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AbstractIn this communication a history of early investigations of

the Caspian Sea is given, with special emphasis on theexpedition in 1904 headed by N. M. Knipovich. The con-tribution of this expedition to the study of biodiversity ofthe sea, particularly newly discovered fish taxa, is exam-ined. The specificity of the work, the new methods of col-lecting and examining samples, the routes of the expedi-tion, and the contribution of each participant to the expe-dition are discussed. Emphasis is placed on the continuityof research conducted on the Caspian Sea. Illustrations forthe paper are original photographs made during the 1904expedition from the archive of Vasily Kononov, a grandfa-ther of the third author. No photographs, except one, havebeen published before.

ZusammenfassungIn vorstehenden Artikel werden frühe Erforschungen des

Kaspischen Meeres im Überblick dargestellt, mit beson-derem Schwerpunkt auf der Expedition, die im Jahre 1904unter der Leitung von N. M. Knipovich durchgeführtwurde. Untersucht wird der Beitrag der Expedition zur Ken-ntnis von der Biodiversität des Kaspischen Meeres mit Blickauf neuerdings entdeckte Fischtaxa. Die Besonderheiten derdamaligen Forschungsarbeit, die neuen Methoden beimSammeln und Untersuchen der Fangproben, die Routen derExpedition und die Beiträge der einzelnen Teilnehmer wer-den diskutiert. Herausgearbeitet wird die Kontinuität in derErforschung des Kaspischen Meeres. Für die Abbildungenwurden die Originalfotos der Expedition von 1904 verwen-det, die sich im Archiv von Vasily Kononov befanden, demGroßvater des an dritter Stelle genannten Autors. Mit einerAusnahme sind die Bilder alle bisher unveröffentlicht.

RésuméCet article est consacré à l’histoire du début des

recherches en Mer Caspienne, avec une insistance parti -culière sur l’expédition de 1904, sous la conduite de N. M.Knipovich. On examine la contribution de cette expédi-tion à l’étude de la biodiversité de cette mer, surtout en cequi concerne les taxons de poissons nouvellement décou-verts. On aborde les aspects originaux du travail, les nou-

velles méthodes de collecte et d’analyse d’échantillons, letrajet de l’expédition et la contribution de chaque partici-pant de l’expédition. On met en évidence la continuité dela recherche menée en Mer Caspienne. Les illustrations del’article se composent de photos originales prises lors del’expédition de 1904, provenant des archives de VasilyKononov, grand-père du troisième auteur. Aucune desphotos, à l’exception d’une seule, n’a été publiée aupara-vant.

SommarioIn questa comunicazione è presentato un resoconto storico

delle prime ricerche sul Mar Caspio, con particolare enfasial la spedizione del 1904 guidata da N. M. Knipovich. È sta -to esaminato il contributo di questa spedizione allo studiodel la biodiversità di questo mare, specialmente le nuove spe -cie di pesci scoperte. Sono discussi vari aspetti tra cui la spe -cificità del lavoro svolto, i metodi di raccolta e dell’esame de -gli esemplari, i percorsi della spedizione e il contributo diogni partecipante alla spedizione stessa. Particolare rilievo èda to alla continuità delle ricerche condotte sul Mar Caspio.Le illustrazioni dell’articolo sono fotografie originali scattatedu rante la spedizione del 1904 e recuperate dall’archivio diVasily Kononov, il nonno del terzo autore. Ad eccezione diuna sola fotografia, nessuna delle altre è mai stata pubblicataprima.

INTRODUCTIONWhile undertaking freshwater and brackish water

fish studies (Bogutskaya & Naseka 2004), particu-larly on the fishes of the Caspian Sea, it becameapparent that there was a need to revise the vastcollection of material from scientific expeditions tothe region, some of which are of great historicalimportance, especially with respect to taxonomicnomenclature. Material collected in many expedi-tions to the Caspian Sea since the 1770s has beendeposited at the Zoological Institute of the RussianAcademy of Sciences, including unique materialcollected during the expedition of 1904. All the

aqua, International Journal of Ichthyology

A brief history of the study of fishes of the Caspian Sea and scientific results of the Caspian Expedition of 1904 headed by N. M. Knipovich

N. G. Bogutskaya1, A. M. Naseka1 and P. A. Tikhonov2

1) Zoological Institute of the Russian Academy of Sciences, Universitetskaya emb. 1, 199034 St. Petersburg, Russia. E-mail: [email protected]

2) Institute of Silicate Chemistry of the Russian Academy of Sciences

Received: 23 March 2007 – Accepted: 20 September 2007

aqua vol. 14 no. 1 - 28 January 20081

aqua vol. 14 no. 1 - 28 January 2008 2


fish collections from the Caspian Sea were exam-ined by us, which stimulated our interest in obtain-ing additional information on their origin and sur-rounding circumstances. The historical context offish collections in the Caspian Sea, clarification ofroutes of the expeditions and data on the locationsof all historically important discoveries seem veryimportant both for general knowledge and tounderstand specific data on collected specimens. The objective of this communication is to provide

a brief history of zoological and fishery surveys andexpeditions to the Caspian Sea, with a focus on theachievements of the expedition in 1904 withrespect to the new methods of collecting and exam-ining samples employed. We tried to assess thecontribution of the 1904 expedition to the study ofbiodiversity of the sea, particularly newly discov-ered fish taxa, the specificity of the work and thecontribution of each participant to the expedition.Emphasis is placed on the continuity of subsequentresearch conducted in the Caspian Sea.Illustrations for the paper (except for Fig. 9) are

original photographs taken during the 1904 expe-dition from the archive of Vasily Kononov, agrandfather of the third author. We do not knowthe name of the photographer. Some of the pho-tographs have very short handwritten inscriptionsexplaining the depicted objects, some do not haveinscriptions and, in these cases, we based our leg-end for the figures on our own knowledge.

