современные проблемы изучения морского биологического
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;BHEH=BYFHJYlhfk3–19 M>D FHJKD:Y;BHEH=BY KH<J?F?GGU?IJH;E?FUBAMQ?GBY FHJKDH=H ;BHEH=BQ?KDH=HJ:AGHH;J:ABY1 ]:<:^jbZgh\ Bgklblml[bheh]bbfhjy><HJ:G<eZ^b\hklhd e-mail: inmarbio@mail.primorye.ru ClZlvyijbgylZdi_qZlb] H[km`^Zxlkyl_jfbgheh]bybr_klvij_^eh`_gguomjh\g_c[bheh]bq_kdh]hjZaghh[jZabyKbkihevah\Zgb_fkh\j_f_gguo^ZgguobjZkq_lh\ijh\_^_ghkjZ\g_gb_\b^h\h]h[h]Zlkl\ZblZdkhghfbq_kdh]hjZaghh[jZaby gZa_fguo ij_kgh\h^guo b fhjkdbo hj]Zgbafh\ D gZklhys_fm \j_f_gb hibkZgu hdheh feg \b^h\ gZa_fguobluk\b^h\\h^guohj]Zgbafh\G_kfhljygZ^ebl_evgmxbklhjbxbamq_gbyhibkZguebrvhdheh luk fhjkdbo \b^h\ ba dhlhjuo luk \b^h\ ij_^klZ\e_gu [_kiha\hghqgufb `b\hlgufb Ba lbih\ fgh]hde_lhqguo`b\hlguolbigZc^_g\fhj_ ba gbo ij_^klZ\e_gu bkdexqbl_evgh fhjkdbfb nhjfZfb Ij_^klZ\bl_eb lbih\ fgh]hde_lhqguo `b\hlguo gZc^_gu \ij_kguo \h^Zo b ebrv lbih\ \dexqZxl gZa_fguo h[blZl_e_c Ij_^klZ\bl_eb ^\mo lbih\ ± ij_kgh\h^gu_ Micrognathozoa b gZa_fgu_ Onychophora ± g_ gZc^_gu\fhjkdbo[bhlhiZoH[km`^ZxlkyjZkq_lufhjkdh]h[bheh]bq_kdh]hjZaghh[jZabydhjZeeh\uojbnh\ ijb[j_`guowdhkbkl_ffhjkdh]hfZdjh[_glhkZbf_chnZmgu>eydZ`^h]hlbiZfgh]hde_lhqguoijb\_^_gukh\j_f_ggu_^Zggu_ihdhebq_kl\mhibkZgguo\b^h\h[km`^Zxlkyijh]ghauhdhebq_kl\_h`b^Z_fuo\b^h\JZaghh[jZab_]em[hdh\h^gh]hfZdjh[_glhkZhp_g_ghgZmjh\g_feg\b^h\\f_chnZmg_ij_^iheZ]Zxlkyhdheh ±feg\b^h\ijbq_f[he__fegijb^_lkygZfhjkdbog_fZlh^JZkkfZljb\Zxlky]bihl_auh[tykgyxsb_ \ukhdh_\b^h\h_jZaghh[jZab_fhjkdh]h]em[hdh\h^gh]hfZdjh[_glhkZbf_chnZmgu Dexq_\u_keh\Z[bheh]bq_kdh_jZaghh[jZab_[bhjZaghh[jZab_fhjkdh_[bhjZaghh[jZab_fgh]hde_lhqgu_fZdjh[_glhkf_ch[_glhkieZgdlhgfhgblhjbg][bhjZaghh[jZaby Contemporary problems of the study of marine biological diversity. A. V. Adrianov (Institute of Marine Biology, Far East Branch, Russian Academy of Sciences, Vladivostok 690041) Terminology and six main levels of biodiversity are suggested and discussed. Species richness and taxonomic diversity of terrestrial, freshwater, and marine organisms are compared based on recent estimates. To date, about 1.5 million terrestrial species and 320 000 species of aquatic organisms have been described. Over more than 200 years of biological research, only about 280 000 marine species, including 180 000 marine invertebrates, have been described. Of 33 phyla of Metazoa, 31 phyla are found in the sea; of these, 13 are represented by exclusively marine forms. Seventeen metazoan phyla are found in fresh water, and 11 phyla include terrestrial organisms. Representatives