AN ABSTRACT OF THE THESIS OF
Gordon Richard Bilyard for the degree of
in Oceanogra
Title:
presented on
Doctor of Philosophy
May 21, 1979
Zoogeography and Ecology of Western Beaufort Sea Polychaeta
(Annelida)
Redacted for Privacy
Jthstract approved:
rey, Jr.
Dr. Andrew G.
The western Beaufort Sea polychaete fauna may be divided into sublittoral and bathyal components.
Sublittoral species occur at depths
of less than 300 in, and may be stenobathic or eurybathic.
Bathyal
species are found exclusively below 300 in.
A dominance of axnphiboreal-Arctic species, a near absence of en-
demic species, and a relatively low number of species in the sublittoral environment are attributed to invasion of the sublittoral environment during interglacial intervals.
Lower sea level, grounded ice,
and low nutrient supply during glacial intervals of the Pliocene and
Pleistocene are suggested agents of faunal extinction.
Within the bathyal polychaete fauna the prevalence of endemic and
Atlantic-boreoarctic species and the absence of Pacific-boreoarctic
species reflect the relative isolation of the bathyal and abyssal
Arctic Ocean.
The bathyal depth of the deepest portion of the North
Atlantic Transversal Ridge (830 m) has permitted some faunal exchange
between the Atlantic and Arctic Oceans since the Miocene, while the
shallowness of the Bering Strait (70 m) has precluded a similar
exchange of Pacific and Arctic bathyal faunas since the Late
Cretaceous.
Isolation of the Arctic Ocean basin below 830 m for
approximately 65 m.y. has permitted the evolution of a strongly
endemic fauna.
Depth and depth-related processes appear to exert primary control
over the distributional patterns of species of polychaetous annelids
in the western Beaufort Sea (Cape Halkett to Barter Island, Alaska).
Species richness and total polychaete abundance are maximal along the
outer continental shelf and upper continental slope.
Stations exhibit-
ing the most similar polychaete assemblages are located at similar
depths.
However, some species distributions appear to correlate better
with sediment type than with depth.
Maximum abundance occurs deeper on
the continental slope to the west, and station clusters generated by
non-hierarchical clustering of the dominant polychaete species data are
not sorted strictly by depth.
in a canonical analysis of discriminance,
the station clusters were projected onto a two-dimensional plane in
"species space."
The first and second canonical variables of the
station clusters correlate with sediment grain size distributions,
suggesting a relationship between polychaete distributional patterns
and the sedimentary environment.
This relationship is further substan-
tiated when sediment grain size distributions for each station are
plotted on a tertiary diagram:
the stations are grouped and ordered
in a pattern similar to that generated by the canonical analysis of
discriminance.
Zoogeography and Ecology of Western
Beaufort Sea Polychaeta
by
Gordon Richard Bilyard
A THESIS
submitted to
Oregon State University
in partial fulfillment of
the reguirements for the
degree of
Doctor of Philosophy
Completed May 21, 1979
Commencement June 1980
APPROVED:
Redacted for Privacy
Associate Prdgessor o)Oceanography in charge of major
Redacted for Privacy
Dean of School of Oceanography
Redacted for Privacy
Dean of Graduate School
Date thesis is presented
May 21, 1979
Typed by Joan A Neuman for Gordon R. Bilyard
Ie
The field and laboratory assistance of R.E. Ruff is gratefully
acknowledged.
I thank J.J. Dickinson and J.G. Castillo for their help
in the laboratory, K. Fauchald (University of Southern California) for
assistance in resolving taxonomic problems, and C.D. Mclntire for his
suggestions concerning statistical analyses.
I am indebted to A.G.
Carey, Jr. for his assistance in the field and guidance throughout the
course of this study.
I also thank A.J. Boucot, C.B Miller and W.G.
Pearcy for their helpful comments and criticisms of this thesis.
The
financial support of the Humble Oil Company by a grant through the
Smithsonian Institution and the National Science Foundation by grant
numbers GA-36679 and DES75-14703 is gratefully acknowledged.
This
study was also supported in part by the Bureau of Land Management
through interagency agreement with the National Oceanic and Atmospheric
Administration, under which a multi-year program responding to the
needs of petroleum development of the Alaskan continental shelf is
managed by the Outer Continental Shelf Environmental Assessment
Program (OCSEAP) Office.
I am indebted to the U.S. Coast Guard for
ship support and the helpful cooperation of the officers and crew of
the USCGC GLACIER.
TABLE OF CONTENTS
Pgf
I.
II.
III.
INTRODUCTION
MATERIALS AND METHODS
PALEOGEOGRAPHY AND PALEOBIOGEOGRAPHY OF THE ARCTIC
OCEAN
Introduction
Mesozoic Era
Cenozoic Era
A.
B.
C.
IV.
ZOOGEOGRAPHY OF WESTERN BEAUFORT SEA POLYCHAETA
Introduction
Results
Discussion and Conclusions
A.
B.
C.
V.
ECOLOGY OF WESTERN BEAUFORT SEA POLYCHAETA
Introduction
Results
Discussion and Conclusions
A.
B.
C.
BIBLIOGRAPHY
3
12
12
12
14
20
20
20
42
51
51
51
61
68
APPENDIX
Stations selected for analysis of the polychaete
fauna.
80
LIST OF FIGURES
Page
Figure
1
Stations (1-58) in the western Beaufort Sea.
4
2.
Ecological stations (1-24).
7
3
Dominant polychaete species, including feeding
type and depth range within the study area for
each species.
9
4
Polychaeta/m2, contoured (solid lines).
53
5
Number of species encountered at each station,
contoured (solid lines).
54
6
Station affinities estimated by Pearson ProductMoment Correlation Coefficients.
56
7
Four clusters of stations (A, B, C, D) in the
western Beaufort Sea.
57
8
Station clusters plotted on the plane described
by the first two canonical variables which were
identified by the canonical analysis of
discriminance.
59
9
Sediment grain size distributions for all stations
except 6 and 7 are plotted following the method
outlined by Shepard (1954).
65
LIST OF TABLES
Page
Table
1
Distribution of samples, by depth interval.
2
Polychaete species listed alphabetically, by
Family.
21
3
Geographic distributions and depth distributions
within the study area of Polychaeta collected in
the western Beaufort Sea.
28
4
Zoogeographic affinities of the sublittoral and
bathyal faunas, summarized from Table 3.
41
5
Larvae of polychaete species for which larval
type is known.
46
6
Significant correlations (P<.Ol) of the first
and second canonical variables (Figure 8) with
dominant polychaeta species, log transformed
data (ln(x+1)). Significant correlations
(P<05) of those species with sediment grain
size, arcsin transformed data (arcsin
v'oportion), are also included.
60
5
ZOOGEOGRAPHY AND ECOLOGY OF WESTERN BEAUFOP SEA POLYCHAETA (ANNELIDA)
I. INTRODUCTION
The Western Beaufort Sea Ecological Cruises, multidisciplinary
oceanographic research cruises aboard the United States Coast Guard
icebreaker "Glacier," were undertaken during the summers of 1971
(WEBSEC-71) and 1972 (WEBSEC-72).
Since 1975, multidisciplinary
research cruises in the western Beaufort Sea have been carried out
aboard the United States Coast Guard icebreakers "Glacier" and
"Northwind," under the auspices of the Outer Continental Shelf Environmental Assessment Program (OCSEAP).
The primary objective of these
cruises was to further understanding of the western Beaufort Sea,
including its physical, chemical, geological, and biological attributes.
This dissertation presents some of the results of research on
the marine biota of the region, in particular the polychaetous annelid
component of the benthic infauna, and is part of a continuing research
effort in the region by the benthic ecology group at Oregon State
University (Carey et al., 1974; Carey and Ruff, 1977; Montagna and
Carey, 1978; Montagna, 1979).
Prior to the WEBSEC cruises, knowledge of the benthic invertebrates of the region was sketchy, contained primarily in the results
of the Canadian Arctic Expedition (1913-1918) and the writings of
MacGinitie (1955).
A key taxonomic paper on the polychaetes of the
Point Barrow region (Pettibone, 1954) was one product of MacGinitie's
interest in the benthos.
The WEBSEC and OCSEAP cruises were thus the
first comprehensive efforts to study the western Beaufort Sea,
2
including the benthic invertebrates.
The zoogeography of the western Beaufort Sea will be studied by
interpreting apparent zoogeographic affinities of the polychaete fauna.
Consideration of the evolution of the Arctic Ocean basin and present
and past attributes of the western Beaufort Sea marine environment
will be an integral part of the interpretive process.
The ecology of the western Beaufort Sea polychaetes will be
investigated through delineation of patterns inherent in the distributions and abundances of the dominant species of polychaetes. Assessment of possible influences of environmental factors on the distributional patterns of the polychaete species will be facilitated through
statistical analyses.
The polychaetous annelids were chosen as the indicator group for
research on the zoogeography and ecology of the western Beaufort Sea
benthic fauna because they are the dominant group of macrobenthic
organisms in soft bottom marine environments (Day, 1967).
Polychaetes
typically occupy a large number of niches in the sedimentary environment and employ, amongst the various species, a large variety of
feeding strategies (see Jumars and Fauchald, 1977).
3
II. MATERIALS AND METHODS
A Smith-McIntyre grab sampler (0.1 m2; Smith and McIntyre, 1954)
and a Sandia-Hessler box corer (Model Mk-3; 0.25 m2) were used to
quantitatively sample the benthic infauna on the continental shelf and
slope.
Qualitative samples of the benthic infauna and epifauna were
collected by Dr. A.G. Carey, Jr. using semi-balloon shrimp trawis
(otter trawis) lined with 1.3 cm stretch mesh.
Trawis having 6.7 m and
3.7 m apertures at the head-rope were used during the WEBSEC-71 and
WEBSEC-72 cruises, respectively.
Samples collected at 58 of the WEBSEC and OCSEAP stations across
the continental shelf and slope (Figure 1, Appendix) were selected for
detailed analysis of the polychaete fauna.
Included were 151 Smith-
McIntyre grab samples, 19 otter trawl samples, and three box core
samples (Table 1).
The depth of the continental shelf break was
defined as approximately 70 m by Carsola (1954) and Carsola et al.
(1961).
The WEBSEC-71 grab samples were carefully washed on a sieve having
a mesh aperture of 0.42 mm using a modified flotation technique aboard
ship.
Grab samples and box core samples collected during later cruises
were sequentially washed on sieves having mesh apertures of 1.00 mm
arid 0.42 mm, while otter trawl samples were washed free of sediment on
various sieves having larger mesh apertures.
Aboard ship all samples
were fixed in a buffered ten percent aqueous solution of formaldehyde.
The samples were returned to the laboratory, where all grab samples
and box core samples were sequentially washed on sieves having mesh
apertures of 1.00 mm and 0.42 mm.
All samples were picked
72
I-u,w
Figure 1.
Stations (1-58) in the western Beaufort Sea.
in meters.
Depth contours are
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GEAR SAMPLING
S
and sorted to major taxonomic categories, and the resultant faunal
fractions were preserved in 70 percent ethanol.
The polychaetes in the 1.00 mm fractions of the WEBSEC-71 grab
samples, and the polychaetes in the 1.00 mm and 0.42 mm fractions of
the OCSEAP grab samples and box core samples were identified to species.
In contrast to grab samples and box core samples, wherein the identif ication of all polychaete specimens was attempted, only selected speci-
mens collected by otter trawl were examined and identified.