PRE-1904 EXPEDITIONSThe Caspian Sea has long excited the interest of

the countries in its proximity, the Slav nationsincluded. Arabic, Iranian and Armenian recordschronicle the marine voyages of the Russians alongthe Iranian coast of the Caspian as early as the endof the ninth century and beginning of the tenthcentury. In the second half of the fifteenth centurythe renowned merchant and explorer AfanasyNikitin, travelling to Iran and India, visited theCaspian cities of Derbent, Shemakha and Baku.The fishery resources of the Volga River and theCaspian Sea were highly sought after and attractedRussians outside the boundaries of their state tofish in the lower reaches of the Volga delta beforethat region became part of Russia. In 1554 Ivan theTerrible drove Khan Yanchurgey out of Astrakhan’(formerly Adjiderkhan, or Adji Darkhap), andcharged the new Khan, a Russian puppet, to pay atribute to the Tsar’s court, and, moreover, to supplythe court of Moscow State with 3,000 large stur-

geon, fresh or salted. At the same time Russianfishermen were permitted unrestricted access tofish in the region from Kazan down to the CaspianSea. In the reign of Peter the Great, a “FisheryOffice” was established in Astrakhan’ and all the“Astrakhan’ Fisheries Areas”; the north-westernriver and sea waters of the Caspian basin weregiven over to the Astrakhan’ municipal society(Strubalina 1990).Under Peter the Great Russia became, for the first

time in its history, a significant maritime nation,resulting in an expansion in its maritime trade.Peter the Great became interested in the report ofHodca Nefis in 1714 about the possibility ofrestoring the former flow of the Amu Daryathrough the Uzboy Valley, which formerly con-nected the Caspian and Aral Seas. Such a connec-tion would have made a southern trade route toPersia and India much shorter. Peter the Great wasalso interested in projects connecting the Caspianwith the Black Sea through the Sea of Azov; workwas undertaken for three years but was not com-pleted. In 1715, on the orders of the Tsar, Alexan-der Bekovich-Cherkasski (Devlet Kizden Murza,before his conversion to Christianity) initiated aseries of expeditions to the Caspian Sea. The 1715to 1717 expedition was headed by AlexanderIvanovich Kozhin and Fyodor Ivanovich Soimonovand surveyed the eastern coast of the Caspian. By1720 they had compiled the first map of theCaspian Sea (its western and northern coasts),while Soimonov continued hydrographic studies ofthe coastline in 1726. In the second half of theeighteenth century hydrographic expeditions,which refined information on the outlines of thecoasts, were undertaken by Ilya Vasilyevich Tok-machev and Mark Ivanovich Voinovich. The expe-dition of 1763 to 1765 described the Asian regions(southern and eastern) of the coast (Lukina 1984).The extensive studies organized by the Russian

Academy of Sciences, by order of Empress Cather-ine the Second (e.g. Bogdanov 1875), were of greatimportance in understanding the geology and bio -logy of the Caspian. Five expeditions were orga-nized. Each of the scientists listed below was com-missioned to conduct an expedition: AcademicianSamuel Georg Gottlieb Gmelin from Tübingen;adjunct of the Academy Johann Anton Gülden-städt from Riga; Academician Peter Simon Pallasfrom Berlin; botanist of the St. Petersburg MedicalGarden Johann Peter Falk from Sweden andadjunct of the Academy Ivan Lepekhin. All five

Table I. List of species (valid names) described by Pallas,Güldenstädt, Eichwald, de Filippi and Kessler based onfish materials collected by expeditions in the Caspian basinbefore 1904.

Pallas Acipenser stellatus Pallas, 1771Ballerus sapa (Pallas, 1814)Benthophilus macrocephalus (Pallas, 1787)Hucho taimen (Pallas, 1773)Vimba vimba persa (Pallas, 1814)

Güldenstädt Alburnus chalcoides (Gueldenstaedt, 1772)Barbus mursa (Gueldenstaedt, 1773)Capoeta capoeta (Gueldenstaedt, 1773)Luciobarbus capito (Gueldenstaedt, 1773)Stenodus leucichthys (Gueldenstaedt, 1772)

Eichwald Aspius aspius taeniatus (Eichwald, 1831)Neogobius affinis (Eichwald, 1831)Neogobius caspius (Eichwald, 1831)Sabanejewia caspia (Eichwald, 1838)

de Filippi Acanthalburnus microlepis (Filippi, 1863)Alburnoides eichwaldii (Filippi, 1863)Barbus cyri Filippi, 1865Capoeta capoeta sevangi Filippi, 1865Proterorhinus nasalis Filippi, 1863Sabanejewia aurata (Filippi, 1863)

Kessler Alburnus filippi Kessler, 1877Alburnus hohenackeri Kessler, 1877Barbus ciscaucasicus Kessler, 1877Barbus goktschaicus Kessler, 1877Benthophilus baeri Kessler, 1877Benthophilus ctenolepidus Kessler, 1877Benthophilus granulosus Kessler, 1877Benthophilus grimmi Kessler, 1877Benthophilus leptocephalus Kessler, 1877Benthophilus leptorhynchus Kessler, 1877Benthophilus spinosus Kessler, 1877Caspiomyzon wagneri (Kessler, 1870)Caspiosoma caspium (Kessler, 1877)Chondrostoma cyri Kessler, 1877Chondrostoma oxyrhynchum Kessler, 1877Clupeonella grimmi Kessler, 1877Cobitis hohenackeri Kessler, 1877Knipowitschia longecaudata (Kessler, 1877)Mesogobius nigronotatus (Kessler, 1877)Neogobius bathybius (Kessler, 1877)Neogobius cyrius Kessler, 1874Neogobius eurystomus (Kessler, 1877)Neogobius goebelii (Kessler, 1874)Neogobius macrophthalmus (Kessler, 1877)Oxynoemacheilus brandti (Kessler, 1877)Salmo caspius Kessler, 1877Salmo gegarkuni Kessler, 1877


N. G. Bogutskaya, A. M. Naseka and P. A. Tikhonov

expeditions started from St. Petersburg in June1768. From 1768-1774 the expedition of Gmelin trav-

elled through the River Don area and the Caucasusregion and, importantly, along the Caspian Sea

coasts, including the southern coast. On 16 (5)June1 1770 Gmelin started from Astrakhan’ to theCaspian Sea and Persia. During that trip (up to 21(10) April 1772) he visited Derbent, Baku, She-makha (= Shamakhy, Samaxi), Salyany, Enzeli (=Enceli, Bandar-e Anzali, Bandar-e Pahlavi), Resht(= Rast, Rasht) and many localities eastwards up toAstrabad (= Asterabad, Esterabad, Gorgan, Gur-gan, Hyrcania). In July 1773 Gmelin undertook asecond journey to the Caspian Sea; he visited theeastern Caspian coast - the islands of Kulaly andSvyatoi (= Artema, Pirallakhi), cape Tyub-Karagan(= Tupqaraghan Muyis), the bays of Aleksanderbay(= Aleksandra Bekovicha-Cherkasskogo Zaliv) andKara-Bugaz-Gol (= Garabogazkol, Kara-Bogaz),the Neftyanoy and Cheleken islands, and thesouthern Caspian coast. Gmelin was captured byUsmey-Khan who held him to ransom. He died in1774 in captivity in the village of Akhmakent nearDerbent. Gmelin presented the results of the expe-dition in his work entitled “Journey across Russiafor studying the three kingdoms of nature” pub-lished posthumously (Gmelin [1770]-1784). Inaddition to other findings, Gmelin described forthe first time the organization of the fisheries in theVolga-Caspian region.Peter Simon (Petr Semenovich in Russian) Pallas

