sea. Perhaps each large gyre (Weddell, Bellinsgausen and
Ross) have own subpopulation of the krill (Latogursky, 1979; Makarov, Spiridonov, 1993). In
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
193
Eastern part of the Weddell Gyre apart from the main drift of krill back to the Weddell Sea part of
krill brings out from the Gyre eastward with the surface waters (Makarov, Spiridonov, 1983).
High biomass of krill was noted in many regions of the Weddell Gyre especially in northern
part that is free from ice in summer period. High biomass of krill was found under ice in the
Weddell Sea as well (Elbrachter et al., 1987). Taking into consideration the domination of krill
(49-77% of whole biomass of zooplankton) in the Antarctic waters that are riches with krill
(Voronina, 1984) there appears to be krill play one of the main roles in feeding of benthos of the
Weddell Sea. The rich settlements of sessile benthos like a giant sieve that filter water mass and
feed on alive and dead phyto- and zooplankton (including krill) and their faices. Average biomass
of benthos in the Weddell Sea (400-500 g/m2) is characterized as one of the highest. Bottom
communities accumulate huge quantaty of organic and nutrients in their biomass. Cold waters
from north-western part of the Weddell Sea bring a lot of nutrients that are results of destruction
organics of the Weddell Sea communities. Moreover in the Weddell Sea only there is the dome-
shaped raising of bottom Antarctic water that is a result of cyclonic circulation (Klepikov, 1963).
The raising of bottom water riches of surface water with nutrients as well. In spring time when
light regime became better in the north part of the Weddell Gyre the bloom of phytoplankton
starts. Owing to the circulating system which is the Weddell Gyre the most part of energy that
formes in its north part are kept into the Gyre accumulating in alive biomass of zoobenthos in
south part of the Gyre. From the preceding the quantitative riches of the Antarctic benthic
communities become understandable. The bottom communities that consume a live and dead
phyto- and zooplankton play a part of accumulator of energy and temporal depository of nutrients.
Table 1
Species diversity of several invertebrates groups of from Antarctic waters
the Weddell Sea and Magellan region
Taxon
Antarctic
waters
Weddell
Sea
Magellan
region
Source
1 2 3 4 5
Porifera 3501 1591 442* 1 Bartel et al., 1997
2 Pansini, Sara, 1999
Hydrozoa 1041 412 1261 1 Canterro, Carascosa, 1999
2 Stepanjants, Sloboda, 2000 with addition
Actiniaria - 16 14 Grebelnyi. 2000 with addition
Polychaeta 650+1 602 2231 1 Knox, Lowry, 1977
2 Gamby, in expedition, 1996
Sipuncula - 9 16 Saiz-Salinas, Pagola-Carte, 1999
Pycnogonida 200 85 46 Turpaeva, in letter, 2002
Decapoda
(Reptantia)
- 0 30 Arntz et al.,1999
Decapoda
(Natantia)
- 5 11 Arntz, Gorny, 1996
Mysidacea 372 323 311 1 Brandt et al., 1998
2 Brandt, 1999
3 Wittmann, 1991
Gammaridea 4701 1162 2061 1 De Broyer, Rauschert, 1999
2 Rauschert, De Broyer, 2000
Cumacea 52 29 31 Mühlenhardt-Siegel, 1999
Isopoda 3481 682 1012 1 Brandt et al., 1998
2 Brandt, 1999
Polyplacophora 13 4 15 Sirenko, Smirnov (in press)
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194
1 2 3 4 5
Gastropoda
(Prosobranchia)
- 2212 2101 1 Linse, 2002,2 Linse et al., 2006
Bivalvia - 802 1311 1 Linse, 2001,2 Linse et al., 2006
Bryozoa 3101 1842 2053 1 Bullivant, 1969
2 Gontar, 2000 with addition
3 Moyano, 1999
Holothuroidea - 341 272 1 Gutt, 1988
2 Deichmann, 1947
Echinoidea - 42 71 1 Larrain et al., 1999
2 Sirenko, Smirnov (in press)
Asteroidea - 502 211 1 Larrain al., 1999
2 Sirenko, Smirnov (in press)
Ophiuroidea 56 26 Smirnov in letter, 2002
Ascidiacea 1291 512 351 1 Kott. 1969, 1971
2 Romanov, 2000 with addition
* - only Demospongiae which compose about 80% of all species of sponges of region.
High diversity of faces in the Magellan Region which depends on the diversity of sediments,
different depths and presence of small and large islands and straits gives many niches. The
diversity niches in the Magellan Region promote high biodiversity. There is a low diversity of
faces in the Weddell Sea but a lot of niches which are reveal by high diversity of many groups of
invertebrates and primarily dominated sessile invertebrates (Spongia, Bryozoa, Ascidiacea, and
Actiniaria) and some movable forms of animals (Holothuroidea, Asteroidea, Ophiuroidea,
Picnogonida and others). The explanations of the increase of number of niches in the Weddell Sea
are in composition of bottom communities and in structure their settlements.
