aqua1).pdf · aqua International Journal of Ichthyology Vol. 14 (1), 28 January 2008 Aquapress ISSN...

aquaInternational Journal of Ichthyology

Vol. 14 (1), 28 January 2008

AquapressISSN

0945-9871

aqua vol. 14 no. 1 - 28 January 2008

Managing Editor:

Heiko BleherVia G. Falcone 11,27010 Miradolo Terme (PV), ItalyTel.: +39-0382-754707Fax: +39-0382-754129E-mail: [email protected]

Scientific Editor:

Friedhelm KruppCurator of FishesSenckenberg Research Instituteand Natural History MuseumSenckenberganlage 2560325 Frankfurt am Main, GermanyTel: +49-69-7542.1255Fax: +49-69-7542.1253E-mail: [email protected]

Editorial Board:

Gerald R. AllenDepartment of Aquatic Zoology,Western Australian Museum, Perth, Australia

Nina G. BogutskayaZoological Institute of the Russian Academy ofSciences, St. Petersburg, Russia

Wilson J. E. M. CostaLaboratório de Ictiologia Geral e Aplicada,Departamento de Zoologia,Universidade Federal do Rio de Janeiro, Brasil

Axel MeyerLehrstuhl für Zoologie und Evolutionsbiologie,Universität Konstanz, Germany

Paolo ParentiDepartment of Enviromental Sciences,University of Milano-Bicocca,Milan, Italy

Mário de PinnaMuseu de Zoologia da USP, São Paulo, Brazil

John E. RandallBishop Museum, Honolulu, Hawaii, U.S.A.

Richard WinterbottomCentre of Biodiversity & Conservation Biology,Royal Ontario Museum,Toronto, Canada

Scopeaqua is an international journal which publishes originalscientific articles in the fields of systematics, taxonomy,biogeography, ethology, ecology, and general biology offishes. Papers on freshwater, brackish, and marine fisheswill be considered. aqua is fully refereed and aims at pub-lishing manuscripts within 2-4 months of acceptance. Inview of the importance of color patterns in species iden-tification and animal ethology, authors are encouraged tosubmit color illustrations in addition to descriptions ofcoloration. It is our aim to provide the international sci-entific community with an efficiently published journalmeeting high scientific and technical standards.

Call for papersThe editors welcome the submission of original manu-scripts which should be sent in digital format to the scien-tific editor. Full length research papers and short notes willbe considered for publication. There are no page chargesand color illustrations will be published free of charge.Authors will receive one free copy of the issue in whichtheir paper is published and an e-print in PDF format.

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Special PublicationSince 2003 Aquapress publishes a series of Special Publi-cations, which are produced at irregular intervals. All Spe-cial Publications have about 100 or more pages and areavailable separately from regular issues of aqua. Enquiriesabout subscriptions and prices should be sent to the pub-lisher at the address given here above or by e-mail to:[email protected]

ISSN 0945-9871Publisher: Aquapress, Redazione aqua,I-27010 Miradolo Terme (Pavia), ItalyPrinter: PRINTO s.r.o. – Czech RepublicCopyediting and layout: Rossella Bulla© 2008 aqua, International Journal of Ichthyology

aqua - International Journal of Ichthyology

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 enfasialla spedizione del 1904 guidata da N. M. Knipovich. È sta-to esaminato il contributo di questa spedizione allo studiodella 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 èdato alla continuità delle ricerche condotte sul Mar Caspio.Le illustrazioni dell’articolo sono fotografie originali scattatedurante 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 scientificresults 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 2008 2

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

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, hiszoological 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 (nowMakhachkala) 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 roublesThe Caspian Sea: 30.0 million roublesLakes of the North-European part: 7.0 millionroublesWaters of Middle Asia and Siberia: 2.5 millionroublesSea-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, backtoMangyshlak, 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).

DISCUSSIONAs 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.

REFERENCESALIYEV, N. A. 2003. Marine Fortress at Caspian. Geopo-litical victory of Russia at the Caucasus: from the his-tory of creation of the Caspian fleet in middle 19th –beginning of 20th century. Voyenno-istoricheskii zhurnal[Military Scientific Journal] 7: 38-41. [In Russian].

ARNOL’D, N. 1904. News about the Caspian Expedition: aletter by N.A. Arnol’d. Vestnik rybopromyshlennosti,rybovodstva i rybolovsta [Bulletin of Fish Industry, Fish Cul-ture and Fisheries] 19: 458-460.

ARNOL’D, N. 1907. To biology of Caspian puzanok shadClupea caspia Eichw. In Knipovich, N. M. (Ed.). TrudyKaspiyskoy Ekspeditsii 1: 222-242. [In Russian].

BAER, K. M. 1860. Fisheries in the Caspian Sea and its tribu-taries. Investigations of the state of fisheries in Russia, 2. Min-isterstvo zemledeliya, St. Petersburg, 216 pp. [In Russian].

BOGDANOV, M. 1875. Review of expeditions and naturalhistory research in Aral-Caspian area in 1720 to 1874.Obschestvo Estestvoispytateley, St. Petersburg, 53 pp. [InRussian].

BOGUTSKAYA, N. G. & NASEKA, A. M. 2004. Catalogue ofagnathans and fishes of fresh and brackish waters of Russiawith comments on nomenclature and taxonomy. KMK Sci-entific Press, Moscow, 389 pp. [In Russian].

BOLDYREV, V. S. & BOGUTSKAYA, N. G. 2007. Revision ofthe tadpole-gobies of the genus Benthophilus (Teleostei:Gobiidae). Ichthyological Exploration of Freshwaters 18:31-96.

BORODIN, N. A. 1904a. News about the Caspian Expedi-tion (from letters to Editor). Vestnik rybopromyshlennosti,rybovodstva i rybolovsta [Bulletin of Fish Industry, Fish Cul-ture and Fisheries] 19: 320-322. [In Russian].

BORODIN, N. A. 1904b. News about the Caspian Expedi-tion: a letter by N.A. Borodin. Vestnik rybopromyshlen-nosti, rybovodstva i rybolovsta [Bulletin of Fish Industry,Fish Culture and Fisheries] 19: 331-337. [In Russian].

DANILEVSKY, N. Y. 1869. Rossia i Evropa. Vzglyad na kul-turnye i politicheskie otnoshenia slavyanskogo mira kgermano-romanskomu. [Russia and Europe: AnApproach to Cultural and Political Attitudes of the SlavicWorld to the German-Romanic One]. Zarya (1869) nos1-6, 8-10. [In Russian].

EICHWALD [C.] E., [VON] 1831. Zoologia specialis quamexpositis animalibus tum vivis, tum fossilibus potissimumRossiae, in universam, et Poloniae in specie, in usum lec-tionum publicarum in universitate caesarea vilnensi haben-darum. Pars posterior specialem expositionem spondylozoo-rum continens. Joseph Zawadski, Vilna [Vilnius], 404pp., 2 pls. [Nona Classis. Pisces, pp. 10-116.]

EICHWALD [C.] E., [VON] 1838. Faunae Caspii marisprimitiae. Bulletin de la Societé Impériale des Naturalistesde Moscou 11:125-174.

EICHWALD [C.] E., [VON] 1841. Fauna caspio-caucasianonullis observationibus novis illustravit Edwardus Eich-wald. Petropoli [St. Petersburg], 233 pp.

FILIPPI F., DE. 1863. Nuove o poco note specie di animalivertebrati raccolte in un viaggio in Persia nell’estate del-l’anno 1862. Archives de Zoologie, Anatomie et Fisiologie(Genova) 2: 377-394.

FILIPPI F., DE. 1865. Note di un viaggio in Persia nel 1862.Milano, i-viii+396 pp.

GEINEMAN, A. 1904. Report on activities of Imperial Russ-ian Society of Fish Culture and Fisheries for the year1903. Vestnik rybopromyshlennosti, rybovodstva i rybolovsta[Bulletin of Fish Industry, Fish Culture and Fisheries] 19:363-384. [In Russian].

GENKEL, A. G. 1909. Materials to phytoplankton of theCaspian Sea. Botanicheskiye zapiski [Botanical memoirs]27: 47-87. [In Russian].

GERASIMOVA, N. G. & TIKHONOV, P. A. 2003. Researcherand restorer of monuments of culture V.N. Kononov.Relikvia 3: 60-63. [In Russian].

GMELIN, S. G. [1770]-1784. Reise durch Russland, zur Unter-suchung der drei Naturreiche. Kaiserliche Akademie derWis-senschaften, St. Petersburg, 1-4, 182+260+508+ 268 pp.

GÖBEL [GOEBEL], C. CH. F. 1837-1838. Reise in die Step-pen des südlichen Russland. Dorpat, 1-2, 325+593 pp.

GRIMM, O. A. 1876. Caspian Sea and its fauna. In:Grimm, O. A. (Ed.). Trudy Aralo-Kaspiyskoy Ekspeditsii,2 (1). Obschestvo estestvoispytateley, St.Peterburg, iii-v +7-168 pp. [In Russian].

GRIMM, O. A. 1877. Caspian Sea and its fauna. In:Grimm, O. A. (Ed.). Trudy Aralo-Kaspiyskoy Ekspeditsii,2 (2). Obschestvo estestvoispytateley, St.Peterburg,i-ii+1-105 pp. [In Russian].

GRIMM, O. A. 1896. Caspian-Volga Fisheries. Ministerstvozemledeliya, St. Petersburg, 154 pp. [In Russian].

GRIMM, O. A. 1899. The international conference onexploration of North seas held in Stokholm in June 1899.Vestnik rybopromyshlennosti, rybovodstva i rybolovsta [Bul-letin of Fish Industry, Fish Culture and Fisheries] 14: 405 -420. [In Russian].

GUELDENSTAEDT [GÜLDENSTÄDT] A. I., [VON]. 1772.Salmo leucichthys et Cyprinus chalcoides descripti. NoviCommentarii Academiae Sciantiarum Imperialis Petropoli-tanae 16 (1771): 531-547.

GUELDENSTAEDT [GÜLDENSTÄDT] A. I., [VON]. 1773.Cyprinus capoeta et Cyprinus mursa descripti. Novi Com-mentarii Academiae Sciantiarum Imperialis Petropolitanae17 (1772): 507-521.



GUELDENSTAEDT [GÜLDENSTÄDT] A. I., [VON]. 1781.Cyprinus barbus et Cyprinus capito descripti. Acta Acade-miae Scientiarum Imperialis Petropolitanae 2 (1778): 239-260.

IGNATIEV, S. M. 2006. Russian navy and hydrobiologicalexpeditional exploration of South seas (end of 19th –beginning of 20th century). Electronic publication:http://www.navycollection.narod.ru/fleets/Russia/flot.html. [In Russian].

IGUMNOV, A. 1907. Meteorological journal performed onboard the steamer Geok-tepe in 1904 from March 12 tillJune 11 of New Style. In: Trudy Kaspiyskoy Ekspeditsii1904 goda [Reports of the Caspian Expedition in the year1904], 1 (Ed. N. M. Knipovich.): 115-138. GlavnoyeUpravleniye Zemleustroystva I Zemledeliya,St.Peterburg. [In Russian].

KESSLER, K. F. 1874. Description of fishes belonging tocommon families for the Black and Caspian seas. TrudySankt-Peterburgskogo obschestva estestvoispytateley [Proceed-ings of Saint-Petersburg Society of Naturalists] 5: 191-324.[In Russian].

KESSLER, K. F. 1877. [Fishes of the Aralo-Caspio-Pontianichthyological region]. Trudy Aralo-Kaspiyskoy Ekspeditsii,4. Obschestvo estestvoispytateley, St.Peterburg, xxviii +360 pp., 8 pls. [In Russian].

KNIPOVICH, N. M. 1904a. News about the Caspian Expe-dition: from letters by N. M. Knipovich. Vestnik rybo-promyshlennosti, rybovodstva i rybolovsta [Bulletin of FishIndustry, Fish Culture and Fisheries] 19: 255-259. [InRussian].