headed the expedition to Transbaikalia, and also toBashkiria, to the Urals and regions of the Volga andCaspian lowlands. The expedition was described inhis work entitled “The Journey to differentprovinces of the Russian Empire” and materials col-lected served as a basis for a wide range of publica-tions (Pallas 1771, 1773a, 1773b, 1776a, 1776b,1787, 1814). Pallas only visited the Caspian Seafrom 4 September (24 August) to 11 September(August 31) 1769 during his short trip from Guriev(now Atyrau) to the delta of the Urals and to theshoreline. A participant of the expedition was a stu-dent of Guriev Nikita Sokolov who visited theCaspian in July 1770 (west of the Urals) and inMarch 1773 (from Kizlyar to Astrakhan’). Pallas col-lected extensive samples and also described fivefishes inhabiting the basin of the Caspian Sea, nowspecies or subspecies (Table I).Johann Anton (Anton Ivanovich in Russian)

Güldenstädt travelled to the Eastern Caucasus in1770. He visited the Terek River, Tiflis (now Tbil-isi), and spent a long time travelling throughKakhetia, Ossetia and Imeretia (historical provinces

1 Here and below, the date in brackets refers to the so-called "Old Style" (Julian Calendar) used in Russia at that time.



of the modern Georgia). He described five riverand anadromous fishes of the Caspian basin (TableI), providing excellent descriptions (Gueldenstaedt1772, 1773, 1781). In 1793-1794 Pallas again vis-ited the Ural and Volga region and the NorthernCaucasus, in particular Kumo-Manych steppe.In 1825 Karl Edward (Eduard Ivanovich in Russ-

ian) Eichwald, who at that time was professor ofzoology of Kazan University, was sent on a mission tostudy the Caspian Sea and the Caucasus. Havingreceived the corvette “Hercules” from the govern-ment, he spent more than three months at sea. FromAstrakhan’, Eichwald travelled by sea to Cape Tyub-Karagan, surveyed the Mangyshlak Peninsula, trav-elled to the western coast (to Tarki, Derbent andBaku) and again to the eastern coast where he stud-ied Balkhan Bay, Cheleken Island and the old estuaryof the Amu Darya River; he also visited Astrabad,Mazanderan and Gilyan. In September 1826 hereturned to Kazan via Baku and Tiflis. Eichwald col-lected rich material on paleontology and ethnogra-phy (Eichwald 1831, 1838, 1841). However, hiszoo logical collections were scanty and he called theCaspian Sea the “dead sea”. Notwithstanding, he des -cribed some new species, including five fish species(Table I), and also established the genus BenthophilusEichwald, 1831.Three expeditions to the Caspian Sea were under-

taken from Orenburg by Grigory Silych Karelin. In1832, the first expedition to study the north-easternpart of the sea took place. On the initiative of Kare-lin, fort Novo-Alexandrovsky was built in the Kyzyl-Tash tract on the eastern coast of the Kaidak (=Kay-dak, Qaydaq) Bay in 1834; it was later transferred tothe western coast of Mangyshlak (=Mangghystau,Mangislak) Peninsula. In 1836 Karelin headed theexpedition on board the vessel “St. Gabriel”, con-ducting a survey of eastern and south-eastern coastsof the Caspian. Karelin collected large amounts ofmaterial, which remained unexamined, and pub-lished almost nothing (Bogdanov 1875).The expeditions of Karelin coincided with the

expeditions of the chemist and geologist Karl Kris-tian Traugott Friedemann (Ferdinand Ivanovich inRussian) Göbel who defined the chemical compo-sition of the Volga and Don waters, waters of theCaspian, Black and Azov seas. Moreover, he visitedGuriev, from where he travelled to Astrakhan’(Göbel 1837-1838). The zoological observationsand collections during that expedition were carriedout by a companion of Göbel, whose surname wasKlaus (his first name is unknown).

In 1851 the Ministry of Agriculture and StateProperty, which was in charge of organizingresearch into natural resources and fisheries withthe participation of the Russian Geographical Soci-ety (established in 1845), embarked on a long-termscientific fishery expedition throughout the terri-tory of the Russian empire. The purpose of theexpedition was twofold: firstly, it would assess thefishery resources of Russia and, secondly, it wouldobtain information that would allow the organiza-tion to target exploitation of the most valuablespecies, particularly of sturgeons in the Volga-Caspian basin and pike-perch in the Sea of Azovbasin. The lack of information on the biology andhabitat conditions of fishes in Russian waters pre-vented the implementation of informed measuresto regulate fisheries in those water-bodies where itwas most needed at that time (Baer 1860).On 5 March (21 February) 1852 Nicholas (Niko-

lay) the First ordered the Minister of State PropertyP. D. Kiselev to send an expedition to the VolgaRiver and the Caspian Sea. In April, the Presidentof the Russian Geographic Society, Grand PrinceKonstantin (Konstantin Nikolayevich Romanov),visited Astrakhan’ on his way abroad. TheAstrakhan’ merchant F. G. Golikov gave him3,000 roubles for studying the Caspian fisheries.The Ministry of State Property allocated an addi-tional sum of 5,000 roubles. In spring, the Geo-graphic Society convened a commission for thedevelopment of a plan for the Caspian expedition.The official task of the expedition was to study

fisheries on the Volga River and in the Caspian Seain terms of technology, statistics and natural his-tory. The minister, P. D. Kiselev, offered the Acad-emician Karl Ernst (Karl Maksimovich in Russian)Baer the leadership of the expedition. On 11 (23)March 1853 a decree was issued designating theCaspian expedition for three years and within twodays the Vice President announced the start of theexpedition; a month later Baer left St. Petersburg.Between then and 1857 he made four scientificjourneys to the coasts of the Caspian Sea, later con-sidered as a single expedition (Lukina 1984). Niko-lay Yakovlevich Danilevsky, a botanist by educa-tion, was appointed as a statistician for the expedi-tion. He studied demography, statistics, agronomy,geography, climatology and hydrography. The main output from Baer’s expedition was the

Regulations of Caspian Fisheries and Seal Hunt-ing, which were confirmed by the government in1865. The regulations mostly dealt with the orga-



nization of trade in the Volga-Caspian fisheryregion. Additionally, during the Caspian expedi-tion of 1853-1857 extensive geological, zoological,palaeontological and craniological material wascollected, which was deposited in the museums ofthe St. Petersburg Academy. Baer presented a par-ticularly large number of specimens to the Zoolog-ical Museum (Lukina 1984).Baer (1860) presented the main results of the

