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Presentation by Catherine W. Mecklenburg T. Anthony Mecklenburg
The RUSALCA mission and detection of change in the
diversity and distribution of fishes
Lowell Wakefield Symposium Anchorage, Alaska 27 March 2013
Presentation by Catherine W. Mecklenburg T. Anthony Mecklenburg
The RUSALCA mission and detection of change in the
diversity and distribution of fishes
Lowell Wakefield Symposium Anchorage, Alaska 27 March 2013
RUSSIAN–AMERICAN LONG-TERM CENSUS OF THE ARCTIC
Cooperative program, started in 2003:
National Oceanic & Atmospheric Administration
& Russian Academy of Sciences
Objectives •Document the state of the Arctic climate and marine life
— particularly in regions of projected rapid change.
•Detect change in Arctic climate and marine life.
(RUSALCA)
Arctic Marine Fish Museum Specimens
Database, 2010 Catch localities for 10,285 catalog lots
Otter trawl stations, RUSALCA 2004, 2009, 2012
East Siberian Sea Beaufort Sea
Chukchi Sea
U.S. 200-mile limit
Chukchi Borderland
Collection localities for samples in the Arctic fish DNA barcode database (C.W. Mecklenburg, PI)
1, 649 CO1 sequences (barcodes)
~200 species
Greenland
Norway Russia
Russia
Alaska
Canada
Hudson Bay
Baffin Bay
Svalbard
Washington
SE Alaska
• Catherine W. Mecklenburg, Research Associate, California Academy of Sciences, San Francisco; Private Consultant, Point Stephens Research, Auke Bay, Alaska
• Peter R. Møller, Head of Vertebrate Zoology and Curator of Fishes, Natural History Museum of Denmark, University of Copenhagen
• Dirk Steinke, Lead Scientist, Barcoding of Marine Life, Biodiversity Institute of Ontario, University of Guelph, Ontario, Canada
Biodiversity of Arctic Marine Fishes: Taxonomy and Zoogeography
(Marine Biodiversity 41(1):109–140 + Online Resources 1–5 )
Mar Biodiv (2011) 4l:109-140
DOI 1 0. 1 007 1s12526-0 I 0-0070-z SENCKENBERG
and zoogeography
Catherine W. Mecklenburg . Peter Rask Msller
Dirk Steinke
Received: 3 June 2010/Revised: 23 September 2010/Accepted: 1 November 2010/Published online: I December 2010i() Senckenberg, Gesellschaft flir Naturforschung and Springer 2010
Biodiversity of arctic marine fishes: taxonomy
Abstract Taxonomic and distributional information on each
fish species found in arctic marine waters is reviewed, and a
list of families and species with commentary on distributional
records is presented. The list incorporates results from
examination of museum collections of arctic marine fishes
dating back to the 1830s. It also incorporates results from
DNA barcoding, used to complement morphological charac-
ters in evaluating problematic taxa and to assist in identifica-
tion of specimens collected in recent expeditions. Barcoding
results are depicted in a neighbor-joining tree of 880 COI
(cytochrome c oxidase 1 gene) sequences distributed among
165 species from the arctic region and adjacent waters, and
discussed in the family reviews. Using our definition of the
arctic region, we count 242 species with documented
presence, if 12 species that likely are synonyms are
excluded. The 242 species are distributed arnong 45 families.
This article belongs to the special issue "Arctic Ocean Diversity
Synthesis"
Electronic supplementary material The online version of this article(doi: I 0. I 007/s 12526-A I 0-0070-z) contains supplementary material,which is available to authorized users
C. W. Mecklenburg (X)
Department of Ichthyology, California Academy of Sciences,
mail: Point Stephens Research, P.O. Box 210307 , Auke Bay, AK
99821, USA
e-mail : ptsteph [email protected]
P. R. Msller
Natural History Museum of Denmark, University of Copenhagen,
Universitetsparken I 5,
2100 Copenhagen O, Denmark
D. Steinke
Biodiversity Institute of Ontario, University of Guelph,
50 Stone Road East,
Guelph, Ontario N1G 2Wl , Canada
Six families in Cottoidei with 72 species and five in
Zoarcoidei with 55 species account for more than half
(52.5%) the species. This study produced Col sequences for
106 of the 242 species. Sequence variability in the barcode
region permits discrimination of all species. The average
sequence variafion within species was 0.3% (range 0-3 .5oh),
while the average genetic distance between congeners was
4.7% (range 3.7-13.3%). The coL sequences support
taxonomic separation of some species, such as Osmerus
dentex and O. mordax and Liparis bathyarcticus and L.
gibbus,' and synonymy of others, like Myoxocephalus
verruco,stts in M. scorpius and Gymnelus lmipowitschi in
G. hemtfasciatus. They sometimes revealed the presence of
additional species that were not entirely expected, such as an
unidentified species of Ammodytes in the western Gulf of
Alaska, most likely A. personatus,' and an unidentified lcelus
species of the I. spatula complex with populations in the
western Gulf of Alaska and the northern Bering and Chukchi
Seas which could be a new species or a species in synonymy.
