theoldestdomesticatedfishes,andtheconsequencesof ...2)_ciprinus.pdf ·...

KeywordsDomestication, origin, common carp, goldfish,guppy, neon tetra, exploited captives, alternatives infish culture, altricial and precocial

AbstractIt is a myth that the common carp was originallydomesticated in China, maintained throughunfounded repetition but with no evidence. The coloraberrations called nishikigoi appeared en masse inthe 1950s from the Niigata Prefecture in Japan.These ornamental common carp called koi becamethe most expensive fish, with some prize-winning indi-viduals valued at more than one million dollars. Thecriteria for domestication or exploited captives areexplained, with special attention to fishes. No attemptis made to resolve the taxonomy of the common carpor goldfish, but some nomenclature data are includedto assist future systematic revisions. About 2000years ago, wild common carp were most abundant inthe inland delta of the Danube River at the northernedge of the important Roman province Pannonia. TheRomans kept fish in specially built piscinae. Commoncarp were the hardiest fish available, therefore themost capable of surviving the primitive methods oftransport of that time. Keeping carp in ponds becamemore popular in medieval times, when fasting wasenforced and fasting food was in demand. The cultureof common carp and the building of special pondsgradually became the most profitable branch of agri-culture in central Europe. Some unintentional artificialselection had already taken place between the twelfthand the mid-fourteenth century. Deep bodied and var-iously scaled or scaleless domesticated formsappeared in nearly every pond system. A domestica-tion of a fish that did occur in China was of the muchsmaller cyprinid called chi, a fish captured there forfood since early times. This silver-grey chi, betterknown as goldfish, Carassius auratus, occasionallyappeared as a xanthic form. These red goldfish havebeen documented since the beginning of the SungDynasty in 960 AD. Releasing such rare xanthic formsinto Buddhist ponds of mercy was considered a betterdeed than releasing an ordinary chi. In the 1200s the

fish had become tame and were used as ornamentalanimals in the garden pools of rich landowners. Fromthe late 13th to the mid 15th century, the Chinesebegan keeping the golden chi in aquarium-like ves-sels, and soon rich and poor alike became breedersof fancy domesticated goldfish. The variously shapedmonstrosities and color aberrants were freaks, butthey became very fashionable at that time and stillare. More recently other species, such as the guppy,Poecilia reticulata, and the neon tetra, Paracheirodoninnesi, became domesticated in the aquarium hobby.Many other fishes kept as ornamentals such asswordtails and platies, discus and angelfishes, as wellas those cultured for food such as rainbow trout,channel catfish and sturgeon are merely exploitedcaptives. The occurrence of alternative phenotypes insome species is explained. These altricial (less spe-cialized) or precocial (more specialized) alternativesare caused by differences in the provision of endoge-nous food (e.g., yolk) in early development, amongother factors, and are a response to changing envi-ronments. Life history – from indirect, with an exter-nally food gathering larva, to direct without larvae – isconstantly adjusted to fit the fluctuating environmentor system created by humans. Recognized in time,these adjustments can have important implications forculture systems.

ZusammenfassungEs ist ein Irrtum, daß der Karpfen ursprünglich in Chinadomestiziert wurde. Die Farbabweichungen mit demNamen Nishikigoi tauchten in den 1950er Jahrenmassenhaft aus der japanischen Präfektur Niigata auf.Diese Zierkarpfen mit der Bezeichnung Koi sollten zuden teuersten Fischen der Welt werden; einige preis-gekrönte Exemplare erzielten mehr als eine Million Dol-lar. Die Kriterien der Domestikation oder Nutztierhaltungwerden im Hinblick auf die Fische erläutert. Vor rund2000 Jahren kam der wildlebende Gemeine Karpfen imBinnendelta des Flusses Danubius – der Donau –, amNordrand der wichtigen römischen Provinz Pannonia,sehr häufig vor. Die Römer hielten die Fische in speziellangefertigten piscinae. Der Gemeine Karpfen wardamals der widerstandfähigste Fisch, den man kannte,

aqua, Journal of Ichthyology and Aquatic Biology

The oldest domesticated fishes, and the consequences ofan epigenetic dichotomy in fish culture

Eugene K. Balon

Department of Integrative Biology, University of Guelph, Guelph, Ontario N1G 2W1, Canada.E-mail: [email protected]

Accepted: 30.01.2006

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und er war deshalb am besten geeignet, die primitivenTransportmethoden jener Zeit zu überstehen. Die Hal-tung von Karpfen in Teichen wurde erst im Mittelalterüblich, als das Fasten durchgesetzt worden war und derBedarf an Fastenspeisen anstieg. Die Zucht desGemeinen Karpfens und der Bau der Teiche entwickel-ten sich zum profitabelsten Zweig der Landwirtschaft inMitteleuropa. Ein gewisses Maß an unbeabsichtigterkünstlicher Auslese hatte zwischen dem zwölftenJahrhundert und der Mitte des vierzehnten bereitsstattgefunden. In fast jedem Teichsystem tauchten bere-its Zuchtformen mit dickem Körper, mit variablemSchuppenmuster und ganz ohne Schuppen auf. InChina hatte man eine Zuchtform aus einem vielkleineren Cypriniden namens chi entwickelt, der dortvon altersher zu Nahrungszwecken gefangen wurde.Dieser silbergraue chi, besser als Goldfisch bekannt,Carassius auratus, trat ab und zu in einer gelblichenFarbvariante auf. Diese sogenannten roten Goldfischesind bereits seit Beginn der Sung-Dynastie um 960 A.D.belegt. Im 13. Jahrhundert waren die Fische bereitszahm geworden und wurden als Zierfische in den Gar-tenteichen reicher Landbesitzer gehalten. Vom Endedes 13. bis zur Mitte des 15. Jahrhunderts hielten dieChinesen den goldenen Chi in aquarienähnlichenGefäßen, und bald züchteten arme wie reiche Leutemodische Zierformen des Goldfischs. Die verschieden-sten exzentrischen Körperformen und Farbabweichun-gen fanden damals ihre Liebhaber, teilweise bis heute.In neuerer Zeit wurden in der Hobby-Aquaristik andereArten Gegenstand der Domestikation, wie der Guppy,Poecilia reticulata, und der Neonsalmler (-tetra), Para-cheirodon innesi. Viele andere Fische werden nur auswirtschaftlichen Gründen, wie Nutztiere also, gehalten,sei es als Zierfische wie Schwertträger, Platys, Discus,Kaiserfische, sei es zu Nahrungszwecken wie Regen-bogenforelle, Wels und Stör. Das Auftreten abweichen-der Phänotypen bei einigen Arten wird erklärt. Die altri-cialen (weniger spezialisierten) oder precocialen (mehrspezialisierten) Abweichungen werden durch Unter-schiede in der Versorgung mit endogener Nahrung (z.B.Eidotter) und andere Faktoren in der frühen Entwicklungverursacht und sind eine Reaktion auf Umweltverän-derungen. Die Entwicklung von Lebewesen, ihre Natur-geschichte – die indirekte mit Larven mit äußererNahrungsaufnahme bis hin zur direkten ohne Larven –wird ständig nachkorrigiert, um zu der sich wandelndenUmwelt oder Systemen, die von Menschen geschaffensind, zu passen. Wenn man diesen Wandel rechtzeitigerkennt, kann man wichtige Schlussfolgerungen darausfür Aufzucht Systeme ziehen.