of two phyla, freshwater Micrognathozoa and terrestrial Onychophora, are thought to be non-marine endemics. The estimates of marine biological diversity of coral reefs, near-shore environments, deep-water macrobenthos, and meiofauna are discussed. For each metazoan phylum, recent data on the number of described species are provided and prognoses of species richness are discussed. Diversity of deep-sea macrobenthos is estimated at approximately 25 million species; and of meiofauna, at about 20–30 million species, including more than 10 million species of marine nematodes. The hypotheses invoked to explain the high species diversity of deep-sea macrobenthos and meiofauna are summarized. (Biologiya Morya, Vladivostok, 2004, vol. 30, no. 1, pp. 3–19). Key words: biological diversity, biodiversity, marine biodiversity, Metazoa, macrobenthos, meiofauna, plankton, biodiversity monitoring. \Zgghf GZpbhgZevghc ZdZ^_fb_c gZmd b KfblkhgbZgkdbf bgklblmlhf \ <Zrbg]lhg_ G_heh]baf ihy\beky dZd khdjZs_gguc \ZjbZgl khq_lZgby [bheh]bq_kdh_ jZaghh[jZab_ bkoh^gh bkihevam_fh]h ebrv ^ey hibkZgby qbkeZ \b^h\ Lovejoy <ihke_^kl\bb ^ey hibkZgby qbkeZ \b^h\ [ueh \\_^_gh ihgylb_ \b^h\h_ [h]Zlkl\h beb h[beb_ Fiedler, Jain dhlhjh_ \ kh\j_f_gghfihgbfZgbbebrvh^gZbadhfihg_gl[bheh]bq_kdh]hjZaghh[jZabyDZkvygh\ J_amevlZlu \Zrbg]lhgkdh]h nhjmfZ him[ebdh\Zgu \ ] \k[hjgbd_ BioDiversity g_lhqgh_ pblbjh\Zgb_ dhlhjh]h b ^Zeh `bagv l_jfbgm biodiversity" L?JFBGHEH=BYKHKLHYGB?IJH;E?FU HP?GDBBIJH=GHAU L_jfbg [bhjZaghh[jZab_ g_kfhljy gZ hq_gv rbjhdh_ ijbf_g_gb_ \[bheh]bb wdheh]bb b ijbjh^hhojZgghciheblbd_kjZ\gbl_evghfheh^±_]hg_lgb\ wgpbdehi_^bb;jblZgbdZThe New Encyclopedia Britannica _ ba^ gb \ kh\j_f_gghf ;bheh]bq_kdhf wgpbdehi_^bq_kdhfkeh\Zj_ 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<_^v k_cqZk \k_o ba\_klguo \b^h\ ± gZk_dhfu_ ± a_e_gu_ jZkl_gby Baf_gblky eb lZdh_khhlghr_gb_\[m^ms_f" Kdhevdh `_ \b^h\ `b\uo hj]Zgbafh\ h[blZxl gZ gZr_c ieZg_l_" Ih jZaebqguf hp_gdZf gZ ieZg_l_ h[blZxl hl ± ^h feg \b^h\ l_ \gZklhys__ \j_fy hibkZgu hl ± ^h \ emqr_f kemqZ_ ± j_Zevgh]h[bhjZaghh[jZabyLovejoy, 1997; Stork, 1997). Wglhfheh]b hp_gb\Zxl j_Zevgh_ jZaghh[jZab_ gZk_dhfuo\±feg\b^h\Erwinijbbamq_gghklb kh\j_f_gghc wglhfhnZmgu f_g__ ± Erwin T.L., Wlh \iheg_ \hafh`gh mqblu\Zy kh\j_f_ggu_ f_lh^u ih^kq_lZ b kl_i_gv wg^_fbafZ wglhfhnZmgu ^h`^_\uo e_kh\ ijb dhlhjhf fgh]b_ \b^u gZk_dhfuo \b^hki_pbnbqgu ^ey [hevrbgkl\Z ba luk \b^h\ ljhibq_kdbo ^_j_\v_\ [he__ \k_o gZk_dhfuo jZklbl_evghy^gu_Erwin, 1997). Ijh]gha h`b^Z_fh]h [bhjZaghh[jZaby hl^_evguo