Many hard particles (gravels and boulders of various sizes) in sediments of the Weddell Sea
is favorable for the development of climax bottom communities of sessile suspension feeders
where Spogia, branchy Bryozoa, Ascidiacea and others dominate. Most of dominant species trying
to grow upwards. Large sponges, high coelenterates, ascidians and sponges arranged on high shaft.
All of them strive upwards closer to food-seston that is drifted with water. A lot of movable
species (Holothuroidea, Ophiuroidea, Crinoidea and Bivalvia) try to climb on to top of large
sponges, gorgonarians, and ascidians closer to food-seston. They (high sessile and some movable
organisms) compose the highest level of bottom settlements. Smaller sponges, gorgonarians,
branchy bryozoan, tube worms and others form a lower level of settlements, and so up to soft
sediments where lowest level consisted of infauna animals are situated.
As a result the multilevel settlements are formed. Each level has different conditions for life
and feeding. The animals of each lower level of settlements have the rest of seston and fecal
masses of animals of higher levels. As far as seston sinks not vertical but at an acute angle owing
to a current the effect of multilevel settlement keeps in rarefied settlements as well. Seemingly
multilevel settlements of bottom communities compensate low diversity of faces in the Weddell
Sea and increase the number of ecological niches. The increase of niche numbers promotes
increase of biodiversity.
During temperature fall in Antarctic the increase of species number was going on in dominant
groups of sessile benthos: Spongia, Bryozoa, Ascidiacea and in other groups which were ready to
low temperature.
Seeming in many cases the increase of the number of species in the Weddell Sea was owing
to sympatric speciation. This is confirmed by a great number of species in the Weddell Sea
belonged to one genus (Table 2). The most number of endemic species (endemism more 70%,
Brandt 1991, Hayward, 1995) characteristic for groups-edificatiors (Spongia, Bryozoa,
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195
Ascidiacea) and movable groups (Pycnogonida, Cummacea, Amphipoda, Tanaidacea, Isopoda,
Echinoidea, Holothuroidea, Ophiuroidea, Asteroidea and Pisces). Probably new species were
formed as a result of food specialization and spatial divergence on different level of bottom
settlements.
Abundant development of sponges, bryozoans, ascidians and other sessil organisms was a
reason of appearance rich fauna of predators that feed on these sessil animals and a lot of numbers
of symbionts that live with them. Abundance of sea stars, brittle stars, sea spiders and other
movable animals in trawls is easily to explain by abundance and diversity of their preys.
Pantopods feed on sponges, bryozoans, hydroids, sea anemons and other animals and even detritus
(Arnaud and Bamber, 1987). Brittle stars and sea stars feed on different crustaceans (including
krill and copepods), sponges, polychaetes, bivalves and other animals and detritus (Deaborn, 1977;
Sokolova, 1993). Some mollusks (chitons and nudibranchs) feed on bryozoans and sponges
(Sirenko, 1997).
Symbiosis of antarctic animals is in the beginning of its study. We know only some Antarctic
symbiotic pairs: gastropod Harpovoluta charkoti+ sea anemon Isosicyonis alba (Arnaud, 1978);
gastropod Dickdelia labioflecta+ sea spider Nymphon isabellae (Sirenko, 2000); gastropod
Capulus subcompressus+ polychaete Serpula narconensis (Schiaparelli et al., 2000; Sirenko and
Schrödl, 2001); brittle stars Ophiurolepis brevirima+sponge Iophon radiatus (Smirnov and Koltun,
1996); brittle stars Ophiurolepis spp. and Theodoria relegata+hydroid Hydractinia vallini
(Smirnov and Stepanjants, 1980) and others. Apparently a symbiosis is more wide spread among
the Antarctic animals than we know that can testify about a long way of development of Antarctic
fauna. Some animals are necrophaguses (some gastropods, brittle stars, sea stars, pantopods, sea
anemones and others).
Table 2
Number of species of different genera the Magellan Region and in the Weddell Sea
Group, source Genus Magellan
region
Common
species
for two regions
Weddell Sea
Sipuncula
Saiz-Salinas, and
Pagola-Carte, 1999
Golfingia 2 2 3
Nephasoma 4 3 5
Cumacea
Mühlenhardt-Siegel,
1999
Hemilamprops 2 1 2
Leucon 6 1 5
Campylaspis 7 5 7
Diastylis 5 0 6
Echinoidea
Mironov, in letter, 2003
Amphipneustes 0 0 4
Antrechinus 0 0 2
Ctenocidaris 0 0 5
Notocidaris 0 0 2
Sterechinus 1 0 2
Ophiuroidea
Smirnov, in letter, 2003
Astochlamys 0 0 2
Amphioplus 2 0 2
Amphiura 4 3 10
Ophiocten 1 0 4
Ophiosteira 1 1 4
Ophiura 1 0 7
Ophiurolepis 1 1 5
Ophiacantha 3 1 4
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196
Seemingly the trophical factor is one of the main one that can explain why Antarctic bivalves
have so small size and are not abundant like in Arctic shelf. They were not able to compete with
large sessil suspender feeding animals in feeding. Moreover gastropods many of which feed on
mainly bivalves have small size of body as well.
