2 Biodiversity Research Center, Academia Sinica, Taipei 11529;
3 Fisheries College of Jimei University, Xiamen 361012
0 Introduction Climate changes are considered the major threat to biodiversity [1, 2]. Responses to climate change include species range expansions[3, 4],the advancement of spring phenology due to increasing temperatures[5, 6],and decreases in aquatic body sizes[7, 8]. Evidence of the changing Arctic includes temperature increases,shrinking sea ice coverage,and shifts in the ocean ecosystem. It is crucial to develop effective conservation and adaptation strategies to hedgeagainst the changing Arctic. In order to gain a better understanding of environmental changes to Arctic biodiversity and ecological impacts,the fourth Chinese National Arctic Research Expedition (4th CHINARE-Arctic) was conducted by the State Oceanic Administration of China in 2010. The present report is a part of the project documenting the effects of climate change on the nekton community and its distribution in the Pacific Arctic,including the Bering Sea,Bering Strait,and Chukchi Sea in the Arctic Ocean. Our perspective was to establish a long-term baseline of Arctic fish,including the species composition,abundances,and zoogeography.
1 Materials and methods Specimens were collected using a middle-water multinet,a French-type beam trawl (2.5 m wide,0.5 m high,and 9 m long; mesh size 10 mm),an otter trawl (1.6 m wide,0.5 m high,and 3 m long; mesh size 20 mm),and a triangular bottom trawl (2.2 m wide,0.65 m high,and 6.5 m long; mesh size 20 mm) aboard the Chinese R/V XUE LONG icebreaker from 11 July to 1 September 2010. Stations sampled were distributed from the Bering Basin through the continental rise,the continental shelf of the Bering Sea,northern and southern Bering Strait,to the shelf of the Chukchi Sea and the Chukchi slope (Table1,Figure 1). Every tow was operated for 10—60 min. The standard length (mm) and weight in grams (g) of all fish specimens were measured. Then specimens were fixed in a 10% buffered formalin-sea water solution and transferred to 70% ethyl alcohol. Specimens were identified using Coad et al. and Mecklenburg et al. Vouchers were deposited in the Biodiversity Collections in Third Institute of Oceanography,State Oceanic Administration,Xiamen and the Biodiversity Research Museum,Academia Sinica,Taipei.
|*MMT is multinet middle-water trawl; FBT is French-type beam trawl; OT is otter trawl; TT is triangular trawl.|
|SL01-SL02, SL05-SL09||01/09/2010||170°54′W||62°33′N||40||sand, mud||OT|
2 Results of fish collections and composition Specimens were collected from 36 stations (one multinet middle-water trawl and 35 bottom trawls) including the Bering Sea,Bering Strait,and Chukchi Sea,at depths ranging 24—626 m. Successful samples were obtained from 1 (2.8%) multinet middle-water trawl,7 (19.4%) French-type beam trawls,21 (58.3%) otter trawls,and 7 (19.4%) triangular beam trawls. Forty-one fish species belonging to 14 families in 7 orders were collected during the expedition (Table2). Among them,the Scorpaeniformes,including 17 species,accounted for almost one third of the total number (34.8%,427 individuals),followed by 14 species in the Perciformes (27.0%,331 individuals),5 species in the Pleuronectiformes (22.3%,273 individuals),and 2 species in the Gadiformes (15.4%,189 individuals). In addition,the most abundant family was the Cottidae (25%,307 individuals),followed in order by the Pleuronectidae (22.3%,273 individuals),Gadidae (15.4%,189 individuals),Zoarcidae (13.6%,167 individuals),and Stichaeidae (13.2%,162 individuals,Table2).