RésuméQue la carpe commune ait été, à l'origine, domes-tiquée en Chine est un mythe. Les aberrations chroma-tiques, appelées nishikigoi, sont apparues en massedans les années 1950, originaires de la préfecturejaponaise Niigata. Ces carpes communes ornemen-

tales, nommées koi, sont devenues les poissons lesplus coûteux, certains exemplaires vainqueurs de con-cours étant estimés à plus d'un million de dollars. Onexpose ici les critères de domestication ou d'exploitationd'espèces captives, avec une attention spéciale con-sacrée aux poissons. Il y a 2000 ans environ, les carpescommunes sauvages abondaient surtout dans le deltaintérieur du Danube, à la limite nord de la grandeprovince romaine de Pannonie. Les Romains mainte-naient les poissons dans des piscinae construites àcette fin. Les carpes communes étaient les poissonsdisponibles les plus résistants et donc à même de sur-vivre aux méthodes de transport primitives de cetteépoque. Gerder des carpes dans des étangs est devenupopulaire au moyen âge, quand le carême était bienobservé et qu'il y avait demande de nourriture maigre.L'élevage de la carpe commune et l'aménagement d'é-tangs ad hoc ont constiué petit à petit le secteur le plusflorissant de l'agriculture en Europe centrale. Une cer-taine sélection artificielle non intentionnelle a déjà eu lieuentre le 12e et la moitié de 14e siècle. Des formesdomestiquées au corps plus haut et à écailles variablesou sans écailles sont apparues dans presque chaquecomplexe d'étangs. La domestication d'un poisson quivivait effectivement en Chine a concerné le cyprinidébien plus petit, appelé chi, une espèce qu'on y pêchaitpour se nourrir depuis des temps reculés. Ce chi grisargenté, mieux connu sous l'appellation de poissonrouge, Carassius auratus, présentait à l'occasion uneforme xanthique. Ces poissons rouges figurent dansdes documents qui datent du début de la dynastie Sung,en 960. Introduire de telles formes xanthiques, rares,dans des étangs bouddhistes d'action de grâce étaitconsidéré comme plus bénéfique que d'y lâcher un chiordinaire. Dans les années 1200, les poissons ont étéapprivoisés et utilisés comme ornement dans les étangsdes jardins de riches propriétaires terriens. Depuis la findu 13e jusqu'à la moitié du 15e siècle, les Chinois ontcommencé à maintenir les chi dorés dans des récipientsen guise d'aquariums et bientôt, riches et pauvres sontdevenus éleveurs de poissons rouges de fantaisiedomestiqués. Les formes monstrueuses très variéesétaient accidentelles, mais elles ont été très demandéesà cette époque et encore maintenant. Plus récemment,d'autres espèces, comme le guppy, Poecilia reticulata,et le tétra néon, Paracheirodon innesi, ont été domes-tiquées pour l'aquariophilie. Beaucoup d'autres pois-sons d'ornement, les porte-épées et les platys, les Dis-cus et les scalaires, aussi bien que ceux qu'on élèvepour l'alimentation comme la truite arc-en-ciel, le pois-son-chat et l'esturgeon, ne sont en fait que des espècescaptives exploitées. On explique l'occurrence de phéno-types alternatifs pour certaines espèces. Cesphénomènes de type altriciel (moins spécialisé) ou pré-cociel (plus spécialisé) sont provoqués par une alimen-tation endogène (p. ex., du vitellus) à un stade préco-coe, parmi d'autres facteurs, et constituent une réactionà des changements environnementaux. L'histoire de la

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Oldest domesticated fishes and epigenetic dichotomy

vie – indirectement, avec un alevin collectant une nour-riture externe, ou directement, sans alevins - s'adapteconstamment aux fluctuations de l'environnement ouaux systèmes créés par l'homme. Reconnues à temps,ces adaptations peuvent avoir des implications impor-tantes sur les systèmes d'élevage.

