Fishing Down Marine Food Webs Daniel Pauly; Villy Christensen; Johanne Dalsgaard; Rainer Froese; Francisco Torres Jr. Science, New Series, Vol. 279, No, 5352 (Feb. 6, 1998), 860-863. Stable URL: hutp//links jstor.org/sici?sic!=0036-8075% 28 19980206%293%3A279%3A5352%3C860%3AFDMFW%3E2.0,CO%3B2-C Science is currently published by American Association for the Advancement of Science, ‘Your use of the ISTOR archive indicates your acceptance of JSTOR’s Terms and Conditions of Use, available at hhup:/www.jstororg/about/terms.hml. JSTOR’s Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at hup:/www jstor.org/journalsaaas.huml Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the sereen or printed page of such transmission, STOR is an independent not-for-profit organization dedicated to creating and preserving a digital archive of scholarly journals, For more information regarding JSTOR, please contact support @jstor.org. hupslwww jstor.org/ ‘Thu Aug 19 17:09:32 2008pean 20. 2 22 28 2. 26 2 20. 960 eR Le a RECS regent or 9 1446 Apa Se PAPZA Ps frag (15 Bn consucts overexpress Fip2to who iraslomod sto PRES ste Inedby Wlenemtiotng wih air atonal soos, ‘Thre tes ofeach sain, PRYTTS AYES wth GI-PA24 a0 ele PRP2A gone) aed PRYTT6 (Prvoe itn pOT-PRPZE.Pa 99 slo PRPZG (re, were hanested in at loprthmc pase Aol al seract as popared 6), and 160 m9 of each war subjected 0.3010 S58 anmeniom Sulfate procetaion The precates wero reas Bonded nab ml of AK 200 [10 mM Hopes pH 9), 200 eM KC. 1.8 mil MgCl, 10% gyearl 02m EDTA, 0.1 Trtan 10005 mANOTT. Initpnesjmetnyjeutory! org, ? mi beam ‘ina, pepe {Tai lwoepin (49 rinosbated vith 180 plot rote G-Sapharocs iat i AKC 200) ona rtator at 4° for 30 Min. (This top removed prea hat Bourn Spoatealy 1h rear THe suparatarts wore ‘25800 mcf prtein @-Sepneresecavlad fo ‘nt-poyoraambeies ead: AGK 200 = 1) 5m G2) Afr 15 row nutsing a €°, the Speratanie neve removes, a he beads were ‘saa 8 20 mie" GK 70D AK but ih TOOmMAKCI, 1» 18m ot AGK2ON, ar! 1S 9f AGK 50 fAGK Bor wh 80M KCI. The ‘wastes Boade wore eluted tco by nating at 25:0 tor 10 min wn 2 400 @ mar of paptige EVMPME (au Tyr Met Pro-Met ty 92) [GAGK 0, Each eluste woe Saeed sont 2%? Hereorbuiterb atch Shours The daeates ‘wer mieocorntuged 10 min athe ern: {anisworo colaciod. Approsimatey 15 ng. of Prp2e was covered, and no soRNAs coprog (Gar hese conations tha sanetty tot mo) ‘FUG no US wae detected In pmol of Prpae Beat. B.S. Horwitz ard J. Abelson, ares Oo. 7,320 fro. ‘Arya sling reacton miu may cartan ~3 Fad and “6M Us seRNAS, compare wth 02 {2OSrMpremPNA fcnothnomnnat racana {heart scompetnt er speg) Levels U6 {SRNR are conta over n PRY112 (19) Fp. Srepeenatty toa RP rojelng deacts mayo Fd. ine lt Bit Chom, 260, 4780 (195) U.Wayragtaan, Mt Conpary J. Ace, Genes ev. 1208 (1080; 5 Barcaues ana tN. Aa fn, Mal Cal 89,9710 (1089, 3.0. Bron and 0. Bagpe EMBO 1,272" 1602 Fr Geson iaP. agra Mica Ade Aes 21,1555 (1085, SW Ruby 74. rang, Ablson, Gone De. 7, 1805/1989). 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Gui, Gansies 149,67 1996 ‘EU et al, 2460. 45, 4001 (196) aL guint, tne, Unversty of San Fan (350 (1887, Ona ghee grader, he wast marty of PZ S31 comerate vith UB ans UU en PNP 3 avay fon tpn eoRNPs 2d epicaosoes (7) bite Prg2e i nl cotactdPrp rm restates tat contin U8, US. ane US SPAS (07), Tus, we have no eudence racating that Prgat sa component of UA UBS SARNS oo ‘he aplosocore “I aesice: RK. Gaur, R. Sgn, MR, Geen, Solon 273, 1708906, Iara, B. itner, N. Mth, N. Sone, roche, E660! 14, 98201065, KBr Semaser A LS & O'Shea Soence 286,122 1008, ha Note wd C. Que, EMBO J 18,4268 (996) F Berdona, J Baryeques, J Abdson ‘Gusta, Gores Do. 4, 85 00, Sits were srppedior eur at Cin OREN fd 009 sadn eta cartaring 2% SOS femove probes. Subsequent roohgs Wer per ‘omod a desorbed (3 Z bend. A Brow, Misbe Aes Ros 21,4685 esa, UNOS enANPs wera coimmuncerecitod wih pajoraagges G2 Ga Tre beure comple Sree ashed wt @ S00 plot NET 80 an hen Ineubstod for *0.mn a 220 wth 160 yo er (40% ter D, 2.5 mid Mig, 9 PE 8000, 6 Ink passur phoaprat. wh oF wet 2rd [AVP the aapematate we seprate fom te ead, anetne beads were washed wth 500 ‘INET 50. Far ig 4, tho samples wore cecty Ahegoterized fr nonaanatang gt says of Us Sea U8 s¥NAS.ForFg 4, Band, sorted pat was aaseato 1 plotaiparatartcomtaneg a . 