Huso husoBeluga(Also: Caviar; European Sturgeon; Giant Sturgeon; Great Sturgeon)

Ge­o­graphic Range

The bel­uga stur­geon, Huso huso, is en­demic to the Ponto-Caspian Sea re­gion that in­cludes the Caspian Sea (the largest in­land body of water in the world) as well as the Sea of Azov and the Black Sea (Bemis & Ky­nard 1997, Berg 1948).

Acipenser­i­formes, in­clud­ing fos­sil species, are re­stricted to the North­ern Hemi­sphere; closely tied to for­mer Laurasian land­masses (Eura­sia, North Amer­ica) (Bemis et al. 1997). The lone con­gener of the bel­uga stur­geon, the kaluga (Huso dau­ri­cus) is en­demic to the Amur River, which runs along part of the Chi­nese/Russ­ian bor­der (Krykhtin & Svirskii 1997). Al­though these two species are the only mem­bers in the genus Huso, there are some doubts whether they are sis­ter species (Doukakis 2000). (Bemis and Ky­nard, 1997; Bemis, et al., 1997; Berg, 1948; Doukakis, 2000; Krykhtin and Svirskii, 1997)

Habi­tat

Bel­uga stur­geon are con­sid­ered eu­ry­ha­line, ca­pa­ble of mov­ing freely be­tween fresh­wa­ter and es­tu­ar­ies. Species of the order Acipenser­i­form often enter rivers for mi­gra­tion to other areas or for spawn­ing.

Bel­uga stur­geon spawn at a water tem­per­a­ture of 9-11 de­grees C and they are sen­si­tive to these water con­di­tions dur­ing pe­ri­ods of spawn­ing (Levin 1997). How­ever, other than spawn­ing times, adults are fairly re­silient to many en­vi­ron­men­tal fac­tors, par­tic­u­larly con­sid­er­ing that they live in some of the most pol­luted and al­tered wa­ter­ways in the world.

  • Aquatic Biomes
  • pelagic
  • lakes and ponds
  • rivers and streams

Phys­i­cal De­scrip­tion

Acipenser­i­formes are prim­i­tive actinoptery­gians that have a fos­sil record dat­ing back to the Lower Juras­sic (200 mya), with an ori­gin per­haps even ear­lier in the Meso­zoic (Bemis et al. 1997). Acipenser­i­formes pos­sess a unique blend of teleost-like char­ac­ter (high fe­cun­dity small egg size) and elas­mo­branch-like char­ac­ters (car­ti­lagi­nous en­doskele­ton, hyostylic jaw sus­pen­sion). They also pos­sess a het­e­ro­cer­cal cau­dal fin and rem­nants of ganoid scales; both are un­com­mon among ex­tant actinoptery­gians. Stur­geon species are noted by a spin­dle-shaped body with five rows of bony scutes and a long snout with sen­sory bar­bels.

The genus Huso can be dis­tin­guished from other stur­geon by pos­ses­sion of bran­chioste­gal mem­branes that are joined and form a flap (Berg 1948). The two species in the genus also pos­sess a rel­a­tively large cres­cent shaped mouth (Berg 1948). Berg (1948) gives the fol­low­ing key for dis­tin­guish­ing be­tween Huso con­geners:

  1. H. dau­ri­cus First dor­sal scute the largest. Bar­bels with­out fo­li­ate ap­pendages. Fewer than 60 rays in dor­sal fin. Found in the Amur Basin.
  2. H. huso First dor­sal scute the small­est. Bar­bels with fo­li­ate ap­pendages. Usu­ally not less than 60 rays in dor­sal fin. Found in the basins of the Caspian, Black, and Adri­atic Seas.

Among fresh­wa­ter fish species, H. huso may have be the largest that has ever lived (Freed­man 1999). Based on Debus's (1997) re­ports on bones ex­ca­vated from the 14th cen­tury, one fifth of all H. huso catches of that time were be­tween 4-6 me­ters in length. Bel­uga this size today are ex­tremely rare. Bemis et al. (1997a) re­port that this species may have reached nearly ten me­ters in length. The largest bel­uga ever recorded mea­sured six me­ters in length and weighed 3,200 kg (Berg 1948, Birstein 1993, Freed­man 1999). The de­creased size of in­di­vid­u­als today may re­flect the shift to catch­ing younger in­di­vid­u­als be­cause of over­fish­ing of older and larger fish, pol­luted liv­ing con­di­tions for bel­uga in the Caspian and other parts of its range, and the pos­si­ble loss of large in­di­vid­u­als from the gene pool. Large bel­uga stur­geon have been tar­geted by fish­er­man for cen­turies.