SommarioSecondo la leggenda i primi allevamenti dellacomune carpa furono intrapresi in Cina. L’aberrazionecromatica nota come nishikigoi apparve improvvisa-mente negli anni ‘50 nella Prefettura di Niigata Pre-fecture in Giappone. Queste varianti ornamentali,chiamate koi, sono oggi molto costose ed alcuniesemplari sono stati valutati anche a prezzi superiorial milione di dollari. In questo articolo sono descritti icriteri su cui si basa l’allevamento di questi pesci, conparticolare attenzione alla specie. Circa 2000 anni fala carpa comune era molto abbondante nel delta delDanubio a nord dell’importante provincia romana del-la Pannonia. I romani, infatti, mantenevano questi pe-sci in particolari vasche (piscinae). La carpa era il pe-sce più resistente, in grado di sopravvivere ai primi-tivi metodi di trasporto di quei tempi. Il mantenimentodella carpa in laghetti artificiali divenne popolaredurante il Medio Evo, quando le carestie erano fre-quenti. L’allevamento della carpa con la costruzionedi apposite vasche divenne gradualmente una delleattività più redditizie dell’Europa centrale. Alcunevarietà erano già selezionate non intenzionalmentetra il XII e la metà del XIV secolo. Forme domestichedal corpo elevato e senza scaglie apparvero un po’ovunque in questi stagni artificiali. Anche la domesti-cazione di un’altra specie di ciprinide, più piccolo,noto come chi e oggetto di pesca da secoli, cominciòin Cina. Il chi è in genere di colore grigio-argento, maspesso si presenta con una vivace colorazione rossaed è quindi più conosciuto come pesce rosso. Il suonome scientifico è Carassius auratus. Queste variantirosse sono note sin dalla dinastia Sung del 960. Laloro immissione nei laghetti sacri buddisti era consi-derato di miglior auspicio del rilascio del comune chi.Nel 1200 il pesce entrò nella sfera domestica dive-nendo un animale ornamentale negli stagni dei giar-dini di ricchi possidenti. Dal XIII al XV secolo, i cinesicominciarono a mantenere il pesce rosso in vaschesimili ai moderni acquari e ben presto sia ricchi chepoveri furono in grado di mantenere allevamenti divarietà di pesci rossi. Le varie mostruosità e aberra-zioni cromatiche divennero fenomeni da baraccone,ma al tempo stesso ricercati come oggetti di moda elo sono tuttora. Recentemente altre specie, come ilguppy, Poecilia reticulata, e il pesce neon, Parachei-rodon innesi, sono state addomesticate per gli amantidell’acquario. Molte altre specie sono allevate sia afini ornamentali, come ad esempio lo swordtail e ilplaty, i discus e i pesci angelo, sia a scopi alimentaricome le trote, i pesci gatto e gli storioni. In questo arti-

colo si dà spiegazione anche della comparsa di feno-tipi alternativi. Queste varianti altricial (meno specia-lizzate) o precocial (più specializzate) sono causate,soprattutto, da differenze nelle riserve endogene dicibo (ad esempio, il tuorlo) nelle prime fasi di sviluppoe rappresentano una risposta all’ambiente mutevole.La storia naturale – variabile da una forma indiretta,con una larva attiva nel procacciarsi il cibo ad unadiretta senza stadio larvale – è costantemente aggiu-stata per adattarsi alle fluttuanti condizioni ambientalio anche in seguito all’intervento dell’uomo. Ricono-sciuti da tempo, questi adattamenti possono avereimportanti implicazioni per l’allevamento.

Introduction

Adomestic animal can be defined as one that has beenbred in captivity for purposes of economic profit to ahuman community that maintains total control over itsbreeding, organization of territory, and food supply.

Juliet Clutton-Brock (1999: 32)in A Natural History of Domesticated Mammals

Domestication by humans, particularly of fishesThe definition of domestication by Clutton-Brockshould apply also to fishes subject to some amend-ments (e.g., Diamond, 2002). First, allow me to repeatsome of my earlier musing on that subject (Balon,1995b: 5): “The questions of where and how organ-isms were first domesticated have occupied manyscholars. Definitive answers are rarely produced(Zeuner 1963, Isaac 1970) because most of thesedomestications started already in the Neolithic(roughly 14,000 years ago). ‘In the study of domesti-cation the scientist and the cultural historian joinforces, each playing a role which the other discipline,by its very nature, cannot fill’ (Isaac 1962). (...) In atruly domesticated organism, (a) the individual is val-ued and kept for a specific purpose, (b) its breeding issubject to human control, (c) its behaviour is differentfrom that of the wild ancestor, and (d) its morphology(including size, coloration and physiology) exhibitsvariations never seen in the wild, (e) some of whichwould not survive without human protection.”The best known examples of domestication are thetransformations of wolves into breeds of dogs, theaurochs into cattle, the guanaco into llama and alpaca,the wild boar into domestic swine, Przewalski’s wildhorses into domestic horses, and the red jungle fowlinto domestic chickens. Dependence is the most impor-tant among the more obvious criteria. “All domesticatedanimals depend for their day-to-day survival upon theirowners. The capacities of wild self-sufficiency havinglong since been subtracted from them, they mustdepend upon whatever prosthetic devices their ownerssee fit to provide” (Livingston, 1994: 14).However, the matter of dependence also applies toanimals at the first stage of domestication which Clut-

