SARSIA

BENTHIC FAUNA IN THE PECHORA SEA

SALVE DAHLE, STANISLAV G. DENISENKO, NINA V. DENISENKO & SABINE J. COCHRANE

DAHLE, SALVE, STANISLAV G. DENISENKO, NINA V. DENISENKO & SABINE J. COCHRANE 1998 08 28. Benthicfauna in the Pechora Sea. – Sarsia 83:183-210. Bergen. ISSN 0036-4827.

Quantitative samples of benthic fauna were collected at 15 stations for analysis of speciesdistribution and faunal composition. Multivariate statistics indicate that the sampling stationscover a heterogeneous area with different types of community composition, as might beexpected, since the Pechora Sea encompasses a wide range of depths and oceanographicconditions. However, some distinct station groupings were evident, which are considered torepresent different faunal community types. Around the Island of Kolguyev, the fauna wasdominated by sub-surface detritivorous Polychaeta. On the coast of Novaya Zemlya, theChernaya Fjord contained an opportunistic faunal composition. The deep area immediatelysouth of Novaya Zemlya was characterised by high numbers of large, surface deposit feedingPolychaeta. The Kara Strait area showed some spatial variation in faunal characteristics, butgenerally contained high numbers of surface deposit feeding Polychaeta and Crustacea, as wellas encrusting suspensivores on stones. The stations sampled between Kolguyev and the PechoraBay contained large amounts of macrofaunal Foraminifera and detritivorous Polychaeta,while the fauna sampled in the Pechora Bay was typical of northern, low salinity environ-ments. Canonical correspondence analyses indicate that water depth and sediment type play amajor role in structuring the benthic fauna. The distribution of community types described inthis investigation largely follow those outlined in previous Russian investigations, despite theuse of different analytical strategies. This investigation provides the background for anintercomparison of methodologies in faunal analyses.

Salve Dahle & Sabine Cochrane (corresponding author, e-mail: Sabine.Cochrane@akvaplan.niva.no),Akvaplan-niva AS, N-9005 Tromsø, Norway. – Stanislav Denisenko & Nina Denisenko, MurmanskMarine Biological Institute (MMBI), Russian Academy of Sciences, 17 Vladimirskaya Str. 183010Murmansk, Russia.

KEYWORDS: Pechora Sea; benthic fauna; community; environment; CCA.

INTRODUCTION

The Pechora Sea has been the subject of many Russianbenthic faunal investigations, from the end of the 1800sto the present (reviewed by DENISENKO & al. 1995).Current analyses of biogeography regard the PechoraSea as a transitional zone between Boreal and Arcticfauna (ANTIPOVA & al. 1989). Climatic variations,resulting in changes in the relative influences of thedifferent water masses in the Pechora Sea are thought toresult in variations in the distribution and biomass ofsome benthic species (GALKIN 1980). However, thisdoes not profoundly influence the overall biomass(DENISENKO & DENISENKO 1995). The benthic fauna inthe Pechora Sea is documented to be mainly dominatedby suspension feeding bivalves, and in some areas alsoby motile surface deposit feeding Bivalvia, Polychaetaand Echinodermata (KUZNETZOV 1970). The areas southof Novaya Zemlya and around the Kara Strait areamongst those with the highest biomass (ANTIPOVA 1975;DENISENKO & al. 1997).

Since co-operation and exchange of information be-

tween Russian and other international scientific insti-tutes has increased in recent years, the question ofintercomparability of data has arisen. Faunal analysesin Arctic areas have been carried out by Russian scien-tists for many decades (reviewed by GALKIN 1979), gen-erally describing ‘biocenoses’, or community types,using both biotic and abiotic data (MÖBIUS 1877). Inmost other countries, the benthic analyses have evolvedfrom Petersen’s concept of the faunal community, us-ing species abundance and diversity (PETERSEN 1914).The former analyses use biomass as the main criterionfor describing dominant species, while the latter inves-tigations usually rely on numerical abundance. Our sur-vey carried out in 1992 aimed to initiate a programme ofintercomparison and standardisation of methodologies,with parallel biological samples taken independentlyby both the Norwegian and Russian teams, using theirown sample collection and processing techniques.

As part of a Norwegian-Russian expedition to thePechora Sea in the south-eastern part of the BarentsSea in July 1992, samples of benthic fauna were col-lected for analysis of species distribution and faunal

184 Sarsia 83:183-210 – 1998

Fig. 1. Location of stations sampled for analysis of benthic fauna in the Pechora Sea. Bathymetry in meters (adapted from LORING

& al.1995).

composition. Sediment samples were also taken forcontaminant analysis, including heavy metals, pesticides,other organic hydrocarbons and radionuclides. Theinformation gathered is incorporated into the ArcticMonitoring and Assessment Programme (AMAP)database. The data on contaminants in sediment havebeen reported in LORING & al. (1995) and SMITH & al.(1995).

The present paper primarily aims to further currentunderstanding of the nature of the benthic fauna in se-lected areas of the Pechora Sea. The results of this in-vestigation using a van Veen grab (VAN VEEN 1933) willalso serve as the background for future inter-compari-son of results with those from the Russian survey, us-ing an Ocean grab (LISITSIN & UDINTSEV 1955).

Study areaThe Pechora Sea is bordered by the Russian mainland tothe south, the Islands Vaygach to the east, NovayaZemlya to the north and Kolguyev to the west (Fig. 1).

Almost the entire Pechora Sea area is ice-covered fromNovember until late June, although coastal polynyasare semi-permanent features in many areas. The extentof sea-ice cover can vary considerably from year toyear, according to the inter-annual dynamics of inflowingAtlantic water as well as air temperature and wind char-acteristics (MIDTTUN & LOENG 1987; MATISHOV & al.1993). This inter-annual variation may significantlyaffect physical and biological processes at the sediment-water interface (WASSMANN & SLAGSTAD 1991).

The Pechora Sea acts as a mixing area for differentwater masses: Atlantic water, water from the Kara andWhite Seas, as well as coastal run-off. The water col-umn is strongly stratified in the deep northern parts ofthe Pechora Sea. The bottom water (below 150 m) ismade up of well oxygenated Atlantic water with salinitiesof 34.5-34.95 and temperatures between –1.0 and –1.5 °C (MATISHOV 1992). Through the northern part ofthe Kara Strait, the narrow Litke Current flows in awesterly direction along the coast of Novaya Zemlya

Dahle & al. – Benthic fauna in the Pechora Sea 185

Fig. 2. Distribution of principal sediment types in the Pechora Sea, showing the sampling stations from this study (based on datafrom KLENOVA 1960 and VINOGRADOVA & LITVIN 1960).

(PAVLOV & PHIRMAN 1995). The central part of thePechora Sea is influenced by the Kolguyev-Pechoracurrent, which is primarily of Atlantic origin, and ischaracterised by a salinity of around 34 and tempera-tures between 0.5 and 2 °C. This current flows in aneasterly direction and, mixed with southern Pechora Seawater, enters the Kara Sea through the southern part ofthe Kara Strait (PHIRMAN & al. 1995). The southern partof the Pechora Sea receives a large, seasonally variablecoastal run-off, mainly from the White Sea and thePechora River (ZENKEWICH 1963). In summer and au-tumn, the southern Pechora Sea is dominated by rela-tively warm, low-salinity water (5-8 °C, 18-26 ‰),which largely flows eastwards and into the Kara Sea,through the strait south-east of Vaygach, or northwardsalong the western coast of Vaygach (MATISHOV 1992;MATISHOV & al. 1993; PHIRMAN & al. 1995).

The Pechora Bay is strongly affected by the approxi-mately 1 800 km long Pechora River, which annually

discharges around 130 km3 of freshwater into the PechoraSea, with an estimated annual sediment load of 6.1 × 106

tonnes (MILLIMAN & SYVITSKI 1992). The transportedfine-grained riverine sediments are either trapped in thePechora Bay or transported by offshore currents to bedeposited slowly in the deeper parts of the Barents Sea(ZENKEWICH 1927). The salinity of the Pechora Bay wa-ter varies between 8 and 18

and the summer tempera-

tures are generally between 9 and 12 °C. ChernayaFjord, located on the southern coast of Novaya Zemlya,is 10-12 km long and 3-5 km wide. The sill depth at themouth of the fjord is approximately 15 m and themaximum basin depth is 80 m. The basin water is ofAtlantic origin with a salinity of around 35.0 and atemperature close to –1 °C (MATISHOV & al. 1993).

The heterogeneous surface sediments in the PechoraSea are shown in Fig. 2. The southern parts are charac-terised by sandy sediments, while mixed sedimentsdominate the Kara Strait area. The deep area south of

186 Sarsia 83:183-210 – 1998

species data base. Samples within the dissimilarity matricesgenerated by the Bray-Curtis index (CHEKANOWSKY 1909;BRAY & CURTIS 1957) were grouped together on the basis oftheir resemblances, using the unweighted pair-group averagemethod (ROHLF 1989). Multi-dimensional scaling (MDS)ordination was used to scale the dissimilarity of n samplesin two-dimensional space, placing the most similar objectsclosest together. A preliminary Principal Co-ordinate(PCoA) ordination using double-centred eigenvectorcalculations and a Principal Component Analysis (PCA)was carried out to achieve an optimised and more effectiveMDS outcome (ROHLF 1989).

Canonical correspondence analysis (CCA) was used toassess the relationship between species abundance and thephysical and chemical characteristics of the sediment. Theprinciples of CCA are explained in FIELER & al. 1994. Con-sidered geometrically, each species can be thought of as apoint in the multidimensional space defined by the stations,and each species is given a weight, or ‘mass’ proportional tothe overall abundance of the species (GREENACRE 1984, 1993).Similarly, each station represents a point in the multidimen-sional space defined by the species and receives a mass pro-portional to the number of individuals counted at that sta-tion. Dispersion is defined as the weighted sum-of-squareddistances of the species points (or, equivalently, of the sta-tion points) to their average. This dispersion is termed iner-tia, which is a measure of variance. Species with most inertiaexplained by the first two or three axes are considered to bemost influenced in their distribution by the selected environ-mental variables. Using one of the environmental variablesas a co-variable removes all inertia attributed to that vari-able. Examination of the remaining inertia gives informa-tion on the relationship between species distribution and theother environmental variables.

Based on a preliminary PCA, the following parameters weredesignated as environmental variables and chosen for CCA:depth, % fine sediment (< 63 µm), total organic carbon (TOC),total nitrogen (TN), the radioactive isotopes 239Pu, 240Pu and134Cs, as well as the metals Mn, Fe, Cd, Cr, Pb, Ni, V, Zn, Al

Novaya Zemlya is characterised by very fine mud, whilethe muddy sediments in the central and northern partscontain an admixture of sand and gravel. Underlying thesurface sediments in the Pechora Sea is dense glacio-marine blue-grey clay, particularly in the central, west-ern, northern and north-eastern parts. Table 1 showsthe basic physical characteristics of the surfacesediments at the stations sampled.