KNIPOVICH, N. M. 1904b. News about the Caspian Expe-dition (from letters to Editor). Vestnik rybopromyshlen-nosti, rybovodstva i rybolovsta [Bulletin of Fish Industry,Fish Culture and Fisheries] 19: 315-316, 316-318. [InRussian].

KNIPOVICH, N. M. 1904c. News about the Caspian Expe-dition: a letter by N.M. Knipovich. Vestnik rybopromysh-lennosti, rybovodstva i rybolovsta [Bulletin of Fish Industry,Fish Culture and Fisheries] 19: 337-339. [In Russian].

KNIPOVICH, N. M. 1904d. News about the Caspian Expe-dition: a letter by N.M. Knipovich. Vestnik rybopromysh-lennosti, rybovodstva i rybolovsta [Bulletin of Fish Industry,Fish Culture and Fisheries] 19: 460-464. [In Russian].

KNIPOVICH, N. M. 1907. General review of the CaspianExpedition of 1904. Trudy Kaspiyskoy Ekspeditsii 1904goda [Reports of the Caspian Expedition in the year 1904],1. Glavnoye Upravleniye Zemleustroystva i Zemledeliya,St.Peterburg, 113 pp. [In Russian].

KNIPOVICH, N. M. 1921. Hydrological investigations inthe Caspian Sea in the years 1914-1915. In: Reports of theCaspian Expedition in the years 1914-1915, 1. (Ed. N. M.Knipovich.): 1-943. Gosudarsvennoye Izdatel’stvo, Peter-burg. [In Russian].

KNIPOVICH, N. M. (Ed.) 1936. The gulfs of the Caspian SeaKomsomolets (Mertvyi Kultuk) and Kaidak (Compex Stud-ies of the Caspian Sea, Vol. 1, part 1). Izdatel’stvoAkademii Nauk SSSR, Moscow-Leningrad, 231 pp. [InRussian].

LEBEDINTSEV, A. A. 1904a. News about the Caspian Expe-dition (a letter of Lebedintsev A. A.). Vestnik rybo-promyshlennosti, rybovodstva i rybolovsta [Bulletin of FishIndustry, Fish Culture and Fisheries] 19: 251-255.

LEBEDINTSEV, A. A. 1904b. News about the Caspian Expe-dition (from letters to Editor). Vestnik rybopromyshlen-nosti, rybovodstva i rybolovsta [Bulletin of Fish Industry,Fish Culture and Fisheries] 19: 319-320.

LEBEDINTSEV, A. A. 1904c. News about the Caspian Expe-dition (a letter of A.A. Lebedintsev). Vestnik rybopromysh-lennosti, rybovodstva i rybolovsta [Bulletin of Fish Industry,Fish Culture and Fisheries] 19: 414-420.

LUKINA, T. A. (Comp.) 1984. Scientific Heritage. Vol. 9.Caspian expedition of K.M. Baer in 1853-1857. Diariesand materials. Nauka, Leningrad, 558 pp. [In Russian].

PALLAS P. S. 1771. Reise durch verschiedene Provinzen desRussischen Reichs. Erster Theil. Physicalische Reise durchverschiedene Provinzen des Russischen Reichs im 1768 und1769sten Jahre. Kaiserliche Akademie der Wis-senschaften, St. Petersburg, [10] +504 pp.

PALLAS P. S. 1773a. Reise durch verschiedene Provinzen desRussischen Reichs. Zweyter Theil. Erstes Buch vom Jahr1770. Physicalische Reise durch verschiedene Provinzen desRussischen Reichs im 1770sten Jahr. Kaiserliche Akademieder Wissenschaften, St. Petersburg, S. [4]+1-368 pp.

PALLAS P. S. 1773b. Reise durch verschiedene Provinzen desRussischen Reichs. Zweiter Theil. Zweites Buch vom Jahr1771. Kaiserliche Akademie der Wissenschaften, St.Petersburg, 371-744 pp.

PALLAS P. S. 1776a. Reise durch verschiedene Provinzen desRussischen Reichs. Dritter Theil. Vom Jahr 1772 und 1773.Kaiserliche Akademie der Wissenschaften, St. Petersburg,[20]+1-454 pp.

PALLAS P. S. 1776b. Reise durch verschiedene Provinzen desRussischen Reichs. Reise aus Sibirien zurück an die Wolga in1773sten Jahr. Das Dritten Theils. Zweytes Buch. Kaiser-liche Akademie der Wissenschaften, St. Petersburg, 457-760+[26] pp.

PALLAS P. S. 1787. Piscium novae species descriptae. NovaActa Academiae Sciantiarum Imperialis Petropolitana 1:347-360.

PALLAS P. S. 1814. Zoographia rosso-asiatica, sistens omniumanimalium in extenso Imperio Rossico et adjacentibusmaribus observatorum recensionem, domicilia, mores etdescriptiones anatomen atque icones plurimorum. Vol. 3.Animalia monocardia seu frigidi sanguinis. Academia Sci-entiarum, Petropolis [St. Petersburg], 428 pp. [Index onpages i-vii before the main text and Content on pages i-cxxv after the main text were added later and publishedin 1831.]

SMIRNOV, N. 1907. Report on activity of Mangyshlak sta-tion of the Caspian Expedition. In: Trudy Kaspiyskoy Eks-peditsii 1904 goda [Reports of the Caspian Expedition in theyear 1904], 1. (Ed. N. M. Knipovich.): 200-214.Glavnoye Upravleniye Zemleustroystva i Zemledeliya, St.Petersburg. [In Russian].

STRUBALINA N. K. 1990. From history of the exploration offish resources of Caspian and Astrakhan’ Region. Nizh-



nevolzhskoye knizhnoye izdatel’stvo, Volgograd, 95 pp.[In Russian].

SUVOROV E. K. 1907. Sketches on study of Caspian shads.In: Trudy Kaspiyskoy Ekspeditsii 1904 goda [Reports of theCaspian Expedition in the year 1904], 1. (Ed. N. M.Knipovich.): 139-199. Glavnoye Upravleniye Zem-leustroystva i Zemledeliya, St. Petersburg. [In Russian].

SUVOROV E. K. 1948a. Fishery water bodies of the USSR.Izdatel’stvo Leningradskogo Gosudarsvennogo Univer-siteta, Leningrad, 238 pp. [in Russian].

SUVOROV E. K. 1948b. Obschaya Ikhtiologii (General Ichthy-ology). Sovetskaya Nauka, Moscow. 579 pp. [In Russian].

SVETOVIDOV, A. N. 1952. Herrings: Clupeidae. The Faunaof the USSR II, 1. USSR AS Press. Moscow-Leningrad,223 pp. [In Russian].

SVETOVIDOV, A. N. 1953. To memory of NikolayMikhailovich Knipovich. Voprosy Ikhthiologii 1: 128-132.[In Russian].

YAKOVLEV, V. E. 1870. On the new and little-knownspecies of fish found in the mouths of the Volga. ProtocolZasedanii Obshchestva Estestvoispytateley Kazanskogo Uni-versiteta (1870): 101-111. [In Russian].

VODYANITSKIY, V. A. 1975. Memoirs of the naturalist.Nauka, Moscow, 57 pp. [In Russian].



AbstractTwo populations of the schooling zooplanktivore,

Chromis viridis (Teleostei: Pomacentridae), observed inMadang Lagoon, Papua New Guinea, exhibited a highdegree of localized spawning synchrony. The reproductiveactivities associated with synchronized spawning fell intothree distinct phases, including an initial aggregationphase, a subsequent spawning phase, and a final embryo-guarding phase. These activities lasted a total of five daysat each site. The aggregation-spawning sequence wasclosely associated with a semi-lunar cycle at the time of thenew and full moon, indicating that lunar cues are used togenerally synchronize activities among interacting individ-uals at a particular site. However, the time of spawning forWongad and Tab populations were not precisely coinci-dent. This absence of temporal alignment in two, relativelyclosely situated C. viridis populations indicates that morelocalized, possibly site-specific, cues determine the fine-scale timing for the onset of the reproductive sequence.

ZusammenfassungZwei Populationen des schwarmbildenden, Zooplankton

fressenden Chromis viridis (Teleostei: Pomacentridae), diein der Madang-Lagune, Papua-Neuguinea, beobachtetwurden, zeigten einen hohen Grad an ortsbezogener Syn-chronizität beim Laichverhalten. Das Fortpflanzungsver-halten, das zum synchronisierten Ablaichen führte,bestand aus drei Phasen: der einleitenden Gruppierungs-phase, einer anschließenden Ablaichphase und einerabschließenden Bewachungsphase. Diese Aktivitätendauerten an beiden Plätzen insgesamt fünf Tage. DieGruppierungs-Ablaich-Sequenz war eng mit einem Halb-mondzyklus bei Neumond und bei Vollmond verbunden,was darauf hinweist, dass allgemein das Mondlicht beiinteraktiven Tieren vor Ort dazu dient, Akitivitäten zusynchronisieren. Die Ablaichzeiten fielen jedoch bei denPopulationen von Wongad und Tab nicht präzise zusam-men. Das Fehlen der zeitlichen Ausrichtung bei zwei C.-viridis-Populationen in relativ großer räumlicher Nähespricht dafür, dass mehr ortsbezogene oder raumspezifis-che Merkmale die Feinabstimmung der Fortpflanzungsse-quenz beeinflussen.

RésuméDeux populations du zooplanctivore vivant en bancs,

Chromis viridis (Teleostei: Pomacentridae), observées dans lelagon de Madang, Papouasie-Nouvelle-Guinée, ont montréun haut degré de synchronisme dans la localisation de laponte. Les activités de reproduction associées à la ponte syn-chronisée se déroulaient en trois phases distinctes: une phaseinitiale de rassemblement, une phase subséquente de ponteet une phase finale de surveillance des embryons. Ces activ-ités ont duré cinq jours au total dans chaque site. Laséquence rassemblement-ponte était étroitement associée àun cycle semi-lunaire au moment de la nouvelle et de lapleine lune, indiquant que des signaux lunaires sont utiliséspour favoriser les activités synchrones entre individus eninteraction sur un site donné. Pourtant, l’époque de la pontedes populations de Wongad et de Tab n’était pas en parfaitecoïncidence. Cette absence d’alignement temporel pourdeux populations relativement voisines de C. viridis montreque d’autres signaux localisés, front peut-être particuliers ausite, déterminent le moment tout à fait précis du début de laséquence de reproduction.

SommarioDue popolazioni della specie gregaria e zooplanctivora

Chromis viridis (Teleostei: Pomacentridae), studiate nellalaguna di Madang, Papua Nuova Guinea, mostravano unalto grado di fecondazione sincronizzata. Le attività ripro-duttive associate con la fecondazione sincronizzata ricade-vano in tre fasi distinte: un’iniziale aggregazione, una suc-cessiva fase di fecondazione e una fase finale di protezionedell’embrione. In ogni sito queste attività si protraevanoper un totale di cinque giorni. La sequenza di aggrega-zione-fecondazione era strettamente associata con la fase diluna nuova e luna piena, indicando che le fasi lunari sonodeterminanti per sincronizzare le attività generali tra gli in-dividui che interagiscono in un sito particolare. Tuttavia, itempi di fecondazione per le due popolazioni di Wongad edi Tab non erano coincidenti in modo preciso. Questa as-senza di allineamento temporale in popolazioni di C. viri-dis situate in stretta prossimità indica che segnali più loca-lizzati, probabilmente sito-specifici, determinano l’esattatempistica per l’inizio della sequenza riproduttiva.