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ton-Brock (1999) calls exploited captives and otherscall tamed or cultured. Most recently wapiti deer havebecome such captives on farms neighbouring our resi-dence in Canada. Many attempts at domesticationremain only at that level. Most fish cultured as food andas ornamentals also remain merely exploited captives,especially in cases where wild individuals are occa-sionally brought into the stock bred in captivity. To be atrue domesticate, therefore, the earlier mentioned crite-ria (d) and (e) must apply, i.e., shape, size and col-oration exhibit variations never or rarely seen in the wildancestor. Some of these variants would not survivewithout human protection in captivity. Furthermore, theoriginal criterion of Clutton-Brock (1999), that of eco-nomic profit, may only partially apply to domesticatesas pets. Fishes that are introduced by humans intowaters other than their native waters and becomeestablished there are termed naturalized by Lever(1996). We are not interested in these in this treatise.Fishes became objects of domestication much laterthan the domestication of many other animals andplants, now fixed by Diamond (2002) at around10,500 years ago at the earliest. Even adding theirtime as exploited captives, in a strict sense only twofish species qualify: the wild common carp, Cyprinuscarpio Linnaeus, 1758, acquired by humans about2000 years ago, and the wild chi better known asgoldfish, Carassius auratus (Linnaeus, 1758) selectedand released into ponds of mercy about 1000 yearsago; the former by the Romans in south centralEurope (Balon, 1974, 1995a, b), and the latter inChina (Chen, 1956; Wang et al., 2000). Only recentlysome other fish species may qualify, although confu-sion reigns because different criteria are used. Atbest, statements like “only the common carp, the cru-cian carp (Carassius carassius) and the goldfish(Carassius auratus) have been truly domesticatedwith the management of all stages in the life cycle,particularly reproduction for a longtime” (Billard, 1999:4) are justified, but the crucian carp does not belonghere (see later). This study is about domestication.The taxonomy of the common carp and goldfish is notattempted but significant data related to nomenclatureof these species, so far ignored by systematicists, areincluded. A putative and brief synthesis of these datais designed as a guideline for the future solution of thecomplex problem.The two closely related cyprinids, the wild commoncarp and the wild goldfish, began to be changed intotrue domesticated animals in early medieval times,though in different parts of the world and for differentpurposes (Chen, 1956; Balon, 1969, 1974, 1995b). Theother, very few modern domesticated fishes are ratherpoorly defined and of questionable status (Zeuner,1963). In contrast to the extinct aurochs, ancestors ofdomestic cattle or the nearly extinct in nature Przewal-ski horse, only the wild common carp may be close toextirpation at its area used for initial domestication,

replaced in the wild by the feral progeny of domesti-cated forms (Fig. 1).The loss of the ancestral wild forms of any domesti-cated organism means a tragic loss of epigeneticpotential. Clutton-Brock (1999: 30) believes “that ani-mals bred under domestication evolve into newspecies as a result of reproductive isolation from theirwild progenitors...” (my italics). Domesticatesreleased into the wild to fend for themselves usuallyperform poorly and, while in time some revert theirshape to resemble their wild ancestors, the process isnearly always incomplete. All such feral forms remainpoor facsimiles of their progenitors (Livingston, 1994).Following the above criteria, I will classify fishes keptby humans first as exploited captives that are littlechanged from the wild ancestral forms, and can usuallybe returned to the wild. Nevertheless, exploited cap-tives are often the first step in domestication with pro-found changes in behaviour (Price, 1999). Animalsbecome domesticates when they change in form, func-tion, color and behavior from their wild ancestors, andif returned or accidentally released into the wild withouthuman protection they survive poorly as feral forms,often only partially resembling their ancestors.Several more domesticates have been produced inthe aquarium industry for ornamental (e.g., guppy,swordtails, platies, mollies, neon tetras, discus,angelfishes) or research (medaka, zebrafish) pur-poses. These are few and mostly of very recent originor of questionable status. Fishes reared in captivity forhuman food like the channel catfish, salmon, the Chi-nese and Indian pelagic spawning carp, tilapia or stur-geon, to mention just a few, are exploited captives.Artificial propagation of rainbow trout, Oncorhynchusmykiss, for food or stocking has led to the creation ofhatchery forms that Behnke (2002: 103) considersdomesticates. In spite of color changes and “a loss ofthe ability (...) to compete with non-game fishes for acommon food supply” these trout are merely exploitedcaptives. However, the few that are undergoing pur-poseful artificial selection as color aberrants will qual-ify as domesticates (Goryczko and Dobosz, 2004).The epigenetic processes that act in domesticationalso modify the life histories of the fishes, allowingsurvival of both altricial and precocial forms (Balon,1990, 2004a,b). The existence of alternative formsand their life-history modifications are explained in thefinal section, including some aquaculture implications.

Domestication of the common carp did notoriginate in China!The origin of common carp domestication is still con-fusing authors as evidenced in the recent work by Liu(2003), in spite of much more precise data given byBanarescu, Grzimek and Ladiges in the first edition ofthe same encyclopedia. The reason for the frequentclaims that common carp domesticates came toEurope from China is that common carp and goldfish

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were not only misidentified in the past but also occur inplaces that are far apart, and so it is “one of thosethings which somebody said a long time ago, having noreal or precise knowledge of the subject, and that it hasbeen slavishly copied ever since” (Burton, 1969, insidecover of issue 48), from one book to another (e.g.,Leonhardt, 1906; Tamura, 1961; Hickling, 1962;Rudzi˜ski, 1961; Steffens, 1967, 1980; Borgese, 1980;Liu, 2003). Bloch (1782: 93) clearly stated: “Das Vater-land des Karpfen ist ohnstreitig in den südlicherenTheilen Europens zu suchen”, and so earlier authors(e.g. Day, 1865 or Taylor, 1884: 7) were correct in stat-ing “it is all but certain that the carp was brought fromsouthern Europe to the more northerly parts; its greatsize and esteemed flavour rendering it a favourite.”Details of this argument concerning the commoncarp have already been given by Balon (1974: 21):“The latest Chinese study (the following quotationsare from the Russian translation, Anonymous, 1961)on pond culture stated that ‘thanks to the creativeefforts of the Chinese people for many generations,breeding of the carp in this country has proceededsuccessfully for more than 2,000 years. From Chinathe breeding of this fish spread all over the world. (...)From Asia the rearing of the carp spread to Europeand later to America, Australia, and Africa’ [my trans-lation from Russian].”Let us finally put this myth to rest (Balon, 1995b: 8-9): “ ‘Kwai Sin Chak Shik, a book written during the