The absence of whole group of brachyuran crabs in Antarctic has another reason. The
distribution of brachyuran crabs is limited by low temperature that influence on the metabolism
crabs owing to Magnesium regulation (Frederich et al., 2001). In spite of the fact of presence of
crabs and helmit crabs in Arctic Ocean (Vassilenko, 2001; Petrjashov, 2001) all of them inhabit
waters with positive temperatures connected with warm Atlantic or Pacific currents. The most
western find of the crab Chionocoetes opilio was in the Laptev Sea (RV "Polarstern" ARK IX/4,
st. 40, depth 231-233 m) where the bottom temperature was +1.44°C (Rachor et al., 1994). From
two orders of Decapoda (Natantia and Reptantia) the only first one had been luck. Shrimps in
contrast to crabs lithodids and anomurans gained an advantage from the Nature and could live in
cold waters with the negative temperature. Species of suborder Natantia must have a high
physiological plasticity and thus the adaptive potential to colonize different environments of the
Arctic and Antarctic Oceans.
It is necessary to note that biodiversity of the Magellan Region can be less because of late
glaciations when south part of South America was covered by ice during several thousand years up
to 12000 years ago (Fig.5). Moreover Magellan Strait was formed about 2000 years ago only
(Moore, 1975). During the late glaciations marine fauna of Magellan Region moved northward
where it was able to survive.
Fig. 5. Reconstruction KLIMAP of earth surface in August during the maximal stage of latest
glaciations 18000 years ago. High of surface of glaciers in meters, isotherms in C°. Contours of
continents is coincide with the present-day isobate 85 m. The boundary of continent ice in the
Southern America is showed with dotted line. 1 – sea ice, 2 – surface free from ice, 3 – internal
waters, 4 – snow and ice (Monin, Shishkov, 1979).
In conclusioin we remind the main reasons of the peculiarities of fauna of the Weddell Sea.
First of all it is the history of origin and development of the Antarctic fauna. Antarctic fauna was
formed in condition of permanent glaciaton. Antarctic glacier had rolled down and carried out
B.I. Sirenko: SOME BREACH OF THE WALLES’ RULE AND OTHER PECULIARITIES OF THE …
197
mainly very poor sorting deposits with a lot of various size rough stones, in the contrary from the
Arctic where many large rivers carry out mainly soft terrigenous materials. As a result of presence
of rough materials on bottom in the Weddell Sea during several million years the stable climax
epifaunal communities with sessile suspension feeders are formed, whereas in the Arctic the
infaunal communities with domination of representatives small buried animals are formed.
The permanent Weddell Gyre supplies bottom communities near Antarctic continent in its
south part by seston and krill that are produced mainly in north part of the Gyre. Bottom
communities are the accumulators that accumulate and keep organic and nutrients inside of
themself.
The main differences of Antarctic and Arctic ecosystems implies that in Arctic Ocean in spite
of a huge input of organics and nutrients from large river they are burring in thick deposits of
shelves and abyssal basins and are removing out from the biological cycle whereas in Antarctic
Ocean in spite of poor receipt of organic and nutrients from outside most of them are permanently
involved in the biological cycle and losses of organic and nutrients are minimum.
Acknowledgement
Authors express gratitude to Wolf Arntz and Julian Gutt from Alfred-Wegener-Institut
fur Polar-und Meeresforschung (Germany) for granting of an opportunity to collect rich
faunistic material from the Weddell Sea and support of our researches.