|Boreogadus saida||BB02, BS08, C02, C05, Co1, Co10, M06, M07, NB08, NB08B, NB09-NB10, R06, R08, SL04, SL05, SL06, SL07, SR07, SR09, SR10, SR11, SR12, NB09-NB10B||179||14.6|
|Perciformes||Ammodytidae||Ammodytes hexapterus||BS02, BS08||2||0.2|
|Zoarcidae||Gymnelus hemifasciatu||B14, SL08, SL09, SR07||11||0.9|
|Lycodes mucosus||BS05, SL09||5||0.4|
|Lycodes palearis||B14, BB02, BB05, BB06||51||4.2|
|Lycodes polaris||BB06, Co10, R08, SR11||5||0.4|
|Lycodes raridens||BB06, BS05, NB09-NB10, SR09, NB09-NB10B||19||1.5|
|Lycodes sagittarius||M06, M07||9||0.7|
|Lycodes seminudus||M06, M07||55||4.5|
|Stichaeidae||Anisarchus medius||B14, R06, R08, SR10, SR11||16||1.3|
|Eumesogrammus praec||BS05, SL08, SL09||15||1.2|
|Leptoclinus maculatus||B14, BB06||6||0.5|
|Lumpenus fabricii||BB06, BS08, CC08, NB08, NB08B, NB09-NB10B||100||8.2|
|Pleuronectiformes||Pleuronectidae||Hippoglossoides robust||B14, BB02, BB06, BS05, BS08,NB08, NB08B, NB09-NB10, R06, R08, SL01, SL02, SL03, SL04, SL05, SL06, SL09, SR03, SR09, NB09-NB10B||191||15.6|
|Lepidopsetta polyxystra||BS02, BS05, SL06||6||0.5|
|Limanda aspera||CC08, NB08, NB08B, NB09-NB10, SL01, SL02, SL04, SL05, SL07, NB09-NB10B||50||4.1|
|Pleuronectes quadritubtus||NB08, NB08B, NB09-NB10, SL01,SL02, SL05, SL06, SL07, SL08, NB09-NB10B||25||2.0|
|Rajiformes||Bathyrajiidae||Bathyraja parmifera||SL01, SL02, SL03||3||0.2|
|Scorpaeniformes||Agonidae||Aspidophoroides olrikii||BS05, BS08, CC08, NB09-NB10,R06, SL01, SL02, SL03, SL04, SL05, SR03, SR09, NB09-NB10B||38||3.1|
|Artediellus scaber||C02, CC08, R08||64||5.2|
|Gymnocanthus tricuspis||BS05, BS08, C02, C05, CC08, Co10, NB08, NB09-NB10, R06, R08, SL01, SL04SR03, SR07, NB09-NB10B||59||4.8|
|Hemilepidotus papilio||BS08, NB09-NB10, SL09, NB09-NB10B||7||0.6|
|Icelus spatula||B14, BS08||18||1.5|
|Myoxocephalus scorpiu||BS05, BS08, CC08, Co1, NB08, NB08B, SR03, SR07||143||11.7|
|Cyclopteridae||Eumicrotremus orbis||BS05, NB09-NB10, NB09-NB10B||3||0.2|
|Hemitripteridae||Nautichthys pribilovius||CC08, SR07||4||0.3|
|Liparidae||Careproctus reinhardti||B14, M07, SR11||6||0.5|
|Liparis fabricii||B14, BB06, BS05, BS08, M06, NB08, NB08B, NB09-NB10, SL01, SL02, SL04, SL05, SR11, SR12, NB09-NB10B||51||4.2|
|Liparis tunicata||BS08, SL04, SL09, NB09-NB10B||5||0.4|
|Psychrolutidae||Cottunculus microps||M06, M07||11||0.9|
Bathyraja parmifera (Bean,1881)
Three specimens were collected from 3 stations to the south of St. Lawrence Is. at a depth of 40 m. This is close to the northernmost record for B. parmifera. In addition,egg pouches were collected at Chukchi slope stations (M06 and M07) at depths of 269—626 m. This is the 1st record of its spawning ground,and the northernmost record has significant biological meaning.
Clupea pallasii Valenciennes,1847
The otter trawl collected 2 specimens only from station NB08 of east of St. Lawrence Island at depths of 35—45 m (Table1).