Caspian studies of 1854-1857 in his book pub-lished in 1860 “Fisheries in the Caspian Sea and itstributaries”, which is the second volume of thenine-volume edition “Studies of the State of Fish-eries in Russia”. The lack of organization of fish-eries and frequent changes in legislation werecaused by “the absence of scientific studies of thelife of fishes in the Caspian Sea and its tributaries,of conditions affecting their reproduction andelimination and of relations of those conditions tofishing techniques, already existing and decreesconcerning them” (Baer 1860: 9).It should be emphasized that the study of fisheries

and what is today termed applied fishery problemsconnected these two giants of thought, Baer, one ofthe most versatile and outstanding naturalists ofthe time, a famous embryologist and a founder offundamental zoology, and Danilevsky, not only anaturalist (and ardent anti-Darwinist), but primar-ily a publicist, theoretician in the field of thenational economy, who in his main literary work“Russia and Europe” (Danilevsky 1869) presenteda peculiar theory of Panslavism.In 1862 the Professor of Zoology of the Univer-

sity of Milan, Filippo de Filippi, travelled through-out the Caspian region, through the northernCaspian part of Persia and Transcaucasia (Mugan-skaya steppe) up to Tiflis. In the description of hisjourney (Filippi 1863, 1865), numerous zoologicalobservations are given and in particular hedescribed six fish species (Table I). Evgeniy Dmitrievich Pelzam, appointed by Kazan

University, worked on the eastern coast of theCaspian Sea in 1867 for six months and again in1868. Alexander Onufrievich Kovalevsky visited theCaspian Sea in 1869. He spent only a few days inPetrovsk (now Makhachkala) and Baku, but col-lected vast numbers of marine animals. From about1867 Vladimir Evgenievich Yakovlev carried outentomological and ichthyological observations inthe vicinity of Astrakhan’. He later published anumber of separate essays describing new species,Rutilus caspicus (Yakovlev, 1870) and Chondrostoma

variabile Yakovlev, 1870 among them (Yakovlev1870).In 1874 and 1876 Oskar Andreyevich Grimm

participated in the Aral-Caspian expedition to theCaspian Sea. He studied the middle and southernpart of the Caspian, concentrating on aspects ofzoological interest. He determined the temperatureand salinity of the waters, zones of dispersal ofaquatic organisms to a depth of 60 m, and accu-mulated a vast collection of materials (Grimm1876, 1877).Grimm was an exceptional personality in Russian

fishery biology, both from a theoretical and practi-cal perspective, and in terms of fishery manage-ment; he had a direct input to the expedition of1904. During journeys throughout the CaspianRegion, Grimm worked as a conservator at theZoological Cabinet of the St. Petersburg Universityand was an assistant professor (“Privatdozent”) atthe Department of Zoology. Material of many of the above expeditions was

examined by Karl Fedorovich Kessler, who, amongother outstanding works, published two books thatlaid the foundation of systematics of fishes of theAral-Ponto-Caspian basin (Kessler 1874, 1877).Among the more than 85 fish species described byhim, 28 are from the Caspian basin (Table I).Kessler proposed the first ecological classificationof fishes (saltwater, brackish water, “inhabiting dif-ferent waters” [euryhaline], anadromous, semi-anadromous and freshwater).In 1879 Grimm submitted a note to the St.

Petersburg Naturalists Society, indicating that asailing ship and a station were needed to conductobservations over a period of one to two years onChetyrekhbugornyi (“Four-hill”) Island or onBiryuchya Spit on the Northern Caspian coast, inorder to study the faunal composition of theCaspian region and the biology of commercialfishes. In his opinion there was an urgent need forobservations on temperature and salinity at differ-ent depths throughout the entire sea area, on thegas content of the water, for studying the impact ofriver flows on salinity of marine water, etc. From1885 Grimm served as Inspector of Agriculture atthe Ministry of Agriculture and State Property, wasInspector of Fisheries at the Main Administrationof Agriculture and Land Use, was Senior Inspectorof Fisheries (an equivalent of present Minister ofFisheries) and in his official capacity was the headof the Nikolsky fish hatchery, which in 1862, afterthe death of its owner and founder Vladimir



Pavlovich Vrassky, became state property. Grimmwas the founder and editor of the journal “FisheriesBulletin” and an active participant of the Interna-tional Council for the Exploration of the Sea.In 1885 Grimm inspected Astrakhan’ shad fishery

factories. In 1887 at the Astrakhan’ Administrationof Fisheries and Seal Hunting, a small library wasestablished comprising volumes on fisheries,ichthyology, zoology, chemistry, bacteriology andmedicine. At the same time, the foundations of theichthyological museum were laid. For this purpose,in 1888, stuffed skins, ship models and an albumof photographs of the Volga fishery were purchasedusing funds provided by Astrakhan’ fishery man-agers. In 1889, N. N. Pushkarev established asmall chemical laboratory and M. I. Arustamovestablished a bacteriological laboratory at thelibrary and museum. Money for the organizationof these laboratories was allocated by the Commit-tee of the Caspian-Volga Fisheries from fundsraised on a voluntary basis from fishery managersat the request of the committee manager S. I.Marshev. Thus, the sea fishery station inAstrakhan’, the first in Russia, was established. In1891 a suitable building (Vorobyev House) wasrented for the laboratories, where they were trans-ferred together with the museum and the library.In 1904 it became “The Ichthyological Laboratory

at the Administration of Caspian-Volga Fisheriesand Seal Hunting” (Strubalina 1990).Here a small digression is needed in order to

describe briefly the state of fisheries, primarily shadfisheries, on the Caspian Sea in the period preced-ing the organization of the expedition in 1904.It is common knowledge that until the beginning

of the nineteenth century fish trade exploited onlysturgeons (“krasnaya ryba” in Russian) and alsoCaspian inconnu, carp, pike-perch, wels, salmon,while all the other fishes were disposed as unfit foruse, leaving it to those who had no gear for catch-ing sturgeons (Suvorov 1948). Salting of shad (sev-eral species of the genus Alosa) in the Astrakhan’Province did not begin until Baer’s propagandafrom 1854, even though Peter the Great and Lep-ekhin struggled against the bias against shad,which for some reason was regarded to be harmful.By the 1870s salting became predominant over oilmelting from fresh shad.The centre of the shad fishery, initially scattered

throughout the entire Volga delta, moved down-stream to the sites where drag seining for shadmigrating from the sea took place. From the early1890s the marine shad fishery started to increase.Buyers and representatives of fisheries set out to seafollowing fishermen. The shad trade becameextremely profitable and highly competitive. By

Fig. 1. Seining shad, draught.



Fig. 2. Shad fishery factory.

Fig. 3. Inside a shad fishery factory, processing of shad.