Reviewing distribution, we found that for 24 species thepatterns assigned by authors understated historical presence
in the arctic region, and for 12 species they overstatedpresence. For instanc e, Hippoglossoides robustus is counted
as an arctic-boreal species rather than predominantly boreal,
and Artediellus uncinatus as predominantly arctic rather than
predominantly boreal. Species with arctic, predominantly
arctic, or arctic-boreal distributions composed 4l% of the
242 species in the region, and predominantly boreal, boreal,
and widely distributed species composed 59%. For some
continental shelf species, such as the primarily amphiboreal
Eumesogrammus praecisus and Leptoclinus maculatus,
distributions appear to reflect changes, including reentry into
Arctic seas and reestablishment of continuous ranges, that
zoogeographers believe have been going on since the end of
land bridge and glacial times.
6 Springer
Arctic Region (green)
Mecklenburg, Møller, Steinke. 2011. Marine Biodiversity 41(1):109–140 + Online Resources 1–5
Head spines large, typically doubled or wedge-like
Head spines absent or small and single
Myoxocephalus scorpius Shorthorn Sculpin
Northern Bering Sea
Southeastern Gulf of Alaska
Myoxocephalus scorpius
Shorthorn Sculpin
arctic–boreal, circumpolar
Geographic distribution
Theragra finnmarchica Norway Pollock
(Koefoed 1956)
Theragra chalcogramma Walleye Pollock
(Pallas 1814)
Norway Pollock and Walleye Pollock are the same species�(e.g., Ursvik et al. 2007, Byrkjedal et al. 2008) �
Gadus chalcogrammus Pallas, 1814�Walleye Pollock�
Gadus chalcogrammus�Walleye Pollock�
predominantly boreal Pacific; also North Atlantic
392 mm TL
Chukchi Borderland: 1 at 365–370 m (74°07’N,166°01’W)
Overall depth range: Surface to 1,200 m Demersal & pelagic
Norway pollock
RUSALCA 2009 74°07’N, 166°01’W
Very close in appearance, yet they are different species (Chernova 2008)
Liparis bathyarcticus Nebulous Snailfish
Liparis gibbus Variegated Snailfish
Liparis bathyarcticus Nebulous Snailfish
Liparis gibbus Variegated Snailfish
Liparis fabricii Gelatinous Seasnail
Liparis tunicatus Kelp Snailfish
DNA sequences (barcodes) indicate 4 distinct species of Liparis are present in the Chukchi and Beaufort Seas
Myoxocephalus stelleri Frog Sculpin
Hippoglossoides robustus�
Bering Flounder�
Buchan Bay, 1965 Bathurst Inlet,1975 Eggs at
Long Strait 1934
East Siberian Sea RUSALCA 2009
arctic–boreal Pacific
Hippoglossoides robustus�
= H. elassodon�Flathead Sole�
(Kartavtsev et al. 2007
arctic–boreal Pacific
Hippoglossoides robustus�
= H. elassodon�Flathead Sole�
(Kartavtsev et al. 2007
arctic–boreal Pacific
Fish species in Arctic Region (n = 242 species)
Arctic 41%
Boreal 59%
Fish species in Arctic Region Fish species in Chukchi Sea
Arctic 41%
Boreal 59%
Arctic 40%
Boreal 60%
1939 (Andriashev 1939)
2011 (Mecklenburg, Møller, Steinke 2011)
Arctic 40%
Boreal 60%
Marine fish species in the Chukchi Sea
Boreal 54%
Arctic 46%
RUSALCA Bottom Trawl Stations & Chukchi Borderland
Bering Strait
Wrangel I.
