BRYOZOAN DIVERSITY IN THE CHUKCHI SEA 1 Historical development of research and current state of bryozoan diversity in the Chukchi Sea Nina V. Denisenko,* and Piotr Kuklinski# § * Department of Marine Research, Zoological Institute of the Russian Academy of Sciences, Saint-Petersburg, Russia. (ndenisenko@zin.ru) # Institute of Oceanology, Polish Academy of Sciences, ul. Powstancow Warszawy 55, Sopot 81-712, Poland. § Natural History Museum, Cromwell Road, London SW7 5BD, UK. 1. Introduction 2. Pioneer studies by Swedish and Norwegian researchers: 1800s to early 1900s 3. Studies by American scientists: 1913–1978 4. Studies by Russian scientists: 1910–2001 5. Recent studies of bryozoans in the Chukchi Sea: 1995–present 6. Current state of bryozoan diversity in the Chukchi Sea with some remarks 7. Acknowledgements 1. Introduction The Chukchi Sea is a marginal sea of the Arctic Ocean between Chukotka and Alaska. The sea is named after the Chukchi people, who are indigenous inhabitants on its shores. It is bounded by Wrangel Island on the west and Beaufort Sea on the east (Figure 1). It is connected to the Bering Sea and the Pacific Ocean by the narrow (approximately 92 km wide) and very shallow (depth of 30–59 m) Bering Strait. Because of its high latitude position and very shallow character (more than half of the total area is less than 50 m) the sea is covered by ice for long periods of time and is navigable for only four months of the year between July and October. Chukchi Sea is an extremely interesting and important area for several reasons. It is one of the most abundant marine ecosystems in the world owing to nutrient-rich waters, supplied from the south 1. This makes the area an important source of nutrients, heat and freshwater for the nearby Arctic Ocean. Arctic marine benthic communities are composed of species of Pacific and/or Atlantic origin as well as endemics.2, 3 The last major glaciation of the Arctic occurred approximately 18,000 years ago and ended 6000 years ago. Ice sheets covered much of the Arctic continental shelf and only the present areas of the Chukchi, Beaufort Seas and East Siberian Seas were largely unglaciated yet almost 2 ANNALS OF BRYOZOOLOGY 2 Figure 1. Location of Chukchi Sea within the Arctic. entirely emergent.4 This resulted in the recolonisation of many parts of the Arctic shelves by benthic fauna and flora over the last 6000–14000 years.5 Thus, the Arctic is considered to be a geologically young environment, which is still being colonised by marine organisms from lower latitudes.6 The Chukchi Sea, because of its link to Pacific Ocean via Bering Strait, can be thought of as a gateway to the Arctic Ocean. All colonizers of the Arctic of Pacific origin first have to establish themselves in this region in order to progress with colonization of more distant areas. Consequently, knowledge about the biodiversity of this area is very important and can provide us with the answers to many scientific questions, particularly with respect to the origin and evolution of Arctic biota. This historical compilation presented here focuses on bryozoans, one of the most diverse rocky bottom organisms of the Chukchi Sea and in general of the Arctic. In spite of the Bryozoa being one of the most species-rich groups in the Chukchi Sea, this group has not been the focus of many investigations. Such investigations as have been undertaken, have over one hundred years of history. Yet, our knowledge of species diversity in this sea is still far from being complete. The first effort to collect bryozoans of the Chukchi Sea was undertaken by the Swedish expedition on R/V Vega in 1878–1889. Later in the 20th century, studies on bryozoans BRYOZOAN DIVERSITY IN THE CHUKCHI