Sung Dynasty in A.D. 1243, describes how carp frywere transported in bamboo baskets in much thesame way as they are transported and traded today.The fry were collected in rivers and reared in ponds;this is recorded in A Complete Book of Agriculture,written in A.D. 1639.’ But if the fish ‘fry were collectedin rivers’ it is doubtful whether they were the commoncarp, Cyprinus carpio, which is not a riverine pelagicspawner. It is more likely that they were the young ofother carps, such as the grass carp Ctenopharyn-godon idellus (Valenciennes, 1844), the silver carpHypophthalmichthys molitrix (Valenciennes, 1844),the bighead carp Aristichthys nobilis (Richardson,1845), and the black carp Mylopharyngodon piceus(Richardson, 1846), of which, some are true riverinepelagic spawners and cannot be bred in pond or ricepaddy conditions.” Furthermore, no domesticatedforms of common carp, like the mirror or leather, oreven the high-bodied forms (Pokornø et al., 1995),were ever reported from China prior to known intro-ductions from Europe (Wohlfarth, 1986). Kottelat(1997: 57) writes that “Balon (1974, 1995) discussedthe origin of domesticated carps and concluded thatthe European ones originated from a rheophilic stockfrom the Danube. I agree with him and Hoffmann(1995a) that it is unlikely to have been imported fromChina, as is often assumed” (see also Zeuner, 1963).And, therefore, in addition to the above “It may benoted [concluded Wohlfarth, 1984: 377] that in most

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Fig. 1. (a) Drawing of the wild common carp from the spawning school on the Danube near Medved’ovo (1954); (b) draw-ing of the last (1627) surviving auroch, the ancestor of cattle, based on a painting found in a shop in Augsburg.

a

b

of the traits in which the races differ, the Chinese carpare more similar to wild carp than the European. TheChinese common carp should be regarded as a semi-domesticated animal, since wild fish are frequentlyadded to breeding stocks.” So the myth of commoncarp domestication in China more than two thousandyears ago is just that – a myth. In China the commoncarp remains at best as an exploited captive or a feralform of the introduced European domesticates.The European domestic forms were later introducedto eastern Asia (e.g., Basavaraju et al., 2003). Thestill lingering myth that carp came to Europe fromChina seems impossible. “Common carp were knownand used in the fish culture of western Europe wellbefore any direct European contact with China[emphasizes Hoffmann, 1995a: 75]. The fish culturemethods used in the west since at latest around 1200were, with their emphasis on special ponds kept per-manently filled for several years growth of a singleage class and then dry for a time after harvest, arequite the reverse of a fish culture integrated with ricepaddy cultivation. And finally, given the known loca-tions of common carp in Europe and of the earliestEuropean fish culture enterprises, we can rule out theindirect route from China via the Middle East...”

The common carp, Cyprinus carpio

“Aristocrat! Aristocrat!” he cried. Luckily, knowinghim better, I was able to explain. Far from wishing tosend anyone to the guillotine, what he meant was thattrout were aristocratic fish, the plaything of the rich,whereas effort would be better devoted to popularis-ing carp culture, the ideal pond fish, he averred, forhungry villagers.

Peter B. N. Jackson (2001: 40-41)in Freshwater Fishery Research Organisations in

Central and Eastern Africa. A Personal Recollection.

The earliest domesticate and the most important fishin aquaculture, the common carp, Cyprinus carpioLinnaeus, 1758, was made taxonomically also themost confusing species. According to the latestreview by Baru et al. (2002: 87) it remains without aholotype and was described “using specimens from apond culture, designating Europe as the terra typica.Presuming that all domestic European forms origi-nated as wild carps from the Danube, specimens fromthe Danube River can be considered as typical.”Most of the confusion comes from giving taxonomicnames to local or feral specimens or populations. Thecommon carp is very variable and forms easily distinctlocal morphotypes (see fig. 6-8 in Balon, 1995b). Sinceearliest times, these forms have tempted authors to givethese deviants separate names (e.g., Bloch, 1782;Heckel and Kner, 1858; Antipa, 1909; Zhou and Chu,1986). These morphotypes were later proven todevelop relatively fast from the wild common carp in

pond culture (see e.g., Tu a, 1958; Bastl, 1961; Mi íkand Tu a, 1965). For example, Zhou & Chu (1986) list12 “species” from Yunan Province, China, and Baruet al. (2002) ended up with a rough count of more than30 synonyms and over 10 subspecies, varieties andmorphs from across its range in comparison with Kotte-lat’s (1997) about 15 and 8, respectively. Baru et al.(2002: 115), after a tortuous review using the best dataavailable (e.g., Mi ík, 1958), concluded that at the pre-sent time only three subspecies of wild C. carpio can berecognized: “1. The European and central Asian wildcommon carp, Cyprinus carpio carpio (...); 2. The eastAsian wild common carp, Cyprinus carpiohaematopterus (...); [and] 3. The south-east Asian wildcommon carp,C. carpio viridiviolaceus (...).” None of thedata are convincing, nor is the claimed occurrence of C.c. haematopterus throughout east Asia from the AmurRiver across China, Korea and Japan (Svetovidov,1933). C. haematopterus was described as a separatebut now invalid species (Eschmeyer, 1998: 701) fromthe area of Nagasaki, Japan, in von Siebold’s FaunaJaponica by Temminck and Schlegel (1845); the#2400 holotype parades among the “stuffed” specimensin the Rijksmuseum van Natuurlijke Historie, Leiden,Netherlands.More than 200 years earlier, “the first Portugueseand Dutch colonies were established at Nagasaki. Bythen common carp culture was widespread in theEuropean homelands of these colonists and it is there-fore possible that the common carp was introduced bythem to Japan” (see Balon, 1995b, caption to fig. 5 onp. 10). The Nagasaki Municipal Museum is below andnext to the Atomic Bomb Museum. If one escapes thehorrors above, tableware decorated with common carpimages once belonging to these early Dutch colonistscan be seen on display (e.g., Fig. 2). At the very leastthis demonstrates an interest in the common carp byDutch colonists of the sixteenth and seventeenth cen-turies. While the species of common carp describedfrom the area as “haematopterus” erroneouslybecame the nominal subspecies for the huge area ofChina more than two centuries later, it is more correctto consider the common carp even there as feraldescendants of European domesticates that escapedor were introduced to Japan and possibly China. Sofar the results from molecular markers have beenlargely inconclusive (e.g., Froufe et al., 2002;Kohlmann et al., 2003). Hence, before this taxonomicconfusion can be resolved, if ever, it would be moreprudent to consider the wild ancestor of the commoncarp as a single species, Cyprinus carpio, widely dis-tributed from the Danube to the Amur rivers, as feralforms elsewhere (but see Kottelat, 1997) and as natu-ralized “wherever suitable conditions prevail” (Lever,1996: 92). Do books like Guidon’s (1983) The WorldAccording to Carp or Bradley’s (1997) This Book is Fullof Carp merely illustrate unjust American disgust withthe fish or its popularity?