This is probably a synonym of Clupea harengus Linnaeus,1758; however,that species is considered an Atlantic Arctic species. Genetic divergence between them needs to be elucidated to determine the species boundary.
Arctogadus glacialis (Peters,1872)
The French-type beam trawl collected only 10 individuals at station M06 in the northern Chukchi Sea at depths of 529 —626 m (Table1). It was accompanied by abundant Arctic cod,Boreogadus saida,in the 2 harvests.
Specimens were collected from 23 stations at depths of 28 m (station C05)—695 m (station M06) by all 3 types of bottom trawls (Table1). In addition,2 other specimens were collected at a temporal ice camp (173°32.36′W,86°50.05′N) by cage. Boreogadus saida was the 2nd mostabundant species collected during the expedition (179 individuals,14.6%).
Ammodytes hexapterus Pallas,1814
Only 2 specimens were collected at stations north of St. Lawrence Island and south of Bering Strait at stations BS02 and BS05 at depths of 28—34 m (Table1).
Gymnelus hemifasciatus Andriashev,1937
Eleven specimens were collected on the shelf of Bering Sea (stations B14,SL08,and SL09),and the Chukchi Sea (station SR07),at depths of 30—131 m.
Lycodes adolfi Nielsen & Fosså 1993
Twelve specimens were collected from station M06,at depths of 529—695 m in the northern Chukchi Sea (Table1). The 1st record on the Chukchi slope was in 2009. This was the 2nd time to confirm a distributional expansion. However,the depth range on the Chukchi slope was shallower than bathymetric records (1 371—1 880 m) in the Atlantic Arctic.
Lycodes mucosus Richardson,1855
Only 5 specimens were collected from stations at depths of 32—40 m between the south Bering Strait and St. Lawrence Is. (Table1).
Lycodes palearis Gilbert,1896
Fifty-five specimens were collected in a restricted area on the western margin of the Bering shelf at depths of 70—131 m (Table1).
Lycodes polaris (Sabine,1824)
Five specimens were collected at 4 stations at depth of 40— 171 m including the western St. Lawrence Island,north Bering Strait,eastern Chukchi,and off Barrow,Alaska.
Lycodes raridens Taranetz & Andriashev,1937
Nineteen specimens were collected from 5 stations including the Bering Sea,south Bering Strait,and Chukchi Sea (Table1).
The record from station SR09 extends the range more northerly to almost 72°N than an earlier record.
Lycodes sagittarius McAllister,1976
Nine specimens were collected from only 2 stations in the northern Chukchi Sea at depths of 269—626 m (Table1). The records extend northward to 75°N beyond the original reference in the Beaufort Sea (71°59.89′N,168°59.54′W,357 m depth) and are deeper[15, 16]. This is also the 1st record in the Chukchi Sea.
Lycodes seminudus Reinhardt,1837
Fifty-five specimens were collected from only 2 stations in the northern Chukchi Sea at depths of 269—626 m. Lycodes seminudus was previously recorded in the Atlantic Arctic and Beaufort Sea; however,this is the second record off the Chukchi Sea since 2009. According to collection localities,it is distributed in deeper water than other congeneric species.
Hippoglossoides robustus Gill & Townsend,1897
This was the most abundant species (191 individual,15.6%) of the expedition,and was collected at 20 stations including the Bering Sea,south Bering Strait,and Chukchi Sea at depths of 24 m (station NB08B)—131 m (station B14). Two specimens were collected at station SR09 (71°59.89′N,168°59.54′W),close to the northernmost limit of its distributional range.
Lepidopsetta polyxystra Orr & Matarese,2000
Limanda aspera (Pallas,1814)
Fifty (4.1%) specimens were collected at 10 stations at depths of 24—40 m.
Pleuronectes quadrituberculatus Pallas,1814
Twenty-five (2.0%) specimens were collected from stations south of St. Lawrence Is. at depths of 24—40 m.