3. 2, 2. [ATP (0. each i and Usa inst 2 frinat23'Cuness ohawae states Reactors ere ‘Soppad by te aaton of 200 of 08M eodurn Seale, 6 mid EDTA, ara 0.5% SOS one. PNAS ‘tere sce by pcan K cgeaton (oO) penal herotomatacion ans hana pects fer To dgetarize Ut and Us siNPs bole a nealing, 100 wl of superstar coxanng ATP was Ieee at 26 fr 90 min win 10mg ta te Koad Sgr washed mute D. The cep fenaed superstar’ was eae colectes ar reg reactors Wa thane, het and. Alon fr aceon Prgat pretar: ML Larbag, 6. ONal, and (Sea acon animus puriaton rower ne rt pelyora hyeraeralnC- Co: line and Roger for ven or sostanos; part land resent marberso! he Gh aberstn fot Gongucivectioam: and Scour, cco, ‘x rank, Hi. Maden, &, Rader, ©: Sioa Silay, Unenbece and K. Zr torcommaris ‘rhe manus. We we ndctea 10 Esperas, sow, andr. Rone tor mats tegen asstance PLLA, was supports by a Howard Flares Medical nttte proctor telows'. CG isan escan Canoor Sosy Pesoareh Pr ‘ester of Manetr Gonotics. Supported by Ni gan cueri9tece ar. 18 September 1997; acepted 18 Daconber 1957 Fishing Down Marine Food Webs Daniel Pauly,” Villy Christensen, Johanne Dalsgaard, Rainer Froese, Francisco Torres Jr. ‘The mean trophic level of the species groups reported in Food and Agricultural Orga- rization global fisheries statistics declined from 1950 to 1994. This reflects a gradual ‘transition in landings from long-lived, high trophic level, piscivorous bottom fish toward short-lived, low trophic level invertebrates and pianktivorous pelagic fish. This effect, also oundto be occurring in inland fisheries, is most pronounced in the Northern Hemisphere. Fishing down food webs (that is, at lower trophic levels) leads at first to increasing catches, then to a phase transition associated with stagnating or declining catches, ‘These results indicate that present exploitation patterns are unsustainable. Exploitation of the ocean for fish and ma- rine invertebrates, both wholesome and valuable products, ought to bea prosperous Sector, given that capture isheries—in con trast tO agriculture and aquaculture—eeap harvests that didnot need to be sown. Yet raring fisheries ate in a global crisis mainly die to open access polices and subsidy- diven overcapitalizaion (1). It may be argued, however, that the global risis is tainly one of economics or of governance, ‘whereas the global esource hase itself ue. tutes naturally. Contradicting this more optimistic view, we show het tht landings ftom global fisheries have shifted in the last 1D. Pauy ard Dagan, Fufotee One, 204 Ne Nal. Unbersiy of Bren Coumoa, Vanoaver, teh Celina, Garada VT 125 \V. Chveteeon, Peas, F. Tores J, Itmationa Contr orig Agate Resources Wansgamet MI, Post Otis Box 26,0718 Maa, Phipps. To whom enreependerce ious bo adsetsed Era palyeroerescor 45 years from larg piscivorous fishes toward smaller invertebrates and planksivorous fishes, especially in. the Northern Hem sphere. This may imply major changes in the structure of marine food webs, Two data sets were used. The first has estimates of trophic levels for 220 different species or groups of fish and invertebrates, covering all statistical categories included in the offical Food and Agricultural Orga- nization (FAO) landings statisti (2). We ‘obtained these estimates from 60 published mass-balance trophic models that covered all major aquatic ecosystem types (3, 4). The models were constructed with the Eco- path software (5) and local data that in- cluded detailed diet compositions (6). In such models, factional trophic levels (7) are estimated values, based on the diet com= positions ofall ecosystem components rath- cr than assumed values; hence, their