100 m
2004, 2012 2009
Chukchi Cap 588 m Slope Sta 1 370 m
Slope Sta 2 236 m
Chukchi Borderland
Six species collected in the Chukchi Borderland are also found on the continental shelf
Liparis fabricii Gelatinous Seasnail
Reinhardtius hippoglossoides Greenland Halibut
Boreogadus saida Arctic Cod
Arctogadus glacialis Polar Cod
Gadus chalcogrammus Walleye Pollock
Leptoclinus maculatus Daubed Shanny
Six species were found only in the Chukchi Borderland
Careproctus reinhardti Sea Tadpole Artediellus atlanticus
Atlantic Hookear Sculpin
Triglops nybelini Bigeye Sculpin
Cottunculus microps Polar Sculpin Lycodes adolfi
Adolf’s Eelpout
Lycodes seminudus Longear Eelpout
Atlantic Water circulation in the Arctic, 200–800 m
Fram Strait
Barents Sea
arctic, mainly Atlantic, probably circumpolar
Lycodes adolfi Nielsen & Fosså, 1993
Adolf’s Eelpout
193 mm TL
Chukchi Borderland: 4 at 580–588 m
Overall depth range: 386–1,880 m Benthic
Yermak Plateau
RUSALCA 2009 76°33’N, 164°27’W
Artediellus atlanticus Atlantic Hookear Sculpin
arctic–boreal, mainly Atlantic, possibly circumpolar
Chukchi Borderland: 5 at 227–236 m 2 at 365–370 m
Overall known depth range: 11–1,366 m Benthic
142 mm TL
Pacific-Arctic Marine Fishes�A distributional atlas and identification guide to the
fishes of the RUSALCA study area�
(northern Bering Sea, East Siberian, Chukchi, and Beaufort Seas, and the adjacent deep waters of the Arctic Ocean)�
Mecklenburg and others�In preparation
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>D#">$)$-/M*40-*C"(#DG-*G")"C"/-*$,#>@G-/*C"(#DG-/*D%*!"#$%&'(''*%(D=*)0-*N"(-,)/*<-"*",G*)0-H*"(-*)0-*/"=-*"/*)0D/-*
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3,4,-,(+,5!4'-!!"#$%&#'&(&$%)"*&+($,$$!Andriashev AP (1954) Fishes of the northern seas of the USSR. Akad
Nauk SSSR, Zool Inst, Opredeliteli po Faune SSSR 53. [In Russian; transl 1964, Israel Prog Sci Transl] [pp 468–471 of transl.]
Andriashev AP (1986) Agonidae. In: Whitehead PJP, Bauchot ML, Hureau JC, Nielsen J, Tortonese E (eds) Fishes of the North-eastern Atlantic and the Mediterranean. Unesco, Paris, pp 1265–1268
Backus BH (1957) The fishes of Labrador. Bull Am Mus Nat Hist 113 (4):273–338
Bean TH (1885) Description of a new species of Aspidophoroides (A. güntherii) from Alaska. Proc US Natl Mus 8:74–75
Busby MS (1998) Guide to the identification of larval and early juvenile poachers (Scorpaeniformes: Agonidae) from the north-eastern Pacific Ocean and Bering Sea. NOAA Tech. Rep. NMFS 137
Frost KJ, Lowry LF (1983) Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort seas, 1976–77. NOAA Tech Rep NMFS SSRF–764
Hunter JG, Leach ST, McAllister DE, Steigerwald MB (1984) A distributional atlas of records of the marine fishes of Arctic Canada in the National Museums of Canada and Arctic Biological Station. Syllogeus (Ottawa) 52
Jensen AS (1942) Contributions to the ichthyofauna of Greenland 1–3. Spolia Zool Mus Haun 2 [pp 34–36].
Jordan DS, Evermann BW (1896) A check-list of the fishes and fish-like vertebrates of North and Middle America. Rep US Fish Comm 21, Append 5:207–584
Jordan DS, Evermann BW (1898) The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part II. US Natl Mus Bull 47: i–xxx, 1241–2183 [pp 2088–2091]
Kanayama T (1991) Taxonomy and phylogeny of the family Agonidae (Pisces: Scorpaeniformes). Mem Fac Fish Hokkaido Univ 38(1.2) [pp 86–87]
Lütken (1877) Korte bidrag til nordisk Ichthyographi. I. Foreløbige Meddelelser om nordiske Ulkefiske (Cottoidei). Videnskabelige
Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn, Aaret 1876-1877: 355-388 + French translation, pp 72-98
Lütken CF (1886) Et bidrag til kundskab om Kara-havets fiske. Pp. 117-154, Pls. 15-17 In: Dijmphna-togtets zoologisk-botaniske udbytte. Copenhagen. i-xxi + 1-515, Pls. 1-41. [As a separate, pp. 1-40, pls. 15-17, dated 1886. Main work may be 1887. In Danish.]