SEA 3 Figure 2. Anton Stuxberg. have been carried out by mainly due to the efforts of both Russian and American scientists. The nationality of these latter scientific expeditions reflects the geopolitical structure of the area. Coastal waters of the Chukchi Sea belong territorially to both Russia and the United States of America. In the main part of the last century, studies by US and Russian scientists were conducted independently. American bryozoologists studied coastal areas of the Northern coast of the Alaskan Peninsula. Russian expeditions covered coastal areas of the Siberia, the Bering Straight area and the eastern and central parts of the sea. In the beginning of the 21st century the efforts of Russian and American researchers were combined to study of this important area, focusing on the southern part of the Chukchi Sea. This study aims at presenting a brief historical account of bryozoan studies undertaken in the area. More importantly we present current knowledge of bryozoan diversity in the Chukchi Sea, combining results of both Russian and American scientist as well as additional new data. 2. Pioneer studies by Swedish and Norwegian researchers: 1800s to early 1900s Swedish zoologist Anton Stuxberg (Figure 2) provided the first published information about bryozoans from the Chukchi Sea. This first record was the result of his research conducted during the Vega (Figure 3) expeditions along the Siberian Coast of the Chukchi Sea between 1875 and 1878, led by the famous Swedish explorer Nordenskiöld. Unfortunately, only a small part of the material from the Chukchi Sea was identified by him. In his comprehensive publication7 he mentioned only Alcyonidium mamillatum Alder (st. 96, 97) from the area of Chukchi Sea. The remaining bryozoan samples from Vega expedition were delivered to the Zoological Institute in Saint-Petersburg (Russia) where they were subsequently identified by the scientist/taxonomist German Kluge. A list 4 ANNALS OF BRYOZOOLOGY 2 Figure 3. R/V Vega. of his determinations was published in 19528 containing 39 species. The species were collected at eight stations from the Vega expeditions and all of them were located near the Siberian coast at a depth range of one to eight metres (Figures 4, 5). Six species belonged to order Cyclostomata, 25 species to Cheilostomata, and two species to Ctenostomata (Table 1). Figure 4. Map of benthos stations sampled by Swedish expedition on R/V Vega in 1878– 1879, Canadian Arctic Expedition (1913) and Norwegian Polar Expedition on R/V Maud in 1922 (after Ushakov37). Figure 5. Locations in Chukchi Sea where bryozoan individuals were found during Russian expeditions between years 1911 and 1938 (after Kluge38). BRYOZOAN DIVERSITY IN THE CHUKCHI SEA 5 Figure 6. Maud – the ship used by Norwegian Polar Expedition between years 1918 and 1925. In 1929 Nordgaard9 published further lists of bryozoans collected by Norwegian polar expeditions on board the vessel Maud (1918–1925) (Figure 6). Although the scientific effort of this expedition was not small (see Figures 4, 5, 7) he recorded only three species from the northern part of the Chukchi Sea: Eucratea loricata L., Serratiflustra serrulata Busk and Myriozoella crustaceum (Smitt). 