While data from all samples were used in the zoogeographical
analysis of the polychaete fauna, only grab samples collected between
Cape Halkett and Barter Island during WEBSEC-7l (stations 1-24; Figures
1,2; Appendix) were selected for ecological analysis of the polychaete
fauna because the sampling gear deployed collected quantitative samples.
Statistical analysis of the resultant data set was thus simplified.
Five grab samples were collected at each of the ecological stations
(stations 1-24) except station six, where four were collected.
More
than 97 percent of the 22,399 specimens collected at the ecological
stations were identified at the species level; most of the remaining
specimens were too damaged to identify.
A complete listing of data
gathered in this study is found in Carey (1979).
A copy of this
report is available through the National Technical Information
Service.
Of the 133 polychaete species encountered in this study, 16 are
new and will be described in forthcoming publications.
Undescribed
taxa bear letter designations in the following discussion (e.g.
Eclysippe sp. A).
Assistance in resolving difficult taxonomic problems
£6
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Figure 2.
150°
Ecological stations (1-24).
48°
46°
Depth contours are in meters.
440
8
was kindly provided by Dr. Kristian Fauchald at the University of
Southern California.
Generic assignments for polychaete species
mentioned in the following discussion follow Fauchald (1977).
To facilitate some of the statistical analyses, a subset of 39
dominant species was selected (Figure 3).
Dominant species were
defined as those which occurred at 12 or more of the 24 stations,
and/or accounted for at least five percent of the total polychaete
abundance at one or more stations.
The U.S. Geological Survey collected sediment samples at all
stations except six and seven.
Particle sizes were measured by the
U.S.G.S. using standard techniques (Barnes, 1974).
These data
(pea 'entage gravel, percentage sand, percentage silt, percentage clay)
were used to elucidate possible relationships between polychaete
species and the sedimentary environment.
Patterns
n the distributions and abundances of polychaetes, and
the possible relationships of those patterns to environmental factors
were defined with the aid of multivariate statistical techniques.
Although linear models were applied to non-linear biological data in
the multivariate data analysis, studies (e.g., Cassie and Michael,
1968; Bernstein et al., 1978; Mclntire, 1978) have demonstrated the
usefulness of these models for identifying and interpreting patterns
in marine benthic data.
Transformations that improve the linearity
of the data were applied to the polychaete and sediment data prior to
statistical analyses.
We applied the transformation ln(x+1) to the
dominant polychaete data to correct for "departures from normality
arising from the Poisson error component which causes pronounced skew
9
Cistenides hyperborea
SS
Lysippe labiota
Scolibregmo inflotum
SS
Typosyllis Cornuta
Anaitides groenlondica
Pholoe minuta
Prionospia steenstrupi
Antinoella sarsi
Baroritolla americana
Chaetozone setoso
Chone murrnanico
Cossuro longocirrata
Eteone long a
S
C
C
C
S
S
SS
S
F
SS
C
Nephtys ciliata
0?
Touberia gracilis
SS
Al I ia suec ca
SS
All a sp. A
SS
Ophelino cylindricaudalus
SS
Scoloplos aCutus
Sternaspia
fossor
Terebellides stroerni
Capitella capitala
Heteromastus tiliformis
Lumbrineris minuta
Maldane sarsi
Micronephthys minuto
Mirruspio cirrifera
Myriochele
Than y x
been
? acut us
Spiochaetopterus typicus
quadricuspis
Loonice cirrof a
Owenia fusiforrnis
Onuphis
Eclysippe up. A
Amage ouriculo
Ophelina abronchiata
Sigarnbra tentaculata
Ophelina up. A
Lumbrineris sp. A
Depth Interval (m)
No. of Smith -McIntyre Grab Samples
SS
SS
S
SS
SS
C,SS?
SS
0?
S
SS
S
F,S
S,0?
S
F
S
S
SS
0?
SS
C,SS?
C?
0 2525 0 ID IS 0 5 10 0 0 10 0
t
Figure 3.
oJ
5
54
I
Dominant polychaete species, including feeding type and depth range within the study
area for each species. Feeding type designations are based on information given in
Pettibone (1963), Day (1967), HarthannSchröder (1971), Ushakov (1972), and Jumars
and Fauchald (1977). SS, subsurface deposit
feeder; S, surface deposit feeder; C, carnivore; F, filter feeder; 0, omnivore. Bars
indicate depths of actual collection; arrows
indicate depth scale changes.
10
We
when the mean count is small" (Cassie and Michael, 1968, p. 18).
applied the transformation arcsin /roportion to the sediment particle
size data to remove the non-linearity associated with proportional
data (Cassie and Michael, 1968).
Distributional patterns of dominant polychaetes were studied
using a two-step procedure described by Mclntire (1978).
The first
step was to cluster stations in "species space" (39 species, 39
dimensions).
Program CL[JSB standardized the data (x0; s.d.l) and
then generated station clusters by solving for the least sums of
squares of distance about a designated number of cluster means (see
Mclntire, 1973).
CLt.JSB employs a non-hierarchical clustering algorithm
and sets no priorities on cluster separations.
The second step was to
identify factors which might be responsible for cluster integrity.
A
canonical analysis of discriminance of station clusters in "species
space" (from CLUSB) was performed for this purpose.
The discriminant
model found the linear functions in "discriminant space" (canonical
variables) in which the between-cluster differences were best
expressed, and projected the station points onto those functions.
The
dimensionality of the data set was reduced because two clusters were
separated by one canonical variable (Cooley and Lohnes, 1971).
All
sample variance was captured in c-i dimensions, where c is the number
of designated clusters (Cooley and Lohnes, 1971).
Station positions on the canonical variables were identified in
the canonical analysis of discriminance.
Cluster integrity was
visualized by plotting station points in "discriminant space."
Station
positions on the canonical variables were correlated with species
11
abundances at stations to determine which species were likely responsible for the assignments of stations to clusters by CLUSB.
The
species-canonical variable correlation matrix is analogous to a factor
pattern matrix in principal components analysis (see Cooley and Lohnes,
1971).
Station positions on the canonical variables and species
abundances at stations were also correlated with all other benthic
station data (number of polychaete species, total polychaete abundance,
percentage gravel, percentage sand, percentage silt, percentage clay,
depth, longitude) to elucidate environmental factors possibly influencing the distribution and abundance of polychaetes.
12
III,, PALEOGEOGRAPHY AND PALEOBIOGEOGRAPHY OF THE ARCTIC OCEAN
A. INTRODUCTION
A knowledge of regional history is helpful for the interpretation
of biogeographic data.
The following discussion synthesizes present
knowledge of the hydrological, climatological, and tectonic evolution
of the Arctic Ocean, and aids the interpretation of polychaete data
from the western Beaufort Sea.
As the earliest developmental phases
are unclear, only the development of the Arctic Ocean since the Late
Mesozoic will be discussed.
B. MESOZOIC ERA
The earth's land masses formed one continuous continent in the
early Mesozoic, but by Late Cretaceous no fewer than six separate land
masses existed (Dietz and Holden, 1970).
The rifting between Europe
and North America which began in the Mesozoic continued into the
Cenozoic, separating the two continents, and resulting in the westward
movement of North America relative to Asia (Dietz and Holden, 1970).
The clockwise rotation of North America caused "a phase of compressional motion between eastern Eurasia and Alaska from 81 m.y. to 61
m.y." (Pitman and Talwani, 1972, p. 640), which culminated in the
uplifting of the Bering Strait region in the Late Cretaceous (Hopkins
and Scholl, 1970).
Uplifting ceased in Late Cretaceous-Early Tertiary
time.
Throughout the Early and Middle Cretaceous the primordial Arctic
Ocean was little more than an indentation in the northern coastline of
13
Laurasia (Dietz arid Holden, 1970, Figure 5).
Kauffman (1973, Figure 1)
includes the northernmost coastline of Laurasia (which stretches across
northern North America, Europe, and northern Asia) in the North
Temperate Realm on the basis of fossil bivalve data.
Much of the North
Temperate Realm marine fauna was shared between the North Pacific,
primordial Arctic, North American, European, and Asian coastlines, as
there were no major biogeographic barriers to restrict migration.
Movement of the North American continent westward and the consequent uplift of the Bering Strait region in the Late Cretaceous and
Early Tertiary created the first major marine biogeographic barrier to
migration between the Arctic Ocean and other marine regions.
Study of
fossil marine molluscs indicated that no exchange of mollusc genera
occurred between the Pacific and Atlantic Oceans during the Early and
Middle Tertiary (MacNeil, 1965).
However, "the Upper Cretaceous
reptilian faunas of northeastern Asia are certainly closer to those of
North America than to any others.
Indeed, there is a close identity
of faunas between these two areas, indicating a free interchange of
faunal elements back and forth, presumably across a Bering land bridge"
(a personal communication to A. Hallam, by E. Colbert:
Hallam, 1967,
p. 219).
Further isolation of the primordial Arctic Ocean during the Late
Cretaceous resulted from the reduction of the Western Interior Seaway,
an epicontinental seaway between the Gulf of Mexico and the Alaskan
Arctic.
By the Early Tertiary, the only marine connection between the
primordial Arctic Ocean and the remainder of the World Ocean was an
epicontinental seaway connecting the Arctic Ocean with the Tethys
14
(Berggren and Hollister, 1974).
Pitman and Taiwani (1972) concluded from North Atlantic gephysica1
data that all tectonic motion in the Arctic region during the Late
Cretaceous (81 to 63 m.y.B.P.) was compressional, and no sea-floor
spreading was in progress at that time.
that study.
No arctic data were used in
Alternatively, the Alpha Cordillera Ridge may have been an
ancient spreading center which was active in the Mesozoic and Early
Tertiary as an extension of the Mid-Atlantic Ridge and the Labrador Sea
Ridge (Vogt and Avery, 1974; Berggren and Hollister, 1974).
The
Labrador Sea Ridge probably became quiescent about 60 m.y, ago, after
which spreading resumed in a new location, between Greenland and Europe
(Vogt and Avery, 1974).
In either case, the Late Cretaceous Arctic
Ocean must have been much smaller than at present, since sea-floor
spreading which resulted in the splitting of the Lomonosov Ridge from
the Eurasian plate and the creation of the Eurasian Basin of the Arctic
Ocean did not commence until the Tertiary (Vogt and Avery, 1974;
Talwani and Eldholm, 1977).
C. CENOZOIC ERA
Paleogene
In the Paleocene and Early Eocene the Arctic Ocean was linked to
the Tethyan region via the Turgai Stratis.
This epicontinental seaway
east of the TJral Mountains formed an effective barrier to terrestrial
faunal and floral migrations between Europe and Asia (Berggren and
Hollister, 1974).
15
Mammalian data strongly suggest that Europe and North America were
linked by a land bridge during at least the Late Paleocene and Early
Eocene (few data are available for the Early and Middle Paleocene).
Simpson (1947), Kurt4n (1973) and West et al.
(1978) consider the
European and North American mammalian faunas of the Early Eocene to be
within the same zoogeographic region.
The Asian mammalian fauna is
much more similar to the European fauna than to the North American
fauna, a point which favors a North Atlantic land bridge, rather than a
Bering land bridge for faunal exchange between Europe and North America
(Kurtn, 1973).
Geophysical and paleobiogeographical data (Hopkins and
Scholl, 1970; Scholl et al., 1975) indicate, however, that the Bering
land bridge was in existence without interruption from the Late
Cretaceous to the Middle or Late Miocene.