Observations on spawning behavior and periodicity inthe Bluegreen Chromis (Pomacentridae: Chromis viridis),

in Madang Lagoon, Papua New Guinea

Kathleen S. Cole

Department of Zoology, 2538 McCarthy Mall, University of Hawaii at Manoa, Honolulu,HI 96822, USA. E-mail:[email protected]

Received: 18 September 2007 – Accepted: 12 October 2007

INTRODUCTIONChromis viridis (Cuvier, 1830), also known as the

blue chromis, or bluegreen chromis, is an Indo-Pacific member of the family Pomacentridae. It’srange extends from the Red Sea and the east coastof Africa in the west to the Line Islands, Marque-san Islands and Tuamoto Archipelago in the east,the Ryukyu Islands in the north, and to New Cale-donia in the south (Myers, 1999). Chromis viridisis a zooplanktivore that forms large shoals in thewater column, a behavior that appears to increaseforaging efficiency (Smith & Warburton, 1992).During the day, schools typically retain a closeproximity to branching thickets of staghorn coral(Acropora spp.) and at night, these same branchingcorals act as protective retreats from nocturnalpredators (Allen 1991). The generalized body colorof C. viridis is an intense, almost neon, blue (Fig.1). Against this blue background, greenish and/oryellowish hues can appear due to the presence andorientation of melanophores, xanthophores andpartially motile iridophores within the dermis(Fujii et al. 1989). Among nesting males, the bodytypically has a pronounced yellow-green col-oration, a progressive darkening of the dorsumgoing from anterior to posterior and a partially orcompletely darkened caudal peduncle and tail (Fig.2) (also see Allen 1991, Myers 1999). The habit ofshoaling behavior, coupled with a brightly visiblecoloration, makes this species highly conspicuousin reef habitats, and it has been the subject ofnumerous reef-based studies (Swerdloff 1970, Sale1971, Sweatman 1985, Amorin 1996, Öhman et.

al. 1998, Coughlin & Strickler 1990, Lecchini et.al. 2005, Lecchini et. al. 2006).In terms of reproductive biology, C. viridis is typi-

cal among pomacentrids in ovipositing demersaleggs and demonstrating paternal egg-guardingbehavior (Swerdloff 1970; Sale 1971). However, as aschooling zooplanktivore, the production of demer-sal eggs requires temporary access to suitable sub-strate for egg deposition and subsequent defense ofembryos until hatching, as well as a means of syn-chronizing male and female reproductive activitiesin order to maximize reproductive opportunitieswhile minimizing substrate-associated time. I hadthe opportunity to note a number of synchronousspawning episodes involving C. viridis while carry-ing out a reproductive study on another damselfishspecies, Amblyglyphidodon curacoa, in MadangLagoon, Papua NewGuinea. The following is a briefdescription of spawning activity and periodicity intwo populations of C. viridis located on the north-east coast of Papua New Guinea.

METHODS AND MATERIALSThe observations involving C. viridis took place at

two study sites located in Madang Lagoon, PapuaNew Guinea. Madang Lagoon is located on thenortheast coast of Papua New Guinea in MadangProvince (5°S, 145°30’E). The lagoon is 25-30 m indepth and contains numerous patch reefs and coralislands (Pandolfi & Minchin 1995). The two sites atwhich spawning observations of C. viridis were car-ried out were Tab and Wongad. Tab (Pig) Island ison the narrow barrier reef that forms the seaward


Observations on spawning behavior and periodicity in the Bluegreen Chromis (Pomacentridae: Chromis viridis), in Papua New Guinea

Fig. 1. Chromis viridis with non-reproductive coloration, Batangas, Luzon. Photo by J. E. Randall.

margin of Madang Lagoon, while Wongad Island islocated in the more exposed, central portion ofMadang Lagoon and is subject to considerable waveaction (Kohn 2001). For a detailed map of the studyarea, which includes both Tab and Wongad Islands,see Jebb & Lowry (1995) or Langer & Lipps (2003).Both study sites were located on the leeward side oftheir respective islands and were characterized by ashallow sandy slope with both branching and mas-sive (e.g. Porites spp., Montastrea spp.) corals. Atboth locales, there were large thickets of staghorncoral (Acropora spp.) broadly skirted by sand andcoral rubble substrate at a depth of 3-4 m. Using finkick counts to estimate distance, theWongad site inwhich observations were made measured 10 m x24 m and the Tab site measured 10 m x 42 m.Morning observations, carried out on SCUBA from9:00 AM to 12:30 PM, consisted of noting the pres-ence or absence of non-feeding aggregations of C.viridis at the two study sites and, when appropriate,the general nature of reproductive behaviorsobserved. Each of the two sites was visited near-dailyfor a total of 55 days over a 63-day period fromNov.18, 1994 to Jan.19, 1995.

RESULTSOver the 63-day period, the development of largeaggregations of C. viridis occurred five times ateach site (Fig. 3). These aggregations were conspic-uous by a generalized lack of observed feedingbehavior and by their absence from these sites dur-ing intervening periods. On each of the five occa-sions of non-feeding aggregation formation, the

progression of activities followed the same patternconsisting of three distinct, sequential phases.The first phase, referred to as the aggregation-for-

mation phase, was characterized by the appearanceof large, non-feeding aggregations of C. viridisadjacent to prominent stands of Acropora spp.These groupings consisted of near-stationaryschools positioned approximately 1.5-2 m oversand and coral rubble substrate. Individuals withinan aggregation exhibited a form of milling behav-ior, with individuals oriented and moving in dif-ferent directions, rather than having synchronizedmovements and orientation typical of feedingshoals. Directly below the hovering aggregation,numerous additional C. viridis were seen clusteringdirectly above the sand and coral rubble substrate.No fish were collected for measurements, but sub-strate-associated individuals and those in the hov-ering aggregations generally appeared to be simi-larly sized. The attentions of the substrate-associ-ated C. viridis were often focused on pieces ofalgae-covered coral rubble, and frequently involvedinspecting and mouthing substrate elements. Theyalso exhibited aggressive behavior characterized bychasing other substrate-associated conspecificsaway from the immediate vicinity of the resident-occupied area. The size of a single defended areaappeared to be relatively small, as large numbers ofsubstrate-associated individuals always appeareddensely packed over the substrate (Fig. 4). Individ-uals in the hovering aggregations located wellabove the substrate were generally blue or blue-green in color. However, at any one time, approx-


Kathleen S. Cole

Fig. 2. Chromis viridis with yellow-green coloration and dorsal darkening associated with male spawning behavior, Eniwetok.Photo by J. E. Randall.

imately 5% of substrate-associated individualsexhibited a differing color pattern consisting ofyellow-green coloration similar to that shown inFig. 2. This color variation was stable over the 1-2h periods per day during which aggregations wereobserved. During the aggregation-formation phase(starting on what is here referred to as Day 1 of thereproductive cycle), spawning was never observed.However, as observations were only carried outduring morning hours, the possibility that spawn-ing may have occurred later on Day 1 cannot beruled out.The second phase of reproductive activity,

observed on the day following aggregation forma-tion (i.e. Day 2), was characterized by widespreadspawning activity. A spawning event was initiatedwhen an individual from within the hovering

aggregation descended to the substrate. This typi-cally elicited a strong response from one or more ofthe substrate-defending fish, including a rapidapproach, conspicuous posturing in either a head-down or head-up position, occasional signal jumps(sensu Sale 1971) followed by rapid movementsback to the defended area. Site-defending fish wereidentifiable as male by the sustained distention of apointed genital papilla, distinct from that of visit-ing conspecifics from within the hovering aggrega-tion. The distended papilla of the latter was rela-tively short and had a blunt terminus broadenough to allow for egg release, indicating that vis-iting fish from the hovering aggregation werefemale. Upon entering a defended site, a visitingfemale typically moved around the male territorybriefly, then initiated repeated linear passes directly



Table I. Occurrence of colonial, synchronized spawning and lunar cycle pattern, where known, among Chromis spp. x indi-cates affirmative; – indicates negative.

Semi-lunar Other Degree ofLunar- once (at new and reported form Synchrony

Chromis species Reported synchrony on lunar cycle full moons) of periodicity Reported Reference

Chromis caeruleus, Eniwetok yes (single observation) unknown unknown 3 d after full unknown Swerdloff 1970moon

Chromis caeruleus, Eniwetok not reported (single unknown unknown 1 d after full unknown Maciolek (pers. comm)observation moon reported in Swerdloff

1970

Chromis cyanea yes x _ from week before moderate de Boer 1978to week after fullmoon

Chromis dispilus (temperate) yes _ _ every 8-10 days high Tzioumis & Kingsford1995

Chromis dispilus (temperate) yes (single observation) unknown unknown 3 d after full unknown Russell 1971moon

Chromis hypsilepis yes _ x peaks 1 d after moderate Gladstone 2007new moon and1 d before fullmoon

Chromis multilineata, yes (single observation) unknown unknown 8 d post-full unknown Myrberg et al. 1967south Bimini moon

Chromis multilineata, yes x _ 5 d after full high Robertson et al.Punta de San Blas moon 1990

Chromis multilineata, yes unknown unknown every four weeks reported as Albrecht 1969Punta de Betin, Santa Marta, high butColumbia details not

provided

Chromis notata (temperate) yes, particularly at height _ _ variable variable, Ochi 1986Mukaishima Island, Japan of spawning season pulses with

broadshoulders

Chromis notata (temperate) yes _ x more tighty high Nakazono et al. 1979Koinoura (northern Kyushu), lunar-synchronousJapan during mid-

spawning season

over the substrate (i.e. skimming, sensu Myrberg etal. 1967), accompanied by conspicuous lateralquivering of the body. These movements appearedhighly stereotypic, and were typical of ovipositionobserved by the author in other damselfishes inwhich egg deposition was visually confirmable (e.g.Cole & Sadovy 1995; Cole, unpublished data).The focus of these passes was usually a piece ofcoral rubble or other substrate-associated promi-nence within the male-defended area.The male either simultaneously moved with, or

directly followed, the path of movement of the vis-iting female until she finished spawning move-ments and left the male-defended area. Pieces ofrubble removed from the site after the completionof tandem female-male passing behavior and exam-ined using a stereo microscope showed the pres-ence of numerous small, near-transparent eggswhich in this (Swerdloff 1970) and other (Myrberget al. 1967) Chromis species are typical of recentlyspawned eggs. During the spawning phase,observed spawning activity was widespreadthroughout the site and typically, when observa-tions were made on several consecutive days, per-sisted over a two-day period (i.e. Days 2-3 of thereproductive cycle) (Fig. 3) .The third phase of reproductive activity was

marked by a lack of spawning activity and by thecontinued presence of site-defending males overtheir respective nest sites. During this phase,mouthing behavior of the nest site was occasionallyobserved. In contrast, head-up and head-down dis-

plays, which were observed during the aggregationand spawning phases were rarely observed after thespawning phase. In addition, the number of fishforming hovering aggregations over the nestingarea was markedly reduced. Male activities duringthe third phase included hovering directly over thenest site, chasing out intruders, and making peri-odic visual inspections of the nest site. This phaseof nest defense typically lasted for 1-2 days follow-ing widespread spawning. The end of the nestdefense phase was marked by the abandonment ofthe spawning area by both overhead aggregationsand substrate-defending fish.The sequence of large aggregation formation,

widespread spawning and subsequent site defensebehaviors was observed five times on each site overthe 63 day period and coincided with both the newand full moons (Fig. 3). During widespread spawn-ing, reproductive activities were observed bothwithin the site and extending into contiguous visi-ble areas. However, as other locations at Tab andWongad were not visited, it is unknown how local-ized, or extensive, such spawning activity was ateach island. Within a site, spawning was highlysynchronous. For example, aggregations alwaysappeared on a site within a 24 h period (i.e. fromone days’ observations to the next) and similarly,after the nest defense phase, abandoned the sitebetween one days’ observations and the next. Inthree instances where observations included thespawning peak and at least one day before and after(Dec. 16-21, 2004 for Tab and Wongad, and Jan.


Kathleen S. Cole

Fig. 3. Frequency of widespread (i.e. aggregation) and isolated spawning events exhibited by Chromis viridis at Tab and Won-gad Islands, relative to the lunar cycle, between November 18, 1994 and January 19, 1995. Grey bars are non-observationaldays, black rectangles are days of observed widespread, or isolated, spawning events, respectively.