2. Studies by American scientists: 1913–1978 New data about bryozoan fauna of this region appeared more than 30 years after Stuxberg’s publication. In 1923,10 Osburn (Figure 8) published the results of collections by Canadian Arctic Expeditions (1913–1918) in the American sector of the Chukchi Sea and at one station near the Siberian coast. Twenty two bryozoan species were identified along the American coastal sector: five species of cyclostome, one ctenostome, and 16 cheilostomes. Tubulipora flabellaris (Fabricius), Crisia eburnea (L., 1758), Crisia denticulata (Lamarck), Entalophoroecia deflexa (Couchin), Eucratea loricata (L.), Tegella armifera (Hincks), Bidenkapia spitsbergensis (Bidenkap), Cauloramphus cymbaeformis Hincks, Caberea ellisi Fleming, Scrupocellaria scabra (Van Beneden), 6 ANNALS OF BRYOZOOLOGY 2 Figure 7. Route of the Maud in the Chukchi Sea area during 1918–1921 (A) and 1922–1925 (B) (after Sverdrup39). Cystisella saccata Busk, Rhamphostomella bilaminata (Hincks), Hippoporina reticulatopunctata (Hincks), Celleporina incrassata (Lamarck) were found opposite the Icy Cape. Securiflustra securifrons (Pallas) and Arctonula arctica (M. Sars) were recorded near the Cape Sabine. Other species were found close to the north-western coast of Alaska (69˚ 40’ N, 163˚ 10’ W); Fungella dalli Kluge, 1955, Eucratea loricata (L.), Dendrobeania murrayana (Johnston) and Celleporella hyalina (L.). Only four species were listed (Eucretea loricata L., Carbasea carbasea (Ellis and Solander), Smittina bella (Busk) and Myriapora subgracilis (D’Orbigny) at Cape Franklin, located near Point Barrow. In a later publication, Osburn11 remarked that the Arctic bryozoan fauna on the northern coast of Alaska (near Point Barrow) were considerably more diverse than previously recorded. He identified 110 species of bryozoans (Table 1), collected by Professor G.E. MacGinitie from the Arctic Research laboratory at Point Barrow. Identifications of the material were presented in the famous monographs published by Osburn in 1950, 1952 and 1953.12-14 Among the 110 species, six were new to science (Tubulipora hastingsae, Plagioecia ambigua, Euritina arctica, Bidenkapia spitsbergensis alaskensis, Hincksina gotica, Parasmittina alaskensis). Based on his studies of the Arctic-American area, Osburn15 concluded that the bryozoan fauna of this region share an affinity with both with Atlantic and Pacific Bryozoa. Moreover, he noticed that some species have a circumpolar distribution. In 1978 a few new records of bryozoans were added by Feder and Jewett.16 During that 7 BRYOZOAN DIVERSITY IN THE CHUKCHI SEA Figure 8. Raymond Osburn. Figure 9. German Kluge. investigation seven bryozoan taxa were found (Kotzebue Sound, located to the north of the Seward Peninsula) however, only three species were identified to species level: Alcyonidium disciforme Smitt, Alcyonidium vermiculare Okada and Flustrellidra gigantea Silen. The presence of both Pacific (A. vermiculare) and Arctic species (A. disciforme) established as a fact the influence of both Arctic and Pacific biotas on the origin of bryozoan fauna of the Chukchi Sea. 3. Studies by Russian scientists: 1910–2001 The most detailed and comprehensive bryozoan study in the Chukchi Sea and in the Arctic as a whole was carried out by the Russian bryozoologist G. Kluge (Figure 9). The material used for his investigations was collected from various localities in the Chukchi Sea during eight Russian expeditions between 1910 and 1938 (Figure 5). His broad investigation of the area resulted in a taxonomic count of 125 species and subspecies inhabiting the Chukchi shelf area, of which 16 were new to science. These were Borgella tumulosa, Boverbankia composita, Tegella inermis, T. arctica var. retroversa, T. anguloavicularis, Callopora amisavicularis, C. obesa, Dendrobeania orientalis, D. pseudolevinseni, Bugulopsis peachi var. beringi, Hippoporella fastigiatoavicularis, Smittina tuberosa, S. beringia, Schizoporella stylifera var. perforata, Hippodiplosia tchukotkensis, Porella tumida, Cheilopora sincera var. orientalis, Celleporina nordenskjoldi. In his study,17 Kluge mentioned, that the presented species list of the Chukchi Sea is most likely not complete. As he