1895 sea fisheries grew so much that they exceededriver fisheries, which was an impetus for its spread-ing further out to sea and along both coasts – downto Petrovsk (now Makhachkala) on the western coastand Dolgiye islands, near Aleksandrovsky Fort (laterFort Shevchenko) on Mangyshlak Peninsula andfurther near Krasnovodsk along the eastern coast.Fishing was conducted at sites for drag seining

(Fig. 1) situated around the “promysly’ (fishery fac-tories, Fig. 2) where fish were landed and processed(Fig. 3). An important part of such fishery areaswas the “vykhod” (“gate”), i.e. room for cool salt-ing of fish with large barrels, stocks of ice, andaccommodation for workers.At the end of the nineteenth century more fishes

were caught in the Caspian basin than in any othersea off Russia (300,000 tons in 1900). Accordingto the data of the “Bulletin of Fisheries” for 1895the value of the Russian fish yield at the end of thenineteenth century was 53.4 million golden rou-bles, exceeded only by USA and Great Britain. Thevalue of the fish yield by region was distributed asfollows:Murman and the White Sea: 1.4 million roublesThe Baltic Sea: 2.0 million roubles

The Black Sea: 3.0 million roublesThe Sea of Azov: 6.0 million roubles The Caspian Sea: 30.0 million roublesLakes of the North-European part: 7.0 millionroublesWaters of Middle Asia and Siberia: 2.5 million roublesSea-hunting and mollusc trade: 1.5 million roublesThe Caspian Sea, at that time the major fishery

basin in Russia and rapidly growing, required mea-sures for regulating the fisheries. Causing particu-lar alarm to fishery managers was the decline ofyields of Volga shads [Alosa caspia (Eichwald,1838), Alosa kessleri (Grimm, 1887) and Alosa vol-gensis (Berg, 1913)], constituting at that time morethan 100,000 tons. The catch of shads hadincreased rapidly but in the late 1890s declinedcatastrophically. It became evident that the causesof the decline needed to be identified and regula-tory measures be taken. Some of the Astrakhan’fishery managers adhered to the most rigorousmeasures, demanding the prohibition of fishing inthe sea completely, since sea fishing depleted fish-eries in the lower reaches of the Volga River. Theyasserted that all shads migrated for spawning in the

Fig. 4. Steamer “Geok-Tepe”



Volga. No precise scientific data to refute that viewwere available at that time (Grimm 1896).The Imperial Russian Society of Fish Culture and

Fisheries, founded in 1881 when Grimm occupieda key position, played an important role in theorganization of studies of biological resources inmarine and fresh waters. In the regulations of the

society the main focus was on “promoting studiesof fishes and other representatives of the fauna”. Atthe meetings of the society and in publications,primarily in the Bulletin of Fisheries, discussionswere conducted on the causes of the state of fish-eries in the lower reaches of the Volga River. Thenecessity of “joint work on the organization of an

Fig. 5a-b. a. Steamer “Krasnovodsk”; b. Steam launch “Provornyy”.

b

a



expedition for the study of the Caspian Sea” wasdeclared repeatedly.Grimm worked in close cooperation with Nikolay

Mikhailovich Knipovich. Thus, in spring 1885Knipovich and Grimm surveyed shad fishery fac-tories; Knipovich, a zoology student, made his firststeps in the field of scientific fishery research. In1899 the Stockholm Conference of the Interna-tional Council for the Exploration of the Sea(ICES) was held (Grimm 1899). Archive docu-ments provided information on the joint efforts ofGrimm and Knipovich aimed at promoting Rus-sia’s membership of ICES. The Stockholm Confer-ence was followed by the conference in Christiania(Oslo) in May 1901. Grimm could not attend forreasons of health, and therefore Knipovich was theonly delegate from Russia.Grimm, by virtue of his position and his own sci-

entific background, was always more interested in

the study of the southern seas of Russia than thenorthern. Thus, in 1899 he wrote to the Ministerof Agriculture and State Property A. S. Ermolovconcerning the participation of Russia in ICES,stating that it was more important to study thoseseas where the main Russian fishery was concen-trated, i.e. the Caspian, Azov and Black Seas.Grimm suggested that studies of the Caspian Seawere to be conducted using the same methods asthose used for the Barents Sea, i.e. by a combina-tion of oceanographic and biological studies; there-fore it appears quite natural that Knipovich wasappointed the head of the Caspian expedition.

THE 1904 EXPEDITIONThe question of this study was raised in 1900 by the

firm “Sapozhnikov Brothers” together with otherAstrakhan’ fishery managers who addressed the Soci-ety with a statement about the necessity of sending

Fig. 6. The main members of the expedition, from left to right: Sergei A. Mitropol’sky, Arseny A. Lebedintsev, Vasiliy N.Kononov, Nikolai M. Knipovich and Alexander G. Genkel.



Fig. 7. In the centre, Alexander K. Igumnov, senior navigator captain.

Fig. 8. On deck of “Geok-Tepe“: second row, right, commander of the ship Dmitry E. Nidermiller



Fig. 9. Map of the Caspian Sea and routes of the voyages. Red line – the first voyage of Geok-Tepe; blue line – the third voy-age of Geok-Tepe; green line – the fourth voyage of Geok-Tepe; pink line – the fifth voyage of Geok-Tepe; numbers showlocations of coastal stations, 1 – in Buinak near Petrovsk, 2 – in Belidzhi near Derbent, 3 – on the Agrakhan Peninsula, 4 –in Kilyazi, 5 – in Oranzhereinyi, 6 – at Sineye Mortse, 7 – in Nikolskoye on Magyshlak Peninsula.

an expedition in the interests of Astrakhan’ fisheries.At the same time the Geographic Society informedthe Minister of Agriculture and State Property aboutthe advisability of such an expedition.In February 1901 a representative of Sapozhnikov

Brothers, Alexander Evaldovich Meisner, fisherymanager Ivan Vasilyevich Bezzubikov and otherscontributed 5,000 roubles to the treasury of theSociety “for the initial expenditure on the organi-zation of the appointed expedition by the society”.



Fig. 10. Port Petrovsk (now Makhachkala).

Fig. 11. Persian fortress at Ashur-Ade.