16-19, 2005 at Tab), mass spawning continued fortwo successive days, but never longer (Fig. 3).Although interruptions in observations (indicated inFig. 3 by gray vertical bars) precluded confirming arestricted 2-day spawning phase in the remainingseven widespread spawning events, partial observa-tions were consistent with the occurrence of nomore that a 2-day period for spawning. The onlyexceptions to this pattern included three instances(Nov. 25 and Jan. 5 at Wongad and Nov. 22 at Tab)when isolated spawning events, consisting of one ortwo spawning pairs, were observed outside of thetime of widespread spawning. In each of these cases,spawning was only observed on a single day andoccurred anywhere from 3-6 days following wide-spread spawning at the same site (Fig. 3).Although reproductive activities appeared to be

highly synchronous within a site, the timing ofspawning for C. viridis observed at Wongad Islandwas non-coincident with that of C. viridis observedat the Tab Island site (Fig. 3). In four of the fivereproductive cycles, C. viridis at Wongad were seenspawning at least one day prior to any spawningobserved at Tab Island (Fig. 3).Initial observations of semi-lunar spawning

occurred during the dry season when water claritywas good. The onset of the wet season, which

started on Dec. 5, 1994, brought substantiallyreduced water visibility (see Fig. 4). However, therewas no evidence that the deterioration in visibilitydisrupted the continued occurrence of bi-monthlyspawning aggregations (Fig. 3).Discussion: Chromis viridis observed at two sites

in Madang Lagoon, Papua New Guinea exhibited ahigh degree of localized spawning synchrony, as evi-denced by the occurrence of widespread spawningover a two-day period close to the time of the newand full moon.The reproductive activities associatedwith spawning fell into three distinct phases: aggre-gation formation; spawning; and embryo defense.Based on the behaviors observed, the aggregationformation phase defined a time of nest preparationon the part of males and of attaining close proxim-ity to male nests on the part of females. This wouldnot be a necessary pre-spawning phase for territorialand/or substrate-associated pomacentrids thatalready occupy, or are in close proximity to, poten-tial nest sites. However, for roving-shoaling zoo-planktivores such as C. viridis, males need to estab-lish short-term nest sites and females need to maxi-mize proximity to nesting males prior to ovulation.Thus, the aggregation formation phase is a necessarypreamble to successful demersal spawning. A wide-spread spawning phase directly followed the aggre-



Fig. 4. Chromis viridis spawning aggregation at the Tab Island site at approximately 3 m depth. Reduced visibility was dueto the recent onset of the rainy season, an event that had no apparent effect on the formation of spawning aggregations.Photo by K. S. Cole.

gation phase and lasted no more than two days. Thethird and last phase of embryo defense was alsocompleted within two days, marked by the depar-ture of substrate-defending males. It is assumed thatthe disappearance of the guarding males was coinci-dent with hatching. The entire reproductive cycle,from aggregation formation to nest abandonment,lasted a total of five days.The duration of embryonic development reported

here for C. viridis is relatively short, compared toother pomacentrids. Among damselfishes (i.e.Pomacentrinae), reports of embryonic developmen-tal periods range from three days for Dascyllus spp.(Thresher 1984) up to six days for some Eupoma-centrus and Pomacentrus spp. (Thresher 1984);among anemonefishes (Amphiprioninae), develop-ment takes six to seven days (Allen 1991). Amongtropical chromines (Chrominae), developmentaldurations include two days for C. caeruleus (Sale1971), three days for C. lineata (Robertson et al.1990), C. multilineata and C. cyanea (Albrecht1969), and four and a half days for C. hypsilepis(Gladstone 2007). Seasonal differences in watertemperatures have been shown to have a significanteffect on the developmental time for C. notata,which can be as short as three days (Nakazono et al.1979) and as long as 12 days (Ochi 1986) depend-ing on seasonal water temperatures. Consequently,the relatively short developmental period for C.viridis reported here may reflect, at least in part,Papua New Guinea’s proximity to the equator andconsistently warm water temperatures, which rangeon average from 27-30 EC (Langer & Lipps 2003).There have been numerous reviews of temporal

spawning patterns among tropical and subtropicalpomacentrids (Thresher 1984; Gladstone & West-oby 1988; Robertson et al. 1990; Tzioumis &Kingsford 1995; Asoh 2003). Documented patternsof damselfish spawning show a broad range of tem-poral expression and include lunar, semi-lunar, andcontinuous with or without lunar-cycle-associatedpeaks (Robertson et al. 1990). Among planktivo-rous, schooling pomacentrid species, spawning syn-chrony is prevalent, frequently with a lunar cycle-associated component (Gladstone &Westoby 1988;Robertson et al. 1990; Tzioumis & Kingsford1995). The two populations of C. viridis observed inthis study exhibited a synchronized semi-lunarreproductive cycle. A pattern of spawning syn-chrony, frequently coupled with a lunar component,has been documented for several other Chromisspecies (Table I). A review of these reports indicates

that reproductive cycling according to lunar cuesappears to be widespread, but is not universal (e.g.C. notata,Table I). Moreover, the temporal nature oflunar-associated spawning not only varies amongspecies, but also occasionally among conspecificpopulations at different locales (e.g. C. hypsilepis,Table I). These observations, as well as similar onesfor non-Chromis pomacentrids (e.g Robertson et al.1990) suggest that for many pomacentrid species,the fine-scale coordination between reproductionand particular phase(s) of the lunar cycle has astrong site-dependent component.Based on available reports provided in Table I, at

least some form of spawning synchrony may beuniversal among Chromis species. Such synchro-nous bursts of spawning tied to lunar maxima andminima may be an adaptation to a schooling modeof life, as suggested by Robertson et al. (1990) andTzioumis & Kingsford (1995). For shoaling zoo-planktivores, obligatory substrate association canonly be transient, and cues used for initiating andcoordinating spawning activity likely need to bestrong and unambiguous. The absence of tempo-rally matching synchrony between the two sitesdescribed here, which are relatively closely located(i.e. both in Madang Lagoon and less than 4 kmapart) suggests that while lunar cues act to syn-chronize activities within the observed C. viridispopulations, other more localized cues appear toinfluence the exact timing of the initiation of thereproductive sequence. For C. viridis, these local-ized cues may involve social cues (Robertson et al.1990), trophic cues (Tyler & Stanton 1995), orsome as-yet unidentified cues that can vary amongpatch reefs yet play an important role in the fine-scale timing of lunar-associated spawning for manysynchronously spawning fish species. In MadangLagoon, Wongad Island typically receives strongerwave action due to its’ unprotected central locationin the lagoon (Kohn 2001). This environmentalcondition may promote rapid aggregation forma-tion and accelerated initiation of spawning,although it’s unclear what advantage advancedspawning under these conditions might provide.

ACKNOWLEDGEMENTSI thank the former Christensen Research Institute

for in-kind support, L. J. Orsak for logistical help, J.Masey for diving assistance and F. Pezold, J. E. Ran-dall, J. S. Stimson and two anonymous reviewers forhelpful comments on the manuscript. Portions ofthis study were supported by a National Science and


Kathleen S. Cole

Engineering Research Council operating grant(Canada), NASA Fundamental Space Biology Pro-gram NAG 2-1600, and additional support moneyfrom the University of Hawaii at M~noa. I’m par-ticularly grateful to J. E. Randall for numerousenlightening discussions regarding Chromis and forproviding closeup images of C. viridis.

REFERENCESALBRECHT, H. 1969. Behaviour of four species of Atlanticdamselfishes from Columbia, South America (Abudefdufsaxatiles, A. taurus, Chromis multilineata, C. cyanea; Pisces,Pomacentridae). Zeitschrift für Tierpsychologie 26: 662-676.

ALLEN, G. R. 1991. Damselfishes of the World. Mergus Pub-lishers Hans A. Baensch, Melle, Germany, 271 pp.

AMORIM, M. C. P. 1996. Sound production in the blue-green damselfish, Chromis viridis (Cuvier, 1830) (Poma-centridae). Bioacoustics 6: 265–272.

ASOH, K. 2003. Reproductive parameters of female Hawai-ian damselfish Dascyllus albisella with comparison to othertropical and subtropical damselfishes. Marine Biology 143:803–810.

COLE, K. S. & SADOVY, Y. 1995. Evaluating the use ofspawning success to estimate reproductive success in aCaribbean reef fish. Journal of Fish Biology 47: 181–191.

COUGHLIN, D. J. & STRICKLER, J. R. 1990. Zooplanktoncapture by a coral reef fish: an adaptive response to evasiveprey. Environmental Biology of Fishes 29: 35-42.

DE BOER, B. A. 1978. Factors influencing the distribution ofthe damselfish Chromis cyanea (Poey), Pomacentridae, on areef at Curaçao, Netherlands, Antilles. Bulletin of MarineScience 28: 550-565.

FUJII, R., KASUKAWA, H., MAYAJI, K. & OSHIMA, N. 1989.Mechanisms of skin coloration and its changes in the blue-green damselfish, Chromis viridis. Zoological Science 6:477-486.

GLADSTONE, W. 2007. Temporal patterns of spawning andhatching in a spawning aggregation of the temperate reeffish Chromis hypsilepis (Pomacentridae). Marine Biology151: 1143-1152.

GLADSTONE, W. & WESTOBY, M. 1988. Growth and repro-duction in Canthigaster valentini (Pisces, Tetraodontidae):a comparison of a toxic reef fish with other reef fishes.Environmental Biology of Fishes 21: 207-221.

JEBB, M. H. P. & LOWRY, J. K. 1995. Natural history ofMadang Lagoon with an appendix of collecting localities.Records of the Australian Museum, Supplements 22: 1-24

KOHN, A. 2001. Maximal species richness in Conus: diver-sity, diet and habitat on reefs of northeast Papua NewGuinea. Coral Reefs 20: 25-38.

LANGER, M. R. & LIPPS, J. H. 2003. Foraminiferal distrib-ution and diversity, Madang Reef and Lagoon, Papua NewGuinea. Coral Reefs 22: 143-154.

LECCHINI, D., SHIMA, J., BANAIGS, B. & GALZIN, R. 2005.Larval sensory abilities and mechanisms of habitat selec-tion of a coral reef fish during settlement. Oecologia 143:326-334.

LECCHINI, D., PLANES, S. & GALZIN, R. 2006. The influ-ence of habitat characteristics and conspecifics on attrac-tion and survival of coral reef fish. Journal of ExperimentalMarine Biology and Ecology 341: 85-90.

MYERS, R. F. 1999. Micronesian Reef Fishes (Third edition).Coral Graphics, Barrigada, Guam, 3308 pp.

MYRBERG, A. A., JR., BRAHY, B. D. & EMERY, A. R. 1967.Field observations on reproduction of the damselfish,Chromis multilineata (Pomacentridae), with additionalnotes on general behavior. Copeia 1967: 819-827.

NAKAZONO, A., TAKEYA, H. &TSUKAHARA, H. 1979. Stud-ies on the spawning behavior of Chromis notata (Tem-minck et Schlegel). Science Bulletin of the Faculty of Agri-culture, Kyushu University 34: 29–37.

OCHI, H. 1986. Breeding synchrony and spawning intervalsin the temperate damselfish Chromis notata. Environmen-tal Biology of Fishes 17: 117-123.

ÖHMAN, M. C., MUNDAY, P. L., JONES, G. P. & CALEY, M.J. 1998. Settlement strategies and distribution patterns ofcoral-reef fishes. Journal of Experimental Marine Biologyand Ecology 225: 219-238.

PANDOLFI, J. M. & MINCHIN, P. R. 1995. A comparison oftaxonomic composition and diversity between reef corallife and death assemblages in Madang Lagoon, Papua NewGuinea. Palaeogeography, Palaeoclimatology, Palaeoecology119: 321-341.

ROBERTSON, D. R., PETERSEN, C.W. & BRAWN, J. D. 1990.Lunar reproductive cycles of benthic-brooding reef fishes:reflections of larval biology or adult biology? EcologicalMonographs 60: 311-329.

RUSSELL, B. C. 1971. Underwater observations on thereproductive activity of the demoiselle Chromis dispilus(Pisces; Pomacentridae). Marine Biology 10: 22–29.

SALE, P. F. 1971. The reproductive behaviour of the poma-centrid fish, Chromis caeruleus. Zeitschrift für Tier-psychologie 29: 156-64.

SMITH, M. F. L. & WARBURTON, K. 1992. Predator shoal-ing moderates the confusion effect in blue-green chromis,Chromis viridis. Behavioral Ecology and Sociobiology 30:103-107.