pointed out this was mostly due to low sampling effort (not all samples in Kluge’s disposal were being treated). His suggestion proved to be right. In studies of the bryozoan fauna from the American side of the Chukchi Sea conducted by Osburn18-21 a few new species records previously not known from the area were added. Kluge was not familiar with Osburn’s investigations due to later receipt of his publications into Russian libraries. As a result Osburn’s data are not included in the 8 ANNALS OF BRYOZOOLOGY 2 most comprehensive monograph of Arctic bryozoans Bryozoa of the Northern Seas of the USSR, the posthumous work of Kluge.22 In the first half of the last century, a large number of Russian expeditions were carried out in the Chukchi Sea. Investigations of different systematic groups in the collected material, including bryozoans, confirmed a strong influence of Arctic Seas and the Pacific Ocean on the Chukchi Sea fauna.23 In 1975 benthic samples were collected along the Siberian Coast by the expedition conducted on vessel R/V Academician Korolev. The expedition worked in offshore waters. In 1976 a voyage of Zoological Institute of the Russian Academy of Sciences led by A.N. Golikov also undertook biological studies in the Chukchi Sea. Benthic samples were collected by scuba divers in shallow waters around Wrangel Island and in the Roger Inlet. In 1989 a large expedition on R/V D. Laptev provided biological samples from the open part of the Chukchi Sea. Strong influence of Arctic Seas and the Pacific Ocean on the Chukchi Sea bryozoan fauna was confirm during studies of distribution and their biogeography in the Arctic region (based on Kluge’s data) conducted by Gontar and Denisenko24 as well as Gontar.25 Unfortunately, as in Kluge’s case26 (see above) Osburn’s data27 from the Alaskan coast was not taken into account during preparation of two mentioned contemporaneous studies.28, 29 4. Recent studies of bryozoans in the Chukchi Sea: 1995–present The first international expedition with participation of both Russian and American scientists was held on the American ship, R/V Alpha-helix, in 1995. Specimens of bryozoans collected during those trips were identified by V. Gontar at the beginning of 2000s. The results of her identifications are kept in an archive at the Zoological Institute. In 2004, a collaborative project between Russian (Zoological Institute, St Petersburg) and American (NOAA) scientists conducted shared investigations in the Chukchi Sea. This project will continue until 2012. At the present, the material is in the process of being sorted and bryozoan identifications are still not completed. A map with recent sampling effort by those expeditions is presented on Figure 10. 5. Current state of bryozoan diversity in the Chukchi Sea with some remarks A survey of the literature revealed the known bryozoan fauna of Chukchi Sea to be 165 species (20 cyclostomes, 12 ctenostomes, and 133 cheilostomes) (Table 1). However, current knowledge of bryozoan diversity in the Chukchi Sea including additional studies of our own unpublished data and supplementary sampling revealed higher levels of diversity than previously recorded. At the present, a tally of the bryozoan fauna indicates that there are 189 species (26 cyclostomata, 15 ctenostomata, 148 cheilostomata) (Table 1). BRYOZOAN DIVERSITY IN THE CHUKCHI SEA 9 Figure 10. Sampling stations of various expeditions carried out by Russian and multinational expeditions between years 1976 and 2005. Numerous expeditions to the area collected bryozoan material sporadically without any special focus on bryozoan fauna. This resulted in low numbers of bryozoan being identified at sampling stations.30-33 Moreover, in many cases bryozoans