Sex­ual di­mor­phism is weakly ex­pressed, and may only be no­table in over­all size and weight. This di­mor­phism is par­tic­u­larly no­table in gravid fe­males that bear the weight of heavy egg masses (Berg 1948, Raspopov 1993).

  • Average mass
    1.1396e+06 g
    40162.11 oz
    AnAge
  • Range length
    6 (high) m
    19.69 (high) ft

De­vel­op­ment

Since fish­er­man or poach­ers cap­ture nearly all bel­uga stur­geon be­fore they can die of nat­ural causes, the de­vel­op­ment of this species is lit­tle known. Length and weight in­creases much more rapidly ear­lier in the life his­tory of these fish than dur­ing later in on­togeny (Raspopov 1993). Length and weight can range greatly in a given age class de­pend­ing on en­vi­ron­men­tal con­di­tions of the stur­geon habi­tat (Raspopov 1993).

After hatch­ing, down­stream mi­gra­tions of bel­uga lar­vae occur be­tween May and June (Levin 1997). From hatch­eries data, it is ob­served that the weight of bel­uga after 28-45 days av­er­ages around 2.5-3 g (Levin 1997). By the time they enter the sea they are 6-11 cm in length. Re­cently hatched lar­vae are also pur­ported to move at about 24 km per day dur­ing their mi­gra­tion to the sea. By the forth year, bel­uga may reach a meter in length. In the spring, ma­ture fish enter the rivers to breed, while younger in­di­vid­u­als (less than 10 years) dis­perse near the North­ern Caspian, their dis­tri­b­u­tion based mainly on eco­log­i­cal fac­tors like water tem­per­a­ture and ben­thic silt (Levin 1997).

Bel­uga stur­geon ma­ture later than most other acipenser­i­forms, usu­ally after their 12th year (Lelek 1987). Raspopov (1993) es­ti­mated that bel­uga stur­geon in the Caspian be­come sex­u­ally ac­tive every 4-8 years for fe­males, and every 4-7 years for males. Under fa­vor­able con­di­tions, a bel­uga may spawn up to nine times in its life­time.

From 1987-1997 in the Volga River (which was once the most im­por­tant spawn­ing habi­tat of bel­uga), fe­male spawn­ers have ranged from 199 to 236 cm, and weighed be­tween 48 to 160 kg (Levin 1997). These are much smaller than recorded sizes for pre­vi­ous years.

Re­pro­duc­tion

Bel­uga do not spawn every year and fe­males will re­sorb eggs un­less con­di­tions are suit­able (Artyukhin et al. 1979). Stur­geon can take be­tween 6 and 25 years to reach sex­ual ma­tu­rity, and fe­males of this species may re­pro­duce only every four to eight years (Raspopov 1993).

Bel­uga stur­geon are anadro­mous, mi­grat­ing to fresh­wa­ter spawn­ing grounds in rivers from larger seas (ei­ther the Caspian, Azov or Black). Mi­gra­tions occur in these pop­u­la­tions twice a year in spring and au­tumn (Berg 1948, Hensel & Hol­cik 1997). Some au­thors have di­vided bel­uga pop­u­la­tions into Win­ter or Spring races by the pe­riod they choose to spawn (Berg 1948). Dis­tances for spawn­ing mi­gra­tions for this species are leg­endary, ex­ceed­ing 2500 km in the Danube and Volga basin (Hensel & Hol­cik 1997, Khoderevskaya et al. 1997).

  • Breeding season
    winter and autumn race (varying depending on Caspian, Asov or Black sea population)
  • Range age at sexual or reproductive maturity (female)
    17-29 (high) years
  • Average age at sexual or reproductive maturity (female)
    6-30 years
  • Range age at sexual or reproductive maturity (male)
    11-23 (high) years
  • Average age at sexual or reproductive maturity (male)
    6-30 years

These stur­geon typ­i­cally spawn large num­bers of eggs in shal­low, gravely sites, and do not pro­vide parental care (Bemis et al. 1997, Lelek 1987).