McAllister DE (1962) Fishes of the 1960 “Salvelinus” program from western Arctic Canada. Natl Mus Can Bull 185:17–39
Mecklenburg CW, Mecklenburg TA, Thorsteinson LK (2002) Fishes of Alaska. American Fisheries Society, Bethesda, Maryland [p 552]
Mecklenburg CW, Møller PR, Steinke D (2011) Biodiversity of arctic marine fishes: taxonomy and zoogeography. Mar Biodivers 41:109–140 [Online Resource 1:29–30]
Mecklenburg CW, Stein DL, Sheiko BA, Chernova NV, Mecklenburg TA, Holladay BA (2007) Russian–American Long-term Census of the Arctic: benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004. Northwest Nat 88:168–187
Møller PR, Nielsen JG, Knudsen SW, Poulsen JY, Sünksen K, Jørgensen OA (2010) A checklist of the fish fauna of Greenland waters. Zootaxa 2378:1–84
Nozères C, Archambault D, Chouinard P-M, Gauthier J, Miller R, Parent E, Schwab P, Savard L, Dutil J-D (2010) Identification guide for marine fishes of the estuary and northern Gulf of St. Lawrence and sampling protocols used during trawl surveys between 2004 and 2008. Can Tech Rep Fish Aquat Sci 2866
Rendahl H (1931) Fische aud dem östlichen Sibirischen Eismeer und dem Nordpazifik. Arkiv f Zool 22A(10): 1–81
Scott WB, Scott MG (1988) Atlantic fishes of Canada. Can Bull Fish Aquat Sci 219
Sheiko BA, Mecklenburg CW (2004) Family Agonidae Swainson 1839 — poachers. California Academy of Sciences Annotated Checklist of Fishes 30
Wienerroither R, Johannesen E, Dolgov A, Byrkjedal I, Bjelland O, Drevetnyak K, Eriksen KB, Høines Å, Langhelle G, Langøy H, Prokhorova T, Prozorkevich D, Wenneck T (2011) Atlas of the Barents Sea fishes. IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828
*
Collaborators & Contributors • IngvarByrkjedal,BergenMuseum,UniversityofBergen,Bergen,Norway
• JørgenS.ChrisKansen,DepartmentofArcKcandMarineBiology,UniversityofTromsø,Tromsø,Norway
• OlegV.Karamushko,MurmanskMarineBiologicalInsKtute,KolaScienceCentre,RussianAcademyofSciences,Murmansk,Russia
• ArveLynghammar,DepartmentofArcKcandMarineBiology,UniversityofTromsø,Tromsø,Norway
• PeterR.Møller,NaturalHistoryMuseumofDenmark,UniversityofCopenhagen,Copenhagen,Denmark
• DirkSteinke,BiodiversityInsKtuteofOntario,UniversityofGuelph,Guelph,Ontario,Canada
• NataliaV.Chernova,ZoologicalInsKtute,St.Petersburg,Russia
• BorisA.Sheiko,ZoologicalInsKtute,St.Petersburg,Russia
• BrendaA.Holladay,InsKtuteofMarineScience,UniversityofAlaskaFairbanks,USA
• MorganS.Busby,NOAA,AlaskaFisheriesScienceCenter,Sea\le,Washington,USA
• JamesW.Orr,NOAA,AFSC,Sea\le,Washington,USA
• WilliamN.Eschmeyer,CaliforniaAcademyofSciences,SanFrancisco,California,USA
• DannyKent,VancouverAquarium,BriKshColumbia,Canada
• CamillaO\esen‐Meyer,UniversityofTromsø,Norway
• ElenaVoronina,ZoologicalInsKtute,St.Petersburg,Russia
• DavidCatania,CaliforniaAcademyofSciences,SanFrancisco,CaliforniaUSA
• JohnFong,CaliforniaAcademyofSciences,SanFrancisco,CaliforniaUSA
• J.AndresLopez,MuseumoftheNorth,UniversityofAlaskaFairbanks,USA • KatherineP.Maslenikov,UniversityofWashington,FishCollecKon,Sea\le,Washington,USA
• ArcadyBalushkin,ZoologicalInsKtute,St.Petersburg,Russia
And many others