were ignored or were identified only to phylum level.34 There are examples of investigations where bryozoans are listed but their identification was not determined by a specialist, and therefore may not be reliable.35 As a result of the limited study effort in Chukchi Sea, its physical isolation and the lack of information flow between Russian and American bryozoologists, some investigations have concluded that the bryozoan fauna of the area is impoverished in comparison to other Arctic regions.36 However, as this brief study indicates it might not be the case. This compilation has indicated that our knowledge about bryozoan fauna in the Chukchi Sea is still incomplete yet diversity is higher than previously thought. These results point out the need for taxonomic revision of many taxa with the use of a scanning microscope or even better with the additional use of molecular techniques, as this is likely to result in even higher number of species recorded in that area. This study also demonstrates that lack of communication among scientists working in isolation whether because of personal preference or political complexities, as in the case of previous studies in the Chukchi Sea may lead to biased conclusions. 10 ANNALS OF BRYOZOOLOGY 2 6. Acknowledgements The authors are grateful to the head of laboratory of marine research of Zoological Institute of the Russian Academy of Sciences Professor B.I.Sirenko and Dr V.I.Gontar for making possible usage of archive data of ZIN. We also would like to thanks Suzanne Williams for comments and corrections leading to improvement of the manuscript. The study has been completed thanks to funding provided by grant DYNARC to PK from European Commission’s programme “Structuring the European Research Area”. Table 1. Current knowledge of Chukchi Sea bryozoan species diversity. 1' -species identified by V. Gontar (archive data, 1" -species collected in the Chukchi Sea during “Vega” expedition in 1878-1879, * - species noted for Bering Strait only. † - species identified from recently collected samples. Species Osburn Osburn Kluge Feder & Gontar & Gontar (1923) (1955) (1952, Jewett Denisenko (2001) samples† 1962) (1978) (1989) Crisiella producta (Smitt, 1865) 1" Crisia arctica (M. Sars, 1863) 1 1" Crisia denticulata (Lamarck, 1816) 1 1 1" Crisia eburnea (L., 1758) 1 1 1 Crisia klugei (Ryland, 1967) Diplosolen obelia (Johnston, 1838) 1 Oncousoecia canadensis (Osburn, 1933) 1 Oncousoecia diastoporides Norman, 1869 1 Oncousoecia polygonalis (Kluge, 1915) 1 1 1 1 1 1 1 1 1 Tubulipora intermedia (Donoghue, 1923) 1 Tubulipora hastingsae Osburn, 1952 1 Tubulipora ventricosa Busk, 1875 Plagioecia grimaldi (Jullien, 1903) 1 1 Diplosolen obelia var. arctica Waters, 1900 Tubulipora flabellaris (Fabricius, 1780) Recent 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1" 1 1 1 1 1 1" 1 Plagioecia ambigua Osburn, 1953 1 Entalophoroecia deflexa (Smitt, 1866) 1 Lichenopora radiata (Audouin, 1828) Lichenopora canaliculata (Busk, 1876) 1 Patinella verrucaria (Fabricius, 1780) 1 Disporella crassiuscula (Smitt, 1867) 1" 1 Disporella hispida (Fleming, 1828) 1 Borgella (Borgiola) pustulosa (Osburn, 1953) 1 Borgella tumulosa Kluge, 1952 1 Fungella dalli Kluge, 1955 1 1 1 Fasciculiporoides americana (d’Orbigni, 1853) 1" 1 1 Alcyonidium disciforme Smitt, 1872 1 1 1 Alcyonidium gelatinosum (L.,1767) 1 1" 1 A. gelat. var. anderssoni Abrikossov, 1932 A. gelat. var. pachydermatum Kluge, 1962 1 1 1 1 1' 11 BRYOZOAN DIVERSITY IN THE CHUKCHI SEA Species Osburn Osburn Kluge Feder & Gontar & Gontar (1923) (1955) (1952, Jewett Denisenko (2001) samples† 1962) (1978) (1989) Alcyonidium mamillatum Alder, 1857 Alcyonidium mytili Dalyell, 1847 1 Alcyonidium enteromorpha Soule, 1951 