MATERIALS AND METHODS

Sampling and laboratory proceduresSampling was carried out from the research vessel R/V DalnieZelentsie, MMBI, at 15 stations (Fig. 1). Five replicateswere taken at each station. Station positioning was carriedout using the ship’s radar, supported by GPS (GlobalPositioning System). A 0.1 m2 lead weighted van Veen grabwith hinged, lockable, rubber-covered inspection flaps of0.5 mm mesh was used. Samples showing inadequate oruneven penetration, or a disturbed sediment/water interfacewere rejected. The samples were gently washed through acircular 1 mm diameter round-mesh screen immersed inrunning sea water, and fixed in 15-20 % borax-bufferedformalin. For glacio-marine clay sediments, the fine surfacesediment was first gently washed from the clay, which wasthen processed separately. Samples were rinsed in thelaboratory using 1 mm round mesh sieves immersed in run-ning fresh-water to remove formalin. Animals were sortedfrom the sediment into phyla and subsequently identifiedto species or lowest taxonomic level possible. A referencecollection was kept of all species identified.

Numerical analysesThe replicate sample data were compiled and then summedfor each taxon to give faunal densities for each station (0.5 m2).The community analyses were based on a 2-way station by

Table 1. Background characteristics of the surface sediments sampled: water depth, sediment type, grain size (< 63 µm, %by weight), % total organic carbon (TOC), % total nitrogen (TN) and TOC/TN ratios (data from LORING & al. 1995).

Stn Depth Position Surface sediment type % % % TOC(m) Latitude Longitude < 63 µm TOC T N /TN

3 53 68°34.20'N 49°59.06'E very sandy mud, greyish 51 0.67 < 0.10 -6 88 69°38.36'N 50°45.18'E very sandy mud, greyish 57 0.81 0.12 6.87 76 70°08.83'N 53°24.28'E muddy sand, greyish 12 0.19 < 0.10 -8 193 70°30.98'N 54°38.56'E sandy mud, brownish 84 1.92 0.29 6.6

11 68 70°42.44'N 54°38.49'E sandy mud, brownish 79 1.65 0.23 7.212 188 70°17.00'N 55°36.40'E muddy sand, brownish 81 2.01 0.28 7.213 207 70°24.28'N 55°07.43'E sandy mud, brownish 91 2.00 0.29 6.914 172 70°13.86'N 55°02.49'E sandy mud, brownish 86 1.70 0.24 7.119 83 70°10.36'N 57°12.51'E muddy sand, brownish 26 0.45 < 0.10 -20 126 70°16.56'N 57°32.58'E very sandy mud, brownish 40 0.78 0.13 6.021 85 70°11.69'N 58°08.68'E muddy sand, brownish 9 0.38 < 0.10 -24 16 69°21.08'N 58°56.58'E very sandy mud, greyish 5 0.10 < 0.10 -26 17 69°14.66'N 57°09.00'E sand, greyish 2 < 0.10 < 0.10 -27 8 69°00.21'N 56°01.35'E sand, brownish 2 < 0.10 < 0.10 -29 11 68°35.31'N 55°13.49'E sandy mud, greyish 75 0.96 < 0.10 -

Dahle & al. – Benthic fauna in the Pechora Sea 187

using summary station data. The brackish Station 29was sufficiently unique in its faunal composition towarrant its removal from the MDS analyses, to avoidobscuring the spatial arrangement of the other stations.Goodness of fit (stress) analyses (ROHLF 1989) showedan excellent agreement between the Bray-Curtis indicesand the MDS ordinations. Based on these analyses,combined with an evaluation of the biological, physical,and chemical characteristics, the sampling stations weredivided into seven main groups, each of which showedat least 60 % dissimilarity from each other. These groupsare considered to represent discrete faunal associations,and are herewith referred to as Faunal Associations A-G (Table 2).

Number of individuals, taxa and faunal diversity indi-ces.A total of 16 phyla, 17 classes and 53 taxonomic orderswere recorded. The best represented phyla, in terms ofnumbers of individuals, were the Annelida, Bryozoa,Crustacea, Echinodermata, Mollusca, andSarcomastigophora (a single species of macrobenthicForaminifera). The numbers of individuals and taxa re-corded from each of the sampling stations are shown inTable 3. A full species list is given in the Appendix.

Fig. 5 shows the mean numbers of taxa and individu-

Table 2. Grouping of stations into areas.

Faunal Location Stations Mean depthassociation included (m)

A Proximity to Kolguyev Island 3, 6 54B Chernaya Fjord, Novaya Zemlya 11 79C Deep area south of Novaya Zemlya 8, 12, 13, 14 190D Northern Pechora Sea and Kara Strait 7, 19, 20, 21 91.5E Dolgiy Island 24 16F South-eastern Pechora Sea 26, 27 12.5G Pechora Bay 29 11

Fig. 4. Two-dimensional MDS scaling plot, showing the inter-relationships between the sampling stations, using summaryfaunal data. The brackish Station 29 is omitted. Both axes rep-resent scaled dispersion.

Fig. 3. Cluster diagram, showing the grouping of stations, basedon percent dissimilarity of summary faunal data.

and Li. The radioactive isotopes and metals are recommendedas essential for environmental monitoring (AMAP 1993). Theselected log-transformed environmental variables, togetherwith the untransformed faunal data were directly entered intothe CCA, and those linear combinations of environmental vari-ables that maximise the dispersion of the species scores (i.e.those which explain most of the species variables) were se-lected on the basis of multiple regression analyses (‘forwardselection’). The CANOCO software package was used (TER

BRAAK 1987-1992). The results from the ordinations were plot-ted using the software package CANODRAW (SMILAUR 1992).

RESULTS AND DISCUSSION

Station groupingsCluster grouping showed between 30 and 50 % dissimi-larity between individual replicates, indicating a highdegree of variability in the benthic fauna across the sam-pling area, with the exception of the Pechora Bay sam-ples, which showed lower dissimilarities. Despite this,replicates generally clustered according to sampling sta-tion. Fig. 3 shows the cluster groupings at the stationlevel. MDS analyses also showed a relatively high de-gree of dissimilarity between individual replicates, butthese mostly grouped according to station.Fig. 4 shows the two-dimensional MDS plot obtained

188 Sarsia 83:183-210 – 1998

4.54 to 5.84 (Table 2) and lowest at the brackish FaunalAssociation G (Pechora Bay), with a value of 2.2. Thisvariation in faunal diversity across the sampling area isthought to reflect natural variations in community struc-tures, as a result of the heterogeneous bottom condi-tions. Since some taxonomic groups were more easilyidentified to species level than others, such diversityindices should be interpreted with care (see WU 1982),but do provide a useful across-field comparison.

Dominant speciesTable 4 lists the five numerically dominant taxa withinthe Faunal Associations, as well as the estimated numberof individuals per m² and the percentage contribution ofeach species to the population sampled. The dominantspecies were ranked according to the number of timesthe species were amongst the five numerically dominanttaxa. Thus, species present in high numbers at only oneor a few sampling stations within the faunal associationare ranked lower than those which are dominant at allthe stations, albeit in lower numbers. This avoids plac-ing emphasis on species which are not representative ofthe faunal assemblage as a whole.

Faunal Association A (Kolguyev) was dominated bythe tube-dwelling sub-surface detritivore Maldane sarsi(Polychaeta). The next dominant Chaetozone setosa(Polychaeta), also a burrower, is a selective surface de-posit feeder, using long palps to collect particles at or

Table 3. Numbers of individuals and taxa recorded from thestations sampled, together with the abundance ratio (A/S - no.individuals / no. species present) and the Shannon-Wiener (H’)diversity index.

Faunal St. no. no. A0.5/S0.5 Shannon-association no. individs. taxa Wiener

A0.5* S0.5* H’ index

A 3 943 80 12 4.636 843 103 8 5.16

B 11 1657 61 27 4.18

C 8 375 64 6 4.5612 737 67 11 3.1013 968 57 17 3.6114 731 74 10 4.38

D 7 1381 159 9 5.8419 1134 99 11 4.8820 1070 140 8 5.4021 3413 197 17 5.39

E 24 1475 91 16 4.54

F 26 747 70 11 5.0927 794 45 18 2.48

G 29 434 11 39 2.19

* Sum of five 0.1 m2 replicates

Fig. 5. Graphic representation of the distribution of A: indi-viduals and B: taxa between the faunal associations found inthe Pechora Sea, 1992. The associations are arranged in orderof increasing depth, from the shallow estuarine Faunal Asso-ciation G in the Pechora Bay to Association C in the deep areasouth of Novaya Zemlya. The category ‘others’ includesBrachiopoda, Chelicerata, Chordata, Cnidaria, Echiurida,Nematoda, Nemertini, Priapulida and Sipunculida.

als of the major taxonomic groups recorded at the faunalassociations, arranged in order of increasing depth. Thebrackish Faunal Association G contained relatively lownumbers of both taxa and individuals, while Faunal As-sociations F (South Pechora Sea coast) and E (DolgiyIsland) contained large numbers of individuals ofSarcomastigophora. In Faunal Associations A(Kolguyev), B (Chernaya Fjord) and particularly C(deep area south of Novaya Zemlya), the Annelida com-prised less than half the recorded number of taxa, but amuch greater proportion of the recorded number of in-dividuals, suggesting the presence of a few numericallyover-abundant taxa within the Polychaeta. The relativelyhigh numbers of taxa and individuals in Faunal Associa-tion D possibly reflects the mixed bottom sedimenttype present, with its large variety of biological niches,as well as the high current velocity turbulence in theKara Strait, which is thought to result in highly produc-tive waters.

The Shannon-Wiener (H’) diversity index was high-est at Faunal Association D (Northern Pechora Sea andKara Strait), with individual station values ranging from

NO

. OF

IND

IVID

UAL

S

0

500

1000

1500

2000

2500

G F E A B D C

Annelida

BryozoaMollusca

CrustaceaOthers

SarcomastigophoraEchinodermata

A

(0.5

m2 )

FAUNAL ASSOCIATION

NO

. OF

TAXA

0

50

100

150

200

G F E A B D C

Annelida

Bryozoa

Mollusca

Crustacea

OthersEchinodermata

(11 m) (13 m) (16 m) (92 m) (190 m)(79 m)(54 m)

(0.5

m2 )

B

Dahle & al. – Benthic fauna in the Pechora Sea 189

just below the sediment surface. The third dominanttaxon Lumbrineris spp. (Polychaeta), is an active car-nivore and scavenger, equipped with strong jaws.Pectinaria hyperborea (Polychaeta) inhabits a hardconical tube constructed of sand grains, and adopts ahead-down position, moving through the flocculentsurface sediment. Nuculoma tenuis (Bivalvia) was alsodominant in this faunal association, inhabiting the fine,flocculent surface sediment layers.