Geophysical data indicate that the land bridge from Europe to
North America was probably located between Greenland and Spitsbergen
(Pitman and Talwani, 1972; Talwani and Eldholm, 1977), rather than
between Greenland, Iceland, and the Faeroe Islands (the "Thulean" land
bridge).
Existence of the Greenland-Spitsbergeri land bridge until the
latest Eocene, about 38 m.y.B.P., (Talwani and Eldholm, 1977) has been
suggested on the basis of the geophysical data.
However, an earlier
Middle Eoene divergence of the European and North American mammalian
faunas (Simpson, 1947; West et al., 1978) is strong evidence that the
land bridge was discontinuous after the Early Eocene.
Extension of the Mid-Atlantic Ridge into the Norwegian-Greenland
Sea region began in the Paleocene (Talwarii and Eldholm, 1977), and by
about late Early Eocene (50 m.y.B.P.) an Arctic-Atlantic connection had
16
probably been established (Berggren and Hollister, 1974).
The initia-
tion of this marine barrier between Europe and North America probably
produced the observed divergence of the mammalian faunas of the two
regions in the Middle Eocene (Simpson, 1947).
By Late Eocene Europe
and North America were in different zoogeographic regions (Kurtn,
1973).
The end of the Early Eocene also marked the end of the Turgai
Straits, due to a marine regression (Berggren and Hollister, 1974).
Neogene
From Middle Eocene to Late Miocene time the only marine corridor
to the Arctic Ocean was through the Norwegian-Greenland Sea.
This
corridor deepened as the Iceland-Faeroe Ridge subsided, with substantial
water exchange between the Norwegian-Greenland Sea and the North
Atlantic Ocean probably having been achieved sometime during the
Miocene (Schrader et al., 1976).
Although the deepest point along the
Iceland-Faeroe Ridge is presently relatively shallow (approximately
1,000 rn), ridge submergence during the Oligocene and Miocene would have
permitted faunal exchange at bathyal depths between the Arctic Ocean
and other marine regions for the first time since the uplift of the
Bering land bridge in the Late Cretaceous.
Since the Miocene, no im-
pediment to littoral, sublittoral, or bathyal faunal exchange with the
Atlantic Ocean has occurred.
The first submergence of the Bering land bridge since the Mesozoic
occurred in the Late Miocene, and continued until the end of the
Miocene, 5 m.y.B.P. (Hopkins and Scholl, 1970).
Faunal exchange
between the Atlantic and Pacific Oceans during that time has been
17
documented for the mollusc fauna (Durham and Allison, 1960; MacNeil,
1965).
Subsidence of the Bering land bridge to its present elevation
at 3.5 m.y.B.P.
(Hopkins and Scholl, 1970) resulted in a more dramatic
exchange of North Atlantic and North Pacific molluscs than had occurred
in the Late Miocene.
Why the earlier migration had not been of equal
magnitude is not understood (Einarsson et al., 1967).
Migrants from the Pacific Ocean to the Atlantic Ocean in the Late
Tertiary and Quaternary are more numerous than migrants from the
Atlantic Ocean to the Pacific Ocean, an imbalance which MacNeil (1965)
attributes to the predominantly northward flow of water through the
Bering Strait, and to the narrowness of the passageway in close proximity to the Pacific Ocean.
Pacific taxa entering the Arctic Ocean could
reach the Atlantic Ocean by many routes, but Atlantic taxa entering the
Arctic Ocean could not reach the Pacific Ocean unless their ranges
extended to the Bering Strait.
The global climatic deterioration which began in the Late Eocene
(Devereux, 1967; Wolfe and Hopkins, 1967; Shackleton and Kennett, 1974)
culminated with the onset of intermittent Northern Hemisphere glaciation in the Pliocene about 3.0 m.y.B.P.
(Berggren, 1972).
The present
interglacial Arctic Ocean is probably cooler than was the pre-glacial
Arctic Ocean, since some of the taxa which migrated through the Arctic
Ocean to the Atlantic Ocean or Pacific Ocean prior to 3.0 m.y.B.P. are
not now found there (Einarsson et al., 1967), and since a southward
migration of taxa from the Arctic Ocean has occurred in the Late
Cenozoic (Kauffman, personal communication).
Intensification of Northern Hemisphere glaciation occurred
18
throughout the Late Pliocene and Pleistocene, and probably reached a
maximum about 0.4 m.y.B.P. (Berggren, 1972).
Associated with this
intensification was the formation of the Arctic Ocean permanent pack
ice, which may have occurred as early as 3.5 m.y.B.P. (Clark, 1971).
A more conservative reconstruction suggests that seasonal pack ice was
present between 2.43 and 0.70 m.y.B.P., and that permanent pack ice
was present thereafter (Herman and O'Neil, 1975).
Since glacial intervals are of considerably longer duration than
interglacial intervals (Broecker and van Donk, 1970), a reconstruction
of arctic climatic conditions during the last glacial interval is most
informative in defining the predominant mode of arctic climatic conditions since the inception of the permanent ice pack.
Termination of
the penultimate glaciation at 127,000 y.B.P. (Broecker and Ku, 1969)
was followed by an interglacial interval of 12,000 years duration.
Although glaciation was initiated at 115,000 y.B.P., intensification
to full glacial conditions did not occur until 75,000 y.B.P.
(Shackleton and Opdyke, 1973).
Maximum glacial conditions persisted
for 64,000 years, until the advent of the present interglacial interval
at 11,000 y.B.P. (Broecker and Ku, 1969).
Full glacial conditions in the arctic were characterized by:
1) a sea-level drop of apprcximately 85 m (CLIMAP Project Members,
1976), 2) ice sheets over North america, Greenland, Iceland, Great
Britain, Scandanavia, the Barents Sea, and the Kara Sea (Boulton and
Rhodes, 1974; Hughes et al., 1977; Kvasov and Blazhchishin, 1978), and
3) thick pack ice and floating ice shelves over the Arctic Ocean basin
(Hughes et al., 1977).
A minimum reconstruction of full glacial con-
19
ditions would therefore include thick permanent pack ice over the
Arctic Ocean basin and grounded ice over most of the present Arctic
Ocean continental shelves.
The drastic reduction of the sublittoral
environment would have been accompanied by a substantial reduction in
primary productivity, due to severe ice cover and reduced light transmission.
In summary, since the onset of intermittent Northern Hemisphere
glaciation the arctic environment has been as warm as at present only
during brief interglacial intervals of about 12,000 years duration.
Present conditions, including seasonally open water along the Arctic
Ocean margin and fully marine conditions on the present continental
shelves, have occurred only 12,000 of each 100,000 years in the recent
geological past.
20
IV. ZOOGEOGRAPHY OF WESTERN BEAUFORT SEA POLYCHAETA
A. INTRODUCTION
Evolution of the Arctic Ocean basin and its marine environment
since the Late Mesozoic have been profoundly influenced by two interacting phenomena:
the relative isolation of the basin from the remain-
der of the World Ocean, and the global climatic deterioration that has
characterized the Cenozoic Era.
The effects of these two phenomena on
the marine biota of the Arctic Ocean are clearly reflected in the
present composition of the western Beaufort Sea polychaete fauna.
B. RESULTS
One hundred thirty-two species of Polychaeta were encountered in
this study (Table 2).
Five species, including Barantolla americana,
Allia abranchiata, Aricidea tetrabranchia, Sigainbra tentaculata, and
Ephesiella macrocirrus, have not been previously collected in arctic
waters (north of 66.5°N).
These species are preceeded by an asterisk
(*) in Table 2.
To study the relative contributions of different faunas to the
polychaete fauna of the western Beaufort Sea, a synopsis of geographic
and depth distributions within the study area is presented for each
species (Table 3).
Species are grouped by the similarity of their
geographic distributions.
Terminology consistent with that of Hoithe
(1978) is used to describe the groups.
Arctic species are defined as
those species occurring only north of 66.5° N latitude.
Amphiboreal-
21
Table 2
Polychaete species listed alphabetically, by Family.