Reinhardtius hippoglossoides (Walbaum,1792)
This species is abundant in Bering Sea from the early records,but only 1 specimen was collected from station M07 on the Chukchi slop at a depth of 269—382 m in this survey. This almost reaches its northernmost record.
Ulcina olrikii (Lütken,1877)
Thirty-eight specimens were collected at 14 stations including the Bering Sea,south Bering Strait,and Chukchi Sea at depths of 27—49 m.
Artediellus atlanticus Jordan & Evermann,1898
Eleven specimens were collected at only 1 station (M07) on the Chukchi slope at depths of 269—382 m. Normally distributed in the North Atlantic,it was recently recorded on the Chukchi slope in 2009.
Artediellus scaber Knipowitsch,1907
Sixty-four (5.2% in number) specimens were collected at 3 stations in the eastern Chukchi Sea at depths of 27—43 m. It was the 5th dominant species of the expedition.
Enophrys diceraus (Pallas,1787)
Three specimens were collected from 2 stations (CC08 and SL08) south of St. Lawrence Is. in the Bering Sea and in the eastern Chukchi Sea at depths of 27—40 m.
Gymnocanthus tricuspis (Reinhardt,1830)
Fifty-nine (4.8%) specimens were collected from 15 stations in the Bering Sea and Chukchi Sea at depths of 27— 55 m.
Hemilepidotus papilio (Bean,1880)
Seven specimens were collected from 4 stations in the Bering Sea and south Bering Strait at depths of 28—40 m.
Icelus spatula Gilbert & Burke,1912
Eighteen (1.5%) specimens were collected from 2 stations in the Bering Sea and south Bering Strait at depths of 28— 131 m.
Myoxocephalus scorpius (Linnaeus,1758)
There were 143 (11.7%) specimens collected at 8 stations in the Bering Sea,south Bering Strait,and eastern Chukchi Sea at depths of 24—49 m. It was the 3rd most abundant species of the expedition.
Triglops pingelii Reinhardt,1837
Only 1 specimen was collected from station (SL09) south of St. Lawrence Island at a depth of 40 m.
Eumicrotremus orbis (Günther 1861)
It is not common in the north Bering Sea. However,three specimens were collected from 3 stations (BS05,NB09-NB10,and NB09-NB10B) around St. Lawrence Is. in the Bering Sea and south Bering Strait at depths of 32— 37 m. This almost reaches its northernmost range
Nautichthys pribilovius (Jordan & Gilbert,1898)
Three specimens were collected from 2 stations (CC08 and SR07) in the eastern Chukchi Sea at depths of 27—31 m. The record from station SR07 is close to its northernmost range in the Chukchi Sea.
Careproctus reinhardti (Krøyer,1862)
Six specimens were collected from 3 stations on the Bering slope (station B14),in the eastern Chukchi Sea (station SR07),and on the Chukchi slope (M07) at depths of 131— 382 m. It seems to be an Arctic species; however,1 specimen collected at station B14 on the Bering slope (60°55.27′N,177°41.53′W) at a depth of 131 m could be the southernmost record.
Liparis fabricii Krøyer,1847
Fifty-one (4.2%) specimens were collected from 15 stations in the Bering Sea,south Bering Strait,and Chukchi Sea at depths of 24—626 m. The record from station B14 (60°55.27′N,177°41.53′W) on the Bering slope at a depth of 131 m extends its southernmost range in 2007 to the north of St. Lawrence Is. (64°30′N,170°26′W).
Liparis gibbus Bean,1881
Only 1 specimen was collected from southern St. Lawrence Is. (station SL03) at a depth of 40 m.
Liparis ochotensis Schmidt,1904
Only 1 specimen was collected at station (NB08; 62°39.52′N,167°20.52′W) southeast of St. Lawrence Is. at a depth of 29 m. This is almost the northernmost range of this species.
Liparis tunicatus Reinhardt,1836
Five specimens were collected from 4 stations between south of St. Lawrence Is. and the south Bering Strait at depths of 28—40 m were at its southernmost extent.