Faunal Association B (Chernaya Fjord) contained largenumbers of Chaetozone setosa and Thyasira sp.(Bivalvia), both of which dwell in the flocculent sedi-ment surface layers. Maldane sarsi and Spiochaetopterus

Table 4. Listing of the five most abundant taxa sampled at the five station groups (A-E) sampled in the Pechora Sea, 1992. No.indicates the calculated number of individuals per m2 at the faunal association, while % shows the percentage of all individualsrecorded from the faunal association represented by the species.

Faunal Rank SpeciesAssociation Order/Class No.* %**

A 1 Maldane sarsi Polychaeta 370 20.722 Chaetozone setosa Polychaeta 80 4.482 Lumbrineris spp. Polychaeta 80 4.484 Pectinaria hyperborea Polychaeta 161 9.014 Nuculoma tenuis Bivalvia 77 4.31

B 1 Chaetozone setosa Polychaeta 750 22.632 Maldane sarsi Polychaeta 346 10.443 Thyasira spp. Bivalvia 296 8.934 Spiochaetopterus typicus Polychaeta 226 6.825 Hyperammina subnodosa Foraminifera 210 6.34

C 1 Spiochaetopterus typicus Polychaeta 452 32.152 Lumbrineris spp. Polychaeta 211 15.013 Maldane sarsi Polychaeta 96 6.834 Terebellides stroemi Polychaeta 43 3.065 Paraonis gracilis Polychaeta 59 4.20

D 1 Cirratulidae indet. Polychaeta 142 4.062 Ophiura robusta Ophiuroidea 88 2.522 Spiochaetopterus typicus Polychaeta 318 9.094 Chaetozone setosa Polychaeta 96 2.745 Byblis gaimardi Amphipoda 336 9.60

E 1 Hyperammina subnodosa Foraminifera 632 21.692 Myriochele fragilis Polychaeta 512 17.573 Euclymeninae indet. Polychaeta 222 7.624 Stegophiura nodosa Ophiuroidea 168 5.775 Owenia fusiformis Polychaeta 130 4.46

F 1 Stegophiura nodosa Ophiuroidea 164 10.642 Ophelia limacina Polychaeta 90 5.842 Hyperammina subnodosa Foraminifera 539 34.984 Leitoscoloplos sp. Polychaeta 65 4.225 Myriochele fragilis Polychaeta 58 3.76

G 1 Pontoporeia femorata Amphipoda 330 38.022 Macoma balthica Bivalvia 302 34.793 Halicryptus spinulosus Priapulida 82 9.454 Diastylis rathkei Cumacea 58 6.685 Marenzelleria sp. Polychaeta 54 6.22

typicus (Polychaeta), the latter secreting a horny tubeand feeding from deposited or near bottom suspendedmaterial (BARNES 1963; KUZNETSOV 1970), were alsopresent in large numbers, buried deep within the under-lying glacio-marine clay. Nuculoma tenuis was alsoamong the dominant species. There was a notable lackof Echinodermata in this faunal assemblage, which wererepresented by only a single individual of Ophiacanthabidentata. Interestingly, investigations around Svalbardalso showed a numerical as well as biomass dominanceof Chaetozone setosa , Maldane sarsi, andSpiochaetopterus typicus (LEYBSON 1939; KENDALL &ASCHAN 1993; HOLTE & al. 1996).

190 Sarsia 83:183-210 – 1998

several genera including Aphelochaeta, and were notfurther identified due to taxonomic difficulties. Ophiurarobusta (Echinodermata) is common in the Arctic(D’YAKONOV 1954) and actively moves over hard sub-stratum, feeding on deposited organic material(KUZNETZOV 1970). In common with other stations inthe Pechora Sea where there is underlying glacio-marineclay, Spiochaetopterus typicus is present in highnumbers. Chaetozone setosa and Byblis gaimardi(Crustacea) were also dominant within this faunal asso-ciation, the latter thought to be at least partially reliantupon suspension feeding.

The fauna in shallow sandy sediments sampled inFaunal Association E (Dolgiy Island) was dominatedby the macrobenthic Hyperammina subnodosa(Foraminifera), but the precise role of these animals inthe community is still unclear. Myriochele fragilis(Polychaeta) and Owenia fusiformis (Polychaeta), bothof the family Owenidae, were also present in high num-bers, the latter species being known to utilise both sur-face deposit and suspension feeding. Unidentified mem-bers of the tube-building family Euclymeninae

Fig. 6. Benthic biocenoses, or communities in the Pechora Sea (adapted from ZENKEWICH 1927).

The deep-water Faunal Association C (SouthernNovaya Zemlya) was dominated bySpiochaetopterus typicus, Lumbrineris spp.,Maldane sarsi, Terebellides stroemi (Polychaeta) andParaonis gracilis (Polychaeta). The dominance of thecarnivorous Lumbrineris spp. in both faunal assemblagesB and C indicates a certain similarity in conditions.Terebellides stroemi inhabits a vertically oriented bur-row, selectively feeding from surface deposits by meansof numerous tentacles, while the feeding mode of themuch smaller Paraonis gracilis, which buries horizon-tally in the flocculent surface layers, is somewhat ob-scure (FAUCHALD & JUMARS 1979).

Faunal Association D contained a large number of theencrusting Balanus sp. (Crustacea). The majority of theindividuals were small and mainly concentrated in a sin-gle replicate, indicating a patchy distribution of newlysettled individuals not representative of adultpopulations in the area. The taxon was therefore ex-cluded from the statistical analyses. The burrowing sur-face deposit feeding family Cirratulidae (Polychaeta)was also abundant. These were small individuals, of

Dahle & al. – Benthic fauna in the Pechora Sea 191

(Polychaeta) were also abundant. It should be notedthat Stegophiura nodosa, which was present in largenumbers, is the only taxon of Echinodermata representedwithin this faunal association.

Faunal Association F (South-eastern Pechora Sea),also a sandy area, was dominated byStegophiura nodosa, which was also common withinFaunal Association G. Hyperammina subnodosa waspresent in high numbers, but only at one of the twostations within the faunal association. Ophelia borealis(Polychaeta), a non-selective surface deposit feeder in-habiting sandy sediments (HARTMANN-SCHROEDER 1971),was also common in this area, together withLeitoscoloplos sp. (Polychaeta), also thought to be anon-selective deposit feeder and active burrower. Theprecise feeding strategy of Myriochele fragilis is un-clear, but it is suggested that these worms selectivelyfeed from near-surface sedimentary material (FAUCHALD

& JUMARS 1979).Faunal Association G (Pechora Bay) contained a fauna

typical of low salinity environments, low in both num-bers of individuals and taxa. Pontoporeia femorata(Crustacea) is well documented from Arctic estuaries(LINDSTRÖM 1992), as is Macoma balthica (Mollusca)(ZENKEWICH 1927). Halycryptus spinulosus (Priapulida)was also abundant at this faunal association. In commonwith benthic investigations in the Rivers Ob and Yenisey(COCHRANE & al. 1997), Diastylis rathkei (Crustacea) andMarenzelleria sp. (Polychaeta) were also among the fivemost abundant taxa sampled in the Pechora Bay. Thisfaunal assemblage is very reminiscent of that found in theBaltic Sea and the Gulf of Finland (ANDERSIN & al. 1978;ANDERSIN & SANDLER 1991).

Six different faunal community types, or biocenoses,have previously been identified in the Pechora Sea(ZENKEWICH 1927; BROTSKAYA & ZENKEWICH 1939). Fig.6 shows the distribution of these biocenoses and thedominant species. It should again be noted that, whileZENKEWICH described biocenoses using species biomassas the main criterion, the present investigation usesnumerical abundance to describe faunal associations.Although the small number of stations sampled in thepresent investigation is not sufficient to represent thePechora Sea benthic fauna as a whole, the faunal as-semblages described here show similarities with thedistribution of the previously described biocenoses,or faunal community types. East of Kolguyev, thedominant fauna included Pectinaria hyperborea, inboth this 1992 investigation and that carried out in1927. The deep area towards the south of NovayaZemlya comprised a separate faunal group in bothinvestigations, both of which document a dominance ofMaldane sarsi. Both studies document an abundance ofSpiochaetopterus typicus in the area extending from the

Fig. 7. Canonical correspondence analysis showing the com-bined plot obtained using all species and station data, togetherwith the four significant environmental variables. 33.6 % ofthe total inertia in species distribution, and 64.8 % of the rela-tionship between species distribution and environmental vari-ables were explained by axes 1 and 2.

northern part of the Pechora Sea and westwards to-wards the Kara Strait. The fauna east of Dolgiy Island isconsidered different to that west of the island in bothstudies, although the dominant species differ betweenthe two investigations, possibly due to the differentmethodologies used.

Thus, despite the differences in analytical method-ologies and the passage of almost 70 years betweenthe present investigation and that of ZENKEWICH (1927),the general trends recorded in the benthic fauna aresimilar. This is not unexpected, as the faunalcommunities appear to be strongly influenced byphysical characteristics in the area. It is encouragingthat, despite large differences in methodologies, thefaunal trends identified in this survey are in generalagreement with those previously described. In thesequel to this article, it will be possible to make firmercomparisons, as the analyses are based on identicalsampling stations and times.

Abiotic factors influencing benthic community structure.To test the relationship between biological and physi-cal characteristics, various correspondence analyseswere carried out on the species data and selected envi-ronmental parameters. Oceanographic data was not used,as information was only available for summer conditions.As for MDS, the brackish Station 29 was removed fromthe analyses.

PCA was first carried out to identify closely corre-lated environmental variables. As might be expected,

192 Sarsia 83:183-210 – 1998

mud (defined as sediment granules less than 63 µm indiameter) was closely correlated with TOC, as well asthe metals Cd, Cr, Pb, Ni, Zn, Al, and Li. Thus, in thesubsequent analyses, the term ‘mud’ represents all thesevariables. TN was closely correlated with iron content,thus these two variables are represented under the headingFe. Depth and Mn are independent variables. Fe wasbest accounted for on Axes 1 and 3, and will not beconsidered further on Axis 2.