After that the minister officially recognized the neces-sity of conducting the expedition. A commission wasestablished for the development of a scientific planfor the expedition, which included representatives ofthe above two mentioned societies and ministries.Unfortunately a large number of participants of theCommission were “so engrossed in organizing aninternational fishery exhibition and internationalcongress” that the beginning of the expedition waspostponed until spring 1904. Therefore, the Com-mission planned to begin preparations (to buy equip-ment, to find a vessel etc.) at the end of 1903, usingthe sum of 5,000 roubles plus interest for 2.5 years.The appointment of Knipovich as leader of the

expedition was announced on 16 (29) December1903. This appointment was facilitated by the factthat by 1904 Knipovich had been relieved of hisduties in the study of the northern seas, as he was nolonger the head of the Murmansk expedition, havingbeen replaced by his assistant L. L. Breitfus in 1902.The expedition of 1904 was given the official title

of “Expeditions for studies of shad and shad fisheriesin the Caspian Sea”. It is noteworthy that it was thefirst Caspian expedition in the modern sense; it hadclearly outlined goals and tasks subject to a single

plan; several researchers of equal professional levelparticipated. From experience accumulated duringthe Murman scientific fishery expedition, Knipovichrealised the necessity of conducting integratedresearch, oceanographic studies included. Accordingto Svetovidov (1953), this expedition divided thestudies of the Caspian Sea into two periods, beforeand after Knipovich. The hydrobiologist V. A.Vodyanitsky called Knipovich the best zoologistamong oceanographers and the best oceanographeramong zoologists (Vodyanitsky 1975).Thus, the expedition had two goals: firstly, it was

intended to carry out a general investigation of theCaspian and, secondly, it aimed to study the biol-ogy of the Caspian shad (A. caspia) as far as possi-ble in a short period.A large part of the research was conducted on board

the vessel Geok-Tepe (Fig. 4), provided by theMarine Ministry and on board vessels temporarilyreplacing it; the vessel Krasnovodsk (Fig. 5a) in theAstrabad Bay and steam-launch “Provornyi” (Fig.5b) near Baku, and also on board steamer “Strazha”(estuaries of the Volga River and in the delta). Addi-tional studies were carried out in the northern part ofthe sea, mostly on board the steamer “Kreiser”2.

Fig. 12. View of Ashur-Ade.

2 Here and below we base our narration on information from the publication immediately devoted to describing the organization and realizationof the expedition (Arnol'd, 1904, 1907; Borodin, 1904a, 1904b; Geineman, 1904; Knipovich, 1904a, 1904b, 1904c, 1904d; Lebedintsev, 1904a, 1904b, 1904c; Smirnov, 1907) without special references.



Fig. 14. Dwelling of a Persian.

Fig. 13. Persian village, in the centre, Lebedintsev, Genkel, Kononov.



Fig. 16. View of Krasnovodsk (now Turkmenbashi).

Fig. 15. Lower Kura River – the Bankovskiy fishery factory.

The following researchers worked on board thevessels (Fig. 6): Nikolai Mikhailovich Knipovich,Sergei A. Mitropol’sky, Arseny Arsenyevich Lebed-intsev, Alexander Germanovich Genkel and VasiliyNikolayevich Kononov. Mitropol’sky was an assis-tant of Knipovich’s, later the head of the Astrakhan’Ichthyological Laboratory. Lebedintsev was one ofthe most prominent Russian hydrochemists andparticipated in expeditions to the Black Sea wherehe discovered a hydrogen sulphide zone. He under-took an extensive study of the hydrochemistry offreshwater because his main affiliation was theNikolsky Hatchery Farm under the supervision ofGrimm. The major studies of Genkel dealt withthe lower plants and fungi. Based upon the expedi-tion results he published a monograph entitled“Materials on phytoplankton of the Caspian Seafrom the data of the Caspian expedition” (Genkel1909). Later Genkel became an eminent botanist,a founder of the university and botanical garden inPerm’. During the period in question Kononov wasa student at the University of St. Petersburg. Heand Genkel were commissioned by the St. Peters-burg Naturalists Society. Beginning as a botanist,Kononov later studied chemistry and archaeology.He became a famous researcher of the materialnature of monuments and a chemist restorer(Gerasimova & Tikhonov 2003).The senior navigator on “Geok-Tepe” was

Alexander Konstantinovich Igumnov (Fig. 7).Apart from navigation during the entire expeditionhe also undertook detailed meteorological observa-tions (Igumnov 1907). Knipovich and Lebedintsevpointed out the high quality of Igumnov’s research,who had served under the command of the famousnaval commander and scientist vice-admiralStepan Osipovich Makarov on “Vityaz” in thePacific Ocean. The commander of the vesselDmitry Egorovich Nidermiller (Fig. 8) sailed inthe Caspian Sea since 1876. Participants of theexpedition also highly appreciated the assistance ofthe entire crew.It is noteworthy that the Russian Imperial Navy

played an important role in the organization andperformance of scientific research (particularlyexpedition research). The Marine Ministry of Rus-sia played a decisive role in the organization of sci-entific studies. Increased interest in the region inquestion was determined by the strategic interestsof Russia and the southern seas were regarded as apotential centre of operations. In addition, thearmy and navy supported hydrobiological and

ichthyological research as they were interested inbeing supplied with fresh, salted and dried fish.A number of biological problems (biology of ship

worms, fouling organisms, malaria mosquito, assess-ment of the epidemiological situation) were of greatimportance for the Russian navy. All this ensuredthe support of the navy for hydrobiological research(a part of the multi-disciplinary oceanological inves-tigations) in that region. The First Congress of Russ-ian Naturalists and Physicians in 1868 addressed theMarine Ministry with an application to provide sub-sistence and working places for naturalists on navalships setting out on voyages.This application was met by “enlightened under-

standing” of the Chief Commander of the Navy,Grand Prince Konstantin. The Grand Prince, thereformer of the navy, had a “highly favourable atti-



Fig. 17. Vasily Kononov with a bathometer and hydro-chemical analysis equipment.



Fig. 18a-b. a. Lebedintsev and sailors in the process of hydrochemical sampling from on board the steamer. b. Lebedintsevand sailors at a trawl capstan.

a

b

tude to the needs of science”, not only allowing suchparticipation, but he also “ordered to conduct col-lecting from on board vessels sailing on officialduty”. Collecting was made a duty a of ship’s doctorsand volunteering officers; the Imperial Academy ofSciences was to develop instructions for that pur-pose. Naturalists received “subsistence proper totheir rank” from the Marine Ministry; howevermany declined, “regarding the work as the bestreward” (Ignatiyev 2006).The military and political situation in the Caspian

region, which was of strategic importance for Russia,determined not only the possibility of conductingresearch, but also the existence of the naturalist. Thealternating Russian-Turkish and Russian-Persianwars of the seventeenth to nineteenth centuries, per-manent raids by nomadic “non-peaceful nativetribes” only allowed surveys to be conducted underthe protection of “ship cannons”. Special militaryteams accompanied the expeditions of Pallas, Gmelinand Baer. The Caspian Sea was regarded as an innersea of Russia and hence the treasury had its own spe-cific interests there, i.e. “supplying the army andfleets” with fish production (Aliyev 2003).