preci sion and accuracy are much higher than for the integer trophic level values used in SCIENCE * VOL. 279 + 6 FEBRUARY 1998 + wow sciencemag.onga : ee a aa a me ple | i A t hae te Fig. 1. Global rons of mean trophic level of ‘ishevios ndings, 19500 1994. (A) Marine areas; [B)niand areas earlier global studies (8). The 220 trophic levels derived from these 60 Ecopath appli- cations range from a definitional value of 1 for primary preccers and detritus to 4.6 (= 0.32) for snappers (family Lutjantdae) con the shelf of Yucatan, Mexico (9). The second data set we used comprises FAO slobal statistics (2) of fisheries landings for the years from 1950 to 1994, which are based on reports submitted annually by FAO member councries and other stares and were recently used for reassessing world fisheries potential (10). By combin- ing these data sets we could estimate the ‘mean trophic level of landings, presented here as time series by different groupings of all FAO statistical areas and for the world (11), For all marine areas, the trend over the past 45 years has been a decline in the mean trophic level ofthe fisheries landings, from slightly more than 3.3 in the early 1950s to fess than 3.1 in 1994 (Fig. 1A). A dip in the 1960s and early 1970s occurred because of extremely large catches [>12 X 10° metric tons (t) per year] of Peruvian anchoveta with a low trophic level (12) of 2.2 (042). Since the collapse of the Peruvi- an anchoveta fishery in 1972-1973, the alobal trend in the trophic level of marine fisheries landings has been one of steady decline. Fisheries in inland waters exhibit, con the global level, a similar marine areas (Fig. 1B): A cl average trophic level is apparent from the early 1970S, in parallel to, and about 0.3 units below, those of marine catches. The previous plateau, from 1950 to 1975, is due swonwasciencemag.org + SCIENCE * VOL. 279 + 6 FEBRUARY 1998, Fig. 2. Trends of mean trophic love of heres lancngs innorthem temperate areas, 1950 to 1994. (A) North Pacife FAO areas 61 and 67) (B) Nothmest ana Westem Cental Atlantic FAO reas 21 and 31); (©) Northeast Atlantic FAD area 27): and (D} Medterranean (FAO area 37} to insufficiently detailed fishery statistics for the earlier decades (10). In northern temperate areas where the fisheries are most developed, the mean tro: phic Ievel of the landings has declined steadily over the last two decades. In the North Pacific (FAO areas 61 and 67; Fig, 2A), trophic levels peaked inthe early 1970s and have since then decreased rapid- ly in spite of the recent increase in landings ‘of Alaska pollock, Theragra chalcogramma, which has a relatively high trophic level of 3.8 (+ 0.24). In the Northwest Atlantic (FAO areas 21 and 31; Fig. 2B), the fisher- ies were initially dominated by plankrivo- rous menhaxlen, Brevoortia spp., and other small pelagics at low trophic levels. As their landings decreased, the average trophic lev- el of the fishery initially increased, then in the 1970s it reversed to a steep decline Similar declines are apparent throughout the time series for the Northeast Atlantic (FAO area 27; Fig. 2C) and the Mediterra- nean (FAO area 37; Fig, 2C), although the later system operates at altogether lower trophic levels ‘The Central Eastem Pacific (AO area 77; Fig. 3A), Southern and Central Eastern Atlantic (FAO areas 41, 47, and 34; Fi 3B), and the Indo-Pacific (FAO areas 51, 51, and 71; Fig. 3C) show no clear trends lover time, In the southern Atlantic this is probably due to the development of new fisheries, for example, on the Patagonian shelf, which tends to mask declines of trophic levels in more developed fisheries. In the Indo-Pacific area, the apparent sta- bility is certainly due to inadequacies of Fig. 3. Trends of mean trophic lvls of fshoros landings in the ntertropical bat and adjacent wa ters (A) Central Eastem Paci FAO area 77) (8) Southwest, Central Eastom, and Southeast tian Ic (FAO eras 41, 34, nd 47}; and (} od we PPaote (FAO areas 51, 97, and 71),