CCA was carried out using the full species matrix,combined with depth, ‘mud’, Mn and Fe (Fig. 7). Al-most 52 % of the species data is explained by the firstfour axes. This plot separated the shallow sandy Sta-tions 24, 26, and 27 on the first axes, as being inverselycorrelated with both depth and mud. On the secondaxis, Station 21 was separated as before, but was alsocorrelated with a high Mn content. Stations 8, 12, 13,and 14 were correlated with both depth and mud, whileStations 3, 6, and 20 showed an intermediate relation tomud. Species with more than 50 % inertia explained byaxes 1 and 2, and most strongly associated with Sta-tions 24, 26, and 27 include Ophelina borealis andStegophiura nodosa (Fig. 8). These species are there-fore inversely correlated with muddy sediments andwater depth. Maldane sarsi showed an affinity formuddy sediments and also, to some extent, depth. Thespecies most strongly and uniquely associated with Sta-tion 21 include members of the encrustingsuspensivorous Bryozoa. Using depth as a co-variablegave the same significant environmental variables aswithout a co-variable. The inertia, or variance, of Mn

Fig. 8. Detail of CCA combined plot, showing species withmore than 50 % inertia explained by the first two axes, to-gether with the five numerically most abundant species at eachof the faunal associations. See text for abbreviations.

Fig. 9. Combined CCA plot with depth as a co-variable, show-ing stations (open circles) together with species with more than50 % inertia explained by the first two axes (solid circles). In-ertia and species-environment relationship as Fig. 6.

was not accounted for by Axes 1 and 2 to the sameextent as the other variables, so care should be takenwhen interpreting the contribution of Mn in commu-nity composition.

Since sediment granulometry, and therefore also me-tallic content, may be to a certain degree depth-related,care should be taken to avoid ‘false associations’ be-tween variables. However, certain patterns were evi-dent when removing depth as a variable for the CCA’s(Fig. 9). Only Stations 21 and 11 had strongly associ-ated species, and Station 21 still appeared inverselycorrelated with muddy sediment. Stations 8, 12, 13, 14,19, and 24 are clumped together, without any closelyassociated species, while the remainder of the stationsoccupied scattered positions along the second axis. Thisgrouping of stations of differing granulometric compo-sition indicates that mud content is not the major or soleinfluential factor. Examination of the third axis (notshown) removed Stations 19 and 24 from this group,leaving Stations 8, 12, 13, and 14 in a discrete group,precisely those stations which form Faunal AssociationC. On the third axis, Stations 8, 12, 13, and 14 wereassociated with iron, and therefore also nitrogen-richsediments, but otherwise the cause of this grouping isnot entirely clear. Of the environmental variables ana-lysed, Stations 24, 26, and 27 along the southern coastof the Pechora Sea appeared to be most influenced bywater depth.

In conclusion, the CCA’s indicate that depth and, toa lesser extent, sediment grain size profoundly influ-ence the structure of the benthic communities, although

Dahle & al. – Benthic fauna in the Pechora Sea 193

Stations 11 and 21 appeared to be influenced by otherfactors. This is likely to reflect the peculiarities of thesesites, the former occupying the inner part of a sill fjord,while the latter is located in a rocky area with a highcurrent velocity. This is also indicated by the speciesassociated with these stations, with opportunistic spe-cies such as Thyasira sarsi and Capitella capitata(Polychaeta), which are often associated with a certaindegree of environmental stress (PEARSON & ROSENBERG

1978; PEARSON & al 1982, 1983; PEARSON & al. 1995),strongly associated with Station 11 (Association B),and a variety of suspensivores strongly associated withStation 21.

By analogy with the known habitats of other Lucinacea(DANDO & al. 1985), Thyasira sarsi may inhabit theredox interface between oxic and anoxic sedimentaryconditions, utilising symbiotic sulphate reducingbacteria. In Chernaya Fjord, this is likely to reflect alow bottom water exchange rate, due to the shallow sillat the mouth of the fjord. The stations in the deep areasouth of Novaya Zemlya (Association C) appearedstrongly inter-related, mainly by depth, but possiblyalso by a combination of other variables. Within FaunalAssociation D (Northern Pechora Sea & Kara Strait),there was considerable variation in both dominant andstrongly associated species between the individual sta-tions, which is likely to be due to heterogeneous sedi-mentary conditions. This may not be fully evident fromthe sediment surface granulometry data presented inTable 1, as these samples were taken for contaminantanalyses and excluded larger stones or rocks. Qualita-tive assessment of the samples noted the presence oflarge stones covered by encrusting organisms, with in-terstitial fine sediment, inhabited by the smaller, soft-bodied animals. This uneven substratum is likely togive rise to a patchy faunal distribution.The specimensof Maldane sarsi and Spiochaetopterus typicus found inthis study were often buried deep within the glacio-marine clay which is present over large parts of thestudy area, thus the distribution of these species maybe related to sedimentary conditions. The strong influenceof bottom topography and sediment composition onbenthic community structure in Arctic areas is a welldocumented phenomenon (see GREBMEIER & al. 1989).

Thus it is suggested that the benthic fauna in thePechora Sea is strongly influenced by abiotic factors,such as depth, water masses, temperature, salinity, cur-rent speed, and sediment granulometry. Since there islittle evidence of significant levels of metal or organicmicro-contaminants in the sediments sampled (LORING

& al. 1995), it is suggested that natural physical condi-tions in the area most influence the benthic communitystructure.

Taxonomic difficulties and literature availabilitySome difficulties were experienced in species identifica-tion of the Pechora Sea material. The taxonomic knowl-edge of certain species, or even families is at best in-complete, and several new or little known species maybe present. For example, the Lumbrineris (Polychaeta:Lumbrineridae) specimens collected did not entirely con-form to the widely available species descriptions. Afitting description exists of a species not subsequentlyreported, highlighting the need to research the lesser-known literature. Also, examination of feature variabil-ity in this taxon suggests that the diagnostic charactersshould be revised (E. Oug, pers. commn). Similarly, thespecimens of Chone (Polychaeta: Sabellidae) found inthis study also warrant further taxonomic study. Muchof the taxonomic literature of relevance to the PechoraSea is published in Russian, and has until recently beendifficult to access for non-Russian readers. Some concernhas arisen that ‘double descriptions’ may have occurred,as a result of Russian and other taxonomists working inisolation of each other. Such gaps in knowledge may intime be filled through continued co-operation betweentaxonomists. It is hoped that the full species list presentedin the Appendix will provide a basis for such discussions.

ABBREVIATIONS

Amp_in: Ampharetidae indet., Axi_or: Axinopsidaorbiculata, Byb_ga: Byblis gaimardi, Cal_la: Calloporalata, Cha_se: Chaetozone setosa, Cir_in: Cirratulidaeindet., Cni_in: Cnidaria indet., Cry_cl: Cryptonaticaaffinis, Edw_sp: Edwardsia sp., Esc_sp: Escharella sp.,Ete_sp: Eteone sp. Euc_dr: Euclymene droebachiensis,Euc_in: Euclymeninae indet., Eun_sp: Eunoe sp.,Hyp_su: Hyperammina subnodosa, Lam_fu: Lampropsfuscatus , Nuc_te: Nuculoma tenuis, Lei_sp:Leitoscoloplos sp., Lep_ca: Lepeta caeca, Leu_na:Leucon nasica, Lum_sp: Lumbrineris spp., Mac_ca:Macoma calcarea, Mal_sa: Maldane sarsi, Mon_la:Monoculodes latimanus, Myr_su: Myriaporasubgracilis, Myr_fr: Myriochele fragilis, Nep_lo:Nephtys longosetosa, Onc_ca: Oncousoecia canadensis,Oph_li: Ophelia borealis, Oph_ro: Ophiura robusta,Owe_fu: Owenia fusiformis, Par_gr: Paradoneis gracilis,Pec_hy: Pectinaria hyperborea, Pho_sy: Pholoesynopthalmica, Pol_pa: Polynices pallida, Por_sp:Porella sp., Pro_ma: Proclea malmgreni, Rho_gr:Rhodine gracilior, Sco_ar: Scoloplos armiger, Smi_mu:Smittina minuscula, Spi_ty: Spiochaetopterus typicus,Ste_no: Stegophiura nodosa, Tan_in: Tanaidacea indet.,Ter_st: Terebellides stroemi, Thy_sp: Thyasira sp.,Tor_in: Tornidae indet.

194 Sarsia 83:183-210 – 1998

ACKNOWLEDGEMENTS

Gennady Matishov, director of MMBI, is acknowledged forleading this joint venture. Thanks to Lars-Henrik Larsen,Akvaplan-niva, and the captain and crew of R/V DalnieZelentsie, MMBI, for fieldwork and practical assistance.Harvey Goodwin, Sigurd Jakobsen, Rune Palerud, and LenaRingstad Olsen were involved in various stages of sampleprocessing, data analyses and map compilation. We thankNatalia Anisimova, MMBI, for identifying the Echinodermataand Eivind Oug, NIVA for advice on the Polychaeta. TomPearson and Ole Jørgen Lønne, Akvaplan-niva, are gratefullyacknowledged for constructive comments and discussions aswell as two anonymous reviewers for criticism of the manu-script. This study has received financial support from theNorwegian Ministry of the Environment, under the PolarEnvironmental Centre Programme, and Akvaplan-niva.

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Accepted 8 September 1997

Editorial responsibility: Tore Høisæter

196Sarsia 83:183-210 – 1998

Appendix. Species list of benthic fauna sampled in the Pechora Sea, 1992.