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| id | nasplib_isofts_kiev_ua-123456789-129405 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 1727-7485 |
| language | English |
| last_indexed | 2025-12-02T14:17:28Z |
| publishDate | 2012 |
| publisher | Національний антарктичний науковий центр МОН України |
| record_format | dspace |
| spelling | Sirenko, B.I. Smirnov, I.S. 2018-01-19T15:39:44Z 2018-01-19T15:39:44Z 2012 Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea / B.I. Sirenko, I.S. Smirnov // Український антарктичний журнал. — 2011-2012. — № 10-11. — С. 186-200. — Бібліогр.: 71 назв. — англ. 1727-7485 https://nasplib.isofts.kiev.ua/handle/123456789/129405 591.9+99 According to the rule of de Candolle-Walles a species diversity decreases from low latitudes regions to high latitudes regions. However in the Weddell Sea the biodiversity of the fauna of some groups are approximately the same (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) or more than this in the Magellan region (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea, Ascidiacea). In spite of the rather high species diversity of bivalves in the Weddell Sea, communities where bivalves dominated (like in Arctic seas) practically are absent and average size of gastropods and bivalves is much less than in the Arctic Ocean. Moreover Reptantia from Decapoda crustacean is absent in the Weddell Sea. We will try to explain the reasons of the breach of the Walles rule of the fauna and will try to explain some other peculiarities of the Weddell Sea. For this purpose we will compare the fauna of some groups in the Magellan region and in the Weddell Sea (Table. Number of species of different genera the Magellan Region and in the Weddell Sea). Согласно правилу меридионального изменения разнообразия Декандоля–Уоллеса, разнообразие видов уменьшается от низких широт к высоким. Сравнение фауны моря Уэдделла с фауной соседнего с ним Магелланова района обнаруживает нарушение этого правила для некоторых групп морских донных животных. У актиний, мизид, кумовых раков, брюхоногих моллюсков, мшанок и морских ежей (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) видовое разнообразие в море Уэдделла почти не уступает таковому в Магеллановом р-не, а у губок, морских пауков, голотурий, морских звезд, офиур и асцидий (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea, Ascidiacea) оно даже выше, вопреки правилу Уоллеса. Несмотря на довольно высокое разнообразие видов двустворчатых моллюсков в море Уэдделла, сообщества, где двустворки доминируют (как в арктических морях), фактически отсутствуют, а средний размер гастропод и двустворчатых моллюсков – намного меньше, чем в Северном Ледовитом океане. Кроме того, ракообразные из группы Reptantia (Decapoda) в море Уэдделла отсутствуют. Мы попытаемся объяснить причины нарушения правила Декандоля–Уоллеса и некоторые другие особенности фауны моря Уэдделла. С этой целью мы сравним фауну некоторых групп беспозвоночных Магелланова района и моря Уэдделла. Відповідно до правила меридіональної зміни різноманіття Декандоля–Уоллеса, різноманіття видів зменшується від низьких широт до високих. Порівняння фауни моря Уедделла з фауною сусіднього з ним Магелланового району виявляє порушення цього правила для деяких груп морських донних тварин. У актиній, мізид, кумових раків, черевоногих молюсків, мшанок та морських їжаків (Actiniaria, Mysidacea, Cumacea, Gastropoda, Bryozoa, Echinoidea) видове різноманіття в морі Уедделла майже не поступається такому в Магеллановому районі, а у губок, морських павуків, голотурій, морських зірок, офіур і асцидій (Spongia, Pycnogonida, Holothuroidea, Asteroidea, Ohpiuroidea, Ascidiacea) воно навіть вище, всупереч правилу Уоллеса. Попри досить велике різноманіття видів двостворчастих молюсків у морі Уедделла, угруповання, де двостворки домінують (як в арктичних морях), фактично відсутні, а середній розмір гастропод і двостворчастих молюсків набагато менший, аніж у Північному Льодовитому океані. Окрім того, ракоподібні з групи Reptantia (Decapoda) в морі Уедделла відсутні. Ми спробуємо пояснити причини порушення правила Декандоля–Уоллеса та деякі інші особливості фауни моря Уедделла. З цією метою порівняємо фауну деяких груп безхребетних Магелланового району й моря Уедделла. Authors express gratitude to Wolf Arntz and Julian Gutt from Alfred-Wegener-Institut fur Polar-und Meeresforschung (Germany) for granting of an opportunity to collect rich faunistic material from the Weddell Sea and support of our researches. en Національний антарктичний науковий центр МОН України Український антарктичний журнал Океанографія та біологічні ресурси Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea Нарушение правила Уоллеса и другие особенности фауны моря Уэдделла Порушення правила Уоллеса та інші особливості фауни моря Уедделла Article published earlier |
| spellingShingle | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea Sirenko, B.I. Smirnov, I.S. Океанографія та біологічні ресурси |
| title | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea |
| title_alt | Нарушение правила Уоллеса и другие особенности фауны моря Уэдделла Порушення правила Уоллеса та інші особливості фауни моря Уедделла |
| title_full | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea |
| title_fullStr | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea |
| title_full_unstemmed | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea |
| title_short | Some breach of the Walles' rule and other peculiarities of the fauna in the Weddell Sea |
| title_sort | some breach of the walles' rule and other peculiarities of the fauna in the weddell sea |
| topic | Океанографія та біологічні ресурси |
| topic_facet | Океанографія та біологічні ресурси |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/129405 |
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