SWEATMAN, H. P. A. 1985. The influence of adults of somecoral reef fishes on larval recruitment Ecolological Mono-graphs 55: 469-485.

SWERDLOFF, S. N. 1970. Behavioral observations on Eniwe-tok damselfishes (Pomacentridae: Chromis) with specialreference to the spawning of Chromis caeruleus. Copeia1970: 371-375.

THRESHER. R. E. 1984. Reproduction in Reef Fishes. TFHPublishing, Neptune City, New Jersey, 399 pp.

TYLER, W. A., III & STANTON, F. G. 1995. Potential influ-ence of food abundance on spawning patterns in a damsel-fish, Abudefduf abdominalis. Bulletin of Marine Science 57:610-623.

TZIOUMIS, V. & KINGSFORD, M. J. 1995. Periodicity ofspawning of two temperate damselfishes: Parma micro-lepis and Chromis dispilus. Bulletin of Marine Science 57:596-609.



AbstractVanderhorstia steelei is described as a new species of shrimp-

goby from six specimens collected on silty sand from 5 mdepth in Cook’s Bay, Moorea, Society Islands. The species isdistinguished by 15 dorsal soft rays, 16 anal soft rays, 17 or18 pectoral rays, 55-58 scales in longitudinal series, a slen-der body (depth 6.4-7.4 in standard length), lanceolate andterminally bilobed caudal fin, and a color pattern featuringnarrow dark bars on the body (from dark brown scale edges)that contain scales with bright yellow centers.

ZusammenfassungBeschrieben wird Vanderhorstia steelei als neue Art auf der

Grundlage von sechs Exemplaren, die in einer Tiefe ab fünfMetern in der Cook-Bucht, Moorea, an den Gesellschafts-Inseln über schlammigem Sand gesammelt wurden. Unter-scheiden lässt sich die Art durch 15 weiche Rückenflossen-strahlen, 16 weiche Afterflossenstrahlen, 17 oder 18 Brust-flossenstrahlen, 55 bis 58 Schuppen in Längsreihen, einenschlanken Körper (Tiefe 6,4 bis 7,4 in Standardlänge), einelanzettförmige, am Ende zweilappige Schwanzflosse sowiedurch die Farbgebung: schmale dunkle Streifen auf demRumpf (durch dunkelbraune Schuppenränder bewirkt),wobei die Schuppen jeweils in der Mitte leuchtend gelb sind.

RésuméVanderhorstia steelei est décrit en tant que nouvelle espèce

de gobie crevette sur base de six spécimens collectés sur dusable envasé à 5 m de profondeur, dans la Baie de Cook,Moorea, Îles de la Société. L’espèce se distingue par 15rayons mous à la dorsale, 16 rayons mous à l‘anale, 17 ou 18rayons pectoraux, 55-58 écailles en séries longitudinales, uncorps élancé (6,4-7,4 de hauteur par rapport à la longueurstandard), une caudale lancéolée et bilobée à l’extrémité, etun patron de coloration fait d’étroites barres sombres sur lecorps (de bords d’écailles brun foncé) qui contiennent desécailles à centre jaune clair.

SommarioVanderhorstia steelei è descritta come nuova specie di go-

bide simbionte di gamberetti sulla base di sei esemplari rac-colti su fondali sabbiosi a 5 m di profondità presso la baia di

Cook, a Moorea, Isole della Società. La specie si distingueper avere 15 raggi molli dorsali, 16 raggi molli anali, 17 o 18raggi pettorali, 55-58 scaglie in serie longitudinale, un corpolongilineo (altezza 6.4-7.4 in lunghezza standard), la pinnacaudale lanceolata e bilobata distalmente e una colorazioneche mostra sottili barre scure sul corpo (derivanti dal mar-gine marrone scuro delle scaglie) che racchiudono scagliedall’area centrale giallo brillante.

INTRODUCTIONThe Indo-Pacific gobiid fish genus Vanderhorstia

was proposed by Smith (1949) for Gobius delagoaeBarnard, 1937 from Mozambique. Smith (1959),provided a brief diagnosis for the genus and added asecond species, V. ornatissima Smith, 1959, alsofrom Mozambique. Five other species of gobiesdescribed before 1959 are now recognized in Van-derhorstia: V. atriclypeus (Garman, 1903) from Fiji;V. longimanus (Weber, 1909) from Indonesia; V.macropteryx (Franz, 1910) from Japan (redescribedby Ikeda et al., 1995); V. auropunctata (Tomiyama,1955) from Japan; and V. ambanoro (Fourmanoir,1957) from Madagascar. Five more new species weredescribed in the period 1974-1995, four of theminitially classified in the genus Vanderhorstia: V.mertensi Klausewitz, 1974 from the Red Sea; V.lanceolata Yanagisawa, 1978 from Japan; V. puncti-ceps (Deng & Xiong in Xu et al., 1980) from China;V. praealta Lachner & McKinney, 1981 from theSeychelles; and V. flavilineata Allen & Munday,1995 from New Britain, Papua New Guinea. Therehave been a surprising 12 descriptions of new speciesof Vanderhorstia between 2004 and the present,therefore the same number in the last four years asin the previous 100 years: V. papilio Shibukawa &Suzuki, 2004 from Japan; V. nannai Winterbottom,Iwata & Kozawa, 2005; V. bella Greenfield & Lon-genecker, 2005 from Fiji; V. nobilis Allen & Randall,2006 from the Philippines; V. opercularis Randall,



Vanderhorstia steelei, a new shrimpgoby from the Society Islands

John E. Randall1 and Philip L. Munday2

1) Bishop Museum 1525 Bernice St., Honolulu, HI 96817-2704, USA.E-mail: [email protected]

2) ARC Centre of Excellence for Coral Reef Studies and School of Marine and Tropical Biology, JamesCook University, Townsville 4811, Australia

Received: 22 September 2007 – Accepted: 04 October 2007

2007a from the Red Sea; V. hiramatsui, V. kizakura,and V. rapa Iwata, Shibukawa & Ohnishi, 2007from Japan; V. attenuata Randall, 2007b from theSolomon Islands; V. auronotata Randall, 2007bfrom Indonesia; V. belloides Randall, 2007b fromPapua New Guinea, and V. dorsomacula Randall,2007b from New Britain. Here we describe thetwenty-fifth species from six specimens collected inMoorea, Society Islands.

MATERIALS AND METHODSThe type specimens of the new species are

deposited in the AustralianMuseum, Sydney (AMS);the Natural History Museum, London (BMNH);Bernice P. Bishop Museum, Honolulu (BPBM);National Museum of Nature and Science, Tokyo(NSMT); and the United States National Museumof Natural History, Washington, D.C. (USNM).Methods of counting and measuring specimens fol-

low Randall (2007b). Morphometric data presentedin Table 1 are given as percentages of the standardlength (SL). Data in parentheses in the descriptionapply to paratypes. Proportional measurements inthe text are rounded to the nearest 0.05.

Vanderhorstia steelei n. sp.(Figs 1-4; Table I)

Vanderhorstia ornatissima (non Smith, 1959). Bac-chet et al. 2006: 440-441, identification on p. 459(lagoon of Pirai, Tahiti).

Holotype: BPBM 40665, male, 51.9 mm SL, Soci-ety Islands, Moorea, Cook’s Bay, Gump ResearchStation, boat mooring, 17°29’30”S 149°49’33”W,silty sand, 5 m, clove oil, P. L. Munday, 19 July 2007.

Paratypes: AMS I.44300-001, male, 31.0 mm SL;BMNH 2007.8.21.1, female, 33.9 mm SL; BPBM40666, female, 40.7 mm SL; NSMT-P 77287,male, 42.2 mm SL; USNM 391935, female,45.0 mm SL, all with same data as holotype.Diagnosis:Dorsal rays VI-I,15; anal rays I,15 or 16

(rarely 15); pectoral rays 17 or 18; scales in longitu-dinal series 55-58; no scales on head, or predorsal orprepectoral areas; body elongate, the depth 6.4-7.25in SL; orbit diameter 3.55-4.0 in head length;fourth dorsal spine longest, 3.85-4.3 in SL; caudalfin lanceolate and terminally bilobed, the sixth andtenth branched rays longest, 2.3-2.5 in SL; pectoralfins reaching to above or slightly beyond anus, 4.25-4.3 in SL; pelvic fins reaching beyond anus, 4.15-4.35 in SL; color in alcohol pale yellowish, the edgesof scales brown; upper side of body with narrowdark bars from darker brown scale edges; a series ofdark spots dorsally on body from dark brown scaleedges; cheek and opercle light gray-brown with palespots, some nearly as large as pupil; a blackish streakin groove above upper lip; median fins spotted; lifecolor as in Figs 2 and 3.Description: Dorsal rays VI-I,15; anal rays I,16

(one paratype with 15); all dorsal and anal soft raysbranched, the last to base; pectoral rays 17 (twoparatypes with 17, and three with 18), the upper-most unbranched; pelvic rays I,5, all soft raysbranched, the fifth rays joined medially; segmentedcaudal-fin rays 17, 14 branched; upper and lowerunsegmented caudal rays ?; longitudinal scale series56 (55-58), progressively smaller anteriorly; trans-verse scale rows 19; no median predorsal scales;median prepelvic scales 9; circumpeduncular scales12; gill rakers 7 + 13 (4-7 + 13-14); pseudobranchwith 7 (7-8) short fleshy lobes; branchiostegal rays 5.Body elongate, the depth at origin of pelvic fins



Fig. 1. Holotype of Vanderhorstia steelei, BPBM 40665, 59.2 mm, Moorea, Society Islands. Photo by J. E. Randall.


John E. Randall and Philip L. Munday

Fig. 2. Cephalic sensory pores and papillae of holotype of Vanderhorstia steelei. Drawing by A. Field.

Table I. Proportional measurements of type specimens of Vanderhorstia steelei as percentages of the standard length.

Holotype Paratypes

BPBM AMS BMNH BPBM NSMT- USNM40665 I.44300 07.8.21.1 40666 P 77287 391935

Standard length (mm) 51.2 31.0 33.9 40.7 42.2 45.0Sex male male female female male femaleBody depth (at P2 origin) 14.3 13.8 15.0 15.5 14.3 15.6Body depth (at A origin) 13.0 12.9 13.3 13.5 13.4 13.3Body width 9.6 9.5 8.8 10.1 9.4 11.0Head length 25.2 25.7 25.1 25.3 25.0 25.1Head width 10.5 11.1 10.9 10.2 9.7 9.9Snout length 5.5 5.1 5.6 5.2 5.2 5.4Orbit diameter 6.3 6.9 6.8 7.1 6.9 7.1Interorbital width 1.3 1.6 1.5 1.5 1.7 1.4Cheek depth 1.6 1.3 1.6 1.6 1.5 1.3Upper-jaw length 10.7 10.5 10.8 10.7 10.8 10.9Caudal-peduncle depth 7.8 7.6 7.4 7.8 7.6 7.8Caudal-peduncle length 12.9 12.9 12.8 11.8 13.0 11.8Predorsal length 28.4 29.6 30.0 29.3 28.3 30.1Preanal length 50.8 50.0 52.7 51.2 49.7 52.4Prepelvic length 28.5 29.1 29.7 28.8 28.0 29.9Base of dorsal fins 59.2 60.6 59.3 60.5 59.7 60.5First dorsal spine 22.4 20.1 17.7 19.5 24.0 18.0Longest dorsal spine broken 24.8 23.4 25.8 25.9 24.5Fifth dorsal spine 19.3 15.8 12.6 16.4 21.9 16.3Spine of second dorsal fin 8.9 aberrant 9.5 9.6 9.1 8.8Longest dorsal ray 15.6 14.5 15.0 14.8 16.1 15.6Base of anal fin 38.4 36.8 36.3 38.6 38.2 37.6Anal spine 6.7 7.5 5.8 7.1 5.0 6.8Longest anal ray 15.5 15.4 15.1 14.7 16.8 15.5Caudal-fin length 43.6 40.4 43.6 42.7 42.4 43.2Pectoral-fin length 23.2 23.5 23.3 23.3 23.5 23.2Pelvic-spine length 7.9 8.6 8.7 7.6 7.2 8.5Pelvic-fin length 23.8 22.9 23.0 23.2 24.1 24.0



Fig. 3. Underwater photograph of Vanderhorstia steelei, Moorea. Photo by M. H. Steele.