lZdkhgh\`b\uohj]Zgbafh\kihabpbclZdgZau\Z_fuo :>JB:GH< 6 LZ[ebpZJZaghh[jZab_jZkl_gbcPlanta\h^hjhke_cAlgaebijhkl_crboProtozoa) G_dhlhju_\ukrb_ lZdkhgulbiu PLANTA* Fhj_ \kdh[dZo±dhebq_kl\hfhjkdbo \b^h\ Ij_kgu_\h^u \kdh[dZo±dhebq_kl\hij_kgh\h^guo\b^h\ KmrZ Dhebq_kl\hhibkZgguokh\j_f_gguo \b^h\\kdh[dZo± bkdhiZ_fuo + + + 270 000 +(50) +(1000) + 235 000 36 000 > 4000 – > 40 000 l. Cyanophyles l. Rhodophyles l. Cryptophyles l. Euglenophyles l. Dinophyles l. Bacillariophyles l. Chrysophyles l. Xanthophyles l. Phaeophyles l. Chlorophyles lZygnematophyles l. Charophyles FUNGI PROTOZOA* lMicrospora** l. Myxozoa** l. Apicomplexa** lRhyzopoda + +(4850) + + + + + + +(2000) + – – + +(36 000) + + + + + + + + + + + + + + + + + + (< 10 000) + + + + – – – – – – – – – – – – + + + + + + 2000 5000 200 800 2000 (+2000) 10 000 (+12 000) 2000 600 > 2000 8000 6000 80 70 000 > 45 000 (+40 000) 1000 1200 4000 2500 l. Foraminifera l. Radiolaria l. Ciliophora + + + – + – – + 10 000 (+30 000) 800 8000 l. Magnoliophyta ALGAE* Bklhqgbd^Zgguo h\b^h\hfh[bebb Palmer., 1997; Lambshead, Schalk, 2001 Coleman, 2001; Whitton, Rajakaruna, 2001 Palmer, 1997; Reaka-Kudla, 1997 Sand-Jensen, 2001 –"– –"– –"– –"– –"– –"– –"– –"– –"– –"– –"– Volk, 2001 Corliss, 1991; Coleman, 2001 Finlay, 2001 –"– –"– Vickerman, 1992 ihKdZjeZlhb^j FboZe_\bq, 2000 Finlay, 2001 Lynn, Corliss, 1991 LZdkhgumdZaZgu\u[hjhqgh LZdkhgij_^klZ\e_glhevdhiZjZablbq_kdbfb\b^Zfb mf_j_gguo wdki_jlh\ hp_gb\Zxsbo h[s__ jZaghh[jZab_ gZ ieZg_l_ gZ mjh\g_ feg \b^h\ Hammond, 1992; Stork ij_^klZ\e_g gZ jbkmgd_ Ih fg_gbxkbkl_fZlbdh\aZf_lgucijhp_glgh\uo\b^h\ihfbfhhkgh\guohljy^h\gZk_dhfuo^h[Z\yl]jb[u^h fegZ\hl\bjmkub[Zdl_jbb\kmff_g_ij_\ukyl feg\b^h\kh\j_f_ggZybamq_gghklv[Zdl_jbchp_gb\Z_lky hl ^h b f_g__ ijb dhebq_kl\_ ba\_klguo \b^h\ g_ [he__ luk Pace, 1999; Dunlap Qbke_gghklv\b^h\a_e_guojZkl_gbcihfg_gbxmf_j_gguo fh`_l m\_ebqblvky ihqlb \^\h_ Z lj_lv_ f_klh ihke_ gZk_dhfuo b ]jb[h\ aZcfml gZa_fgu_ o_ebp_jh\u_jZaghh[jZab_dhlhjuohp_gb\Z_lky\feg\b^h\ GZ wlhf nhg_ ijh]gha ^ey fhjkdbo lZdkhgh\ \u]ey^bl\_kvfZ i_kkbfbklbqgh jbk b ih fg_gbx fgh]bowdki_jlh\\h[s_f[ZeZgk_[bhjZaghh[jZabykhhlghr_gb_ gZa_fguo b fhjkdbo \b^h\ \k_o hj]Zgbafh\ [m^_l gZ mjh\g_ : 1 (Briggs Z ih^h[gh_ khhlghr_gb_lhevdh^eyMetazoa – 10 : 1 (Tickel, 1999; Williamson, 1999). D gZklhys_fm \j_f_gb ihfbfh luk \b^h\ [_kiha\hghqguo \ fhj_ hibkZgu luk \b^h\ \h^hjhke_c luk \b^h\ ijhkl_crbo luk \b^h\ ohj^h\uo\hkgh\ghfju[luk\b^h\\bjmkh\b[Zdl_jbc jbk ] Ih hp_gdZf jZaguo wdki_jlh\ kl_i_gv bamq_gghklbfhjkdbo[bhglh\\Zjvbjm_lhlm[Zdl_jbc^hmju[±gZb[he__bamq_gguofhjkdboh[blZl_e_c