Undescribed taxa are given letter designations (e.g.,
Eclysippe sp. A). Described species not previously
collected in arctic waters (north of 66.5°N) are preceded
by an asterisk (*)
AMPHARETIDAE
Amage auricula Malmgren, 1866
Ampharete acutifrons (Grube, 1860)
mpharete arctica Malmgren, 1866
vega (Wirén, 1883)
Miphicteis gunneri (Sars, 1835)
Eclysippe sp. A
Glyphanostomuin pallescens (The1, 1879)
Lysippe labiata Malmgren, 1866
Melinna cristata (Sars, 1851)
Sabellides borealis Sars, 1856
Genus A
APISTOBRANCHIDAE
Apistobranchus tulibergi (Th4e1, 1879)
CAPITELLIDAE
*Barantolla americana Hartman, 1963
Capitella capitata (Fabricius, 1780)
Heteromastus filiformis (Claparède, 1864)
Parheteromastus sp. A
Genus B
CHZET0PTERIDAE
Spiochaetopterus typicus Sars, 1856
22
Table 2. (continued)
CIRRATULIDAE
Chaetozone setosa Malmgren, 1867
Cirratulus cirratus (MUller, 1776)
Tharyx ? acutus Webster and Benedict, 1887
COSSURIDAE
Cossura longocirrata Webster and Benedict, 1887
Cossura sp. A
DORVILLEIDAE
Schistomeringos caecus (Webster and Benedict, 1884)
Schistomeringos sp. A
FLABELLIGERIDAE
Brada incrustata StØp-Bowitz, 1948
Brada inhabilis (Rathke, 1843)
Brada nuda Annenkova, 1922
Brada villosa (Rathke, 1843)
Diplocirrus hirsutus (Hansen, 1878)
Diplocirrus 1oxgisetosus (Maren2eller, 1890)
Pherusa plumosa (MUller, 1776)
GONIADIDAE
Glycinde wireni Arwidsson, 1898
EESIONIDAE
Nereimyra punetata (MUller, 1776)
LUMBRINERIDAE
Lumbrineris !1is (MUller, 1776)
Lunthrineris irnpatiens (Claparède, 1868)
23
Table 2. (continued)
LUMBRINERIDAE (continued)
Lumbrineris latreilli (Audouin and Mime-Edwards, 1833)
Luinbrineris minuta Th4el, 1879
Lwnbrineris sp A
Lumbrineris sp. B
MAGELONIDAE
Magpa longicornis Johnson, 1901
MALDANIDAE
Clymenura polaris (Th4el, 1879)
Lumbriclymene minor Arwidsson, 1907
Maldane sarsi Na].xngren, 1865b
Notoroctus oculatus var, arctica Arwjdsson, 1907
Petaloproctus tenuis (Thgel, 1879)
Praxillella praetermissa CMalmgren, 1865b)
NEPHTYIDAE
g1aophainusmaimgreni CThe1, 1879)
Micronepht
minuta (Thel, 1879)
Nepy ciliata (MUller, 1789)
Nephtys paradoxa Maim, 1874
NEREIDAE
Nereis zonata Maimgren, 1867
Nicon sp. A
ONUPHIDAE
Nothria conchylega (Sars, 1835)
Onuphis quadricuspis Sars, 1872
24
Table 2
continued)
OPHELIIDAE
abranchiata StØp-Bowitz, 1948
Ophelina acuninata Oersted, 1843b
2ina cylindricaudatus (Hansen, 1878)
Ophelina
p0 A
rane s,p
A
ORB INI IDAE
acutu
(Verrill, 1873)
OWEN I IDAE
Myriochele heeri Malmgren, 1867
Owenia fusiformis delle Chiaje, 1841
PARAONIDAE
*j)j
aiDranchiata (Hartman, 1965)
Allia suecica (Elaison, 1920)
Ailia sp, A
*Arjcidea tetrairanchia Hartman and Fauchald, 1971
Arcidea ushakovi Zaks, 1925
Paraonis sp
Tauberia
A
1is (Tauber, 1879)
PECTINARI IDAE
Cistenides
(Malmgren, 186)
PHYLLODOCIDAE
Anaitides citrina (Malingren, 1865a)
Anaitides groeniandica (Oersted, 1843a)
Eteone flava (Fabricius, 1780)
25
Table 2
(continued)
PHYLLODOCIDAE (continued)
Eteone
(Fabricius, 1780)
Mysta barbata (Malmgren, 1865a)
Mystides borealis The1, 1879
Paranaitis wahibergi (Malmgren, 1865a)
PILARGIIDAE
*sjgra tentaculata (Treadwell, 1941)
POLYNOIDAE
Antinoella badia (The1, 1879)
Antinoel.la sarsi (Malmgren, 1865a)
Arcteobia anticostiensis (McIntosh, 1874)
Enipo 91is Verrill, 1874
Eucranta villosa Malmgren, 1865a
Eunoe oerstedi (Malmgren, 1865a)
Gattcirrosa (Pallas, 1766)
Harmothoe imbricata (Linnaeus, 1767)
Lagisca extenuata (Grube, 1840)
Melaenis loveni Malmgren, 1865a
SABELLIDAE
Branchionirna infarcta (Kröyer, 1856)
Chone duneri Malmgren, 1867
Chone infundibuliformis Kröyer, 1856
Chone murinanica Lukasch, 1910
Euchone
pi11osa (Sars, 1851)
Jasmineira schaudinni Augener, 1912
26
Table 2. (continued)
SCALIBREGMIDAE
ia crassa (Oersted, 1843b)
inflatum Rathke, 1843
SERPULIDAE
Apomatus globifer Thdel, 1879
SIGALIONIDAE
Pholoe minuta (Fabricius, 1780)
SPHAERODORIDAE
*Ephesjella macrocirrus Hartinan and Fauchald, 1971
Sphaerodoridiuin claparedii (Greeff, 1866)
Sphaerodoridium sp. A
Shaerodorosis biserialis (erke1ey and Berkeley, 1944)
haerodoropsis minuta (Webster and Benedict, 1887)
Sphaerodoropsis sp. A
Sphaerodoropsis sp
B
Sphaerodorum
(Rathke, 1843)
SPIONIDAE
Laonicecirrata (Sars, 1851)
Minuspio cirrifera (Wirgn, 1883)
Polydora caulleryi Mesnhl, 1897
2nosio steenstrupi Malmgren, 1867
SPIRORBIDAE
Dexiospira
Spirorbis
spirilluin
(Linnaeus, 1758)
granulatus (Linnaeus, 1767)
27
Table 2. (continued)
STERNASPIDAE
Sternaspis fossor Stimpson, 1854
SYLLIDAE
Autyalexandri Malmgren, 1867
fallax Malmgren, 1867
Exogone dispar (Webster, 1879)
Exogone naidina Oersted, 1845
Sphaeryllis erinaceus, (Claparde, 1863)
Typosyllis cornuta (Rathke, 1843)
Typosyllis fasciata (Malmgren, 1867)
TEREBELLIDAE
Artacama proboscidea Malmgren, 1866
Axionice flexuosa (Grube, 1860)
Lanassa nordenskioldi Malmgren, 1866
Lanassa venusta Maim, 1874
Laphani
boecki Malmgren, 1866
Leaena abranchiat
Malmgren, 1866
Nicolea zostericola Oersted, 1844
Polycirrus medusa Grube, 1855
Proclea graff ii (Langerhans, 1884)
TRICHOBRANCHIDAE
Terebellides stroemi Sars, 1835
Trichobranchus glacialis (Ma].mgren, 1866)
TRQCHOCHAETIDAE
Trochochaeta carica (Birula, 1897)
Table 3.
Geographic distributions and depth distributions within the study area of Polychaeta
collected in the western Beaufort Sea. Abbreviations for types of geographic distributions
are: A, Arctic (found north of 66.5°N); abA, amphiboreal-Arctic (found in the boreal
Atlantic, boreal Pacific, and Arctic); AbA, Atlantic-boreoarctic (found in the boreal
Atlantic and Arctic); PbA, Pacific-boreoarctic (found in the boreal Pacific and Arctic).
An asterisk (*) indicates that the taxon is also found in temperate and/or tropical regions.
The number of records equals the number of stations at which a taxon was collected.
Occurrences within depth intervals are based on the depth of collection of individual
saxnples.
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
000000
0000000000tflOLflOLfl
0u-)000000000c'2mc')
0OUDNWO'HI I 11110
H H H H H 0
0I H IHHHHHHHHHH00000U)
0 U 000000000 Lfl 0 U 0 A
U) H I
Brada incrustata
A
1
x
Brada villosa
abA*
6
x
Chone infundibuliformis
abA*
1
x
Cirratulus cirratus
abA*
2
x
Dexiospira piri1lum
abA*
3
x
Eunoe oerstedi
abA*
1
x
A?
5
x
abA*
1
x
Genus A
Harmothoe imbricata
I
I
I
I
I
I
I
I
Table 3. (continued)
GEOGRAPHIC
TAXON
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
0
U)
0 U) 0 0
0000000000 U)0000
N
0 000000000
1110
00flDNO'r-1I
III-I I-IH0
II
U)-1I
0 0000 11)
U)
('.1
0 -1 0 U) 000000000
I
I
Luxnbriclymene minor
AbA
1
x
Melaenis loveni
abA
3
x
Nephtys paradoxa
abA*
2
x
Ophelina acuminata
abA*
3
x
Pherusa pluxnosa
abA*
1
x
Sabellides borealis
abA
3
x
Sphaerodoridium claparedil
AbA
1
x
abA*
3
x
abA*
2
x
haerodoropsis minuta
osyl1is fasciata
rnpharete vega
A
4
Autolytus fallax
AbA
6
Axionice flexuosa
abA
4
xx
xx
xx
-1
-t
-I .-
) Cfl
I
I
I
t
-I .-
U)
0 U') 0 )
A
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
000000
In 0 In 0 In
0000000000
In 000000000 -1 c'i
0
00HLnIDNO'H
I1r00
Lflr41 11111111 I
1
Cv)
I IHHHrr-I00000In
0 -I 0 In 000000000 In 0 In 0 Cv)
I
In
r-I r
Brada inhabilis
abA*
4
xx
Brada nuda
PbA
3
xx
Leaena abranchiata
abA*
2
xx
Nicolea zostericola
abA*
5
xx
Polycirrus medusa
abA*
5
xx
Polyphysia crassa
ZbA*
6
xx
Schistomeringos eaecus
abA*
7
xx
Sphaerosyllis erinaceus
abA*
2
xx
Autolytus alexandri
abA*
2
x
x
Eucranta villosa
abA*
2
x
x
Exogone dispar
abA*
6
Exogone naidina
abA*
10
(N C)
In
ID N CX) Ø'
I
I
rX -I (N (N Cv) A
xxx
xxx
tA)
0
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
000000
Ui
0000000000
0 Ui 000000000 H CN 0(N 0
Ui
Ui
I 0
Lfl 'D N ) Oi H I
00
H (N 11111
II Ii-IHHHr-I0
U)HI
I H H H H H H H H H H 00 0 00 Ui
Ui 0 Ui 0
C')
0 H 0 U) 000000000
I
Gattyana cirrosa
abA*
6
xxx
Glycinde wireni
PbA
2
x
Lagisca extenuata
abA*
9
xxx
Magelona longicornis
PbA*
3
x
Nereis zonata
abA*
12
Polydora caulleryi
abA*
7
Spirorbis granulatus
abA*
5
xxx
xxx
xxx
Trichobranchus glacialis
abA*
2
x
abA*
9
abA*
10
abA*
13
abA*
4
xnpharete arctica
Apis tobranchus tu1lbe
ippe labiata
Artacama proboscidea
C')
I
I
I
)
A
x
x
x
xxxx
xxxx
xxxx
x
xxx
H
Table 3. (continued)
GECAAPHIC
TAXON
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
000000
0000000000
U)
0 U) U)
0U)0000000O0c'r)r
C)
00 H C
U) H I
('.1
I
I
I
I
H H
abA*
14
x
Laphania boecki
abA
10
xxxxx
Praxillella praetermissa
abA*
6
x
Proclea graffii
abA*
7
xxxxx
Axnphicteis gunneri
abA*
6
xx
x
Lanassa venusta
abA
5
xx
x
Chone duneri
abA*
10
xx
x
Euchone papillosa
abA*
13
xxx
x
Glyphanostomum pallescens
abA*
5
xx
x
Lanassa nordenskioldi
abA
3
xx
x
Lumbrineris fragilis
abA*
xxx
x
Mysta barbata
abA
x
x
2
I 0
,-1 H H H H 0
0) H I
I
I
I
I
I
I
I
U) 'D N CO 0) H H c'
Cistenides hyperborea
12
CO
I
I H H H H H H H H H HU) 00000
H 0 U) 000000000 0 U) 0 U)m
0 U)
I
U) 'O N
()
e
A
x
x
x
(A)
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
000000
0000000000 if) 0 If) 0 If)
0 if) 0000 0 0000 H CN ('1
00 H C")
Lf)
O N COO) H I
c
I 0
I IHHHHHHHHHH00000
C')
I H H H H H 0
0 H 0 If) 000000000 if) 0 Lfl 0 Cfl
If) H I
I
I
I
I
I
A?
9
Typosyllis cornuta
abA*
15
xxx
xxx
xxx
Anaitides groen1andica
abA*
20
xxxxx xx
Ampharete acutifrons
abA*
10
Lumbrineris impatiens
abA*
6
Melinna cristata
abA*
12
Notoproctus oculatus
AbA
Pholoe minuta
abA*
Nothria conchylega
Paraonis sp. A
Clyrnenura polaris
abA
A
5
4
16
11
Prionospio steenstrupi
abA*
15
Spiochaetopterus typicus
abA*
11
I
I
A
x
x
x
xxx
xxx
xxx
x
x
x
x
xxx
xxx
I
I
I
x
x
x
x
x
x
x
xxxxxxx xx
xxxxxxxxxx
('3
('3
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
00 U)
00000
0000000000
U)
0 U)
0
U)
000000000
H
(N
(N
00HNC')U)'DNWO)HI
ii
ito
l.flr-1t till liii IHr-IHHHO
Io H
IrIHHHHHr-IHH00000Lfl
0 U) 000000000
U) 0 U) 0
'.0 N U) 0) H H N N C') A
N U) H H
C')
(N
C')
C')
In
c
A?
11
Barantolla americana
abA*
17
xxxxxxxxx
xxxx xxxxxx
Eteone longa
abA*
22
xxx xxx x xxx x
Nephtys ciliata
abA*
13
Chone murmanica
A
17
abA*
15
A?