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Wild common carp, the ancestor of the domesti-cated formsAll data so far point to the wild common carp fromthe Danube River as the most ancestral form (Fig. 3)that was initially kept as exploited captives and laterdomesticated. The inland delta of this river, recentlybadly damaged (Balon and Hol ík, 1998, 1999) by theinfamous Gab ikovo water works, was the area clos-est to cultural events opportune to interest in the carp.Carp remnants found in archeological excavations ofRoman settlements in this area confirm that commoncarp was indeed already then of interest (Balon,1995b). We will therefore focus entirely on describingthe wild common carp from this area before it wasextirpated or replaced by feral forms.Around 1955 large schools of the wild common carpwere still spawning every year over many inundatedmeadows along the Slovak part of the Danube andLesser Danube (see Mi ík, 1957; Bél, 1962). As itwas assumed that these fish were the direct descen-dants of much larger schools of wild common carpexploited by the Celts and Romans in the same areaabout two millennia earlier, I initiated detailed studiesof these fish with my late laboratory assistants VictorMi ík and Julius Bél (Balon and Mi ík, 1956; Balon,1957, 1958a,b; Mi ík, 1958). There follows a briefsummary of these studies already presented earlierby Balon (1995a,b) and again compiled by Baru et

al. (2002).The wild common carp had a long, round, torpedo-shaped body with the transition from head to the trunkforming an almost straight line dorsally, and not show-ing the depression (notch), typical for most domesti-cated or feral carps (Fig. 4). Very regular, large scalescovered the entire body, each scale pigmented dark atits joining edge, giving the body a mesh-like appear-ance. The basic coloration was a golden body, darkeron the dorsum and lighter at the ventrum, head anddorsal and caudal fins greenish-brown, the ventralparts of the caudal fin, anal and pelvic fins, lips and bar-bels light with an orange tinge (Fig. 3). The lateral linepores pierced scales through the middle of the sidesand were complete but barely visible on live individuals.The females have, on average, longer heads thanmales. The average length of the front barbels isabout half that of the rear barbels. The females havea broader forehead than males. At spawning timemales develop breeding tubercles which appear aspinhead sized white granules on the scales aboveand below the lateral line, on the caudal peduncle andon the head, especially on the anterior parts of oper-cles, preopercles and under the eyes. Tuberclesoccur even along the insides of first rays of the pec-toral and pelvic fins, and along the unbranched rays ofthe dorsal, anal and caudal fins. Breeding tuberclesdevelop also in some females, but only on the head;

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Fig. 2. A plate of the early Dutch colonists at Nagasaki (17th century) decorated with a common carp image, exhibited atthe Nagasaki Municipal Museum. Photo by E. K. Balon, 1994.

never on scales or fins.The morphometric characters are given in Table I.The number of spines and rays in the dorsal fin of theDanubian wild common carp was (II) III-IV 18-21 (22);the number of spines and rays in the anal fin was (II)III 4-5 (in both fins the last spine was a large bonestrongly serrated along its posterior edge, one of thebest bones for carp identification in archeologicalsamples); the number of spines and rays in the cau-dal fin was IV - VIII 16-18 IV - VIII; the number ofspines and rays in pelvic fins was II 7-9 with no sig-nificant difference between females and males. Thenumber of scales in the lateral line was (34-35) 36-39

(40) with no difference between left or right side orbetween sexes, and the number of scales above andbelow the lateral line was always 5-7/5-7. Number ofgill rakers outside/inside was 22-28/29-34 with theinside number always higher and no differencebetween sexes. The pharyngeal teeth were moliformwith clear grooves on the crushing surfaces, in threerows as 1.1.3-3.1.1. The differences between thesecharacters of the wild common carp from the Danubeare insignificant in comparison with these from thecommon carp of the Amur River. Overall, the variationin characters compiled by Baru et al. (2002) greatlyoverlap and subspecific differences appear only ifforced statistically. Similarly, the differences in morpho-metric characters between the Danubian and AmurRiver wild common carp are insignificant (Table I). TheDanube females were larger than the males (498.5 ±11.17 versus 447.5 ± 6.95 mm standard length).The karyotype of the wild common carp is 100 (2n)of somatic chromosomes. There is a remarkably largenumber of α chromosomes; no sex chromosomeswere ever found. “Karyotype of the wild common carp,C. carpio, from the Danube is identical with those ofthe cultured domestic Rumanian, Hungarian andUkrainian forms of the carp (Raicu et al. 1972) as wellof the French ones (Hafez et al. 1978). Similarly, nosubstantial differences exist between the karyotypes ofEuropean carps and some Chinese subspecies (Zhan& Song 1980; Ráb 1995). Ráb (1994) stated that thereis no difference between the karyotype of carps fromwild populations in the Danube and that of wild carpsfrom the Amur River” (Baru et al., 2002: 94-95).

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Table I. Morphometric characters of 87 individuals from spawning schools of the wild common carp, Cyprinus carpio, inthe Slovak parts of the Danube River, collected May 1954 and May 1956 (Mi ík, 1958), compared to the wild commoncarp from the Amur River (Gromov, 1973; see also Baru et al., 2002).