  • Parental Investment
  • no parental involvement

Lifes­pan/Longevity

The bel­uga stur­geon is one of the most long-lived of all ver­te­brate species. Re­port­edly it may have reached ages over 100 years (Ono et al. 1983, Birstein 1993, Khodor­evskya 2000). Most acipenser­i­form species ma­ture at a late age (nor­mally greater than 10 years). Fe­males ma­ture later than males, nor­mally after 15 years for fe­males and 10 for males (Berg 1948). In­di­vid­u­als no longer die of nat­ural-age re­lated causes, with most being fished soon after reach­ing re­pro­duc­tive ma­tu­rity. The world de­mand for caviar (stur­geon and pad­dle­fish roe) is to blame for the short­ened lifes­pan of these fish. The de­mand far ex­ceeds the avail­able sup­ply, forc­ing legal fish­ers to catch in­creas­ingly younger and smaller fish, and pro­vid­ing fuel for a grow­ing black mar­ket. In­di­vid­u­als today are not found be­yond their 56th year (Raspopov & Novikova 1997).

  • Typical lifespan
    Status: wild
    56 (high) years

Be­hav­ior

Bel­uga con­gre­gate only dur­ing spawn­ing times and seem to be a less gre­gar­i­ous species then other Ponto - Caspian stur­geon (Levin 1997).

Acipenser­i­form species use en­vi­ron­men­tal pa­ra­me­ters such as water tem­per­a­ture, flow ve­loc­ity, and tur­bid­ity as spawn­ing cues (Beames­der­fer & Farr 1997; May­den & Kuha­jda 1996, Scarnec­chia et al. 1989, Sulak & Clugston 1998). Damming al­ters these cues, lim­it­ing nat­ural re­cruit­ment. A high rate of tumor de­vel­op­ment and sig­nif­i­cant per­cent­age of goan­dal re­sorp­tion near the Vol­gograd dams re­veal that stur­geon species, in­clud­ing Huso huso, are often in a state of chronic phys­i­cal stress caused by habi­tat de­gre­da­tion (Altuf'ev et al 1981).

Com­mu­ni­ca­tion and Per­cep­tion

Food Habits

Adult bel­uga stur­geon are mainly pis­ci­vores, swim­ming at mid­dle depths and prey­ing mostly on pelagic fish species. This is un­like most other stur­geon species, which nor­mally feed on bethic in­ver­te­brates while swim­ming along the bot­tom. In the Black Sea they feed on species such as floun­der (Platichthys fle­sus) and other flat­fish, go­b­ies (Go­b­i­idae), and Black Sea an­chovy (En­graulis en­cra­si­co­lus) (Berg 1948). In the Caspian they are re­ported to feed mainly on the Caspian roach (Ru­tilus ru­tilus), but also on her­ring and na­tive go­b­ies. Lit­tle is known about the diet of lar­val and ju­ve­nile H. huso.

Un­like other Ponto-Caspian stur­geon species bel­uga do not con­gre­gate, but re­main dis­persed when not mi­grat­ing (Levin 1997). Stur­geon species can en­dure long pe­ri­ods of star­va­tion and often do not eat for long pe­ri­ods dur­ing spawn­ing mi­gra­tions (Beames­der­fer & Farr 1997)

  • Animal Foods
  • fish
  • mollusks
  • aquatic crustaceans

Pre­da­tion

The large size of adults makes pre­da­tion on them dif­fi­cult, and a nat­ural preda­tor be­sides hu­mans is un­known. Many ju­ve­niles fall prey to pol­lu­tion and hy­dro­elec­tric dams (Altuf'ev 1997). Most pop­u­la­tions are sup­ported solely by hatch­ery re­leased in­di­vid­u­als. In the 1980's the num­ber of young bel­uga's re­leased from the Volga river per year was 19.4 mil­lion. The sur­vival rate of these in­di­vid­u­als was es­ti­mated to be 0.1% (Khoderevskya 1997). The num­ber killed by nat­ural causes or causes other than fish­ing pres­sures has not been doc­u­mented.

Ecosys­tem Roles

Bel­uga are the largest pis­ci­vores in the Pronto-Caspian re­gion. It dif­fi­cult to de­ter­mine their role in the past given their en­dan­gered sta­tus, but in pe­ri­ods where they were quite abun­dant (be­fore 1950) they would have been among the top preda­tors in the Ponto-Caspian, play­ing a sim­i­lar role as the fish eat­ing seals of the re­gion.