1 Alcyonidium vermiculare Okada, 1925 1 1 1 1 1 1 1 1 1 1 1 1 Flustrellidra cervicornis (Robertson, 1900) 1 Flustrellidra corniculata (Smitt, 1872) 1 Flustrellidra gigantea (Silen, 1947) 1 Bowerbankia composita Kluge, 1952 1 1 1 Bowerbankia gracilis var. aggregata O’D, 1926 1 1 Vesicularia fasciculata (Soule, 1953) Eucratea loricata (L., 1758) 1" 1 Alcyonidium pedunculatum Robertson, 1902 Recent 1 1 1 1 1 Eucratea loricata var. arctica (Kluge, 1915) 1" 1 1 Eucratea loricata var. cornuta (Osburn, 1932) 1" 1 Eucratea loricata var. macrostoma (Ortmann, 1890) 1" 1 1" Electra arctica (Borg, 1931) 1 1 Carbasea carbasea (Ellis et Solander, 1886) 1 1" Carbasea nordenskjoldi (Kluge, 1929) 1" 1 1 1 1 1 1 1 1 1 1 Serratiflustra serrulata (Busk, 1880) 1 1" 1 1 Securiflustra securifrons (Pallas,1766) 1 1 1 1 1 Chartella membranaceotruncata (Smitt, 1868) 1 1 1 1 1 Hincksina gotica (Busk, 1856) 1 1" 1 1 Microporina articulata (Fabricius, 1821) 1 Callopora craticula (Alder, 1857) 1 Callopora lata (Kluge, 1907) Callopora lineata (L., 1767) 1 Callopora obesa Kluge, 1952 Callopora aurita (Hincks, 1877) 1" 1 1 * 1 1 1 1 1" Cauloramphus spiniferum (Johnston, 1832) 1 Caulorhamphus intermedius Kluge, 1962 1 1 1 1 1 1 1 1 1 1 1 Tegella amissavicularis (Kluge, 1952) 1 1* Tegella anguloavicularis Kluge, 1952 1 1 1 Tegella arctica (D’Orbigny, 1850) 1 1 1 1 1" 1 1 * 1 Tegella inermis (Kluge, 1952) Tegella unicornis (Fleming, 1828) 1 1 1 1 1 1 1 1 1 1 1 1 1 Bidenkapia spitsbergensis var. alaskensis Osburn, 1950 1 Septentriopora nigrans (Hincks, 1882) 1 1 1 Tegella retroversa (Kluge, 1952) Bidenkapia spitsbergensis (Bidenkap, 1897) 1 1 Callopora weslawski (Kuklinski et Taylor, 2006) Tegella armifera (Hincks, 1880) 1 1 Callopora smitti Kluge, 1946 Cauloramphus cymbaeformis (Hincks, 1877) 1 1 1 1 1 1 1 Doryporella spatulifera (Smitt, 1868) 1 1 1 1 1 Amphiblestrum trifolium var. quadrata (Hincks, 1880) 1 1 1 1 1 1 1 1 Bugula tschkotskensis Kluge, 1952 Bugula pacifica (Robertson, 1905) 1 12 Species ANNALS OF BRYOZOOLOGY 2 Osburn Osburn Kluge Feder & Gontar & Gontar (1923) (1955) (1952, Jewett Denisenko (2001) samples† 1962) (1978) (1989) Dendrobeania fessa (Kluge, 1955) 1 Dendrobeania flustroides (Packard, 1863) 1" Recent 1 1 1 1 1 1 1 1 1 1 1 1 1 Dendrobeania pseudomurrayana Kluge, 1955 1 1 1 Semibugula birulai Kluge, 1929 1" 1 1 Dendrobeania fruticosa (Packard, 1863) 1 1 Dendrob. fruti. var. quadridentata (Loven, 1834)1 1 Dendrobeania levinseni (Kluge, 1924) Dendrobeania murrayana (Johnston, 1847) 1 1 1 Dendrobeania pseudolevinseni Kluge, 1952 Dendrobeania multiseriata (O’Donoghue, 1925) Caberea ellisi Fleming, 1780 1 1 1 Notoplites sibirica (Kluge, 1929) 1 Scrupocellaria arctica (Busk, 1855) 1 Scrupocellaria minor (Kluge, 1915) Scrupocellaria scabra (Van Beneden, 1848) 1 1 1 S. scabra var. paenulata Norman, 1903 1 Tricellaria gracilis (Van Beneden, 1848) 1 Tricellaria gracilis var. inermis Kluge, 1962 1 Tricellaria erecta (Robertson, 1900) 1 Tricellaria pribilofi (Robertson, 1905) 1 1 1 1" 1 1 1 1 1 1 Tricellaria beringia (Kluge, 1955) 1 Membraniporella crassicosta Hincks, 1888 1 Cribrilina annulata (Fabricius, 1780) 1 1 Cribrilina spitsbergensis Norman, 1903 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Plesiothoa expansa Dowson, 1859 1 1 1 1 Celleporella hyalina (L., 1767) 1 1" 1 1 C. reflexa (Dick, Grischenko, Mawatari, 2006) 1 1 Adeonelopsis tuberculata (Busk, 1854) 1" 1 1 1 1 1 1 1 1" 1 1 1 Ragionula rosacea (Busk, 1856) 1 Posterula sarsi (Smitt, 1868) Arctonula arctica (M. Sars, 1851) Umbonula patens (Smitt, 1868) 1 Escharoides jacksoni (Kluge, 1900) 1 Escharella dijmphnae Kluge, 1929 1 1 1 1 1 1 1" Escharella immersa (Fleming, 1828) Escharella levinseni Hayward, 1994 1 Escharella ventricosa (Hassall, 1842) 1 Escharella labiata (Levinsen, 1886) 1 Escharelloides cancellatum (Smitt, 1868) 1 Hincksipora spinulifera (Hincks, 1889) 1 Hemicyclopora polita (Norman, 1864) 1 Porella acutirostris Smitt, 1868 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Porella patula M.Sars, 1851 Porella belli (Dowson, 1859) 1 1 Tricellaria peachi (Busk, 1851) Hippothoa divaricata var. arctica Kluge, 1906 1 1 1 Tricellaria ternata (Ellis et Solander, 1786) 1 1 1 13 BRYOZOAN DIVERSITY IN THE CHUKCHI SEA Species Osburn Osburn Kluge Feder & Gontar & Gontar (1923) (1955) (1952, Jewett Denisenko (2001) samples† 1962) (1978) (1989) Porella compressa (Sowerby, 1806) 1 1 1 1 Porella concinna (Busk, 1854) 1 1 1 1 Porella smitti Kluge, 1907 Recent 1 1 1' Porella tumida Kluge, 1955 Porella proboscidea (Hincks, 1888) 1 Porella minuta (Norman, 1869) 1 Porella obesa Waters, 1900 1 * 1 1 1 1 Porelloides laevis (Fleming, 1828) 1 1' Cystisella fragilis (Levinsen, 1914) 1 1 1 1 1" 1 1 1 Cystisella beringia (Kluge, 1929) 1 1" 1 1 1 Lepraliella contigua (Smitt, 1868) 1 1 1 1 1 Lepraliella hippopus (Smitt, 1868) 1 1 1 1 1 * Cystisella saccata Busk, 1856 1 Hippoporella fastigiatoavicularia Kluge, 1952 Rhamphostomella bilaminata (Hincks, 1968) 1 1 Rhamphostomella sibirica Kluge, 1929 1" 1 1 1 1 1 1 Rhamphostomella costata Lorenz, 1886 1 1" 1 1 1 Rhamphostomella hincksi Nordgaard, 1906 1 1 1 1 1 Rhamphostomella ovata (Smitt, 1868) 1 1" 1 1 1 Rhamphostomella plicata (Smitt, 1868) 1 1 1 1 Rhamphostomella scabra (Fabricius, 1780) 1 1 1 1 1 1 1 1 1 1 * 1 1 Schizobrachiella stylifera (Levinsen, 1887) S. stylifera var. perforata (Kluge, 1952) Harmeria scutulata (Busk, 1855) Hippoporina harmsworthi (Waters, 1900) 1 Hippoporina murdochi Kluge, 1962 1 Hippoporina reticulatopunctata (Hincks, 1877) 1 1 1 1 1 1 1 1" 1 1 1 1 1 Cheiloporina sincera (Smitt, 1868) 1 1 1 C. sincera var. praelucida (Hincks, 1888) 1 Hippoporina tschukotkensis Kluge, 1952 Hippoporina pertusa (Esper, 1797) 1 Cheiloporina inermis (Busk, 1880) 1 Phylactella pacifica O’Donoghue, 1923 1 Stomachetosella sinuosa (Busk, 1860) 1 1 1 Stomachetosella limbata (Lorenz, 1886) 1 Stomachetosella cruenta (Busk, 1854) 1 Pachyegis princeps (Norman, 1903) 1 Myriapora orientalis (Kluge, 1929) 1 1 1" 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Myriozoella crustacea (Smitt, 1868) 1 1 1 Myriapora subgracilis (D’ Orbigny, 1852) 1 1 Stomachetosella hincksi Powell, 1968 Pachyegis groenlandica (Norman, 1894) 1 1 Myriozoella plana (Dowson, 1868) 1 Cylindroporella tubulosa (Norman, 1868) 1 1 1 1 1 1 1 1 1 1 1 1 Smittina bella (Busk, 1860) 1 Smittina beringia Kluge, 1952 Smittina majuscula (Smitt, 1868) 1 1 1 1' 14 Species ANNALS OF BRYOZOOLOGY 2 Osburn Osburn Kluge Feder & Gontar & Gontar (1923) (1955) (1952, Jewett Denisenko (2001) samples* 1962) (1978) (1989) Smittina minuscula (Smitt, 1868) 1 1 1 Smittina mucronata (Smitt, 1868) 1 1 1 1 Smittina tuberosa Kluge, 1952 1 Smittoidea propinqua (Smitt, 1868) 1 1 Parasmittina jeffreysii (Norman, 1903 ) 1 Parasmittina trispinosa (Johnson, 1838) 1" 1" Parasmittina alaskensis Osburn, 1952 1 Euritina arctica Osburn, 1950 1 1' 1 1 1 1 Schizomavella lineata (Nordgaard, 1896) 1 Schizomavella porifera (Smitt, 1868) 1 Microporella ciliata (Pallas, 1766) 1 1 1 Microporella germata (Dick et Ross, 1988) 1 Microporella arctica (Norman, 1903) 1 Lepralioides nordlandica (Nordgaard, 1905) Cellepora nordenskjoldi Kluge, 1929 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1" 1 Celleporina ventricosa Lorenz, 1886 Buffonellaria biaperta (Smitt, 1873) Celleporina incrassata (Lamarck, 1856) Recent 1 1 Notes 1 J. M. Grebmeier, L. W. Cooper, M. H. Feder, B. I. Sirenko, 2006. Ecosystem dynamics of the Pacific influenced Bering and Chukchi Seas in the American Arctic. Progress in Oceanography, 71, (2006), 331-361. 