6
Tauberia gracilis
abA*
16
x
xxxxxxx
xxxxxxx
xxx
xxxxxxxxxxxx
xxxxxxxxxxxx
xxxxxxxxxxxx
Trochochaeta car ica
abA*
12
xxx
Eteone f lava
abA*
8
Scalibre2ma inflatum
abA*
20
Sphaerodoropsis biserialis
PbA*
11
Allia sp. A
Cossura longocirrata
Sphaerodoridium sp. A
x
xx
x
xxx
xxx
C')
x
x
x
x
x
x
xx
x
x
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTEP'TAL (meters)
RECORDS
0 0000
000000
0000000000 0
0 000000000 H C4 ('4 () ()
00HUNOr-4I I
En 0 Lfl
En
En
I
I
10
I IHHHHHHHHHH00000En
000000000 tfl 0
in H I
0 H 0
En
C'
I
I
I
I
En
H H (N
Antinoella sarsi
abA*
18
xxxxx
Aricidea ushakovi
PbA*
27
x
Diplocirrus hirsutus
AbA
6
x
Diplocirrus longisetosus
abA
9
Laonice cirrata
abA*
20
Onuphis guadricuspis
abA*
16
Ophelina cylindricaudatus
abA*
21
Scoloplos acutus
AbA*
19
Sternaspis fossor
abA*
14
Antinoella badia
abA
8
Capitella captata
abA*
17
Owenia fusiformis
ai*
10
En
I
I
I
I H H H H i-I 0
En 0 (Y)
t.O N CO O' H H (N (N en A
x
x
x
x
I
x
xx xxx
x
x
xxx
xxx
x
xxxxxxxxx
x
xxxxxxxxx
xxxxxx
x
xxx
XXXXXXXXXXXX
x
xxx xxxx
)C
x
xx
xx
x
x
xxxxxxx xx
xxxxxx xx
xxxx
xx
01
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
0 0000
000000
0000000000
LflH
0 Lfl 0) Cfl
0
Lfl
000000000
00HCLfl¼DNHI('1I I 110
In
C'4
I HIHHr-Hr1H00000LU
0
0 In 000000000
In 0 J) 0 (
In 'D 1' CX) (7 -I H C'I Cl ('1 A
CI In .-1 .-I C'I m
Lfl H I
A?
11
Chaetozone setosa
abA*
33
Heteromastus filiformis
abA*
29
Micronephtys minuta
abA*
21
Sphaerodorum gracilis
abA*
11
Maldane sarsi
abA*
28
Minuspio cirrifera
abA*
35
Myriochele heeri
abA*
24
Terebellides stroemi
abA*
33
Allia suecica
abA*
27
Nereimyra punctata
abA*
16
Aglaophamusmalmgreni
abA*
31
Parheteromastus sp. A
I
I
I
I
I
xxx
I
I
I
I
.-I H H H ,- 0
x
x
xxxxxxxxxxxxxxxx
xxxxxxxxxxxx xxx
x
xxxxxxx xx x
xxx
x
xxx xxxxxxxxx
xxx xxxxxxxxxxx
xxx
x
xxx
x
x
x
xxxxxx
xxxxxxxxx xxxxxxx
x
xxxxxxxxx
xxx
xxxx x
xx
xxx
x
x
x
xxxxxx
x
x
0
Table 3. (continued)
GEOGRAPHIC
DISTRIBUTION
TAXON
DEPTH INTERVAL (meters)
RECORDS
00000
000000
00 00 0 00 0 00 LU 0 LU 0 LU
0LUO000o0O0OHc'c'cm
OOHcNmnDNo-I
11110
-4,-4r-IHO
Lfl.1
I
I
I
I
I
I
I
I
I
-4
.- 00000 LU
I
C) r 0 U) 0 0000000 if) 0 LU 0 m
A
I
Lumbrinerisminuta
PbA
37
32
Tharyx ? acutus
r-I
I
r-1
-I
p-I
.-4
.-1
xxxxxxxxxxxxxxxxxx
x x x x x x x
x
x x
Anaitides citrina
abA*
1
x
Mystides borealis
abA*
1
x
Paranaitis wahlbergi
AbA
1
x
Jasmineira schaudinni
AbA
5
x
4
x
8
x
x x x
x
A?
11
x
x x x
x
Arcteobia anticostiensis
abA*
1
x
Enipo
abA*
1
x
abA*
2
x
Sphaerodoropsis sp. B
Amage auricula
Eclysippe sp. A
acilis
Petaloproctus tenuis
A?
abA
x x
X
x x x x x
x
-4
Table 3. (continued)
TAXON
GEOGRAPHIC
DISTRIBUTION
RECORDS
DEPTH INTERVAL (meters)
0000000
0U)0 0000
0000000000
U)
U)(fl
0
U)
000000000
H
C4
I0
0) H
U)
N
00
C)
I IHHHHHO
Lflr-1I
11111$
IHHH
H H H .-4IH H
H 000 C) 0 U)
c'
C')
a)
I
I
I
I
0 U)HHCC')'U)¼0Sa)0)HHNCNC')A
H 0 U) 000000000 U) 0 U) 0
C')
A?
3
xx
Apoinatus globifer
AbA*
1
x
Branchiomma infarcta
abA
1
x
bA*
1
x
Luinbrineris sp. A
A?
4
x
x
Schistomerinos sp. A
A?
6
x
x
Ophelina abranchiata
AbA*
11
xxx
Sigarrthra tentaculata
abA*
23
xxxxxxxxxxx
A?
9
x
xxx
Luxnbrineris latreilli
abA*
3
x
x
Allia abranchiata
AbA*
17
x
A?
15
x
Sphaerodoropsis sp. A
Ephesiella macrocirrus
Genus B
Ophelina sp. A
x
xxxx
x
xxxx xx
xxxxx
l..3
a)
Table 3. (continued)
GEOGRAPHIC
TAXON
DISTRIBUTION
DEPTH INTERVAL (meters)
RECORDS
00000
000000
0000000000LnOLflOtfl
N N m CV
0 in 0 0 0 0in 0'.00S 0Q) 0O OH
I 0
-I
00 H N (fl
in
I H H H H H H -I H H H 00000
in 0 c
0 H 0 in
H
HNNC')
A
1'Lfl'.ON
NLflr-I HN
inHi 11111111 IHHHHr-I0
000000000 in 0
'
I
I
I
I
I
A?
2
xx
AbA*
5
xx
Nicon sp. A
A?
2
x
Cossura sp. A
A?
3
xx
Lunthrineris sp. A
A?
3
xx
Tachytrypane sp. A
Aricidea tetrabranchia
x
0
Arctic species are found in the arctic, boreal Pacific (Bering Sea and
Sea of Okhotsk), and boreal Atlantic (Labrador Sea, NorwegianGreenland Sea, and the waters of Greenland, Iceland, and northern
Europe).
Pacific-boreoarctic species inhabit arctic waters and the
boreal waters of the Pacific, but do not occur in the boreal waters of
the Atlantic, while Atlantic-boreoarctic species inhabit arctic waters
and the boreal waters of the Atlantic, but do not occur in the boreal
waters of the Pacific.
Taxa with ranges extending into temperate
and/or tropical regions are noted by the presence of an asterisk
(*)
in Table 3.
Undescribed species have been designated, albeit questionably
(Table 3), as Arctic species.
Since some of these species may be
collected in subarctic regions in the future, the level of endemism in
the Arctic Ocean po].ychaete fauna may be overestimated in the following
discussion.
The geographic distributions defined above are theoretical constructs applied to a continuum of species distributions, and are
intended to serve only as general guides for the grouping of species.
The author's discretion has been exercised in the assignment of some
taxa to their respective distributional groups.
The polychaete species encountered in this study may be arbitrarily divided into a sublittoral fauna and a bathyal fauna.
The sub-
littoral fauna includes the 115 species with upper depth range limits
of 20 to 150 m (Tables 3, 4) exclusive of Tharyx ? acutus (due to
taxonomic uncertainty).
All degrees of stenobathy and eurybathy are
exhibited by species of the sublittoral fauna.
Assigned to the bathyal
41
Table 4.
Zoogeographic affinities of the sublittoral and bathyal
faunas, summarized from Table 3.
SUBLITTORAL FAUNA
GEOGRAPHIC
DISTRIBUTION
NUMBER OF
SPECIES
PERCENT
OF FAUNA
89
77
11
10
Atlantic-boreoarctic
9
8
Pacific-boreoarctic
6
5
NUMBER OF
SPECIES
PERCENT
OF FAUNA
Arctic
9
53
Atlantic-boreoarctic
5
29
Amphiboreal-Arctic
3
18
mphiboreal-Arctic
Arctic
BATHYAL FAUNA
GEOGRAPHIC
DISTRIBUTION
42
fauna are 17 species with minimum depths of occurrence in excess of
300 m, beginning with Sphaerodoropsis sp. A in Tables 3 and 4.
The absence of species with upper depth range limits of 151 to
300 m depth may largely reflect sampling intensity (Table 1) rather
than a gap in the occurrence of additional species with increasing
depth, yet the assignment of species to a sublittoral or bathyal fauna
may be justified by the difference in the zoogeographic affinities of
the two faunas.
The sublittoral fauna is predominantly composed of
species exhibiting amphiboreal-Arctic distributions (89 species, 77
percent of the fauna), the vast majority of which (78 species) also
occur in temperate and/or tropical latitudes.
Species with Arctic
(four described species, seven undescribed species), Atlantic-boreoarctic (nine species), and Pacific-boreoarctic (six species) distributions account for only 23 percent of the sublittoral fauna.
By
contrast, nine of the seventeen bathyal species are undescribed and
probably endemic to the Arctic Ocean basin.
Five species (29 percent)
are Atlantic-boreoarctic; only three species (18 percent) are amphiboreal-Arctic.
There are no Pacific-boreoarctic bathyal species.
C. DISCUSSION AND CONCLUSIONS
The Sublittoral Fauna
Zoogeographic affinities of the western Beaufort Sea sublittoral
polychaete fauna are similar to zoogeographic affinities of. other
benthlc Launas within the Arctic Ocean basin.
inatei
A polychaete fauna dom-
y amphiboreal-Arctic species, but including some Pacific-boreo-
43
arctic and Atlantic-boreoarctic species was reported from the Canadian
Archipelago (Grainger, 1954).
Similarly, about half the species of
western Beaufort Sea bivalve molluscs with upper depth ranges shallower
than 300 m exhibit an'iphiboreal-Arctic distributions (Bernard, in
press).
The balance of the bivalve fauna includes Atlantic-boreoarctic
(32 percent) and Pacific-boreoarctic (17 percent) species (Bernard, in
press).
A predominance of species with geographic distributions
extending into the Atlantic and/or Pacific Oceans, coupled with a
dearth of endemic species, has also been reported for isopod
crustaceans (Menzies et al., 1973), sea stars (Grainger, 1966),
loricate molluscs (Yakovieva, 1950), and bryozoans (Powell, 1968), as
well as a mixed collection of benthic invertebrates from the Point
Barrow region (MacGinitie, 1955).
The absence of a strong endemic component and the consequent
boreal character of the Arctic Ocean sublittoral fauna have been cited
as evidence of a youthful fauna (Zenkevitch, 1963; Briggs, 1974; Knox
and Lowry, 1977).
Zoogeographic affinities of the sublittoral
polychaete fauna of the western Beaufort Sea, as discussed earlier,
further substantiate the boreal character of the sublittoral fauna
and are supportive of this hypothesis.