3 6 7 8 11 12 13 14 19 20 21 24 26 27 29 sum

SARCOMASTIGOPHORAAstorhizidae

Hyperammina subnodosa Brady, 1884 - 15 - 37 105 32 - - - 9 - 316 28 511 - 1053PORIFERA

Porifera indet. 1 1 - - - - - 2 - - - - - - - 4CNIDARIA

Cnidaria indet. - - - - - - - - - - 1 - - - - 1HYDROZOA

Hydrozoa indet. 9 14 20 3 - 2 - 3 14 10 3 1 2 - - 81Monobranchia parasiticum Mereschkowsky, 1877 5 6 - - - - - - - - - - - - - 11

ANTHOZOAAnthozoa indet. - - - - - - - - - - - 2 - - - 2Actiniaria indet. - 1 - - - - - 3 - 1 - - - - - 5Capnella glomerata (Verill, 1869) - - - - - 3 - - - - - - - - - 3Cerianthus lloydii Gosse, 1859 2 7 4 - - - 1 2 15 - 3 - - - - 34Cerianthus sp. - - - - - - - - - - 2 - - - - 2Edwardsia sp. 1 - - - - - - - - - 7 19 10 10 - 47Edwardsiidae indet. 2 6 5 - - - - - 10 14 1 3 3 - - 44

NEMERTININemertini indet. 5 5 1 4 7 - 2 5 10 3 5 5 9 8 10 78

NEMATODANematoda indet. - - 2 2 - - - - - - 1 23 4 - - 32

SIPUNCULIDASipunculida indet. 4 1 - - 2 - - - - 2 2 - - - - 11Phascolion strombus (Montagu, 1804) - - - - 3 - - - - - - - - - - 3Golfingia margaritacea (M. Sars, 1851) - 9 7 - 2 - - - 9 1 1 - - - - 28Golfingia sp. - - - - - - - - - - 1 - - - - 1

PRIAPULIDAPriapulida indet. - - - - - - - 1 - - - - - - - 1Halicryptus spinulosus Siebold, 1849 - - - - - - - - - 1 - - - - 41 42Priapulus caudatus Lamarck, 1816 5 4 - 3 3 3 - 1 - - 2 1 - - - 22

ECHIURAEchiurida indet. - - - - - - - - - - 2 - - - - 2Echiurus echiurus Pallas, 1780 1 - - - - - - - - - 3 1 - - - 5

ANNELIDAPolychaetaPolynoidae

Polynoidae indet. 3 7 20 1 2 - - - 3 19 - - - - - 54Eunoe sp. - - - - - 2 - - - - 14 - - - - 16Gattyana sp. 1 3 - - - - - - - 1 4 - - - 1 10

Dahle &

al. – Benthic fauna in the Pechora Sea197

Harmothoe sp. - 3 - - - - - - - - 3 1 - - 1 8Nemidia torelli Malmgren, 1866 3 5 - - - - - - 3 1 - - - - - 12Pholoe synopthalmica Claparède, 1868 - - 16 2 8 6 17 9 15 33 69 1 - - - 176

PhyllodocidaePhyllodocidae indet. - 1 - - - - - - - - - - - - - 1Eteone sp. 5 - 6 1 22 - 4 - 3 5 23 10 20 8 - 106Phyllodoce groenlandica F.P. Örsted, 1842 4 1 1 - - - 1 - 8 3 3 8 8 - - 37

HesionidaeKefersteinia cirrata (Keferstein, 1862) - - - - - - 1 - - - 1 - - - - 2Nereimyra punctata (O.F. Müller, 1776) - - - - 103 - - - - - - - - - - 103

SyllidaeAutolytus sp. - - - - - 1 - - - - 1 - - - - 2Eusyllis blomstrandi Malmgren, 1867 - - 1 1 - 1 - - - 1 - - - - - 4Langerhansia cornuta (Rathke, 1843) - - - - - - - - - 1 - - - - - 1

NereidaeHediste diversicolor O.F. Müller, 1776 - - - - - - - - - 1 - - - - - 1Nereis zonata Malmgren, 1867 - - - - - - - - 1 1 - - - - - 2

GlyceridaeGlycera capitata Oersted, 1843 - - - - - - - - - 5 - - - - - 5Glycera lapidum Quatrefages, 1865 - - 14 - - - - - - 6 17 - - - - 37

NephtyidaeMicronephtys minuta (Théel, 1879) - - - - - - - - - - 2 - 3 - - 5Nephtys ciliata (Müller, 1776) 2 1 - - - - - - - 1 - - - - - 4Nephtys longosetosa Örsted, 1842 - - - - - - - - - - - 8 20 11 - 39Nephtys pente Rainer, 1984 - - - - - - - 1 - - - - - - 1

SphaerodoridaeSphaerodorum sp. - - - - - 1 - 1 - 2 1 - - - - 5

LumbrineridaeLumbrineris spp. 27 53 21 65 - 31 199 127 25 4 8 - - - - 560

OnuphidaeNothria conchylega (M. Sars, 1835) - - - - - - - - - 8 2 - - - - 10

DorvellidaeParougia sp. - - - - - - - - - 65 1 1 - - - 67Protodorvillea kefersteini (McIntosh, 1869) - - - - - - - - - - 1 - - - - 1

OrbinidaeLeitoscoloplos sp. 50 24 12 13 - 1 15 5 - 17 56 30 31 34 - 288Orbinia cuvieri (Audouin & M.-Edwards, 1833) - - - - - - - - - - - - - 3 - 3Scoloplos armiger (O.F. Müller, 1776) - - 2 - - - - - - - 3 8 8 - - 21

ParaonidaeParaonidae indet. - - - - - - - - - 1 - - - - - 1Aricidea sp. 2 - 3 3 33 1 14 - - 3 7 - - 1 - 68

Appendix (continued) 3 6 7 8 11 12 13 14 19 20 21 24 26 27 29 sum

198Sarsia 83:183-210 – 1998

Paraonis gracilis (Tauber, 1879) 24 95 18 6 3 16 80 15 4 9 3 - - - - 273Paradoneis lyra (Southern, 1914) - - - - - - - - - - 3 - - - - 3

CossuridaeCossura sp. 1 - - - - 1 - - - - - - - - - 2

ApistobranchidaeApistobranchus tullbergi (Théel, 1879) - 1 1 - - - - - - 1 - - - - - 3Apistobranchus sp. 15 2 - 3 - 1 - 2 - - 7 - 1 3 - 34

SpionidaeSpionidae indet. - - - - - - - - - 1 2 - - - - 3Laonice cirrata (M. Sars, 1851) 1 - - 2 - 1 - - 1 - - - - - - 5Marenzelleria sp. - - - - - - - 2 - - - - 17 5 27 51Polydora caeca/flava - - - - - 3 - - - 2 - - - - - 5Polydora socialis (Schmarda, 1861) - - 1 - - - - - - - - - - - - 1Polydora sp. - - - - 72 - 24 2 1 - 48 - 2 - - 149Spio decoratus Bobretzky, 1871 4 - - 2 - 1 - - 1 35 - - - - 45Spio sp. - - 6 - - - - - - - - - - - - 6

ChaetopteridaeSpiochaetopterus typicus M. Sars, 1856 1 32 206 70 113 407 288 139 193 236 2 - - - - 1687

CirratulidaeCirratulidae indet. 11 9 35 7 97 5 6 11 26 20 203 36 - 4 - 470Chaetozone setosa Malmgren, 1867 49 31 55 5 375 2 16 - 5 10 122 52 20 14 - 756Cirratulus sp. - 2 - - 2 - - 1 - - - - - - - 5

CapitellidaeCapitella capitata (O. Fabricius, 1780) - 1 1 - 25 - - 1 3 - 11 2 - 5 - 49Heteromastus filiformis (Claparède, 1864) 4 8 - 5 1 - - 11 5 28 2 - - - 67Mediomastus sp. - - - - - - - - 1 - - - - - - 1Notomastus sp. - - 10 - - - - - - 2 1 - - - - 13

MaldanidaeMaldanidae indet. 1 3 10 1 12 - - - - - - 6 - - - 33Euclymeninae indet. 9 7 1 - - - - - - - 3 111 4 - - 135Axiothella sp. 1 - - - - - - - - - - - - - - 1Clymenura sp. 5 1 19 - - - - - 10 - 20 7 24 - - 86Euclymene droebachiensis (M. Sars in G.O. Sars, 1872) - - - - - - - - 1 - 50 - - - - 51Lumbriclymene cylindricaudata Sars, 1872 - - - - - 4 - - - - 1 - - - - 5Maldane sarsi Malmgren, 1865 184 136 58 21 173 47 61 62 170 15 8 - - - - 935Nicomache sp. - - 3 - - - - - 18 1 13 - - - - 35Praxillella praetermissa (Malmgren, 1865) 16 1 - - - - - - - - - - - - - 17Praxillella sp. - - - - - 1 - - - - - 4 6 1 - 12Praxillura longissima Arwidsson, 1907 - - 7 - - - - - 1 - - - - - - 8Rhodine gracilior Tauber, 1879 70 2 40 - - 1 - - 4 - 221 - - - - 338

Opheliidae

Appendix (continued) 3 6 7 8 11 12 13 14 19 20 21 24 26 27 29 sum

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al. – Benthic fauna in the Pechora Sea199

Ophelia borealis (Rathke, 1843) - - - - - - - - - - 4 4 44 46 - 98Ophelina acuminata Oersted, 1843 1 - - - - - - - - - 1 - - - - 2Travisia forbesii Johnston, 1840 - - - - - - - - - - - 16 6 1 - 23

ScalibregmidaeScalibregma inflatum Rathke, 1843 7 8 7 11 23 1 4 2 - 5 1 4 - - - 73

OwenidaeGalathowenia oculata (Zachs, 1923) 13 55 30 2 - 2 4 11 8 3 19 258 47 4 - 455Myriochele danielsseni Hansen, 1879 - 2 - 3 2 - - - - - - - - - - 7Myriochele heeri Malmgren, 1867 - - - - - 24 94 82 - - - - - - - 200Owenia fusiformis delle Chiaje, 1841 - - - - - - - 11 1 - 4 65 54 3 - 138

FlabelligeridaeFlabelligeridae indet. - - 1 - - - - - - 1 - - - - - 2Brada sp. - 1 - - 2 1 1 - 1 1 - 19 - - - 26Diplocirrus hirsutus (Hansen, 1879) - - - - - - - - - 1 - - - - - 1Pherusa sp. 1 - - - 2 - - - - - - 6 - - - 9

PectinariidaePectinaria hyperborea (Malmgren, 1866) 150 11 - 2 3 - 1 1 - 1 - 15 - - - 184

AmpharetidaeAmpharetidae indet. - - 1 - - - - 1 - 5 4 1 1 - - 13Ampharete acutifrons (Grube, 1860) 1 1 - 1 - - - - - 2 - 1 - - - 6Ampharete baltica Eliason, 1955 - 3 - - - - 4 2 - - 1 30 3 - - 43Ampharete finmarchica (M. Sars, 1866) - 2 6 4 - 3 3 9 - 1 - - 2 - - 30Ampharete goesi (Malmgren, 1966) - - 5 - - - - - - 2 1 - - - - 8Ampharete lindstroemi Malmgren ,1867 - - - - - - - - - - 5 - - - - 5Ampharete sp. - - - - - - 3 - - 5 - - - - - 8Amphicteis sundevalli Malmgren, 1866 - - - - 2 - - - - - - - - - - 2Anobothrus gracilis (Malmgren, 1866) - 4 48 - - - - 1 3 13 23 1 - - - 93Artacama proboscidea Malmgren, 1866 5 - - - - - - 1 - - - - - - - 6Lysippe labiata Malmgren, 1866 - 3 56 4 2 2 4 10 6 20 32 - - - - 139Melinna cristata (M. Sars, 1851) - - 1 - - - - - - - - - - - - 1Sabellides borealis M. Sars, 1856 - - - - 5 - - - - - - 1 - - - 6