Cottunculus microps Collett,1875
Eleven (0.9%) specimens were collected from stations (M06 and M07) on the Chukchi slope at depths of 269— 626 m. One specimen collected from the Chukchi slope at depths of 227—236 was the 1st record from the western Arctic (Mecklenburg et al. 2011). Our records prove its range extends from the Atlantic Arctic to the Pacific Arctic.
Anisarchus medius (Reinhardt,1837)
Sixteen (1.3%) specimens were collected at 5 stations on the Bering slope and in the Chukchi Sea at depths of 37— 169 m.
Eumesogrammus praecisus (Krøyer,1836)
Fifteen (1.2%) specimens were collected from 3 stations south of St. Lawrence Is. to the south Bering Strait at depths of 32—40 m.
Leptoclinus maculatus (Fries,1838)
Six specimens were collected from 2 stations on the Bering slope and in the Bering Sea at depths of 70—131 m.
Lumpenus fabricii Reinhardt,1836
One hundred (8.2%) specimens were collected from 6 stations in the Bering Sea,south Bering Strait,and Chukchi Sea at depths of 24—70 m. It was the 4th most abundant species of this expedition.
Stichaeus punctatus (Fabricius,1780)
Twenty-five (2.0%) specimens were collected at station CC08 in the eastern Chukchi Sea at a depth of 27 m.
Chauliodus macouni Bean,1890
Only 1 specimen was collected at station (B07,with a depth of 3 873 m) in the Bering Sea by multinet middle-water trawl at a depth range of 300—500 m. This is the 1st record of this mesopelagic deep-sea fish in the Bering Sea and almost in the northernmost of its distribution.
3 Discussion The most abundant families sampled during the expedition using 3 types of bottom trawls and combining the data were the Cottidae,Pleuronectidae,Gadidae,Zoacidae,and Stichaeidae; in terms of the number of species,the Cottidae and Zoacidae each had 8 species,followed by the Pleuronectidae,Liparidae,and Stichaeidae with 5 species each. These results were similar to those of an earlier report in 2004. However,the ranking of the species abundances differed; the most 6 abundant species of our study were Hippoglossoides robustus (vs. 3rd in 2004),Boregadus saida (2nd vs. 4th),Myoxocephalus scorpius (3rd vs. 2nd),Lumpenus fabricii (4th vs. 7th), Artediellus scaber (5th,the same),and Gymnocanthus tricuspis (6th vs. 1st). In addition,the 4 most abundant species comprised half of the fish specimens,which comprised 79% in 2004.
3.1 Numbers of families and species did not differ among the 3 different fishing methods,but the abundant species varied by method Using 3 types of bottom trawls,7,21 and 7 trawls were conducted with the French-type beam trawl,otter trawl,and triangular trawl,respectively. In total,283 individuals belonging to 22 species in 9 families were collected with the French-type beam trawl,659 individuals belonging to 28 species in 11 families with the otter trawl,and 283 individuals belonging to 21 species in 9 families with the triangular trawl. Even though the numbers of families and species collected with the otter trawl were slightly higher than French-type beam trawl and triangular trawl,this was because more-intensive collection efforts were made with the otter trawl. Among them,the abundant species varied with the different fishing methods. The 3 most abundant species collected with the French-type beam trawl were Lycodes seminudus (55 individuals,mainly at stations M06 and M07 on the Chukchi slope),B. saida (54 individuals),and L. palearis (37 individuals); with the otter trawl,they were H. robustus (168 individuals),B. saida (116 individuals),and L. fabricii (93 individuals); with the triangular trawl,they were M. scorpius (126 individuals,mainly at station CC08 with 104 specimens),Artediellus scaber (62 individuals),and Stichaeus punctatus (25 individuals). In addition,the dominant species by French-type beam trawl were mainly collected at stations with deeper depth (M06,M07,and B14); by otter trawl were mainly at stations on north Bering Shelf (NB08,NB09,and NB10); by triangular trawl were mainly at station CC08 in eastern Chukchi Sea. Reasons to the different dominant species by fishing gear were mainly the lake of experimental design due to the unexpected weather situations,and the insufficient sampling efforts on the wide geographic range distribution on Arctic fishes. The average species numbers in catches were 5.6 (with the French-type beam and otter trawls) and 5.7 (with the triangular trawl). However,the average individual numbers were a little lower with the otter trawl (32.6 individuals per catch) than the others (40.4 individuals in both).