That the shallow water fauna appears to be depauperate in numbers
of species has also been presented as evidence in support of an
immature fauna (Zenkevitch, 1963; Dunbar, 1968; Knox and Lowry, 1977).
Knox and Lowry (1977) estimate that 300 species of polychaetes occur
in the Arctic Ocean.
By comparison, more than 650 species probably
occur in the Antarctic region (Knox and Lowry, 1977), more than 750
44
species probably occur in the shallow waters off South Africa (Day,
1967), and about 550 polychaete species have been reported from depths
of less than 200 in off southern California (Hartman, 1969a; Hartman,
1969b)
The collection of 115 species of sublittoral polychaetes in
the present study is, therefore, not inconsistent with the concept of
a depauperate shallow water fauna, despite the collection of some undescribed species and the extension of some species' ranges into the
Arctic Ocean basin.
Seasonal variations in temperature (-2°C to +2°C) and salinity
(27°/
to 35°/©) within the surface waters (0-250 in) of the Arctic
Ocean (Coachman and Aagaard, 1974) are not of great magnitude, and
are, therefore, probably not factors which contribute to the youthful
character of the sublittoral fauna.
Climatic changes during the
Pliocene and Pleistocene are probably responsible for the youthful
character of the sublittoral fauna (Zenkevitch, 1963; Dunbar, 1968).
Under severe glacial conditions, which are thought to have abated less
than 10,000 years ago (Hughes et aL, 1977), many sublittoral species
were probably eliminated (Bernard, in press).
Loss of habitat and low
nutrient supply were the probable agents of faunal extinction, since
Dunbar (1968) has shown that low temperature alone is insufficient to
prevent the development of a rich marine fauna.
Re-invasion of the
sublittoral environment by Atlantic and Pacific species during interglacial intervals would be consistent with the concept of an immature
sublittoral fauna, depauperate in the total number of species and in
the number of endemic species.
A high incidence of direct development, without a pelagic larval
45
stage, is generally displayed by high latitude benthic invertebrates
(Thorson, 1957; Thorson, 1966; Dell, 1972), but because direct develop-
ment limits dispersal, it would be a disadvantage to species invading
new habitats.
An invasion fauna would be expected to contain species
with pelagic larvae.
A relatively high incidence of species with
pelagic larvae (24 of 34 species) is found among those species of sublittoral polychaetes from the western Beaufort Sea for which the mode
of reproduction is known (Table 5).
These data are biased by:
1) few
studies of species with benthic larvae compared to species with pelagic
larvae, and by 2) the collection of most of these data in temperate
latitudes, since some species are known to change their reproductive
patterns with latitude (Thorson, 1946).
Yet the data illustrate that
a pelagic larval phase is not uncommon in Arctic Ocean sublittoral
species.
The Bathyal Fauna
The zoogeographic affinities of the fauna found exclusively below
300 or 400 in in the Arctic Ocean are similar to those for other benthic
taxa.
Few endemic species of isopods are found on the continental
shelf, but endemic and Atlantic-boreoarctic species are prevalent in
the isopod fauna below 425 m (Menzies et al., 1973).
Of the seven
species of western Beaufort Sea bivalves with upper depth range limits
in excess of 300 m, four are endemic and three have Atlantic-boreoarctic distributions (Bernard, in press).
In addition, the bathyal
ostracod fauna (Joy and Clark, 1977) and the total benthic fauna in
bathyal and abyssal depths off Siberia (Zenkevitch, 1963) are dominated
46
Table 5.
Larvae of polychaete species for which larval type is known.
P, pelagic larva; Ps, pelagic larva with short pelagic life
Taxa are ordered
(a few days or less); B, benthic larva.
as in Table 4
TAXON
LARVA
SOURCE
Cirratulus cirratus
B
Stephenson, 1950
Dexiospira spirillum
Ps
Thorson, 1946
Harmothoe iinbricata
P
Thorson, 1936
Nicolea zostericola
B
Thorson, 1946
Sphaerosyllis erinaceus
B
Cazaux, 1972
Gattyana crrosa
P
Thorson, 1946
Nereis zOnata
B
Thorson, 1936
Polydora caulleryi
P
Hannerz, 1956
Spirorbis grariulatus
Ps
Thorson, 1946
Artacama proboscidea
P
Thorson, 1946
Luxnbrineris fragilis
B
Thorson, 1946
Mysta barbata
P
Thorson, 1946
Nothria conchylega
P
Thorson, 1936
Anaitides groenlandica
P
Thorson, 1946
Lumbrneris impatiens
P
Cazaux, 1972
Melinna cristata
Ps
Nyholin, 1950
Pholoeminuta
P
Mileikovsky, 1968
Prionospio steenstrupi
P
Hannerz, 1956
Spiochaetopterus typicus
P
Bhaud, 1966
Eteone longa
P
Thorson, 1946
Nephtys ciliata
P
Thorson, 1946
Eteone f lava
P
Thorson, 1946
47
Table 5
(continued)
TAXON
LARVA
SOURCE
Scalibregma inflatuin
B
Thorson, 1946
Antinoella sarsi
P
Thorson, 1946
Laonice cirrata
P
Hannerz, 1956
Onuphis guadricuspis
B
Fauchald, personal communication
Capitella capitata
P,B
Grassle and Grassle, 1977
Owenia fusifortnis
P
Thorson, 1946
Heteromastus filiformis
P
Rasmussen, 1956
Sphaerodorum gracilis
B
Thorson, 1946
Miriuspio cirrifera
P
Hannerz, 1956
Nyriochele heeri
P
Mileikovsky, 1968
Terebellides stroemi
B
Thorson, 1946
Nereimyra punctata
P
Bhaud, 1967
48
by endemic and Atlantic-boreoarctic species.
Conspicuously absent
from this and other studies (Zenkevitch, 1963; Menzies et al., 1973;
Joy and Clark, 1977; Bernard, in press) is a Pacific-boreoarctic
element.
The composition of the bathyal polychaete fauna reflects the
geological history of the Arctic Ocean basin.
The absence of Pacific-
boreoarctic polychaetes (and other taxa) is due to the effectiveness of
the Bering Strait as a topographic barrier to faunal dispersion.
Subsidence of the Bering land bridge to its present elevation at 3.5
m.y.B.P. during the Pliocene (Hopkins and Scholl, 1970)
has permitted
the exchange of shallow water species at a maximum depth of 70 m only
during interglacial intervals.
No exchange of truly bathyal or abyssal
species between the Pacific and Arctic Oceans has been possible for
over 65 m.y.
Dominance of the Arctic Ocean bathyal fauna in general by endemic
and Atlantic-boreoarctic species is well documented (Ekman, 1953;
Zenkevitch, 1963; this study).
Because the North Atlantic Transversal
Ridge (between Greenland, Iceland, and Great Britain) has never been
deeper than at present (average, 500-600 m; maximum,
830
m), only
those bathyal and abyssal bivalves which occur at 560 in or less are
found in both the Arctic and Atlantic Ocean basins (Clarke, 1963).
Similarly, of the. eight described species of bathyal polychaetes found
in this study, six occur at depths of less than 1,000 m in the Atlantic
Ocean.
The remaining two species (Ephesiella macrocirrus, 2,000 to
2,500 m; Allia abranchiata, 1,500 to 2,000 m) are known from only one
study (Hartman, 1965).
49
Although some bathyal bottom invertebrates have planktonic larvae,
most have benthic larvae (Milelkovsky, 1971) and limited dispersal
ability.
Hence, submarine ridges are probably major topographic
barriers for a majority of the bathyaJ. polychaetes.
No data on larval
types for any of the bathyal polychaete species of the western Beaufort
Sea are available to document this hypothesis.
The existence of many endemic bathyal and abyssal species is
explained by geographic isolation.
The Arctic Ocean has not been in
contact with the Pacific or Atlantic Oceans at depths greater than
1,000 m since possibly the Late Cretaceous (see Chapter 3), and
adequate time for speciation has elapsed.
Furthermore, the narrow
limits within which temperature and salinity vary below 250 m (-1°C to
+1°C; 34.92°/
to 34.99°/; Coachman and Aagaard, 1974) are evidence
of relatively stable bathyal and abyssal environments during interglacial intervals.
The magnitude of glacial-interglacial changes in
temperature and salinity are unknown.
By its presence the endemic fauna further reveals that the Arctic
Ocean has not been anoxic in the recent past.
If permanent ice cover
on the Arctic Ocean and Norwegian-Greenland Sea during glacial
intervals (CLIMAP Project Members, 1976) had prevented a flow of
oxygenated water into the Arctic Ocean at depth, extinction of the
bathyal and abyssal faunas would have ensued.
The extant endemic
fauna is too rich to have evolved since the end of the last glacial
interval 10,000 years ago.
No data on the evolutionary rates of
Polychaeta are available, but Paleogene fossils of many living species
of Brachiopoda, Mollusca, and Echinoidea occur in Arctic and boreal
SQ
regions (Durham and MacNeil, 1967).
Evolutionary rates for other
groups of benthic invertebrates are thus too slow for substantial
speciation to have occurred within those groups since the end of the
last glacial interval.
51
V. ECOLOGY OF WESTERN BEAUFORT SEA POLYCHAETA
A. INTRODUCTION
The interpretation of zoogeographic data for Polychaeta and other
benthic taxa has identified factors that probably influence the composition of Arctic Ocean benthic faunas, present and past.
Since species
are not distributed uniformly within a zoogeographic region, the identification of additional factors influencing species distributions on a
smaller scale is possible.
Statistical techniques have been employed
to define distributional patterns inherent in the polychaete fauna of
the western Beaufort Sea, and to identify characteristics of the
physical environment which may, in part, affect such distributional
patterns.
B. RESULTS
De
distributions of the dominant species
A majority of the dominant species of polychaetes are distributed
over a broad depth range from 20 to 1,500 m or greater (Figure 3), but
a few species that occur in either the shallower or deeper portions
of the study area have narrower depth ranges.
The variety of over-
lapping depth distributions shown in Figure 3 is like that encountered
in other studies (Mills, 1969; Curtis, 1972), and indicates that the
dominant polychaetes of the western Beaufort Sea are not grouped into
mutually exclusive assemblages along depth contours.
however, groups exist in a more general sense.
As will be seen,
52
Trends in numerical abundance
Estimates of polychaete abundance are contoured in Figure 4.
Abundance generally increases with increasing depth from the 20 m
contour to the outer continental shelf in the eastern portion of the
study area and to the upper continental slope in the western portion of
the study area.
Abundance decreases below these depths, and very low
abundance is found at the deepest stations (Z>1500 m).
Trends in species richness
Species richness (number of species collected at each station) is
contoured in Figure 5.
are evident.
Trends similar to those for faunal densities
Deeper than 20 in, species richness increases.
It is
greatest along the outer continental shelf and upper continental slope
(40-140 rn), and decreases with increasing depth on the middle and
lower continental slope.
Relatively few species, as well as numbers
of individuals, were collected at depths exceeding 1500 in.
Caution must be exercised when interpreting data collected at the
deepest stations, however.
Our standard sampling procedure of collect-
ing five Smith-McIntyre grab samples at each station may be inadequate
at these depths because of the low numbers sampled; better estimates
of species richness would be obtained with a larger sample size (i.e.,
a 0.25 m2 box corer and replicate cores).