The absence of a serious marine enemy at the endof the nineteenth century actually reduced theactivities of the Russian fleets to the protection offisheries, seal-hunting and salt industry areas andtransportation functions. A special detachment ofarmed ships was formed for the Caspian fleet, sec-onded to the Astrakhan’ Fishery Administrationfor performing police duties. The navy had also“undertaken obligations” to provide support to theresearch in the region. All the Russian hydrobio-logical expeditions up to 1904 were conducted onboard ships of the Caspian fleet. The steamerGeok-Tepe was a part of the Baku fleet since 1863.It was used as a dispatch-vessel until 1918. Its dis-placement was 1,100 tons and armament compris-ing 47 mm guns (Aliyev 2003).The first voyage of Geok-Tepe (Fig. 9) from 11

March (28 February) to 3 April (21 March) 1904was from Baku to the outskirts of the deepest areas ofthe southern part, thence to the middle part, visitingthe Apsheron Strait and deep areas, and then to Pe -trovsk, Mangyshlak, northwards along the ice, backto Mangyshlak, Petrovsk (Fig. 10) and again to Baku.The second voyage from 15 (2) April to 17 (4)



Fig. 19. Descending of a net from on board the steamer.

April was short because of bad weather. However,launching from “Provornyi”, Knipovich, Genkeland Mitropolsky could visit the fish factory ofMartirosov on the Shikhova Kosa Spit (=Sakhovskaya Kosa, Shakhov Mys, Shikhov Mys,Sah Dili), which is at the tip of Apsheron Penin-sula, and they also observed shad fishing. The third voyage 20 (7) April to 3 May (20 April)

was long, to the south-eastern corner of the sea and

the Astrabad Bay (now Gorgan, or Gurgan, Bay,Fig. 9). In Astrabad Bay, participants of the expe-dition carried out a three-day voyage on board thesteamer “Krasnovodsk”.Astrabad Bay was separated from the sea by the

Miankale Peninsula (= Miyan Kaleh; Potemkin-skaya Kosa in Russian) and Ashur Islands, fromwhich the Ashur-Ade Island was the largest (Figs11-12). Already at that time (before the lowering of



Fig. 20. A card that was placed in a bottle; requests were sent to return found bottles with a filled in card.

the sea level in 1929-1977) it was quite shallow,overgrown by reeds, its depths not more than threemetres. Freshened by the Kara-Su River and manyshallow rivers, it was an excellent place for spawn-ing and foraging by cyprinids, percids and wels.Participants of the expedition also undertook ter-restrial expeditions to examine the local environs(Figs 13-14).Most Moslems in the vicinity of the Caspian Sea

are of the Shiite sect, who follow local fatwas (reli-gious decrees) not allowing them to eat certain fishand shellfish. They were not interested in “kras-naya ryba” (sturgeons), valuable to the Russians.The beginnings of fisheries in the Persian watersorganized by Astrakhan’ fishery managers go backto the late eighteenth century, which was longbefore the Turkmanchai agreement of 1828between Russia and Persia concluded. Persian river



Fig. 21. Examination of shads on board the steamer.

waters were held by leasehold by the Russian fish-ery manager Lionozov since 1872 and he retainedthe lease until 1918. Apart from that, on agree-ment with the Iranian government, which was fre-quently disturbed by Turkmenian tribes, in 1842-1843 the naval station was moved from Sara Islandoff Baku to Ashur-Ade Island, where two navalships with a crew of 130 people and two cannonsstayed permanently (Aliyev 2003).The fourth voyage from 14 (1) May to 18 (5)

May was to the south-western part of the sea (Fig.9). Geok-Tepe passed to the estuaries of the KuraRiver, to the outlet into the Kyzylagach Bay (= Kizil-Agach, Kyzylagadzh, Zaliv Kirova, Qizila-gac), to Lenkoran’ (= Lankaran), Astara and theregion of deeper water in the southern part of thesea. From the estuary of Kura the members of theexpedition team visited the Kura fisheries.Bankovskiy fishery factory (Fig. 15), downstreamfrom Salyany, was well-equipped with fish receiv-ing quays, refrigerators, houses, barracks andstores. The importance of the Bankovskyi fishingindustry is illustrated by the fact that the latter pre-revolutionary leaseholder fishery manager Tagievpaid the Treasury for the right of fishing in thatarea a lease of 937,880 roubles (Suvorov 1948a). The fifth voyage continued from 20 (7) May to

11 June (29 May). Geok-Tepe passed from Bakualong the western coast approximately up to 42°N,then it crossed the sea towards the Peschanyy Cape,passed into the Tyub-Karagan Bay and returned tothe western coast near Petrovsk and passed north-wards to 12-feet roadstead. Geok-Tepe remainedthere on the anchor when Knipovich and Lebed-intsev went to Astrakhan’ and conducted worksfrom on board the steamer Strazha in the lowerreaches of the Volga. Geok-Tepe then passed to thenorth-eastern part of the sea to Mangyshlak,returned to Petrovsk and along the western coast tothe latitude of Derbent, crossed the sea to Kras-novodsk (Fig. 16) and returned to Baku. Addi-tional work in the lower reaches of the Volga Riverand in its mouth was also carried out by Knipovichand Mitropolsky on the steamer “Strazha” from 17(4) June to 19 (6) June.One of the goals of the Caspian expedition was to

organize coastal stations that were established atthe following locations: 1) in Buinak near Petrovsk,N. A. Borodin, 2) in Belidzhi near Derbent (latermoved to the entry into the Agrakhan Bay to theLopatinsky fishery factory on the Agrakhan (=Uch) Peninsula), E. K. Suvorov, 3) in Kilyazi (=

Giliazi) (that station did not have a permanentobserver and was only visited from time to time aswell as some other points of the Baku area, 4) onthe Oranzhereinyi fishery factory in the westernpart of the Volga delta, I. N. Arnol’d, 5) on fisheryfactories of Bezzubikov near Sineye Mortse (in theeastern part of the Volga River), V. Yu. Golynetsand 6) in the village Nikolskoye near Tyub-Kara-gan Bay on Mangyshlak Peninsula (near FortAlexandrovsky), I. N. Arnol’d, N. A. Smirnov.It is necessary to mention the following partici-

pants of the expedition. Nikolay AleksandrovichSmirnov worked in the Murman expedition study-ing seal hunting. From 1903 he was keeper at theAdministration of the Caspian-Volga fish and sealhunting, and in 1905 became the head of theAstrakhan’ Ichthyological Laboratory, but in early1906 he was replaced by his colleague Mitropolsky,who participated in the same expedition; Smirnovwent to participate in the Murman expeditionagain. Evgenii Konstantinovich Suvorov had justgraduated (in 1903) from St. Petersburg Univer-sity, Department of Zoology under the supervisionof the famous hydrobiologist VladimirMikhailovich Shimkevich and was on the staff ofthe Department of Agriculture. Later Suvorovstudied different aspects of applied ichthyology,wrote manuals on fishery and general ichthyology.The textbook “General Ichthyology” written bySuvorov (1948b) still remains an unrivalled work.Nikolay Andreyevich Borodin was an eminent