Kfh]mlebhgbkhklZ\blvdhgdmj_gpbxkmohimlguf \b^Zf" DZd wlh gb m^b\bl_evgh gh kh\j_f_ggu_ f_lh^ubih^oh^udbkke_^h\Zgbxfhjkdhc[bhlu \k_ `_^ZxlgZflZdmxgZ^_`^m JZaghh[jZab_ ieZgdl_jh\ g_\_ebdh <b^h\hc ihl_gpbZe [Zdl_jbc ^Zxsbo h[s_c ijh^mdpbb ieZgdlhgZihhp_gdZfwdki_jlh\g_ij_\ukblluk \b^h\ Hammond <_kv nblhieZgdlhg Fbjh\h]h hd_ZgZ g_kfhljy gZ h]jhfgmx qbke_gghklv b ijh^mdlb\ghklv ij_^klZ\e_g ebrv \b^Zfb h^ghde_lhqguo\h^hjhke_cij_`^_\k_]h^bZlhfh\ufbbi_jb^bg_yfbbwlZpbnjZfh`_lm\_ebqblvky\emqr_fkem;BHEH=BY FHJY lhf 30 1 2004 KH<J?F?GGU?IJH;E?FU 7 LZ[ebpZJZaghh[jZab_fgh]hde_lhqguo`b\hlguoMetazoa) Fhj_ <ukrb_lZdkhgu lbiuih^lbiu bdeZkku METAZOA Ij_kgu_ \h^u \kdh[dZo± F_chDhjZeeh- FZdjh[_glhk- IeZgdlhg dhebq_kl\h \ucjbn [_glhk ij_kgh\h^gu_\b^u guo\b^h\ + + + KmrZ Dhebq_kl\hhibkZgguokh\j_f_g- Bklhqgbd^Zgguo guo\b^h\\kdh[- h\b^h\hfh[bebb dZo±bkdhiZ_fuo + + + > 1 500 000 – – – 1 lPlacozoa + lPorifera lCnidaria + + + + – + – + +(150) +(30) – – 7000 11 000 deHydrozoa + + + + +(30) – 2700 deScyphozoa de. Cubozoa de. Anthozoa l. Ctenophora l Dicyemataria** l. Orthonectida** l. Xenoturbellida l. Plathelminthes + + + + + + – + + + + + – – + – + + – + + + – + – + – – – – – – – + – – – – – – – + 2000 17 6200 100 70 20 2 > 20 000 l Gnathostomulida l. Micrognathozoa + – – – + – – – – + – – 54 1 l. Rotifera + – + + + – 2000 l Acanthocephala** + + – 1100 l Nemathelminthes deNematoda deGastrotricha l Cephalorhyncha + + + + – – – + + + + + – – – – + + +(200) + + + – – > 20 000 20 000 430 567 dePriapulida deLoricifera deKinorhyncha + + + + – – + + + – – – – – – – – – 17 (+5) 80 150 de. Nematomorpha** l. Entoprocta + + – 320 + + + – + – 150 lMollusca + + + + +(6000) +(10 000) de. Aplacophora + + + – – – hl ^h 290 de Polyplacophora de Monoplacophora de. Scaphopoda de. Cephalopoda de. Gastropoda + – + + – – – – – – – – 850 20 + + + + + + – – + – + + – – +(6000) – – +(10 000) deBivalvia + + + – + – 350 660 (+10 000) hl40 000 ^h100 000 (+15 000) 20 000 ;BHEH=BY FHJY lhf 30 1 2004 Kmffbjh\Zghih ]jmiiZf Barnes, Harrison, 1991 Hooper, Soest, 2002 Kmffbjh\Zghih deZkkZf Barnes, Harrison, 1991 LZf`_ –"– –"– K_jZ\bg Orme et al., 2002 Kozloff, 1993 Israelsson, 1999 Caira, Littlewood, 2001 Lammert, 1991 Kristensen, Funch, 2000 Clement, Wurdak, 1991 Dunagan, Miller, 1991 Gardner, 2001 –"– Ruppert, 1991 :^jbZgh\FZeZoh\ LZf`_ –"– Adrianov et al., 2002 :^jbZgh\, FZeZoh\, 1999 Nielsen, Jesperson, 1997 Kmffbjh\Zghih deZkkZf Barnes, Harrison, 1994 Lindberg, 2001 –"– Orme et al., 2002 –"– Barnes, Harrison, 1994; Lindberg, 2001 Orme et al., 2002