Station affinities
Pearson Product-Moment Correlation Coefficients for the polychaete
I
I
I
I
I
£
000
2,000
0 00
BE4VFORT SEA
7I01\
CAPE
.7
A
A
N\
HALKETT
\
N
N
£
N
/
AN
N
A
')
COLVILLE R.
I
/
KIJPARLJK
A
b
I?
SARTER I
.
.
70'
CANNING
ALASKA
I
I
I
I
152°W
Figure 4.
1500
if
14B°
Polychaeta/m2, contoured (solid lines).
lines) are in meters.
I'b
Depth contours (dashed
I'4'+
£
71°N
700
ALASKA
I
52° W
Figure 5.
I
I
150°
CNN/N&
fi
I
I
148°
146°
Number of species encountered at each station, contoured
lines). Depth contours (dashed lines) are in meters.
44°
(solid
Is,
55
fauna were calculated from the dominant species abundance data as a
measure of station affinity.
The strongest affinities approximately
parallel the depth contours (Figure 6).
Cluster analysis
Station clusters from three, four, five and six cluster analyses
were examined in "discriminant space."
Four clusters were chosen for
further analysis because the orientation of five or more clusters in
"discriminant space" showed a loss of cluster integrity.
Furthermore,
the orientation of five or more clusters in "discriminant space"
reduced the variance captured by the second canonical variable to
uninterpretable levels (<1 percent of variance captured).
Stations that were clustered together in four clusters by CLUSB
were mostly aligned with the depth contours, but east-west trends were
evident (Figure 7).
Stations two (45 m), eight (44 m), and nine
(200 m) on the outer continental shelf and upper continental slope
clustered with the shallowest stations (stations 1, 7, 12, 17, 21;
21-33 m) to form cluster A, rather than clustering with the outer
continental shelf-upper continental slope stations to the east
(stations 13, 14, 18, 19, 22, 23; 47-106 m).
Similarly, the deepest
tations 6, 11, 16; 1,870-2,670 m) clustered with the upper continental
slope stations to the east (stations 20, 24; 463-495 m) to form
cluster D.
The westernmost upper continental slope stations, at which
some of the highest faunal densities occurred, clustered separately
(cluster C).
Thus, in comparison with the eastern portion of the study
area, the western portion had one more cluster and in two other
BE4UFOT SEA
7
CAPE
HALKETI
N
-
-
'ELF P
A
B4
TH
KUP4RIJK P
ALASKA
I52 W
Figure 6.
I5O
148
145
Station affinities estimated by Pearson Product-Moment Correlation Coefficients. All station pairings (solid lines) for which
the correlation coefficient is >.645 (the level at which one or
more station pairings occur for each station) are indicated
(P<.00l).
Ui
a.'
I
I
I
I
D
"-
71°
A
N-
C
D
c
D
BEAUFORT SEA
C
A
MA
7C
152°W
Figure 7.
1500
48°
146°
Four clusters of stations (A, B, C, D) in the western Beaufort
Depth contours are in meters.
Sea.
-4
58
clusters, A arid D, deeper stations were included than would be expected
on the basis of the mean depth for the group.
Discriminant analysis
In Figure 8 the station points are projected onto the plane
described by the first and second canonical variables.
The first and
second variables retained 83 percent and 12 percent of the sample
variance, respectively.
The variables were interpreted by correlating
station positions on the variables with species and environmental data
for each station.
Since the environmental data were not very compre-
hensive, the possible influences of many factors (e.g., temperature,
salinity, and dissolved oxygen at the sediment-water interface,
sediment pore water, and food input) upon species distributions cannot
be addressed.
Significant positive correlations (P<.O1) of station positions on
the first canonical variable were found with 1) the number of polychaete species at each station (species richness), 2) the natural
logarithm of the total abundance of the polychaete fauna at each
station, 3) percentage sand, and 4) percentage gravel in the sediments.
(Correlations of the first variable with depth and longitude were nonsignificant,)
Species significantly correlated with the first variable
predominantly demonstrated significant correlations with percentage
sand or percentage sand and percentage gravel (Table 6).
Highest
abundances of those species were generally found at stations in
cluster B (Figure 7) on the outer continental shelf and upper continental slope (40-140 m).
2O
B
-
C
)
A
:,
-20
-40
Figure 8.
I
I
-20
I
I
0
I
20
40
FIRST CANON/CAL VAR/ABLE
Station clusters plotted on the plane described by the first two canonical variables
Station clusters
which were identified by the canonical analysis of discriminance.
station.
correspond to those given in Figure 7; each point represents one
JI
0
Table 6.
Significant correlations (P<.O1) of the first and second
canonical variables (Figure 8) with dominant poiychaeta
species, log transformed data (ln(x+l)). Significant
correlations (P<.05) of those species with sediment grain
size, arcsin transformed data (arcsin tproportion), are also
included.
gr, gravel; sd, sand; st, silt; ci, clay; ns, no
significant correlations with grain size.
POSITIVE CORRELATIONS, FIRST CANONICAL VARIABLE:
sd
Anaitides groenlandica
sd & gr
Pholoe minuta
sd & gr
Antinoella sarsi
sd
Prionospio Steenstri.lpi
ns
Chaetozone setosa
sd & gr
Scalibregma inflatum
sd & gr
Chorie murmanica
ns
Spiochaetópterus typicus
sd & gr
Heteromastus filiformis
sd &
Sd
Lysippe labiata
Sd
Tharyx ? acutus
sd
Micronephthys minuta
sd & gr
Typosyllis cornuta
sd & gr
ophelina cylindricaudatus
gr
Terebellides stroemi
NEGATIVE CORRELATIONS, FIRST CANONICAL VARIABLE:
ci
2\mage auricula
ci
Ophelina abrachiata
cl
Sigaxnbra tentaculata
POSITIVE CORRELATIONS, SECOND CANONICAL VARIABLE:
sd
Anaitides groenlandica
St
Capitel].a capitata
st
Cistenides hyperborea
st
st
Cossura longocirrata
Sternaspis fossor
61
Station positions on the first variable were negatively correlated
with percentage silt (P<.05) and percentage clay (Pz.O1).
Species
negatively correlated with the first variable were correlated with
percentage clay (Table 6), and were most abundant at stations comprising cluster D (Figure 7).
Station positions on the second variable correlated significantly
with percentage silt (P<.Ol).
(Note that because station positions on
the second variable plotted negatively, correlations were reversed in
sign; for simplicity the signs have been changed in Table 6 and the
text.
Note also that depth and longitude correlated non-significantly
with the second variable.)
The second variable was primarily corre-
lated with species associated with silt (Table 6), with one exception.
Anaitides groenlandica correlates with both the first (P<.O1) and
second (P<.05) variables.
Since this species is predaceous (tlshakov,
1972), unmeasured factors such as prey density may influence its
distribution in such a way that there are correlations with both
variables.
Those species associated with silt and the second canonical
variable (Table 6) were generally most abundant at stations in cluster
A (Figure 7).
C. DISCUSSION ?ND CONCLUSIONS
Longitudinal trends
Species richness changed little from east to west along the depth
contours (Figure 5).
(Note that 53, 56, and 58 species were found at
stations two, three, and eight, respectively, on the outer continental
shelf and upper continental slope in the western portion of the study
62
area.)
Numerical abundance was greatest on the upper continental slope
between 200 and 800 m in the western portion of the study area (Figure
4).
Greater abundances of certain species, particularly Lurribrineris
minuta, Maldane sarsi, Minuspio cirrifera, Micronephthys minuta,
Owenia fusiformis, Scoloplos acutus, Tauberia gracilis, and Tharyx ?
acutus, were largely responsible for increased abundance on the upper
continental slope in the western portion of the study area.
The
highest abundances of both the polychaeeous annelids (Figure 4) and
the total benthic invertebrate fauna (Carey and Ruff, 1977) occurred in
this region.
Decreased species richness and greater abundances of the
species noted above resulted in a less equitable dominance structure
within the polychaete fauna.
Warmer waterE that are rich in particulate organics (Johnson,
1956) periodically intrude into the western Beaufort Sea from the
Bering and Chukchi Seas.
istics (Hufford, 1973).
They may be traced by temperature characterIn the summers of 1971 and 1972 these waters
intruded to approximately 148°W longitude and were characterized by
temperatures above 0°C.
Recent measurements by current meter and STD
(Aagaard, 1978) indicate that the Bering-Chukchi Sea water moves
around Point Barrow in pulses whose direction is influenced by the
local ba.thymetry of the shelf edge.
The similarity between the
abundance pattern of the polychaetous annelids (Figure 4) and the
distributions of Bering-Chukchi Sea water in 1971 and 1972 (Hufford
et al., 1974) further supports the hypothesis of Carey and Ruff (1977)
that organic fallout from these warmer waters enriches the benthic
environment on the upper continental slope to the west of l48°W
63
longitude, and thereby fosters high standing stocks of benthic invertebrates in that region.
Faunal trends and the sedimentaryenvironment
Correlation coefficients for station pairs (Figure 6)
and results
of the cluster analysis of stations (Figure 7), both based on the data
of dominant polychaete species, indicate that station affinities are
strongest parallel to the depth contours.
Similar results were
obtained in earlier studies of the Beaufort Sea fauna when Carey et al.
(1974) calculated indices of affinity (Renkonnen, 1944; Sanders, 1960)
for station pairs using megaepifauna data (organisms caught by a 1.3
cm stretch mesh shrimp trawl net), and when Carey and Ruff (1977)
applied recurrent group analysis (Fager, 1957) to both the megaepifa.una
data and the bivalve mollusc data.
High station affinities parallel to the depth contours appear to
contradict the predominance of broad depth distributions exhibited by
a majority of the dominant species (Figure 3).
However, since species
are not uniformly distributed throughout their ranges, and since
correlation coefficients are sensitive to changes in species abundance,
no contradiction exists.
Minuspio cirrifera, for example, was found at
22 stations throughout the entire depth range of the study area, but it
was very abundant (more than 1,400/zn2) only on the continental slope in
the western portion of the study area (stations four, five, ten).
A general downslope extension of the dominant species occurs in
the western portion of the study area, and results in the crossing of
depth contours by the station clusters in the area.
The crossing of
64
depth contours (Figure 7) can be explained largely on the basis of the
sedimentary environment, which has been shown to influence the distribution and abundance of benthic infaunal species (see Thorson, 1957,
1966; Sanders, 1960).
The seaward sequence of silt-sand-clay is wider
in the western portion of the study area.
When sediment grain size distributions for all of the stations
occupied (except six and seven, where no sediment data are available)
are plotted (Figure 9), it is apparent that stations with similar
sedimentary characteristics are grouped together.
The sediments at
cluster A stations are sandy muds (mud = silt + clay) which have a
relatively uniform proportion of silt (36-48 percent).
Cluster B
stations are characterized by the predominance of coarser grain sizes
(sand and gravel).
Gravel is found only at cluster B stations, and
occurs at all but one station in that group (14, underlined in Figure
9).
Two stations (three, with mixed sediments; 23, with silty clay
sediments) in cluster B contain much less than 50 percent coarse
granules, but their inclusion in cluster B is logical since each
contains gravel and therefore exhibits a relatively high degree of
sedimentary heterogeneity.
Muds with low sand content occur at cluster
C and D stations, which may be distinguished by their differing siltclay ratios.
Higher variability of the grain size proportions at cluster A and
B stations, compared to cluster C and D stations (Figure 9), probably
reflects the greater intensity with which physical processes affect the
sediments on the continental shelf, compared to the continental slope.