Measurements Danube Amurand counts range range

1 - 2 in % of standard length 359-635 mm 518.0 mm

1 - 3 head length 22.7 - 26.7 21.6 - 28.51 - 6 snout length 7.6 - 10.8 7.6 - 10.56 - 11 orbit diameter 2.9 - 3.9 2.6 - 4.57 - 9 interorbital width 8.9 - 11.7 7.5 - 8.511 - 3 postorbital head length 11.2 - 13.8 8.6 - 13.51 - 12 predorsal distance 42.1 - 48.1 43.5 - 51.512 - 15 body depth 24.6 - 30.9 26.5 - 35.512 - 26 body width 13.8 - 22.0 –16 - 2 caudal peduncle length 18.1 - 23.2 15.5 - 20.518 - 19 minimum body depth 11.1 - 13.6 10.5 - 13.520 - 13 pectoral origin to pelvic base 20.7 - 26.4 21.5 - 25.513 - 14 pelvic base to anal base 27.5 - 33.1 –12 - 21 base length of dorsal fin 35.8 - 44.9 35.5 - 42.514 - 16 base length of anal fin 7.3 - 10.4 9.0 - 11.520 - 26 longest pectoral ray length 15.6 - 20.9 15.5 - 20.513 - 27 longest pelvic ray length 14.7 - 19.9 14.5 - 18.512 - 28 longest dorsal fin ray length 14.0 - 18.8 10.5 - 14.514 - 29 longest anal fin ray length 14.3 - 17.5 10.5 - 14.5

Fig. 3. Male and female of the wild common carp fromthe spawning school at the Lesser Danube aboveKolárovo (1955) below the wild carp individual caught thefollowing year in September. Photos by E. K. Balon.

The life history and ecology of the wild common carpwas compiled in detail by Balon (1967b, 1974, 1995a,b)and again by Baru et al. (2002). For the sake ofbrevity only some selected highlights will be repeatedhere.In the Danube the wild common carp femalesreleased two or three portions of eggs within 10 to 14day intervals. In 1955 the carp spawned in largeschools on the inundated meadows above the villageof Kolárovo (Fig. 5). The first spawning occurred onMay 6th and 7th, but because of receding water anda sudden drop in temperature the spawning was inter-rupted. It resumed between the 5th and 7th Junewhen the flood returned and the temperature rose to18ºC. Although various forms of the domesticatedcarps – deep bodied and of irregular scalation –stocked into this area were present in the vicinity, onlyrarely were one or two such individuals caught withinthe school of hundreds typical wild common carp. Fur-thermore, such single domestic forms caught with thewild common carp were always immature or not in

spawning condition. Later Lelek (1987; Brunken et al.,1989) established that the feral carp in the Rhine,morphed close to the ancestral wild common carp,also maintained spawning allochrony from the domes-ticated forms. Is the “species recognition concept”advanced by Paterson (1985, 1993) that precise?The age and growth of the wild common carp, deter-mined from scales of the subsample from the spawn-ing school at the inundated meadows near Kolárovo,is shown in Table II (Balon, 1957). Males belonged toage groups 3 to 15, mostly group 5, and females toage groups 5 to 9, most of them in age group 7.The activated eggs of the wild common carp have ayellowish tint and are 1.5 to 1.8 mm in diameter. Theouter egg envelope is adhesive and sticks to grassblades, preventing the eggs from falling onto the anoxicbottom of freshly flooded grassland. At about 20° to23°C, most embryos hatch in three days. For about twodays after hatching the free embryos hang in a verticalposition on the grass blades. Five days after activation,the posterior part of the swimbladder is filled and swim-

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Fig. 4. A typical wild common carp and a feral form of domesticated carp with a characteristic notch at the neck dorsum(below) from the Danube River delta before 1900. From Antipa.

ming horizontally, the larvae start to capture and ingestexternal food (Balon, 1958a, 1995b).“When we subsequently witnessed [writes Balon,1995b: 19] similar turbulent mating of wild carp groupson other freshly flooded meadows along the Lesserand main Danube’s inundation areas, by then alreadygrossly restricted by flood dikes, it became obvious tome that the original floodplain of this piedmont zone ofthe Danube (e.g., Mákkai, 1985) must have beenclearly dominated by this fish during Celtic and Romantimes. (...) Hol ík (1995b) recently noted: ‘The wildcarp was commercially harvested four decades ago(...) At present, however, its distribution is restricted tothe minor part of the main channel of the Danube. Onlysingle specimens are occasionally caught. Of 1536fish taken, tagged and released in the Danube atGab ikovo in 1993, only 5 (0.3%) were a carp but onlyone still displayed the typical characters of the wildform! (...) The extinction of (...) the wild carp isexpected within a decade...’.” (see also Hol ik, 2003).

Lucullan demands, Pannonia and the first carpcaptivesWhereas the genuine wild common carp thrived inthe area described above more than 2000 years ago,still lived in historical times, and was abundantenough only 50 years ago to lend itself for major stud-ies (Fig. 6), none may be left now. Two significant his-torical events coincided and contributed to the factthat for the first time wild common carp from this areabecame not only a major food source but alsoexploited captives:(1) In the last years BC and in the first and secondcentury AD, an extraordinary luxury, even excess, offood consumption, and the associated importation offoreign foods developed among the Romans (Friedlän-der, 1936). Sergius Orata, the teacher of Cicero,