TerebellidaeTerebellidae indet. - - 2 - 3 - - - 1 1 1 3 - - - 12Amaeana trilobata (M. Sars, 1863) - - - - - - - - - - 7 - - - - 7Amphitrite cirrata O.F. Müller, 1771 - - - - - - - - - 1 - - - - - 1Lanassa venusta (Malm, 1874) - - 1 - - - - - - - 1 - - - - 2Laphania boecki Malmgren, 1866 - - 19 - - - 3 - 1 7 3 - - - - 33Leaena ebranchiata M. Sars, 1865 - 1 3 2 2 - - - - 3 5 - - - - 16Lysilla loveni Malmgren, 1865 - - - - - - - - - - 1 - - - - 1Neoamphitrite affinis (Malmgren, 1866) - - 2 - - - - - - - - - - - - 2Nicolea zostericola Ørsted in Grube, 1860 - - - - - - - 1 - - - - - - - 1

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200Sarsia 83:183-210 – 1998

Paramphitrite tetrabranchia Holthe, 1976 - - 1 - - - - - - - 2 - - - - 3Pista cristata O.F. Müller, 1776 - - - - - - - - - - 1 - - - - 1Polycirrus sp. - - - - - - - - - - - 1 - - - 1Proclea malmgreni (Ssolowiew, 1899) 2 2 - - - - - - - - - 4 15 4 - 27Thelepus cincinnatus (O. Fabricius, 1780) - - - - - - - - - 7 - - - - - 7

TrichobranchidaeTerebellides stroemi M. Sars, 1835 4 7 18 16 43 6 28 35 4 16 6 36 - - - 219

SabellidaeSabellidae indet. - - 3 2 3 - - - - - - 1 - - - 9Chone duneri Malmgren, 1867 - - - - 23 - - - - - 48 2 - - - 73Chone filicaudata Southern, 1914 - - 1 - - - - - - - - - - - - 1Chone infundibuliformis Krøyer, 1856 - - 1 - - - - 1 1 - 1 - - - - 4Chone paucibranchiata (Krøyer, 1856) - - - - - - - - - - 4 - - - - 4Chone sp. 1 1 2 - - - - 2 5 3 - - - - - 14Branchiomma sp. - - - - - - - - - 1 - - - - - 1Euchone analis (Krøyer, 1856) - - 2 - - - 4 - - 1 - - - 1 - 8Euchone elegans Verrill, 1873 - - - - - - - - - - 13 - - - - 13Euchone papillosa M. Sars, 1851 - - - - 93 1 - 3 - - - 1 - - - 98Euchone sp. - - - 1 - 1 - 3 - - - - 2 - - 7Jasmineira caudata Langerhans, 1880 - - - - - - - - - 1 - - - - - 1Sabella sp. - - - - - - - - - - - 1 - - - 1

OLIGOCHAETAOligochaeta indet. - - - - - - - - - - 3 - 22 - - 25

CHELICERATAPYCNOGONIDA

Pycnogonida indet. 1 - - - - - - - - - - - - - - 1Pantopoda indet. - - - - - - - - - 1 - - 1 - - 2

CRUSTACEAOSTRACODA

Ostracoda indet. - 1 1 - - - 5 3 1 31 40 - - - - 82Cypridinidae

Philomedes globusus (Lilljeborg, 1853) - - - - - - - - - 2 - - - - - 2COPEPODA

Calanoida indet. - 1 1 1 2 4 1 - 3 - 9 - - - - 21CIRRIPEDIABalanidae

Balanus balanus (Linnaeus, 1758) - - 3 - - - - - - - - - - - - 3Balanus crenatus Bruguière, 1789 - - 41 - - - - - 13 1 4 - - - - 59

MALACOSTRACACumacaea

Cumacea indet. - - - 2 - - - - 1 6 - - - - - 9

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LeuconidaeEudorella emarginata (Krøyer, 1846) 10 2 1 - - - - - 3 9 6 - - - - 31Eudorella truncatula Bate, 1856 - - 2 - - - - - - - - - - - - 2Leucon nasica (Krøyer, 1841) 3 2 2 - - - 2 - 1 1 14 - - - - 25

NannastacidaeCampylaspis sp. - - - - - - - - - - - 7 5 5 - 17

LampropidaeLamprops fuscatus G.O. Sars, 1865 - - - - - - - - - - - 13 16 - - 29

DiastylidaeBrachydiastylis resima (Krøyer, 1846) 4 6 1 1 - - 1 1 - 28 29 8 - - - 79Diastylis goodsiri (Bell, 1855) - - - 1 7 1 1 7 - - - - - - - 17Diastylis rathkei (Krøyer, 1841) 2 2 - - - - - - - - - 17 4 - 29 54Diastylis scorpioides (Lepechin, 1780) - - 6 - - - 1 - 10 4 - - - - - 21Diastylis sp. - - 3 - - - - 3 - - 1 - - - - 7

TanaidaceaTanaidacea indet. - - - - - - - - - - 11 - 8 3 - 22

ApsenidaeSpyraphus anomalus G.O. Sars, 1869 - - - - - - - - 55 3 - - - - - 58

AmphipodaAmphipoda indet. - 2 9 - - - - - 5 1 19 - - 1 - 37

AcanthonotozomatidaeAcanthonotozoma sp. - - - - - - - - - - 2 - - - - 2

AmpeliscidaeAmpelisca eschrichti Krøyer, 1842 1 - - 1 - - 1 - 3 - - - - - - 6Ampelisca macrocephala Liljeborg, 1852 1 - 1 - - 1 - - - - - 2 - - - 5Byblis gaimardi (Krøyer, 1846) 9 3 15 - - 1 - 2 9 5 644 - - - - 688Haploops tubicola Liljeborg, 1855 24 10 - 8 3 - 10 5 89 8 4 - - - - 161

AmphithoidaeAmphithoe sp. - - - - - - - - - - 2 - - - - 2Arctolembos arcticus (Hansen, 1887) - 2 - - - - - - 3 - 4 - - - - 9Unciola leucopis (Krøyer, 1845) - - 2 - - - - - 5 9 20 - - - - 36

AtylidaeAtylus smitti (Goës, 1866) - - - - - - - - - 1 - - - - - 1

CalliopiidaeCalliopiidae indet. - - - - - - 1 - - - - - - - - 1Halirages fulvocincta (M. Sars, 1858) - - - - - - 1 - - - - - - - - 1

CorophiidaeCorophium crassicorne Bruzelius, 1859 - - - - - - - - - - - 1 26 3 - 30

EusiridaeRhachotropis aculeata (Lepechin, 1780) - - - - - - - - 1 - - - - - - 1

Isaeidae

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202Sarsia 83:183-210 – 1998

Isaeidae indet. - - - - - - - - - - 72 - - - - 72Photis reinhardi Krøyer, 1842 - - - - - 2 - - 3 3 1 39 - - - 48Photis sp. - - - - - 1 - - - - - - - - - 1Protomedeia fasciata Krøyer, 1842 - - - - - - - - - - 384 - - - - 384Protomedeia grandimana Brüggen, 1905 4 - 5 - - - - - - - 1 - - - - 10

IschyroceridaeIschyroceridae indet. - - - - - - - - - - 23 - - - - 23Ischyrocerus sp. - - - - 2 - - - - - - - - - - 2

LysianassidaeLysianassidae indet. 1 - 2 1 - - - - 4 2 5 1 3 - - 19Anonyx nugax (Phipps, 1774) - - - - - 1 - - - - - - - - - 1Anonyx sp. - - - - - - 3 1 - 1 1 - - - - 6

MelitidaeMelita dentata (Krøyer, 1842) - - 6 - - - - - - 1 21 - - - - 28

MelphidippidaeMelphidippa sp. - - - - - 3 3 - - - - - - - - 6

OedicerotidaeOedicerotidae indet. - - - - - 2 - - 111 - - - - - - 113Aceroides latipes G.O. Sars, 1866 - - - - - 2 - - - - 1 - - - - 3Arrhis phyllonyx (M. Sars, 1858) 1 - - 1 5 1 1 2 - - - - - - - 11Monoculodes latimanus (Goës, 1866) - - 3 - - - - - - - 8 2 - - - 13Monoculodes tuberculatus Boeck, 1871 - - - - - - - - - - 2 - - - - 2Monoculodes sp. - - - - - - - - - - - - 2 - - 2Westwoodilla caecula (Bate, 1856) - - 1 - - - - - - - - - - - - 1

PhoxocephalidaeHarpinia propinqua G.O. Sars, 1891 - - - - - - - - - - 20 - - - - 20Harpinia serrata G.O. Sars, 1879 - - 5 - - - - - 16 2 2 - - - - 25Phoxocephalus holbolli (Krøyer, 1842) - - 2 - - - - - - - 6 - - - - 8

PleustidaePleusymtes pulchella (G.O. Sars, 1876) - - - - - - 2 - - - - - - - - 2

PodoceridaePodoceridae indet. - - - - - - - - 1 - - - - - - 1Dyopedos bispinis (Gurjanova, 1930) - - - - - - - - - - 3 1 2 - - 6

PontoporeiidaeMonoporeia affinis (Lindström, 1855) - - - - - - - - - - - - - - 7 7Pontoporeia femorata Krøyer, 1842 - 3 - - 2 - 6 13 - - - - - - 165 189Priscillina armata (Boeck, 1861) - - - - - - - - - - - - 9 1 - 10

StegocephalidaeStegocephalus inflatus Krøyer, 1842 - - - - - - - 1 - - - - - - - 1

StenothoidaeStenothoidae indet. - - - - - - - - - - 23 - - - - 23

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SynopiidaeSyrrhoe crenulata Goës, 1866 - - 2 - - - - - - - - - - - - 2Tiron spiniferus (Stimpson, 1853) - - 3 - - - - - - - - - - - - 3

HyperiidaeParathemisto sp. - - 1 - - - - - - - - - - - - 1

IsopodaIsopoda indet. 1 - - - - - - - - - - - - - - 1Gnathidae

Gnathia sp. - - - - - 1 - 8 11 1 - - - - - 21Parasellinae

Munna sp. - - - - - - - - 1 - 16 - - - - 17Pleurogonium inerme G.O. Sars, 1883 - - - - - - - - - - - - 2 - - 2

DecapodaCrangonidae

Brachyura indet. - 1 - - - - - - - - - - - - - 1Sabinea septemcarinatus (Sabine, 1824) - - - - - - - - 1 - - - - - - 1

PaguridaePaguridae indet. - - 1 - - - - - - - - - - - - 1Pagurus pubescens Krøyer, 1838 - 1 3 - - - - 1 - - 1 - - - - 6

MOLLUSCACAUDOFOVEATA

Caudofoveata indet. 4 4 - - - - - - - - - 1 - - - 9Chaetodermidae

Chaetoderma sp. Lovén, 1845 4 - - - 2 - - - - - - - - - - 6POLYPLACOPHORAIschinochitonidae