Ice rafting, ice gouging, waves, currents, and tides have been shown
65
100%
Sand & Gravel
100% Clay
100% Silt
Figure 9.
Sediment grain size distributions for all
stations except 6 and 7 are plotted following the method outlined by Shepard (1954).
Letter designations indicate the cluster
(Figure 7) to which the station belongs.
Gravel occurs exclusively at cluster B
stations; only station 14 (underlined) in
that cluster contains no gravel.
to influence the composition and morphology of the continental shelf
sediments (Barnes, 1974; Barnes and Reimnitz, 1974; Reixnnitz et al.,
1978).
The clustering of stations with similar sedimentary characteristics indicates that the polychaetes are good discriminators of
sedimentary environments (grain size and/or associated factors), and
that the longitudinal trends observed in the station clusters (Figure
7) are probably related to similar trends in the distribution of
sedimentary environments across the continental shelf and slope of the
western Beaufort Sea.
This relationship may be explained more fully by
reference to a canonical analysis of discriminance
The positive
correlations (P<.Ol) of the first canonical variable with species richness and total polychaete abundance indicate that the stations occupied
may be grouped and ordered primarily by the diversity of the polychaete
fauna at each station.
Diversity is highest at group B stations, found
along the outer continental shelf and upper continental slope.
The
predominantly inner shelf stations in group A exhibit somewhat decreased diversity, while lowest diversities are found at group C and D
stations.
The correlation of faunal diversity with sediment grain size
is obvious when Figures 8and 9 are compared:
diversity decrease concurrently.
grain size and faunal
Positive correlations of the second
canonical variable with silt (P< .01) and with species associated with
silt further indicate that characteristics of the sedimentary environment influence the dIstribution and abundance of the polychaetous
anne1ids.
In summary, polychaete data and sediment data were used to compare
67
stations occupied on the continental shelf and slope.
Four station
groups were generated by cluster analysis of the polychaete species
data.
The clusters were ordered in two-dimensional space by a canon-
ical analysis of discriminance with a minimal loss of variance (Figure
8).
Sediment grain size data were also used to compare stations by
plotting station points on a tertiary sediment diagram (Figure 9).
Correlations of the canonical variables in Figure 8 associated
the greatest proportion of sample variance with grain size, and the
second greatest proportion of sample variance with the silt content of
the sediments.
Examination of the station orientation in Figure 9
revealed an analogous pattern of sample variance:
most variance was
associated with grain size, and the station groups were ordered in the
same sequence as they were along the first canonical variable.
Sample
variance in Figure 9 was also attributed to the group A stations (as
in the second canonical variable), since projection of the station
points onto the silt axis would greatly reduce station point scatter.
As the stations were similarly grouped and ordered by analyses
performed on polychaete species data and sediment data, a dependency of
the polychaetes on the sedimentary environment (grain size and/or
associated factors) is highly probable.
The distribution and abundance
of the dominant polychaetes probably resulted largely from variations
in the sedimentary environment on the continental shelf and slope of
the western Beaufort Sea.
68
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APPENDIX
APPENDIX
Station numbers used in the following discusStations selected for analysis of the polychaete fauna.
United
States
Coast
Guard deisgnations (U.SC.G., Oceanographic
sion and in Figure 1 are followed by the
Beaufort Sea Ecological Cruises of
Report No. CG373-64) for those stations occupied during the Western
SMG,
Smith-McIntyre
grab;
OTB,
otter trawl; BxC, Sandia1971 and 1972. Abbreviations used are:
Beaufort
Sea
Ecological
Cruise
of
1971; WEBSEC-72, Western
Hessler box corer; WEBSEC-71, Western
Environmental Assessment
Beaufort Sea Ecological Cruise of 1972; OCSEAP-4, Outer Continental Shelf
Cruise of Summer,
Cruise of Summer, 1976; OCSEAP-7, Outer Continental Shelf Environmental Assessment
1977.
STATION
NUMBER
STATION
NUMBER
NUMBER
OF
SAMPLES
1
7i
5
SMG
2
72
5
3
75
4
C.G.
SAMPLING
GEAR
DEPTH
INTERVAL
(meters)
DATE
LATITUDE
LONGITUDE
WEBSEC-71
9/9/71
71°04.l'N
151°22.3'W
20-21
SMG
WEBSEC-7l
9/9/71
71°10.ltN
l51°08.9'W
45
5
SMG
WEBSEC-71
10/9/71
71°14.8'N
150°27.6'W
132-140
84
5
SMG
WEBSEC-71
12/9/71
71°18.3'N
150°21.6'W
540-831
5
85
5
SMG
WEBSEC-71
12/9/71
71°22.O'N
150°38.0'W
795-997
6
86
4
SMG
WEBSEC-71
14/9/71
71°45.l'N
l50°35.O'W
2139-2461
7
78
5
SMG
WEBSEC-71
11/9/71
70°58.1'N
149°59l'W
27-28
8
82
5
SMG
WEBSEC-71
11/9/71
71°08.3'N
149°47.7'W
44-45
9
83
5
SMG
WEBSEC-71
11/9/71
71012.2*N
149°44.8'W
169-232
10
58
5
SMG
WEBSEC-71
5/9/71
71°15.2'N
149°28.8'W
603-991
11
57
5
SMG
WEBSEC-71
4/9/71
71°21.O'N
149°26.2'W
1618-1926
CRUISE
CG.
STATION
NUMBER
STATION
NUMBER
DEPTH
INTERVAL
NUMBER
OF
SAMPLES
SAMPLING
GEAR
CRUISE
DATE
LATITUDE
LONGITUDE
(meters)
12
63
5
SMG
WEBSEC-71
7/9/71
70°43.O'N
149°00.O'W
23-24
13
44
5
SMG
WEBSEC-71
31/8/71
71°01.O'N
148°22.7'W
46-48
14
30
5
SMG
WEBSEC-71
30/8/71
71°06.O'N
147°57.O'W
85-111
15
29
5
SMG
WEBSEC-71
29/8/71
71°08.5'N
148°O0.O'W
324-430
16
20
5
SMG
WEBSEC-71
25/8/71
71°13.7'N
147°22.6'W
2295-3010
17
12
5
SMG
WEBSEC-71
22/8/71
70°18..O'N
146°05.O'W
26-27
18
9
5
SMG
WEBSEC-71
22/8/71
70°44.O'N
145°52O'W
57-58
19
8
5
SMG
WEBSEC-71
22/8/71
70°48.5tN
145°56.1'W
81-84
20
7
5
SMG
WEBSEC-71
21/8/71
71°00.5'N
145°35,O'W
447-480
21
1
5
SMG
WEBSEC-71
19/8/71
71°15.5'N
143°36.9'W
32-34
22
3
5
SMG
WEBSEC-71
20/8/71
70°27.O'N
143°34.O'W
48
23
5
5
SMG
WEBSEC-71
20/8/71
70°34.6'N
143°380tW
105-109
24
6
5
SMG
WEBSEC-71
20/8/71
70°45.6'N
143°35.4'W
494-498
25
19
3
SMG
WEBSEC-71
24/8/71
71°00.O'N
147°04.O'W
574-700
26
64
1
OTB
WEBSEC-71
6/9/71
70°43.O'N
149°02.O'W
50
C.G.
STATION
NUMBER
STATION
NUMBER
DEPTH
INTERVAL
NUMBER
OF
SAMPLES
SAMPLING
GEAR
CRUISE
DATE
LATITUDE
LONGITUDE
(meters)
27
66
1
OTB
WEBSEC-71
7/9/71
70°43.O'N
149°06.0'W
105-109
28
1
2
OTB
WEBSEC-72
4/10/72
70°14.1'N
143°23.5'W
28-37
29
2
1
OTB
WEBSEC-72
4/10/72
70°22.9'N
143°30.1'W
51
30
4
1
OTB
WEBSEC-72
5/10/72
70°43.l'N
143°42.8'W
464
31
9
1
OPB
WEBSEC-72
7/10/72
70°34.8tN
144°23.1tW
71
32
10
1
OTB
WEBSEC-72
7/10/72
70°20.O'N
144°40.O'W
41
33
11
1
OTB
WEBSEC-72
8/10/72
70°10.9'N
144°30.5'W
27
34
12
1
OTB
WEBSEC-72
8/10/72
70°18.7'N
145°13.O'W
30
35
15
1
OTB
WEBSEC-72
9/10/72
70°33.O'N
145°40.O1W
50
36
16
1
OTB
WEBSEC-72
9/10/72
70°40..8'N
145°24.9'W
79
37
17
1
OTB
WEBSEC-72
9/10/72
70°51.5'N
145°17.O'W
357
38
24
1
OTB
WEBSEC-72
13/10/72
70°35.1'N
146°35.3'W
48
39
25
1
OTB
WEBSEC-72
14/10/72
70°20.O'N
146°28.O'W
34
40
26
1
OTB
WEBSEC-72
14/10/72
70°21.7'N
146°32.7'W
27
41
28
1
OTB
WEBSEC-72
15/10/72
70°31.5'N
147°32.0'W
29
CoG.
DEPTH
INTERVAL
NUMBER
STATION
NUMBER
OF
SAMPLES
42
36
1
OTB
WEBSEC-72
43
37
1
OTB
44
--
3
45
--
46
STATION
NUMBER
SAMPLING
GEAR
CRUISE
LATITUDE
LONGITUDE
18/10/72
71°11.6'N
148°32.1'W
159
WEBSEC-72
19/10/72
71°05.7'N
148°41.O'W
55
BxC
OCSEAP-4
29/10/76
71°43.6'N
151°46.5'W
1643-1738
1
SMG
OCSEAP-7
9/10/77
72°23.7'N
154°37.2'W
2470
--
1
SMG
OCSEAP-7
10/10/77
72°21.5'N
153°37.O'W
2840
47
--
1
SMG
OCSEAP-7
10/10/77
72°21.2'N
153°45.2'W
2650
48
--
2
SMG
OCSEAP-7
20/10/77
72°53.6'N
146°29.O'W
3750-3841
49
--
5
SMG
OCSEAP-7
21/10/77
72°55.0'N
146°30.O'W
3511-4200
50
--
2
SMG
OCSEAP-7
22/10/77
72°47.O'N
146°23.5'W
3569-3570
51
--
1
SMG
OCSEAP-7
22/10/77
72°42.O'N
143°40.0'W
3386
52
--
1
SMG
OCSEAP-7
23/10/77
72°55.O'N
142°05.O'W
3475
53
--
4
SMG
OCSEAP-7
24/10/77
70°52.O'N
141°41.O'W
1958-2086
54
--
2
SMG
OCSEAP-7
25/10/77
70°40.5'N
141°36.,O'W
997-1097
55
--
5
SMG
OCSEAP-7
25/10/77
70°41.5'N
141°41.5'W
640-686
56
--
1
SMG
OCSEAP-7
25/10/77
70°41.O'N
141°27.O'W
DATE
(meters)
1025
C.G.
STATION
NUMBER
STATION
NUMBER
NUMBER
OF
SAMPLES
SAMPLING
GEAR
DEPTH
INTERVAL
CRUISE
DATE
LATITUDE
LONGITUDE
(meters)
57
--
1
SMG
OCSEAP-7
30/10/77
71°12.O'N
145°35.O'W
2104
58
--
2
SMG
OCSEAP-7
31/10/77
71°05.O'N
146°33.O'W
1144