Fig. 4. Underwater photograph of Vanderhorstia steelei with Alpheus rapacida, Tahiti. Photo by P. Bacchet.

7.0 (6.4-7.25) in SL, at origin of anal fin 7.7 (7.4-7.75) in SL; body compressed, the width at pectoral-fin origin 1.35 (1.2-1.5) in body depth; head length3.95 (3.9-4.0) in SL; head compressed, the width1.35 (1.2-1.5) in body depth; snout length 4.6 (4.5-5.05) in head length; orbit diameter 4.0 (3.55-3.75)in head length; interorbital space narrow, the leastwidth 19.4 (15.7-19.8) in head length; caudal-peduncle depth 3.25 (3.2-3.4) in head length; cau-dal-peduncle length 1.95 (1.9-2.15) in head length.Mouth oblique, forming an angle of about 30° to

horizontal axis of body, the lower jaw projecting;mouth large, the maxilla reaching or nearly reachinga vertical at posterior edge of orbit, the upper-jawlength 2.35 (2.3-2.45) in head length; front ofupper jaw with three small, incurved, nearly recum-bent, canine teeth on each side, the lateral pairlargest, the inner pair convergent; side of jaw poste-rior to canines with a row of more than 30 tinyincurved teeth, no larger than a medial row of papil-lae; front of lower jaw with two rows of smallincurved teeth, ending about one-fourth distanceback in jaw by a series of four or five recurved canineteeth, one or two as large as largest canine at front ofupper jaw, followed by a single row of moderate,slightly incurved, conical teeth; no teeth on vomer

or palatines; roof of mouth with prominent well-spaced papillae; edge of lips smooth, the inner sur-face papillose; tongue tip rounded, the upper surfaceand roof of mouth with scattered papillae; no dis-tinct mental flap.Gill opening broad, extending forward to a vertical

at posterior edge of orbit; gill membranes attachedonly anteriorly to isthmus, with no free fold; gill rak-ers slender, the longest at angle about half length oflongest gill filaments.Posterior nostril a moderately large oval aperture

just before fleshy edge of orbit a little above center ofeye, with a slight anterior rim; anterior nostril ashort membranous tube anteroventral to posteriornostril just above upper lip, the internarial distanceabout equal to greatest diameter of posterior nostril;cephalic sensory pores and papillae as in Fig. 2;prominent nasal pore directly anterodorsal to poste-rior nostril; not shown, two medial pores in interor-bital space, one anterior and one posterior.Scales on body progressively smaller anteriorly;

scales weakly ctenoid posteriorly, becoming cycloidanterior to about base of fifth dorsal soft ray, andpartially embedded anteriorly; no scales on head,nape anterior to a line from dorsal-fin origin toupper free end of preopercle, or prepectoral area; no



Fig. 5. Underwater photograph of Vanderhorstia sp. B, Palau. Photo by R. F. Myers.

scales on fins except for three rows of moderatelylarge scales on caudal-fin base.Origin of first dorsal fin over origin of pelvic fins,

the predorsal length 3.5 (3.3-3.5) in SL; dorsalspines slender and flexible; first dorsal spine 4.95(4.15-5.65) in SL; third or fourth dorsal spineslongest (both damaged in holotype), 3.35-4.3 in SLin paratypes; fifth dorsal spine 5.2 (4.6-7.95) in SL;spine of second dorsal fin 2.85 (2.75-2.85) in headlength; thirteenth dorsal soft ray longest, 6.4 (6.2-6.9) in SL; origin of anal fin slightly posterior to avertical at origin of second dorsal fin, the preanallength 1.95 (1.9-2.0) in SL; anal spine 3.75 (3.4-5.0) in head length; fourteenth anal soft ray longest,6.45 (5.95-6.8) in SL; caudal fin lanceolate and ter-minally bilobed, the sixth and tenth rays longest, 2.3(2.3-2.4) in SL; pectoral fins pointed, the tenth andeleventh rays longest, reaching slightly posterior toanus, 4.3 (4.25-4.3) in SL; prepelvic length 3.5(3.35-3.6) in SL; pelvic spine 3.2 (2.9-3.5) in headlength; fifth pelvic ray longest, nearly reaching orextending slightly posterior to origin of anal fin, 4.2(4.15-4.35) in SL; pelvic frenum extending to tips ofpelvic spines, its posterior margin slightly concave.Color of holotype in alcohol pale yellowish, the

edges of scales brown; upper side of body with ninedark bars, from darker brown scale edges, the last atbase of caudal fin broadest; a faint dark bar betweeneach pair of dark bars; a series of 14 dark spots dor-sally on body from dark brown scale edges, the firstbefore origin of dorsal fin, and the last small on edgeof caudal peduncle; cheek and opercle light gray-brown with round to elongate pale spots, somenearly as large as pupil; a blackish streak in grooveabove upper lip; front of snout and upper lip gray-brown; dorsal fins light brown with pale spots, thoseon second dorsal in six rows; anal fin similar butwith irregular rows of pale dashes on membranes;caudal fin with pale rays and gray-brown mem-branes, the dorsal third with a row of pale spots alongeach membrane, the membranes of rest of fin withnarrow longitudinal pale streaks; paired fins withpale rays and light gray-brown membranes, the pec-torals with a large gray-brown spot at base containingtwo vertical rows of pale spots.Life color as in Figs 3 and 4 (these fish not col-

lected).Etymology: This species is named in honor of

Mark H. Steele of California State University,Northridge, California who took the photographof Fig. 3 and alerted us to the location of thespecies in Moorea.

Remarks: The type specimens were evidently pre-served in formalin that was acidic, because a vertebralcount could not be made from an x-ray. However, itwould be expected to be 10 +16, as reported for thegenus; also the dorsal pterygiophore formula is prob-ably 3-22110 (Fig. 1A of Birdsong et al. 1988).Of the described species of Vanderhorstia, V. steelei is

most similar to V. auronotata from the MoluccaIslands, sharing with it a similar color pattern and thesame longitudinal scale and pectoral-ray counts. Itdiffers in having one more dorsal and anal rays thanV. auronotata, a more slender body (depth 6.4-7.25,compared to 5.5 in SL for V. auronotata), and in den-tition, especially in having much larger teeth.Myers (1999: 242, pl. 155, fig. C) illustrated a

species of shrimpgoby by an underwater photographfrom Palau as Vanderhorstia sp. B (reproduced here asFig. 5). It is very similar in color, at least anteriorly,differing in the smaller yellow spots on the head andthe yellow scale centers within each dark bar on thebody forming a more distinct vertical yellow line.Specimens are needed to determine if it is the sameor a very similar species.We also note the similarity of the underwater pho-

tograph of Vanderhorstia sp. 2 of Okamura &Amaoka (1997: 691, middle-right fig.) from Iri-omote-jima, Japan.

ACKNOWLEDGEMENTSWe thank foremost Mark H. Steele and Philippe

Bacchet for their underwater photographs of the newspecies of Vanderhorstia, Robert F. Myers for his pho-tograph of Vanderhorstia sp. B from Palau, and Ash-ley Field for his drawing of Fig. 2. We are also grate-ful to Arthur Anker for the identification of thealpheid shrimp shown in Fig. 4, and KatherineMun-day, Melissa Schmitt, and Andrew Brooks for assis-tance in collecting the type specimens. Gerald R.Allen, David W. Greenfield, and Helen A. Randallreviewed the manuscript.

REFERENCESALLEN, G. R. & MUNDAY, P. 1995. Description of four newgobies (Gobiidae) from the western Pacific Ocean. Revuefrançaise d’Aquariologie 22 (3-4): 99-104.

ALLEN, G. R. & RANDALL, J. E. 2006. Vanderhorstia nobilis,a new species of shrimpgoby from Indonesia and thePhilippines. aqua, Journal of Ichthyology and Aquatic Biology12 (1): 39-44.

BACCHET, P., ZYSMAN, T. & LEFÈVRE, Y. 2006. Guide desPoissons de Tahiti et ses Îles. 607 pp. Au Vent des Iles Edi-tions, Tahiti.



BARNARD, K. H. 1937. Further notes on South Africanmarine fishes. Annals of the South African Museum 32, pt. 2(6): 41-67.

BIRDSONG, R. S., MURDY, E. O. & PEZOLD, F. L. 1988. Astudy of the vertebral column and median fin osteology ingobioid fishes with comments on gobioid relationships.Bulletin of Marine Science 42 (2): 174-214.

FOURMANOIR, P. 1957. Poissons Téléostéens des eaux mala-gaches du Canal deMozambique. Mémoires de l’Institut Sci-enfique de Madagascar, sér. F, 1: 1-316.

FRANZ, V. 1910. Die Japanischen Knochenfische der Samm-lungen Haberer und Doflein (Beiträge zur NaturgeschichteOstasiens.). Abhandlungen der Akademie der Wissenschaften4 (suppl.) (1): 1-13.

GARMAN, S. 1903. Some fishes from Australasia. Bulletin ofthe Museum of Comparative Zoölogy at Harvard College. 39(8): 229-241.

GREENFIELD, D. W. & LONGENECKER, K. R. 2005. Vander-horstia bella, a new goby from Fiji (Teleostei: Gobiidae).Proceedings of the California Academy of Sciences 56: 619-623.

IKEDA, Y., NAKABO, T. & HIRAMATSU,W. 1995. Redescrip-tion of Vanderhorstia macropteryx (Perciformes: Gobiidae)with designation of a neotype. Japanese Journal of Ichthyol-ogy 42 (3/4): 303-310.

IWATA, A., SHIBUKAWA, K. & OHNISHI, N. 2007. Three newspecies of the shrimp-associated goby genus Vanderhorstia(Perciformes: Gobiidae: Gobiinae) from Japan, with re-des-criptions of two related congeners. Bulletin of the NationalMuseum of Natural Science (Tokyo), ser. A, suppl. 1: 185-205.

KLAUSEWITZ, W. 1974. Eilatia latruncularia n. gen. n. sp.und Vanderhorstia mertensi n. sp. vom Golf von Aqaba(Pisces: Gobiidae: Gobiinae). Senckenbergiana biologica 55:205-212.

LACHNER, E. A. & MCKINNEY, J. F. 1981. A new fish speciesof the genus Vanderhorstia (Teleostei: Gobiidae) from theAmirante Islands, Indian Ocean. Proceedings of the Biologi-cal Society of Washington 93 (4): 963-970.

MYERS, R. F. 1999. Micronesian Reef Fishes, ed. 3. vi + 330pp. Coral Graphics, Guam.

OKAMURA, O. & AMAOKA, K. 1997. Fishes of Japan. 783 pp.Yama-kei Publishers, Tokyo (in Japanese).

RANDALL, J. E. 2007a. Vanderhorstia opercularis, a newshrimpgoby from the northern Red Sea. Electronic Journalof Ichthyology 3 (1): 18-25.

RANDALL, J. E. 2007b. Descriptions of four new shrimpgob-ies of the genus Vanderhorstia from the western Pacific.aqua, International Journal of Ichthyology 12 (3): 89-100.

SHIBUKAWA, K. & SUZUKI, T. 2004. Vanderhorstia papilio, anew shrimp-associated goby from the Ryukyu Islands, Japan(Perciformes: Gobiidae: Gobiinae), with comments on thelimits of the genus. Ichthyological Research 51 (2): 113-119.

SMITH, J. L. B. 1949. Forty-two fishes new to South Africa,with notes on others. Annals and Magazine of Natural His-tory, ser. 12, 2: 97-111.

SMITH, J. L. B. 1959. Gobioid fishes of the families Gob-iidae, Periophthalmidae, Trypauchenidae, Taenioididaeand Kraemeriidae of the western Indian Ocean. Ichthy-ological Bulletin 13: 185-225.