Ischnochiton albus (Linnaeus, 1767) - - - - - - - - - - 5 - - - - 5GASTROPODA

Gastropoda indet. - - - 1 - - - - 1 - - - - - - 2Prosobranchia indet. - 1 - - - - - - - - - - - - - 1

LepetidaeLepeta caeca (Müller, 1776) - - 8 - - - - - 9 3 49 - - - - 69

TrochidaeMargarites costalis (Gould, 1841) - - - - - - - - - - 1 - - - - 1Margarites striatus (Leach, 1819) - - - - - - - - - - - - 15 4 - 19Margarites olivacea (Brown, 1827) - - - 3 - - - 2 - - - - - - - 5Solariella obscura (Couthoy, 1838) - 3 8 - - - - - - - 1 5 - - - 17Solariella varicosa (Mighels & Adams, 1842) - - 6 - - - - - 11 - - - - - - 17Solariella sp. 1 - - - - - - - - - - - - - - 1

TurbinidaeMoelleria costulata (Möller, 1842) - - 1 - - - - - - 4 4 - 1 - - 10

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204Sarsia 83:183-210 – 1998

RissoidaeAlvania cruenta Odhner, 1915 - - - - - 2 - 2 - - - - - - - 4

TornidaeTornidae indet. - - 1 - - - - - - - 3 - - - - 4

TrichotropidaeTrichotropis borealis Broderip & Sowerby, 1829 - - 1 - - - - - - 1 - - - - - 2

LamellariidaeVelutina undata Brown, 1838 - - - - - - - - - 1 - 1 - - - 2

NaticidaeNaticidae indet. - - - - - - - - - - - - - 1 - 1Amauropsis islandica (Gmelin, 1791) - - - - - - - - - - - - 3 1 - 4Cryptonatica affinis (Gmelin, 1791) - - - - - - - - - 2 - 6 2 1 - 11Polinices nanus (Möller, 1842) - - 1 - - - - - - - 1 - 3 - - 5Polinices pallidus (Broderip & Sowerby, 1829) - - - - - - - - - 2 1 4 4 1 - 12

BuccinidaeBuccinum glaciale Linnaeus, 1766 - - 1 - - - - - - - - - - - - 1Turrisipho fenestratus (Turton, 1834) - - - 1 - - - - - - - - - - - 1

CancellariidaeAdmete couthouyi (Jay, 1839) - - 2 - - - - - 1 - - - - - - 3

TurridaeCurtitoma trevelliana (Turton, 1834) - - 1 1 - - - - - - - - - - - 2Obesotoma simplex (Middendorff, 1849) - - - - - - - - - - - - 6 - - 6Obesotoma woodiana (Möller, 1842) - - - - - - - - - - 1 - - - - 1Oenopota impressa (Mörch, 1869) - - 2 1 - - - - 6 - 2 - - - - 11Oenopota pyramidalis (Ström, 1788) - - 1 - - - - - - - - - - - - 1Oenopota harpularia (Couthouy, 1838) - 1 - - - - - - 1 - - 1 - - - 3Oenopota rugulata (Moller in Troschel, 1866) - - 6 - - - - - 1 - - - - - - 7Oenopota violacea (Mighels & Adams, 1842) - - 5 - - - - - - - 7 - - - - 12

OpisthobranchiaOpisthobranchia indet. 1 - - - - - - - - - - - - - - 1

TurbonillidaeMenestho truncatula Odhner, 1915 - - - - - - - - - - - - 3 - - 3

DiaphanidaeDiaphana minuta Brown, 1827 - - - 2 17 - 1 1 - - - - 2 1 - 24

RetusidaeRetusa obtusa (Montagu, 1803) - - - - 18 1 - - - - - - - - - 19Retusa sp. Brown, 1827 - 1 - - - - - - - - - - - - - 1Philine finmarchica M. Sars, 1858 - - - - - - - - - - - - 1 - - 1Philine quadrata (S. Wood, 1839) - - - - - - - - - 1 - - 5 - - 6

ScaphandridaeCylichna alba (Brown, 1827) 1 - 1 2 2 3 2 3 - 1 1 1 11 5 - 33

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Cylichna occulta (Mighels, 1842) 1 1 1 3 - - - 2 - - - 1 1 - - 10Cylichna sp. - 2 - - - - - - - - - - - - - 2

BIVALVIABivalvia indet. 6 9 1 - - 1 - 2 1 - - - 10 1 - 32

NuculidaeNuculoma tenuis (Montagu, 1808) 24 53 80 3 8 39 - 33 28 23 20 - 2 3 - 315

NuculanidaeNuculana pernula (Müller, 1779) 3 14 3 8 12 4 3 15 14 5 5 - - - - 85Nuculana sp. Link, 1807 - - 2 - - - 1 - - - - - - - - 3Yoldia hyperborea Torell, 1856 40 9 - 2 - 1 - - - - - - - - - 52Yoldiella frigida (Torell, 1859) - - - - 8 1 - - - - - - - - - 9Yoldiella lenticula (Möller, 1842) - - - 3 7 2 1 11 - - - - - - - 24Nuculanidae indet. - - 1 - - - - - - - - - - - - 1

MytilidaeMytilidae indet. - - - - - - - - - - - 2 - - - 2

Crenella decussata (Montagu, 1808) - - - - - - - - - - 6 - 19 1 - 26Dacrydium vitreum (Holbøll in Möller, 1842) - - - - - - - - - - 1 - - - - 1Musculus corrugatus (Stimpson, 1851) - - - - - - - - - 1 - - - - - 1Musculus niger (J.E. Gray, 1824) 2 - 4 - - - - - 4 - 1 - - - - 11Musculus sp. 1 - 3 - - - - 1 - - 2 - - - - 7Mytilus edulis Linnaeus, 1758 - - - - - - - - - - - - 3 1 - 4

ArcidaeBathyarca glacialis (J.E. Gray, 1824) - - - - 8 - - - - - - - - - - 8

PectinidaePectinidae indet. - - 1 - - - - - - - - - - - - 1

ThyasiraidaeThyasiridae indet. - - 1 - - - - - - - - - - - - 1Axinopsida orbiculata (G.O. Sars, 1878) - - 1 - - - - - - - 3 - 1 - - 5Thyasira equalis (Verrill & Bush, 1898) - 9 - - - - - - - 2 - - - - - 11Thyasira gouldi (Philippi, 1846) - - 9 - - 10 6 - 1 2 1 - - - - 29Thyasira sarsii (Philippi, 1846) - - 4 - - 1 - - - - - - - - - 5Thyasira sp. 15 17 1 2 148 - - - - - - 12 - - - 195

LasaeidaeMontacuta maltzani (Verkrüzen, 1876) - - - 1 - - - - - 1 - - 1 15 - 18Montacuta spitzbergensis Knipovitsch, 1901 11 7 - - - - 1 - 5 2 1 1 - - - 28Montacuta sp. - 1 - - - - - - - - - - - - - 1

AstartidaeAstartidae indet. 1 - 1 - 2 - - - 3 - - - - 1 - 7Astarte sulcata (da Costa, 1778) - 2 - - - - - - - - - - - - - 2Ciliatocardium ciliatum (O. Fabricius, 1780) - 4 2 - - 2 - - 9 1 - 3 - - - 21Serripes groenlandicus (Bruguière, 1789) - - - - - - - - - - - 22 5 3 - 30

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206Sarsia 83:183-210 – 1998

Tridonta borealis Schumacher, 1817 - 8 33 1 - - - - 8 2 1 2 - - - 55Tridonta elliptica (Brown, 1827) - - 34 - - - - - 3 10 2 - - - - 49Tridonta montagui (Dillwyn, 1817) - 10 19 2 10 - - - 8 21 10 6 - - - 86

CardiaceaCardiacea indet. 9 - - - - - - - - - - - - - - 9

TellinidaeMacoma balthica (Linnaeus, 1758) - - - - - - - - - - - - - - 151 151Macoma calcarea (Gmelin, 1791) 21 9 16 3 2 2 - 1 - 34 32 - 8 8 - 135Macoma crassula (Deshayes, 1855) - - 6 - - - - - 1 3 8 15 - - - 33Macoma loveni Jensen, 1904 - - - - - - - - 3 - - - - - - 3Macoma sp. 17 10 - - - - - - - - - 1 - - - 28

ArcticidaeArctica islandica (Linnaeus, 1767) - - - - - - - - - - - - 1 - - 1Liocyma fluctuosa (Gould, 1841) - - - - - - - - - - - 7 5 1 - 13Pillucina sp. - - - - - - - - - - - 2 - - - 2

MyidaeMya truncata Linnaeus, 1758 - - 7 - - - - - 3 5 17 19 - - - 51

HiatellidaeHiatella arctica (Linnaeus, 1758) - - 9 - - - - 1 3 10 2 1 - - - 26Panomya arctica (Lamarck, 1818) - - - - - - - - - 1 - - - - - 1

ThraciidaeThracia myopsis (Möller, 1842) - - 4 - - - - - 3 - 1 - 2 1 - 11

LyonsiidaeLyonsia arenosa (Möller, 1842) - 1 - - - - - 1 3 - 1 1 - - - 7

PandoridaePandora glacialis (Leach, 1819) - - - - - - - - 1 - - 6 - - - 7

BRACHIOPODAHemithiris psittacea Gmelin, 1792 - - - - - - - - - 1 9 - - - - 10

BRYOZOAStenolaemata

Cyclostomata indet. - - 6 - - - - - - - 9 - - - - 15Crisiidae

Crisia eburnea (Linnaeus, 1758) - - - 1 - - - - - - - - - - - 1Crisiella producta (Smitt, 1865) - - - - - - - 3 - - - - - - - 3

DiastoporidaeDiplosolen sp. - - 1 - - - - - - - 2 - - - - 3

LichenoporidaeLichenopora crassiuscula (Smitt1867) - - - - - - - - - - 1 - - - - 1Lichenopora verrucaria (O. Fabricius, 1780) - - 6 - - - - - - 2 7 - - - - 15Lichenopora sp. - 1 - - - - - - - - - - - - - 1

Tubuliporidae

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Dahle &

al. – Benthic fauna in the Pechora Sea207

Oncousoecia canadensis (Osburn, 1933) - - - - - - - - - 1 7 - - - - 8Oncousoecia diastoporides (Norman, 1869) - - - - - - - - - 5 1 - - - - 6Tubulipora sp. - - - 1 2 - - - - - - - - - - 3

GYMNOLAEMATAScrupariidae

Eucratea loricata (Linnaeus, 1758) - 2 - 2 - 1 1 2 1 - - 1 2 - 1 13Membraniporidae