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Due to the in­cred­i­ble value of the roe of this species known com­mer­cially as bel­uga caviar - the most ex­pen­sive food item in the world(Birstein et al. 1998) - this species is among the most sought after nat­u­rally avail­able com­mer­cial prod­ucts in the world. Fe­male bel­uga can pro­duce 12 % of their body weight in caviar and bel­uga caviar can de­mand $200/kg. This species has be­come so de­pleted that its sur­vival is sup­ported en­tirely or nearly en­tirely by hatch­eries through­out its lim­ited range (Secor et al. 2000, Khodor­evskya 1999). With­out nat­ural re­cruit­ment, pop­u­la­tion struc­ture, age com­po­si­tion and ge­netic di­ver­sity of the bel­uga stur­geon have suf­fered (Secor et al. 2000, Raspopov 1993).

Most bel­uga stur­geon were his­tor­i­cally found in the Caspian Sea. This sea is fed by over 100 rivers, the most im­por­tant of these was the Volga in Rus­sia which histror­i­cally sup­plied 75% of the Caspian Seas's stur­geon catch (Artyukhin 1997, Khoder­shaya et al. 1997). Of the other rivers, the Ural River is the only re­main­ing free-flow­ing major river feed­ing the Caspian. It is also the only such river where bel­uga stur­geon can re­pro­duce nat­u­rally, with­out the ben­e­fits of hatch­eries (Khoderoveskya et al. 1997, De­Meu­lanaer & Ray­mak­ers 1996). How­ever, poach­ing and pol­lu­tion prob­lems are higher in the Ural than other parts of the sea, and about 50% of spawn­ing grounds are lost there due to sed­i­men­ta­tion and heavy agri­cul­tural and in­dus­trial pol­lu­tion (Ve­rina and Pe­seridi 1979).

The il­le­gal catch of stur­geon in the Caspian Sea and Volga River may be ten times greater than the legal catch (TRAF­FIC 2000). Poach­ers are re­port­edly re­mov­ing half of the ma­ture bel­uga in­di­vid­u­als every year. From De­cem­ber 1995 to De­cem­ber 2000 U.S. Fish and Wildlife Ser­vices con­fis­cated 3044.19 kg of il­le­gally im­ported eggs of Huso huso (cal­cu­lated from USFWS LEMIS sys­tem). With an av­er­age spawn­ing size of Huso huso in­di­vid­u­als being 70 kg (Khodor­evskya 1999) and a go­na­do­so­matic index (GSI) rang­ing from .07-.15 the num­ber of in­di­vid­u­als killed to ob­tain this caviar is be­tween 290 to 621.

Be­cause of the un­sup­port­able de­mand for caviar, bel­uga stur­geon are being caught at ages closer and closer to sex­ual ma­tu­rity. More than 80% of spawn­ing fe­males are be­tween 17 and 29 years of age, and nearly 90% of males are be­tween 11 and 23 years of age (Raspopov 1993, Rospopov and Novikova 1997). With proper man­age­ment it is said that it may take be­tween 40 to 45 years for stocks in the Caspian Sea just to re­turn to sta­ble lev­els (Artyukhin 1997). The slow mat­u­ra­tion of stur­geon tied with low num­bers have placed es­ti­mates of pop­u­la­tion re­cov­ery for some species at more than a cen­tury (Secor and Wald­man 1999).

  • Positive Impacts
  • food
  • research and education

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There is some ev­i­dence that PCPs and other dan­ger­ous chem­i­cals may be re­tained with the eggs of bel­uga stur­geon (Altuf'ev 1997). Small traces of en­vi­ron­men­tal tox­ins ap­pear in caviar (Boyle 1994). The lev­els of some of these chem­i­cals were con­sid­ered high from sam­ples cho­sen from some major re­tail­ers in the U.S. (be­tween 3.17-3.27 ppm of DDT, 410-640 ppb of Ar­clor 1254, and 2.1 - 2.8 ppm of se­le­nium) but these num­bers are lower than United States Food and Drug Ad­min­is­tra­tion ac­tion lev­els. It is doubt­ful that these traces can be harm­ful to any­one since few in­di­vid­u­als con­sume such large amounts of caviar that the chem­i­cals could ac­cu­mu­late to dan­ger­ous lev­els.