TOMIYAMA, I. 1955. Notes on some fishes, including onenew genus and three new species from Japan, theRyukyus and Pescadores. Japanese Journal of Ichthyology 4(nos. 1/2/3): 1-15.

WEBER, M. 1909. Diagnosen neuer Fische der Siboga-Expedition. Notes from the Leyden Museum 31 (4): 143-169.

WINTERBOTTOM, R., IWATA, A. & KOZAWA, T. 2005. Van-derhorstia nannai, a new species of burrow-associatedgoby from Palau and the Philippines (Pisces: Gobiidae).aqua, Journal of Ichthyology and Aquatic Biology 9 (3):109-114.

XU, C.-Y., DENG, S., XIONG, G. & ZHAN, H. 1980. Twonew fishes from East China Sea. Oceanology and Limno-logy Sinica 11 (2): 179-184 (in Chinese, English summ.).

YANAGISAWA, Y. 1978. Studies on the interspecific rela-tionship between gobiid fishes and snapping shrimp. I.Gobiid fishes associated with snapping shrimps in Japan.Publications of the Seto Marine Biological Laboratory 24(4/6): 269-325.



ERRATAaqua International Journal of Ichthyology Vol. 13 (1) 25 October 2007Opistognathus rufilineatus, a new species of jawfish (Opistognathidae) from the Bird’s Head Peninsula, western NewGuinea by W. F. Smith-Vaniz and G. R. Allenon page 37: Indonesia, Irian Jaya (western New Guinea), Barat Providence, North Triton Bay, 3°47’01”S, 134°9’55”E

The day I first met George W. Barlow was in the 1980swhen we were both invited to give a talk for the Pacific CoastCichlid Association at San Francisco’s Steinhard Aquarium.Until late, we talked about many issues, such as the growingdestruction of freshwater habitats, marine environments(“cichlid people always invite me, but most aquarists don’tknow that I am primarily a marine biologist” he told me),and Central America. Both of us collected and studied fishesthere. When I asked “why Midas”, George, who had startedworking in the area in the 1960s told me “…once upon atime, if stories are true, there lived in ancient Greek KingMidas, whose touch turned everything into gold, but when-ever I see a goldfish I wonder ifKing Midas was not a Chi-nese…”. He called Cichlasomacitrinellum “Midas” because of arare brightly coloured morph,which he studied in LakeNicaragua, Managua and somecrater lakes in Nicaragua. Hiscontribution on cichlids pub-lished in the Investigation of theIchthyofauna of NicaraguanLakes in 1976 is still the stan-dard work on those species.Last November, when I was

back inNicaragua together withAxel Meyer from Konstanz,Germany, G. Barlow’s formerPh.D. student at Berkeley, Irecalled our conversation thatevening. George had told me, that the Midas cichlid repre-sented the largest biomass in the Nicaraguan lakes, whichtoday is no longer the case. I found that now almost 80% ofthe cichlids in Lakes Nicaragua and Managua are exotics,even though as early as 1955 George Myers warned “…thatany introduction of Tilapia mossambica would most likelyhave a disastrous effect upon the native fauna…”.On our second meeting at the American Cichlid Associa-

tion (ACA) on the occasion of their International CichlidCongress in Orlando in 1989, where we both were againinvited, I remember talking about Cichlasoma. George andother leading ichthyologists wanted to see this genus retained,at least in part, and George continued using Cichlasoma in hislast book, The Cichlid Fishes Nature’s Grand Experiment ofEvolution (Barlow 2000). Robert R. (Bob) Miller and Georgehelped me tremendously with my book on the freshwaterfishes of the world, for which I am extremely grateful. Their

views will be reflected in that book. We talked most of thetime about cichlids, their behaviour, the fascinating biologyof theMidas cichlid and why it is found in many of the craterlakes – unlike C. labiatum. We met on many other occasionsand, without a single exception, I always listened to this wise,extremely knowledgeable and humble man. Our lastencounter was in New Jersey in 2001. George proudly pre-sented me with his new book dedicated “To Heiko – a realcichlidot! Alles Gute, George”. I will keep this copy foreverand read it over and over again. George always talked withmein German, which he was quite good at. In the process ofpursuing his PhD studies in ichthyology he was increasingly

drawn to ethology, the study ofanimal behaviour. This culmi-nated in a two year postdocto-ral fellowship (1958-1960),which he spend in the labora-tory of Konrad Lorenz in Bava-ria. He also spent one year inBielefeld, Germany (1977-1978), participating in a re-search programme on the de-velopment of behaviour.George was always open tohobbyists around the world,but his main interest was in be-havioural ecology, ethology, be-havioural mechanisms, evolu-tion and speciation in fishes,reptiles and amphibians and inmany other fields. After his re-

tirement he became Professor Emeritus at the Department ofIntegrative Biology University of California at Berkeley in1993, where he pursued these interests. There is an excellentWebsite by George Barlow(http://ib.berkeley.edu/labs/barlow/cv.htlm), where itbecomes obvious what an incredible person he was. In Feb-ruary 2002, George joined the Editorial Board of aqua, serv-ing for five years as an active advisor. In March 2006 hedecided to resign from the Board because of his failing health.George was convinced that cichlids, because of their “explo-sive” radiation into multiple species, have become celebritiesin the realm of evolutionary biology. With George we all losta unique celebrity, who cannot be replaced. George was ableto tell us more about his beloved fishes than anyone else. Asmentioned in his last book, which everyone should read,cichlids spoke to George and I believe him as fishes also talkto me. Farewell George, I will continue to talk to you as long


George W. Barlow (center) with fish-lovers at ICC 1989

Distinguished ichthyologists and fish collectors left us in 2007

George W. Barlow 1929-2007

For scientists and particularly for fish-lovers around the world 2007 was a sad year. Three outstanding personalities passed away.Jacques Géry’s obituary was published in aqua 12(4). One month later we lost another leading personality in science, George W.Barlow, and before end of the year the greatest collector and scholar of Malawi cichlids, Stuart Grant, suddenly passed away. Theyare remembered for their outstanding fieldwork, ichthylogical and other natural history research, their enthusiasm and their for-midable contribution to the inventory of species on our planet.

A man, who has done more for Lake Malawi cichlids thananyone else, has suddenly passed away. He was a man wholoved fishes and Malawi, the country were he has spent mostof his life. He helped biologists, the press and in particularaquarists from around the world to learn about and under-stand the fishes of his beloved lake, particularly the cichlids.He cared about nature, the environment and all natural crea-tures, while he condemned genetic alterations resulting inman’s own creatures, or rather monsters.I had the privilege to know Stuart since the 1970s, shortly

after he started exporting fishes from Lake Malawi. We firstmet at the International Pet Fair “Interzoo“ in 1976 or 1978.I vividly remember this polite, gentle and very correct Eng-lish Gentlemen, who spoke per-fect Oxford English.“You mind if I call you Heiko”

was one of his first words “youcan call me Stuart”, and “may Isit down”. “You are a major fishcollector, importer and exporterand I heard you are wellrespected in the ornamental fishcommunity; can you tell me whoare the good guys in Ger-many…?”. Our meetings be-came a regular event, every twoyears at the Interzoo. In the early1980s he sent me fishes I hadbeen after for research, zoos andpublic aquaria around the world.Hundreds of mormyrids, realgiants, some almost one meterlong, which live in Lake Malawi.No one ever collected thembefore. I shall never forget onebeautiful spring day in 1990, sit-ting with him in a bistro in Paris,on the corner of Rue Cuvier,looking over 1200 Lake Malawi cichlids and identifyingthem (Stuart helped me tremendously cataloguing thespecies and variants – no one knew them better). Sometimeswe talked in German, in French, but mostly in English. Healso spoke perfect Chichewa and the northern Malawi nativelanguage, Chitumbuka.Stuart always emphasised that his fishermen are very well

trained. Fishing in Malawi for ornamentals never involvesanaesthetics, electricity or explosives, and only a very smallportion of the rocky habitats are fished.The last time I visited his operation in Salima, Kambriri

Point was in January 2003. I drove into this amazing set upof more than 360 cemented vats, each with a capacity of1000 litres (or more) and buildings with aquariums, holdingtanks and offices. Further on were several houses, his man-sion, a guest and a dining house and there was a long sandybeach. Stuart, at the time had over 70 people working forhim. I never forget his incredible hospitality, waiters in white

dresses serving us the most delicious breakfast, overlookingthis fantastic lake, framed by all kinds of exotic plants. Notonly had Stuart chosen a beautiful spot, he also created hisown paradise. Hence it is not surprising that he had guests allthe time, taking advantage of this unique place.Stuart was born on 20 June 1937 in Sattford, England and

already as a young boy developed an interest in tropicalfishes, but kept coldwater fishes he collected in nearbystreams and lakes because of a lack of money. When joiningthe Midland Aquarist Society he would not yet imagebecoming the prime fish collector and exporter in what hecalled “The heart of Africa”. In 1958 he completed his ser-vice in the Royal Air Force in Cyprus and applied for a post

in Nyasaland, were he ended upworking 14 years as an accoun-tant and personnel officer. Hisinterest in fishes continued andeventually he started collecting,keeping a few iron-framed tankswith Lake Malawi fishes. AfterNyasaland became the Republicof Malawi in the late 1960s, heobtained a permit and exportedcyprinodontids (Nothobranchiussp.) to the American KillifishAssociation. In 1973 he wasoffered a fishing license and anofficial export permit and wasmore than happy to be able tostay in the country he loved, pur-suing an activity he had alwaysdreamed of. After Peter andHenny Davies, who began atMonkey Bay in 1963/64, Stuartbecame the second exporter ofMalawi fishes. His business grewcontinuously, until the morningof 11 October 2007 when a

stroke suddenly took him away from us.Stuart has hosted famous ichthyologists, like A. J. Rib-

bink from South Africa; authors, like Hans J. Mayland andAd Konings; countless biologists and aquarists. He wasalways a gentlemen, one of those you cannot find anylonger these days.David Livingstone was the first European to explore Lake

Malawi, but more than anyone else Stuart Grant helpedexplore the aquatic environment. Stuart promoted ourunderstanding of this incredible and unique evolutionaryhotspot. This lake, which was his live, hosts between 700 and1200 species of cichlids, more than all remaining members ofthe family Cichlidae combined. To evolutionary biologistsLake Malawi is the quintessential example of speciation gonewild and Stuart was always there to help them. Who will fol-low in his footsteps? Personally, I feel, that with Stuart Grantone of the most precious, still undescribed species, hasbecome extinct. Heiko Bleher


Stuart Malcolm Mariot Grant 1937-2007

Stuart Grant & author 2003, Kambiri Point, Malawi


Index of aqua Vol. 13 (1-4)(Index by: 1. Author(s); 2. New Taxa; 3. Biology/Ecology/Biography/Reviews)