2 A.N. Golikov and O.A. Scarlato, Evolution of Arctic Ecosystems during the Neogene Period. In: The Arctic Seas. Climatology, oceanography, geology and biology. Y.Herman (ed.). (New York: Van Nostrand Reinhold Company, 1989), 257-279. 3 K. H. Dunton, Arctic biogeography: the paradox of the marine benthic fauna and flora. Trends in Ecology and Evolution, 7, (1992), 183-189. 4 Dunton, note 2 and references therein. 5 Dunton, note 2. 6 Dunton, note 2. 7 A. Stuxberg, Die evertebraten – fauna des Sibirischen Eismeeres. In: Die Wissenschaftlichen Ergerbnisse der Vega-Expedition. Leipzig: Brockhaus, (1883), 481-600. 8 G.A. Kluge, Bryozoans (Bryozoa) of the Chukchi Sea and the Bering Stright. In: Far North-East of the USSR. Vol. 2. Fauna and flora of the Chukchi Sea. (Leningrad: Academy of Sciences of the USSR, 1952), 138-168. (in Russian) 9 O. Nordgaard, Bryozoa. The Norwegian North Polar expedition with the “Maud” 1918-1925. Scientific results. (Bergen: A.S. John Griegs Bokthrykkeri, 1929), Vol. V. No 10, 12 p. 10 R.C. Osburn, Bryozoa. Report of the Canadian Arctic Expedition 1913-18. (FA Acland, Ottawa, VIII, 1923), 1-13. 11 R.C. Osburn, The circumpolar distribution of the arctic-alaskan Bryozoa. Essays in the Natural BRYOZOAN DIVERSITY IN THE CHUKCHI SEA 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 15 sciences in honor of Captain Allan Hancock. (Los Angeles: University of Southern California Press, 1955), 29-38. R.C. Osburn, Bryozoa of the Pacific Coast of America Part 1, Cheilostomata – Anasca. Allan Hancock Pacific Expeditions, 14(1) (University of Southern California, Los Angeles, 1950), 1-270. R.C. Osburn, Bryozoa of the Pacific Coast of America Part 2, Cheilostomata – Ascophora. Allan Hancock Pacific Expeditions, 14(2) (University of Southern California, Los Angeles, 1952), 271-611. R.C. Osburn, Bryozoa of the Pacific Coast of America Part 3, Cyclostomata, Ctenostomata, Entoprocta and Addenda. Allan Hancock Pacific Expeditions, 14(3) (University of Southern California, Los Angeles, 1953), 613-841. Osburn, note 10. H.M. Feder and S.C. Jewett, Survey of the epifaunal invertebrates of northern sound, southeastern Chukchi Sea, and Kotzebue Sound. Report of Inst. of Mar. Sci., Fairbanks: University of Alaska, No R78-1, (1978), 124 p. Kluge, note 8. Osburn, note 11. Osburn, note 12. Osburn, note 13. Osburn, note 10. G.A. Kluge, Bryozoans of the Northern Seas of the USSR. Opriedielitiel po Faune SSSR (Moscow, Leningrad.: Academy of Sciences of the USSR. 1962), 1-584. (in Russian) P.V. Ushakov, Chukchi Sea and its bottom fauna. In: Far North-East of the USSR. Vol. 2. Fauna and flora of the Chukchi Sea. (Leningrad: Academy of Sciences of the USSR, 1952), 5-84. V.I. Gontar and N.V. Denisenko, Arctic Ocean Bryozoa. In: The Arctic Seas. Climatology, oceanography, geology and biology. Y.Herman (ed.). (New York: Van Nostrand Reinhold Company, 1989), 341-371. V.I. Gontar, Bryozoa. In: List of species of free-living invertebrates of Eurasian arctic Seas and adjacent deep waters. Sirenko B.I. (ed). (St.Petersburg: Zoological Institute of the Russian Academy of Sciences, 2001), 115-121. Kluge, note 22. Osburn, note 11. Gontar and Denisenko, note 24. Gontar, note 25. Stuxberg, note 7. Osburn, note 10. Nordgaard, note 9. Kluge, note 8. for example Feder and Jewett, note 16. for example Feder and Jewett, note 16. Gontar and Denisenko, note 23. Ushakov, note 23. Kluge, note 8 H.U. Sverdrup, Results of astronomical observations. The Norwegian North Polar expedition with the “Maud” 1918-1925. Scientific results. (Bergen: A.S. John Griegs Bokthrykkeri, 1929), Vol. I. No 3, 1-24. 16 ANNALS OF BRYOZOOLOGY 2