Con­ser­va­tion Sta­tus

Since it is dif­fi­cult to dis­tin­guish be­tween a male and a fe­male with­out dis­sec­tion, males are as sus­cep­ti­ble to fish­ing pres­sure as fe­males full of roe. To keep up with fish­ing pres­sure, Khoderevskya (1999) re­ports that be­tween 15-20 mil­lion fin­ger­lings need to be re­leased an­nu­ally from hatch­eries in order to main­tain a bel­uga pop­u­la­tion in the Caspian Sea. This fig­ure is not cur­rently being met. Birstein and col­leagues (1997) re­port that in 1995 due to the lack of brood­stock, there was no ar­ti­fi­cial breed­ing or nat­ural re­pro­duc­tion in the Volga River that year. This river his­tor­i­cally held most of the world's bel­uga spawn­ers and re­leases (Artyukhin 1997). In­ter­na­tional trade in bel­uga caviar should be halted in order to re­lieve the pres­sure on this species. Al­though cur­rently placed under CITES Ap­pen­dix II, an up­list­ing to Ap­pen­dix I would pro­vide a work­ing ban on in­ter­na­tional trade. The IUCN has also listed this species as crit­i­cally en­dan­gered. As the U.S. is the sec­ond largest im­porter of bel­uga caviar in the world, place­ment of this species under the U.S. En­dan­gered Species Act would pro­vide im­me­di­ate re­lief from much of the pres­sure to fish this species. De­c­la­ra­tion of any ban must also be en­forced, as a well-es­tab­lished black mar­ket has been suc­cess­fully op­er­at­ing since the fall of the So­viet Union (De­Meu­lanaer & Ray­mak­ers 1996, Khor­dorevskaya et al. 1997).

Habi­tat de­struc­tion also threat­ens this species. Dams block mi­gra­tory routes. In its major spawn­ing river the Volga, only a small per­cent­age (less than 20%) of bel­uga are able to mi­grate up­stream past the major dam on that river the Vol­gograd Dam (Rochard et al. 1990, Gert­sev 1999). Fish lifts and el­e­va­tors must trans­port these in­di­vid­u­als across the dam. Often they find water con­di­tions on the other side of the dam un­suit­able for re­pro­duc­tion be­cause of water tem­per­a­ture and flow rates. Prob­lems of mi­grat­ing lar­vae mov­ing to­ward adult grounds up­stream from the dam are lit­tle stud­ied. Debus (1997) con­sid­ers that the pas­sage of ju­ve­niles down­stream the Volga River as nor­mally lethal, be­cause ju­ve­niles do not sur­vive pas­sage through the hy­dro­elec­tric power tur­bines of the dams.

Other Com­ments

Acipenserids have been hunted for their roe since at least 1100 B.C. (Wei et al. 1997). An­cient Greek, Roman and Chi­nese lit­er­a­ture make ref­er­ence to acipenser­i­form species and caviar; and the Chi­nese re­ported trade as early as the 10th cen­tury A.D. (De­Meu­le­naur & Ray­mak­ers 1998, Bur­ton & Bur­ton 1975).

For more in­for­ma­tion on bel­uga stur­geon and to learn how you can help save this species visit, http://​www.​caviaremptor.​org

Con­trib­u­tors

William Fink (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Prosanta Chakrabarty (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

estuarine

an area where a freshwater river meets the ocean and tidal influences result in fluctuations in salinity.

external fertilization

fertilization takes place outside the female's body

fertilization

union of egg and spermatozoan

food

A substance that provides both nutrients and energy to a living thing.

freshwater

mainly lives in water that is not salty.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

natatorial

specialized for swimming

native range

the area in which the animal is naturally found, the region in which it is endemic.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pelagic

An aquatic biome consisting of the open ocean, far from land, does not include sea bottom (benthic zone).

piscivore

an animal that mainly eats fish

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

Ref­er­ences

Altuf 'ev, Y. 1997. Mor­pho­func­tional ab­nor­mal­i­ties in the or­gans and tis­sues of Caspian Sea stur­geon caused by eco­log­i­cal changes. Pp. 34-45 in V Birstein, A Bauer, A Kaiser-Pohlmann, eds. IUCN Species Sur­vival Com­mi­sion. Stur­geon stocks and caviar trade work­shop. Bonn, Ger­many: Oc­ca­sional Pa­pers of the IUCN Species Sur­vival Com­mis­sion No. 17.