Amphiblestrum auritum (Hincks1877) - 1 - - - - - - - - - - - - - 1Callopora lata (Kluge1907) - - - - - - - - - - 3 - - - - 3Callopora lineata (Linnaeus, 1767) - - 2 - - - - - 1 - - - - - - 3Callopora smitti Kluge, 1946 - 1 - - - - - - - - 1 - - - - 2Doryporella spatulifera (Smitt, 1868) - - 1 - - - - - - - - - - - - 1Electra arctica (Borg, 1931) - - 12 - - - - - - 1 105 - - - 1 121Electra crustulenta (Pallas, 1766) - - - - - - - - - - 3 - - - - 3Tegella arctica (d’Orbigny, 1851) - - - - - - - - - - 1 - - - - 1Tegella armifera (Hincks, 1880) - - 1 - - - - - - - - - - - - 1Tegella armifesoides (Kluge, 1955) - 1 - - - - - - - - 1 - - - - 2Tegella nigrans (Hincks, 1882) - 1 - - - - - - - - - - - - - 1Tegella sp. - - - - - - - - 1 - - - - - - 1

CribrilinidaeCribrilina annuata (O. Fabricius, 1780) - - - - - - - - - 1 - - - - - 1Cribrilina spitzbergensis (Norman1903) - - - - - - - - - - 5 - - - - 5

ScrupocellariidaeScrupocellaria scabra (van Beneden, 1848) - - - - - - - - - 1 - - - - - 1Semibugula birulai (Kluge1929) - - - - - - - - - - - 5 - - - 5Tricellaria gracilis (van Beneden1848) - 1 - - - - - - - - - - - - - 1Tricellaria peachi (Busk, 1851) 1 - - - - - - - 3 - - - - - - 4

BicellariidaeBugula fastigiata (Dalyell1847) - 2 - - - - 1 - - - - 2 - - - 5Dendrobeania elongata (Nordgaard, 1906) - - 1 - - - - - - - - - - - - 1Dendrobeania levinseni (Kluge1929) - - - - - - - 1 - - - - - - - 1Kinetoskias arborescens Danielssen, 1868 - - - - - - - 3 - - - - - - - 3

SmittinidaeArctonula artica (M. Sars, 1851) - 1 - - 2 - - - - - - - - - - 3

HippothoidaeHippothoa divaricata Lamouroux, 1821 - - 6 - - - - - - 2 86 - - - - 94Hippothoa expansa Dowson, 1859 - - 5 - - - - - - - 2 - - - - 7

MicroporellidaeMicroporella ciliata (Pallas, 1766) - - 5 - - - - - - - 49 - - - - 54

EscharellidaeEscharella ventricosa (Hassall, 1842) - 1 - - - - - - - 1 2 - - - - 4

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208Sarsia 83:183-210 – 1998

Escharella sp. - - - - - - - - - - 1 - - - - 1Escharelloides spinulifera (Hincks, 1889) - - 1 - - - - - - 1 18 - - - - 20

SmittinidaeSmittinidae indet. - - - - - - - - - - 2 - - - - 2Cystisella saccata (Busk1856) - - 3 - - - - - - - - - - - - 3Pachyegis groenlandica (Norman, 1894) - - 2 - - - - - - - 6 - - - - 8Pachyegis producta (Norman, 1903) - - 4 - - - - - - - 11 - - - - 15Palmicellaria skenei (Ellis & Solander, 1786) - - 1 - - - - - - - - - - - - 1Porella acutirostris Smitt, 1868 - - 1 - - - - - - - - - - - - 1Porella compressa (J. Sowerby1806) - - - - - - - - - 1 5 - - - - 6Porella fragilis Levinsen, 1914 - - - - - - 1 - - - - - - - - 1Porella minuta (Norman, 1868) - 2 - - - - - - - - - - - - - 2Porella obesa (Waters, 1900) - - - - - - - - - - 1 - - - - 1Porella sp. - - - - - - - - - 1 3 - - - - 4Porelloides laevis (Fleming, 1828) - - 6 - - - - - - - 6 - - - - 12Porelloides struma (Norman, 1868) - - 1 - - - - - - 1 2 - - - - 4Smittina minuscula (Smitt, 1868) - - - - - - - - - - 1 - - - - 1Smittina rigida Lorenz, 1886 - - 2 - - - - - - 1 4 - - - - 7

SchizoporellidaeBuffonellaria biaperta (Michelin, 1841) - - 1 - - - - - - - - - - - - 1Hippodiplosia harmsworthi (Waters, 1900) - - 2 - - - - - - - - - - - - 2Hippodiplosia propinqua (Smitt, 1868) - - - - - - - - - 2 - - - - - 2Hippodiplosia sp. - - - - - - - - - - 2 - - - - 2Porella obessa (Waters1900) - - - - - - - - - - 6 - - - - 6Schizoporella biaperta (Michelin, 1906) - - - - - - - - - - 1 - - - - 1Schizoporella bispinosa (Nordgaard1906) - - 1 - - - - - - - - - - - - 1Schizoporella crustacea (Smitt, 1868) - - - - - - - - - - 1 - - - - 1Schizoporella pachystega (Kluge, 1929) - - 6 - - - - - - - 21 - - - - 27Schizoporella smitti Kluge, 1962 - - 1 - - - - - - - 3 - - - - 4Schizoporella sp. - - 1 - - - - - - - - - - - - 1

CelleporidaeCellepora pumicosa (Pallas, 1766) - - - - - - - - - - 1 - - - - 1Celleporina incrassata (Lamarck, 1788) - - - - - - - - - 2 4 - - - - 6Celleporina ventricosa (Lorenz, 1886) - - 15 - - - - - - - 15 - - - - 30

StomachetosellidaeStomachaetosella cruenta (Busk, 1854) - - - - - - - - - - 1 - - - - 1Stomachaetosella limbata (Lorenz, 1886) - - - - - - - - - - 13 - - - - 13Stomachaetosella magniporata (Nordgaard, 1906) - - 1 - - - - - - - 1 - - - - 2Stomachaetosella producta (Packard, 1863) - - - - - - - - - - 1 - - - - 1Stomachaetosella sinuosa (Busk, 1860) - - 3 - - - - - - 2 11 - - - - 16

Rhamphostomellidae

Appendix (continued) 3 6 7 8 11 12 13 14 19 20 21 24 26 27 29 sum

Dahle &

al. – Benthic fauna in the Pechora Sea209

Escharopsis lobata (Smitt, 1868) - - 1 - - - - - - 1 1 - - - - 3Ragionula rosacea (Busk, 1856) - - - - - - - - - - 3 - - - - 3Rhamphostomella hincksi (Nordgaard, 1898) - - 2 - - - - - - 1 - - - - - 3Rhamphostomella sp. - - - - - - - - 1 - - - - - - 1

MyrioxoidaeMyriapora subgracilis (dOrbigni, 1852) - - 7 - - - - - 4 - 39 - - - - 50

HipponellidaeHippoporella hippopus (Smitt, 1868) - - 6 - - - - - - - 10 - - - - 16Lepraliella contigua (Smitt, 1868) - - - - - - - - - - 1 - - - - 1

SmittinidaePorella concinna (Busk, 1854) - - 11 - - - - - - 3 18 - - - - 32Smittina mucronata (Smitt, 1856) - - - - - - - - - 1 4 - - - - 5

HippodinidaeCheiloporina sincera (Smitt, 1868) - - - - 5 - - - - - - - - - - 5Escharina alderi (Busk, 1856) - - - - 3 - - - - - - - - - - 3

AlcyonidiidaeAlcyonidium disciforme Smitt, 1872 - - - - - - - - - - - 5 3 4 - 12Alcyonidium gelatinosum (Linnaeus, 1761) - - - - - 3 - 1 - - - - - - - 4Alcyonidium proboscideum (Kluge, 1962) - - - - - - - - 1 - - - - - - 1

ECHINODERMATAAsteroideaGoniopectinidae

Ctenodiscus crispatus (Retzius, 1805) - - - 9 - 1 3 13 - 2 - - - - - 28Asteriidae

Asterias rubens Linnaeus, 1758 - - - - - - - - - - 1 - - - - 1Ophiuroidea

Ophiuroidea indet. - - - 1 - 1 - 2 - - - - - - - 4Ophiactidae

Ophiopholis aculeata (Linnaeus, 1767) - - - - - - - - - 8 1 - - - - 9Amphiuridae

Amphiura sundevalli (Müller & Troschel, 1842) - 2 2 - - - - 1 4 24 8 - - - - 41Ophiacanthidae

Ophiacantha bidentata (Retzius, 1805) - - 1 1 2 4 10 1 16 33 1 - - - - 69Ophiocten sericeum (Forbes, 1952) - - - 7 - 20 9 10 - 12 - - - - - 58

OphiuridaeOphiura robusta (Ayres, 1851) - 1 14 - - - - - 19 68 74 - - - - 176Ophiura sarsii Lütken, 1858 - 2 - - - 1 - - - 4 - - - - - 7Stegophiura nodosa (Lütken, 1854) 4 6 - - - - - - 5 - - 84 113 51 - 263

EchinoideaStronglyocentridae

Strongylocentrotus pallidus (Sars, 1871) - - 3 - - - - - - 3 2 - - - - 8

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210Sarsia 83:183-210 – 1998

Strongylocentrotus sp. Brandt, 1835 - - - - - - - 1 - - - - - - - 1HolothuroideaPsolidae

Psolus phantapus (Strussenfelt, 1765) - - - - - - - - - - 4 - - - - 4Psolus sp. Oken, 1815 - - - - - - - - 3 1 - - - - - 4

CucumariidaePentamera caleigera (Stimpson, 1854) - - - - - - - 1 - - - 11 - - - 12

MyriotrochidaeMyriotrochus rinkii Steenstrup, 1852 - - - - - - - - - - - 15 - - - 15

EupyrgidaeEupyrgus scaber Lütken, 1857 - - - - - 2 6 - - 1 - - - - - 9

CHORDATAChordata indet. 1 - - - 3 - - - - - - 1 - - - 5

AscidiaceaAscidiacea indet. - - 1 - - - - - 4 1 - - - - - 6StyelidaeCnemidocarpa rhyzopus (Redikorzev, 1907) - - - - - - - - - - - 9 - 1 - 10Pelonaia corrugata (Forbes & Goodsir, 1841) - 1 - - - - - - - - - 1 20 - - 22

PyuridaeBoltenia echinata (Linnaeus, 1767) 1 - 1 - - - - - - - - 2 - - - 4

MolgulidaeEugyra glutinans (Möller, 1842) - - - - - - - - - - - - 1 - - 1

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