3.2 Species numbers and abundances decreased with depth and latitude Our results indicate decreasing trends in species numbers and abundances with depth with the otter trawl; however,the trend was not clear with either the French-type beam trawl or triangular trawl. In addition,numbers of species and individuals at depths of 20—40 m with the otter trawl were higher than the others due to greater sampling efforts as mentioned earlier. A decreasing trend in species number with depth was also observed at Russian-American Long-term Census of the Arctic stations; however,a fish abundance trend was not seen.
3.3 Extension of previously documented geographic ranges was evident Among the 41 fish species collected during the expedition,17 species had a North Pacific to Arctic region distribution; the remaining 24 species had a Arctic-Boreal distribution. Geographic range extension was observed and could be classified into the following types: northward extension,southward extension,extension from the Atlantic Arctic to the Pacific Arctic,and a significant range extension from historical documentation. In total,10 species,including L. raridenc and L. sagittarius in the Zoarcidae; H. robustus,L. polyxystra,and R. hippoglossoides in the Pleuronectidae; B. parmifera in the Bathyrajiidae; E. orbis in the Cyclopteridae; N. pribilovius in the Hemitripteridae; L. ochotensis in the Liparicae; and C. macouni in the Stomiidae,exhibited a northerly extension beyond their known geographic range or were found at the northernmost limit of their range. However,3 species belonging to the Arctic Boreal,including C. reinhardti,L. fabricii and L. tunicata in the Liparidae,contrarily exhibited a southerly range extension. In addition,for 2 species formally known only to be distributed in the Atlantic Arctic until recently,a trans-Arctic distribution from the Atlantic Arctic to the Pacific Arctic was documented . The mechanism of this trans-Arctic distribution is poorly known. In addition,L. seminudus extended its known western border from the Beaufort Sea to the Chukchi slope.
4 Conclusions Fishes collected during the CHINARE-Arctic in 2010 Arctic expedition included 41 fish species belong to 14 families in 7 orders. The most abundant groups (families) were the sculpins (Cottidae),Bering flounder (Pleuronectide),Arctic cod (Godidae),eelpouts (Zoacidae),and eelblennys (Stichaeidae). The 6 most abundant species were Hippoglossoides robustus,Boregadus saida,Myoxocephalus scorpius,Lumpenus fabricii,Artediellus scaber and Gymnocanthus tricuspis. These results are similar to earlier reports [12, 14]. However,abundant species varied by fishing methods; numbers of families and species recorded did not differ with the fishing method; species numbers and abundances decreased with depth and latitude; and some species were observed to have extended their known geographic range. We could not clarify a connection between range extension and the changing climate in the Arctic region. Further long-term investigation is needed because of a lack of collection efforts of the Arctic benthic fishes.Acknowledgments: This work was supported by the China Program for International Polar Year 2007—2011 and the Special Research Foundation for Public Welfare Marine Program (Grant no. 201105022-2). Samples Information and Data issued by the Resource-sharing Platform of Polar Samples (http://birds.chinare.org.cn) maintained by Polar Research Institute of China (PRIC) and Chinese National Arctic & Antarctic Data Center (CN-NADC). We thank captain Shen Quan and all crew aboard the R/V XUE LONG icebreaker for security and logistical support during the expedition. Grants were received from the Third Institute of Oceanography, State Oceanic Administration, Xiamen and the Biodiversity Research Center, Academia Sinica, Taipei.
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