Altuf 'ev, Y., A. Vlansenko, A. Polenov, N. Shevel­eva. 1981. Jour­nal of Ichthy­ol­ogy, 20(2): 94-103.

Artyukhin, E.N, 1997. The cur­rent sta­tus of com­mer­cial stur­geon species in the Black Sea-Caspian Sea basin. Pp. 9-13 in Birstein, V.J., Bauer, A., Kaiser-Pohlmann, A., eds. . IUCN Species Sur­vival Com­mi­sion. Stur­geon stocks and caviar trade work­shop. Bonn, Ger­many: Oc­ca­sional Pa­pers of the IUCN Species Sur­vival Com­mis­sion No. 17.

Beames­der­fer, R., R. Farr. 1997. Al­ter­na­tives for the pro­tec­tion and restora­tion of stur­geons and their habi­tat. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 407-417.

Bemis, W., E. Find­eis, L. Grande. 1997. An overview of Acipenser­i­formes. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 25-71.

Bemis, W., B. Ky­nard. 1997. Stur­geon rivers: an in­tro­duc­tion to acipenser­i­form bio­geog­ra­phy and life his­tory. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 167-183.

Berg, L. 1948. Fresh­wa­ter Fishes of the U.S.S.R. and ad­ja­cent coun­tries Part I. Is­rael Pro­gram for Sci­en­tific Trans­la­tions; [avail­able from the Of­fice of Tech­ni­cal Ser­vices, U.S. Dept. of Com­merce, Wash­ing­ton] 1962.

Birstein, V. 1997. Con­clud­ing re­marks. Pp. 71-88 in V Birstein, A Bauer, A Kaiser-Pohlmann, eds. Stur­geon Stocks and Caviar Trade Work­shop. Bonn, Ger­many: IUCN.

Birstein, V. 1993. Stur­geon and Pad­dle­fishes: Threat­ened Fishes in Need of Con­ser­va­tion. Con­ser­va­tion Bi­ol­ogy, 7 (4): 773-782.

Birstein, V., P. Doukakis, B. Sorkin, R. De­Salle. 1998. Pop­u­la­tion ag­gre­ga­tion analy­sis of three caviar-pro­duc­ing species of stur­geons and im­pli­ca­tions for the species iden­ti­fi­ca­tion of black caviar. Con­ser­va­tion Bi­ol­ogy, 12 (4): 766-775.

Boyle, R. 1994. The cost of caviar. The Am­i­cus Jour­nal, Spring: 22-27.

Bur­ton, M., R. Bur­ton. 1975. En­cy­clo­pe­dia of Fishes. New York: Oc­to­pus Book Lim­ited.

De­Meu­le­naer, T., C. Ray­mak­ers. 1996. Stur­geon of the Caspian Sea and in­ves­ti­ga­tions of the in­ter­na­tional trade in caviar. Cam­bridge: TRAF­FIC In­ter­na­tional.

Debus, L. 1997. Stur­geons in Eu­rope and causes of their de­cline. Pp. 44-53 in V Birstein, A Bauer, A Kaiser-Pohlmann, eds. Stur­geon Stocks and Caviar Trade Work­shop. Bonn, Ger­many: IUCN.

Doroshov, S., F. Binkowski. 1985. Epi­logue: a per­spec­tive on stur­geon cul­ture. Pp. 160- 163 in F Binkowski, S Doroshov, eds. North Amer­i­can Stur­geons. Dor­drecht, Nether­lands: Dr. W. Junk Pub­lish­ers.

Doukakis, P. 2000. Sys­tem­at­ics and con­ser­va­tion ge­net­ics of stur­geons (Order Acipenser­i­formes). Yale Uni­ver­sity: Un­pub­lished Dis­ser­ta­tion.

Foot, T. 2000. The Gui­ness Book of World Records 2001. Lon­don: Gui­ness World Records Ltd.

Freed­man, B. 1999. En­cy­clo­pe­dia of En­dan­gered Species. De­troit: In­ter­na­tional Lim­ited.

Gert­sev, V. 1999. . A model of stur­geon dis­tri­b­u­tion under a dam of a hy­dro-elec­tric power plant.. Eco­log­i­cal Mod­el­ing, 119: 21-28.

Hensel, K., J. Holèík. 1997. Past and cur­rent sta­tus of the stur­geons in the upper and mid­dle Danube River. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 185-200.