statistician, ichthyologist and public figure. From1899 to 1906 he served in St. Petersburg at theDepartment of Agriculture as a senior expert infish culture. Later he lived in Uralsk, was in chargeof a statistical economic study of the Ural Cossackdetachment. Borodin was a member of the StateDuma. Ivan Nikolayevich Arnol’d, as well as Lebe-dinsky, worked at the Nikolsky factory and was anexpert in the study of plankton. It was Arnol’d whoestablished the spawning areas of the Caspian shadand put forward the possibility of the developmentof its eggs in marine water. V. Yu. Golynets workedat fishery factories in the eastern part of the Volgadelta in the firm of Bezzubikov.According to Lebedintsev, hydrochemical equip-

ment of the expedition was quite good (Fig. 17). Theexpedition had four bathometers (Petterson-Nansenlarge bathometer, Petterson small bathometer, Petter-son bathometer with a screw, Wil-Timchenko bath-ometer) and different thermometers. After takingdepth measurements, a Nansen plankton net was



deployed on the same rope from the forecastle (Fig.18a-b). It should be noted that important assistancewas rendered by famous polar explorer FridtjofWedel-Jarlsberg Nansen, director of the Central Lab-oratory for international marine research in Christia-nia (= Oslo, Norway). That laboratory produced “thelatest hydrological equipment”, ordered byKnipovich, and provided plankton nets.Apart from dredges for the analysis of the benthos,

a specially designed slide trawl was used (Fig. 19).Knipovich emphasized the benefits of using thetrawl. Catches of fishes in the sea were carried out bynets of different kinds, primarily with shad fixed gillnets with mesh size from 8 mm to 45 mm.Current directions were studied using bottles. For

that purpose they sought the assistance of fisherymanagers in Baku, who purchased 1,000 lemonadebottles, small shot, paraffin, sealing-wax, and corksfor them. A small amount of shot was placed onthe bottom of the bottle, which was fixed withmelted paraffin on the bottom. Thus, the bottlewas fixed in a vertical position in the water. A spe-cial card was placed in each bottle. Requests weresent to fishery factories, local fishermen, steamshipsocieties and fishery inspectors to return foundbottles with a filled in card (Fig. 20).

DISCUSSION As a result of the expedition quite extensive data

were obtained, which allowed the conclusion that“hydrological conditions are underlying the bio-logical phenomena of the basin… Productivity ofthe basin is restricted and it is the complex ofhydrological conditions, which is crucial”.The major results of the expedition (Lebedintsev

1904c, Knipovich 1907, Suvorov 1907, Arnol’d1907) can be summarised as follows:1. The boundary of animal life in the plankton

occurs at a depth of 350 to 400 m. The cause ofthis is apparently the low oxygen content at greaterdepths, but not the presence of hydrogen sulphide,which was only detected in the deepest layers (atdepths below 600 m) of the southern hollow.2. Characteristic features of the vertical distribu-

tion of plankton and benthos of different taxo-nomic groups of invertebrates were revealed for thefirst time.3. An important contribution was made into the

study of taxonomy, infraspecies structure and biol-ogy of shad (Fig. 21). Distinct data were obtainedon the distribution and spawning of the Caspianshad [(A. caspia) and Brazhnikov shad (Alosa brash-

nikowi (Borodin, 1904)] in the sea, their divisioninto local stocks and forms or subspecies. It isshown that the decline of the shad fisheries inLower Volga is the result of excessive fishing.Apart from hydrographic, hydrological and

hydrobiological information, large collections ofdifferent animal taxa were made. The rich collec-tion of crustaceans, both benthic and pelagic, wasgiven to the best specialist in crustaceans of theCaspian Sea, G. O. Sars at Christiania; shad wereexamined by Suvorov, Borodin and Arnol’d, finger-lings of shad were examined by Mitropol’sky, otherfishes were examined by Berg. Other groups of ani-mals and algae were also given to specialists.A large collection of fishes was deposited at the

Zoological Museum of the Imperial Academy ofSciences, now the Zoological Institute of the Russ-ian Academy of Sciences. Six new taxa of speciesrank were described from the material of the expe-dition: Alosa caspia knipowitschi (Iljin, 1927), Alosacaspia persica (Iljin, 1927), Alosa sphaerocephala(Berg, 1913), Alosa volgensis (Berg, 1913), Alosacurensis Suvorov, 1907 and Clupeonella engrauli-formis (Borodin, 1904).Concerning other fish taxa, collections of the expe-

dition still retain their importance today. The uniqueendemic monotypic genus and species Anatirostrumprofundorum Berg, 1927 was described from thematerial of the expedition. There are only a few col-lected specimens of the tadpole-goby Benthophilusctenolepidus Kessler, 1877, an earlier describedspecies, but a very rare one; the same can be said alsoabout Benthophilus leptocephalus Kessler, 1877. Addi-tionally, the expedition also procured specimens ofspecies that were not described until later – Ben-thophilus mahmudbejovi Ragimov, 1976, Ben-thophilus leobergius Berg, 1949. Benthophilus pinchukiRagimov, 1982 (Boldyrev & Bogutskaya 2007).One of the results of the expedition was also the

impetus it gave to all subsequent researchers of theCaspian Sea. Knipovich worked there as supervisorof special scientific fishery expeditions in 1912,1913, and 1914 to 1915 (Knipovich 1921). Heparticipated in the “All-Caspian Scientific FisheryExpedition” of the Caspian Commission of theUSSR Academy of Sciences from 1931 to 1935(Knipovich 1936). In those years he cooperatedwith Anatoliy Nikolayevich Svetovidov, a famousRussian ichthyologist, later the author of the clas-sical monograph “Fauna of Russia. Clupeidae”(Svetovidov 1952). The first author of this paperwas the last Ph.D. student of Svetovidov.



ACKNOWLEDGEMENTSWe thank Yu. A. Laius, V. V. Spodareva, T. I.

Igoshina and staff of the Library of the ZoologicalInstitute of the Russian Academy of Sciences fortheir valuable help in various aspects of preparationof this article. We are grateful to C. Smith (Leices-ter University) for linguistic revision of the manu-script and the reviewers for their very helpful com-ments. The study was supported by the Pro-gramme of the Division of Biological Sciences ofthe Russian Academy of Sciences “BiologicalResources of Russia” and approval of the Presidentfor support of St. Petersburg Ichthyological School.

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