Author(s):Allen, Gerald R.and Mark Erdmann: Two new species of bamboo sharks (Orectolobiformes: Hemiscylliidae) from Western New Guinea. aqua 13 (3-4):91-106, 23 January 2008.Allen, Gerald R.and Peter J. Unmack: A new species of rainbowfish (Melanotaenia: Melanotaeniidae), from Batanta Island, western New Guinea. aqua 13 (3-4): 107-118, 23 January 2008.Allen, Gerald R. and Mark V. Erdmann: Corythoichthys benedetto, a new pipefish (Pisces: Syngnathidae) from Indonesia and Papua New Guinea. aqua 13 (3-4): 119-124, 23 January 2008.Allen, Gerald R. and Mark V. Erdmann: Pterois andover, a new species of scorpionfish (Pisces: Scorpaenidae) from Indonesia and Papua New Guinea. aqua 13 (3-4): 125-136, 23 January 2008.Allen, Gerald R. and Mark V. Erdmann: Pseudanthias charleneae, a new basslet (Serranidae: Anthiinae) from Indonesia. aqua 13 (3-4): 137-142, 23 January 2008.Allen, Gerald R., Gill, Anthony C. and Mark V. Erdmann:A new species of Pictichromis (Pisces: Pseudochromidae) from western New Guinea with a redescription of P. aurifrons. aqua 13 (3-4): 143-152, 23 January 2008.Allen, Gerald R., Gill, Anthony C. and Mark V. Erdmann: A new species of Pseudochromis (Pisces: Pseudochromidae) from Irian Jaya Barat Province, Indonesia. aqua 13 (3-4): 153-160, 23January 2008.Allen, Gerald R. andMark V. Erdmann: Pterocaesio monikae, a new species of fusilier (Caesionidae) from western New Guinea (Papua and Papua Barat provinces, Indonesia). aqua 13 (3-4): 161-168, 23 January 2008.Allen, Gerald R. and Mark V. Erdmann: A new species of damselfish (Chrysiptera: Pomacentridae)from western New Guinea and the Togean Islands, Indonesia. aqua 13 (3-4): 169-176, 23January 2008.Allen, Gerald R. and Mark V. Erdmann: Paracheilinus nursalim, a new species of flasher wrasse (Perciformes: Labridae) from the Bird’s Head Peninsula of western New Guinea with a key to thespecies of Paracheilinus. aqua 13 (3-4): 177-186, 23 January 2008.Casaux, Ricardo Jorge and Cecilia Yanina Di Prinzio: The diet of the large puyén Galaxias platei (Galaxiidae) at Rosario Lake, Patagonia, Argentina. aqua 13 (2): 77-86, 8 November 2007.Costa, Wilson J. E. M.: Kryptolebias gracilis n. sp. (Teleostei: Cyprinodontiformes: Rivulidae): a new killifish from the Saquarema Lagoon basin, southeastern Brazil. aqua 13 (1): 7-12, October2007.Costa, Wilson J. E. M.: Five new species of the aplocheiloid killifish genus Rivulus, subgenus Melanorivulus, from the middle Araguaia River basin, central Brazil (Teleostei: Cyprinodontiformes:Rivulidae). aqua 13 (2): 55-68, 8 November 2007.Erdmann, Mark: Blue Auction “Special Edition” – Preface. aqua 13 (3-4): 89, 23 January 2008.Gomon, Martin F.: A new genus and miniature species of pipehorse (Syngnathidae) from Indonesia. aqua 13 (1): 25-30, October 2007.Lima, Flávio C. T., Britski, Heraldo A. and Francisco A. Machado: A new Moenkhausia (Characiformes: Characidae) from central Brazil, with comments on the area relationship between theupper rio Tapajós and upper rio Paraguai systems. aqua 13 (2): 45-54, 8 November 2007.Randall, John E. andKuiter, Rudie H.:Wetmorella tanakai, a new wrasse (Perciformes: Labridae) from Indonesia and the Philippines. aqua 13 (1): 1-6, 25 October 2007.Smith-Vaniz, William F. and Gerald R. Allen: Opistognathus rufilineatus, a new species of jawfish (Opistognathidae) from the Bird’s Head Peninsula, western New Guinea. aqua 13 (1): 35-42,October 2007.Winterbottom, Richard and Margaret Zur: Three new species of genus Trimma from Palau, Western Pacific (Percomorpha: Gobiidae). aqua 13 (1): 13-24, October 2007.Winterbottom, Richard and Laura Southcott:Two new species of the genus Trimma (Percomorpha: Gobiidae) from western Thailand. aqua 13 (2): 69-76, 8 November 2007.Wirtz, Peter: The return of the yellow grouper – annual migration and return to the same site by a xanthistic Mycteroperca fusca (Pisces: Serranidae). aqua 13 (1): 31-34, October 2007.

New Taxa:Chrysiptera giti n. sp. a new species of damselfish (Chrysiptera: Pomacentridae)from western New Guinea and the Togean Islands, Indonesia. aqua 13 (3-4): 169-176, 23 January 2008.Corythoichthys benedetto n. sp. a new pipefish (Pisces: Syngnathidae) from Indonesia and Papua New Guinea. aqua 13 (3-4): 119-124, 23 January 2008.Hemiscyllium galei n. sp. Two new species of bamboo sharks (Orectolobiformes: Hemiscylliidae) from Western New Guinea. aqua 13 (3-4):91-106, 23 January 2008.Hemiscyllium henryi n. sp. Two new species of bamboo sharks (Orectolobiformes: Hemiscylliidae) from Western New Guinea. aqua 13 (3-4):91-106, 23 January 2008.Kyonemichthys n. gen. A new genus and miniature species of pipehorse (Syngnathidae) from Indonesia. aqua 13 (1): 25-30, October 2007.Kyonemichthys rumengani n. gen. A new genus and miniature species of pipehorse (Syngnathidae) from Indonesia. aqua 13 (1): 25-30, October 2007.Kryptolebias gracilis n. sp. (Teleostei: Cyprinodontiformes: Rivulidae): a new killifish from the Saquarema Lagoon basin, southeastern Brazil. aqua 13 (1): 7-12, October 2007.Melanotaenia synergos n. sp. A new species of rainbowfish (Melanotaenia: Melanotaeniidae), from Batanta Island, western New Guinea. aqua 13 (3-4): 107-118, 23 January 2008.Moenkhausia cosmops n. sp. A new Moenkhausia (Characiformes: Characidae) from central Brazil, with comments on the area relationship between the upper rio Tapajós and upper rio Paraguaisystems. aqua 13 (2): 45-54, 8 November 2007.Opistognathus rufilineatus n. sp. a new species of jawfish (Opistognathidae) from the Bird’s Head Peninsula, western New Guinea. aqua 13 (1): 35-42, October 2007.Paracheilinus nursalim n. sp. a new species of flasher wrasse (Perciformes: Labridae) from the Bird’s Head Peninsula of western New Guinea with a key to the species of Paracheilinus. aqua 13(3-4): 177-186, 23 January 2008.Pictichromis caitlinae n. sp. A new species of Pictichromis (Pisces: Pseudochromidae) from western New Guinea with a redescription of P. aurifrons. aqua 13 (3-4): 143-152, 23 January 2008.Pseudanthias charleneae n. sp. a new basslet (Serranidae: Anthiinae) from Indonesia. aqua 13 (3-4): 137-142, 23 January 2008.Pterois andover n. sp. a new species of scorpionfish (Pisces: Scor-paenidae) from Indonesia and Papua New Guinea. aqua 13 (3-4): 125-136, 23 January 2008.Rivulus crixas n. sp. Five new species of the aplocheiloid killifish genus Rivulus, subgenus Melano-rivulus, from the middle Araguaia River basin, central Brazil (Teleostei: Cyprinodontiformes: Rivulidae). aqua 13 (2): 55-68, 8 November 2007.Pseudochromis jace n. sp. a new species of Pseudochromis (Pisces: Pseudochromidae) from Irian Jaya Barat Province, Indonesia. aqua 13 (3-4): 153-160, 23 January 2008.Pterocaesio monikae n. sp. a new species of fusilier (Caesionidae) from western New Guinea (Papua and Papua Barat provinces, Indonesia). aqua 13 (3-4): 161-168, 23 January 2008.Rivulus javahe n. sp. Five new species of the aplocheiloid killifish genus Rivulus, subgenus Melanorivulus, from the middle Araguaia River basin, central Brazil (Teleostei: Cyprinodontiformes:Rivulidae). aqua 13 (2): 55-68, 8 November 2007.Rivulus karaja n. sp. Five new species of the aplocheiloid killifish genus Rivulus, subgenus Melanorivulus, from the middle Araguaia River basin, central Brazil (Teleostei: Cyprinodontiformes:Rivulidae). aqua 13 (2): 55-68, 8 November 2007.Rivulus rubromarginatus n. sp. Five new species of the aplocheiloid killifish genus Rivulus, subgenus Melanorivulus, from the middle Araguaia River basin, central Brazil (Teleostei: Cyprinodon-tiformes: Rivulidae). aqua 13 (2): 55-68, 8 November 2007.Rivulus salmonicaudus n. sp. Five new species of the aplocheiloid killifish genus Rivulus, subgenus Melanorivulus, from the middle Araguaia River basin, central Brazil (Teleostei: Cyprinodon-tiformes: Rivulidae). aqua 13 (2): 55-68, 8 November 2007.Trimma fucatum n. sp. Two new species of the genus Trimma (Percomorpha: Gobiidae) from western Thailand. aqua 13 (2): 69-76, 8 November 2007.Trimma gigantum n. sp. Three new species of genus Trimma from Palau, Western Pacific (Percomorpha: Gobiidae). aqua 13 (1): 13-24, October 2007.Trimma randalli n. sp. Three new species of genus Trimma from Palau, Western Pacific (Percomorpha: Gobiidae). aqua 13 (1): 13-24, October 2007.Trimma sanguinellus n. sp. Two new species of the genus Trimma (Percomorpha: Gobiidae) from western Thailand. aqua 13 (2): 69-76, 8 November 2007.Trimma tauroculum n. sp. Three new species of genus Trimma from Palau, Western Pacific (Percomorpha: Gobiidae). aqua 13 (1): 13-24, October 2007.Wetmorella tanakai n. sp. a new wrasse (Perciformes: Labridae) from Indonesia and the Philippines. aqua 13 (1): 1-6, 25 October 2007.

Biology/Ecology/Biography/Reviews:A new species of Pictichromis (Pisces: Pseudochromidae) from western New Guinea with a redescription of P. aurifrons. aqua 13 (3-4): 143-152, 23 January 2008.The diet of the large puyén Galaxias platei (Galaxiidae) at Rosario Lake, Patagonia, Argentina. aqua 13 (2): 77-86, 8 November 2007.The return of the yellow grouper – annual migration and return to the same site by a xanthisticMycteroperca fusca (Pisces: Serranidae). aqua 13 (1): 31-34, October 2007.Book review: Reef ad shore fishes of the Hawaiian Island. aqua 13 (2): 87, 8 November 2007.Book review: Fishes of the World. aqua 13 (2): 88, 8 November 2007.

BLABER, S. J. M. 1980. Fish of the Trnity inlet systemof North Queensland, with notes on the ecologyof fish faunas of tropical Indo-Pacific estuaries.Australian Journal of Marine and FreshwaterResearch 31:137-46.

DAY, J. H., BLABER, S. J. M., & WALLACE, J. H. 1981.Estuarine fishes. In: Estuarine Ecology with Parti-cular Reference to Southern Africa. (Ed. J.H. Day.):197-221. A. A. Balkema, Rotterdam.

DIMMICH, W. W. 1988. Ultrastructure of North Ame-rican cyprinid maxillary barbels. Copeia 1988 (1):72-79.

TREWAVAS, E. 1983. Tilapiine Fishes of the GeneraSarotherodon, Oreochromis and Danakilia.British Museum (Natural History), London, 583 pp.

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aquaInternational Journal of Ichthyology

Vol. 14 (1), 28 January 2008

Contents:

N. G. Bogutskaya, A. M. Naseka, P. A. Tikhonov: A brief history of the study of fishes of theCaspian Sea and scientific results of the Caspian Expedition of 1904 headed by N. M. Knipovich ................ 1-26

Kathleen S. Cole: Observations on spawning behavior and periodicity in the Bluegreen Chromis(Pomacentridae: Chromis viridis), in Madang Lagoon, Papua New Guinea ..................................................... 27-34

John E. Randall and Philip L. Munday:Vanderhorstia steelei, a new shrimpgoby from theSociety Islands ................................................................................................................................................ 35-41

Obituaries – Distinguished ichthyologist and collecors who died in 2007.George W. Barlow 1929-2007 and Stuart Malcolm Mariot Grant 1937-2007................................................. 42-43

Index of aqua Vol. 13 ......................................................................................................................................................... 44

Papers appearing in this journal are indexed in: Zoological Record;Biolis – Biologische Literatur Information Senckenberg;

www.aqua-aquapress.com; www.aquapress-bleher.com; www.aquageo.com; www.Joachim-Frische.com

Cover photo: Underwater photograph of Vanderhorstia steelei with Alpheus rapacida, Tahiti. Photo by P. Bacchet.

Pseudocoris n. sp., initial-phase male, 90 mm SL, Holmes Reef, Coral Sea. Detailed descriptioncoming up in aqua 14 (2) soon. Aquarium photo by F. Walsh.

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