Khodor­eveskya, R., E. Krasikov, G. Do­gopol, O. Zhu­ravlev. 2000. For­ma­tion of the stock of Caspian Acipenserids under pre­sent day con­di­tions.. Jour­nal of Ichthy­ol­ogy, 40 (8): 602-609.

Khodor­evskaya, R., G. Dov­gopol, O. Zhu­ravl­eva, A. Vlasenko. 1997. Pre­sent sta­tus of com­mer­cial stocks of stur­geon in the Caspian Sea basin. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 209-219.

Krykhtin, M., V. Svirskii. 1997. Stur­geon catch and cur­rent sta­tus of stur­geon stocks in the Amur River. Pp. 29-34 in V Birstein, A Bauer, A Kaiser-Pohlmann, eds. Stur­geon Stocks and Caviar Trade Work­shop­Pro­ceed­ings of a Work­shop Held 9-10 Oc­to­ber in Bonn, Ger­many. Bonn, Ger­many: IUCN Oc­ca­sional Paper No 17.

Lelek, A. 1987. The Fresh­wa­ter Fishes of Eu­rope Vol. 9. AULA-Ver­lag Wies­baden: Threat­ened Fishes of Eu­rope.

Levin, A. 1997. The dis­tri­b­u­tion and mi­gra­tion of stur­geons in the Caspian Sea. Pp. 13-20 in V Birstein, A Bauer, A Kaiser-Pohlmann, eds. Stur­geon Stocks and Caviar Trade Work­shop, Pro­ceed­ings of a Work­shop Held 9-10 Oc­to­ber in Bonn, Ger­many. Bonn, Ger­many: IUCN Oc­ca­sional Paper No 17.

May­den, R., B. Kuha­jda. 1997. Threat­ened species of the world: Scaphirhynchus sut­tkusi Williams & Clem­mer 1991 (Acipenseri­dae). En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 418-419.

Ono, R., J. Williams, A. Wag­ner. 1983. Van­ish­ing Fishes of North Amer­ica. Wash­ing­ton, D.C.: Stonewall Press.

Raspopov, V. 1993. Growth rate of Caspian Sea bel­uga. Jour­nal of Ichthy­ol­ogy, 33(9): 72-84.

Raspopov, V., A. Novikova. 1997. Size and age com­po­si­tion of lar­vae and spawn­ers of the great stur­geon, *Huso huso*, mi­grat­ing in the Volga. Jour­nal of Ichthy­ol­ogy, 37(2): 166-173.

Rochard, E., G. Castel­naud, M. LeP­age. 1990. Stur­geons (Pisces: Acipenseri­dae); threats and prospects. Jour­nal of Fish Bi­ol­ogy, 37 (Sup­ple­ment A): 123-132.

Scarnec­chia, D., T. Gengerke, C. Moen. 1989. Ra­tio­nale for har­vest slot limit for pad­dle­fish in the Upper Mis­sis­sippi River. North Amer­i­can Jour­nal of Fish­eries Man­age­ment, 9: 477-487.

Secor, D., A. Are­fiev, A. Niko­laev, A. Sharov. 2000. Restora­tion of stur­geons: lessons from the Caspian Sea stur­geon ranch­ing pro­gramme. Fish and Fish­eries: 215-230.

Secor, D., J. Wald­man. 1999. His­toric abun­dance of Delaware Bay At­lantic stur­geon and po­ten­tial rate of re­cov­ery. Amer­i­can Fish­eries So­ci­ety Sym­po­sium, 23: 203-216.

Sulak, K., J. Clugston. 1998. Early life his­tory stages of gulf stur­geon in the Suwan­nee river, Florida. Trans­ac­tions of the Amer­i­can Fish­eries So­ci­ety, 127: 758-771.

TRAF­FIC, 2000. Re­view of 10 species of Acipenser­i­formes, pre­pared in the Six­teenth Meet­ing of the CITES An­i­mals Com­mit­tee. 18 pp:

Ve­rina, I., N. Per­sidi. 1979. On the stur­geon spawn­ing grounds con­di­tions in the Ural River. Stur­geon Cul­ture of In­land Wa­ters, Caspian fish­eries In­sti­tute, As­trakhan: 33-34.

Wei, Q., F. Ke, J. Zhang, P. Zhang, J. Luo. 1997. Bi­ol­ogy, fish­eries, and con­ser­va­tion of stur­geons and pad­dle­fish in China. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 48: 241-255.