devised special reservoirs where fishes for the kitchenwere stored. These reservoirs called piscinae soonbecame very fashionable, for they ensured a perma-nent supply of a variety of fresh fishes independent ofweather conditions and fishing success (Zeuner,1963). Fish rearing in piscinae was adopted, for exam-ple, by Lucinius Muraena, who began adding freshwa-ter fishes to the initially stored marine ones (Pliny theElder AD 24 - 79; in translation 1958-1962). The patri-cians soon competed in building such piscinae. Tobring water to his ponds, a reputed gourmet, ConsulLucullus (75 BC), dug a trench that cost more than hisvilla through a hill near Naples. Varro (116-27 BC;1912) and Columella (∼50 AD; 1941-1968) claimed thatthe Roman patrician preferred seawater ponds, andthat freshwater ponds (piscinae dulces) were consid-ered plebeian, but the documented prejudice is proof oftheir existence. Also, the existence of “plebeian”owned fish ponds may signify that the desire to trans-port wild common carp from the Danube to patricianpiscinae in “Italia” was applicable also to plebeiantroops and the accompanying tradesmen and artisans(but see Hoffmann, 1995, Appendix A).(2) In the first years AD, the Roman Empireexpanded its northern boundary beyond the barrier ofthe Alps right up to the banks of the Danube River(Sitwell, 1981). The northernmost province Pannoniafaced formidable forces of Celts and Germans acrossthe Danube and the Empire had to establish a strongmilitary presence quickly (Fig. 6). “In the second cen-tury, the comparatively short stretch of river betweenVienna and Budapest, about 150 miles long, requiredno less than four legions to guard it – X Gemina atVienna itself, XIV Gemina Martia Victrix (...) at Car-nuntum, I Adiutrix at a place called Brigetio (...), and IIAdiutrix at Aquincum (now Budapest). By contrast, allof Roman Britain in the second century required only

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Table II. Back-calculated mean standard lengths (in mm) for males and females of the wild common carp from the spawn-ing school at the Lesser Danube above Kolárovo.

Year of Age Mean SL Back-calculated mean SL at the end of each growth seasonspawning group at time of

capture l1 l2 l3 l4 l5 l6 l7 l8 l9 l10 l11 l12 l131952 males III 422 154 382 4101951 IV 400 158 298 360 3911950 V 428 149 298 354 391 4141948 VII 459 135 228 304 363 398 425 4481947 VIII 462 142 196 296 358 391 426 4451945 X 548 180 203 246 436 450 472 504 522 5411944 XI 478 150 230 283 302 341 375 410 440 460 4701940 XV 598 148 235 278 335 361 410 422 467 485 501 521 536 550

152 259 316 368 392 422 446 476 495 501 521 536 550

1950 females V 418 120 321 366 390 4111949 VI 458 137 228 325 416 430 4501948 VII 491 136 276 343 385 422 453 4751947 VIII 483 150 271 379 404 456 450 468 4781946 IX 527 169 231 323 384 421 452 491 509 526

142 265 347 396 424 451 478 493 526

three legions; Roman North Africa managed with asingle one” (Sitwell 1981, p. 120). “To paraphrase fur-ther from Sitwell’s synthesis [continued Balon, 1995b:23-24], these at least 20 000 fighting legionnaires wereaccompanied by supporting troops, wives, mistresses,children, slaves and tradesmen to a total of more than100 000 Romans, in addition to ‘indigenous communi-ties (civitates) [which] came under the military control ofa high-ranking officer (...) from a neighboring Romanunit. (...) there was at least one auxiliary unit for everytwo civitates’ (Mócsy 1974, p. 49). These togetherformed a human population large enough to establish acarp-eating tradition, if a wild common carp was, as Ibelieve at least seasonally, the most abundant andmosteasily caught fish in the area. The influence of thisyoungest province and its army was so strong that‘instead of Rome controlling the Danube frontier, itwould be nearer the truth to say that men from that fron-tier controlled Rome’ (Sitwell 1981, p. 122).”Movements of legionnaires, troops, auxiliary unitsand civilians from Pannonia across or around the Alpsto Poetovio, Emona via north-eastern Italia to Rome[...] required the construction of relatively good roads(see also fig. 59 in Mócsy, 1974), one of which endedas the Roman branch of the Amber Road at Devin –

Marus (Morava) River entry into the Danube – andcrossed the Danube north into the Celtic, German andSarmatian territory toward the Baltic Sea as a foot-path. Legionary fortresses and Roman towns wereestablished at the edge of the largest floodplain areawithin the piedmont zone of the Danube: castra andcolonia Carnuntum upstream and castella Gerulata(now Rusovce) downstream of the Morava (Marus)River entry into the Danube at the upper end of thefloodplain. Castellas Ad Flexum, Quadrata, Arrabonaand Ad Statuas before the larger castra and coloniaBrigetio (now Szöny) at the Váh (Duria) River entryopposite the castella Celamantia (near today’s villageI a) at the floodplain’s downstream end. This flood-plain area was utilized as the westernmost spawninggrounds of the wild common carp, Cyprinus carpio.So the carp must have been well known to theRomans in Pannonia.Under the rule of caesar Hadrian (117-136 AD) andlater of Marcus Aurelius (161-180 AD), the Romansbuilt a smaller military fort (castellum) Celamantiaacross the river from Brigetio, within the Danubefloodplain. Repeatedly repaired, the stone fort wasfinally abandoned shortly after the death of caesarValentinius I (375 AD). Archeological excavations at

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Fig. 5. Commercial fishermen and their boat constructed for seine-net catches of the spawning wild common carp at theshallow floodplains of the Lesser Danube above Kolárovo. Photo by E. K. Balon, 1955.

Celamantia were resumed over the past 20 yearsbecause of its possible submersion by theGab íkovo-Nagymaros water works (Rajtár, 1990,1992). In earlier such excavations at Carnuntum andBrigetio interest in Roman man-made artefacts pre-vailed. At Celamantia on this occasion fish remainswere also carefully sampled, revealing that the bonesof the wild common carp outnumbered all other fishbones (K. Hensel, personal communication).Together with those from Novae on the lower Danube(Szymczyk, 1987) and Nicopolis ad Istrum from 100-450 AD (Irving, 1994) these samples from Celaman-

tia (Fig. 7) constitute the first direct evidence thatRomans knew the wild common carp of the Danubewell. It is most likely therefore, that, while living on theDanube in Pannonia the Romans, not only consumedthe wild common carp but also transported them aliveto other Roman provinces, because of the popularfashion at that time of keeping fishes in piscinae.Returning to “Italia”, many thousands of soldiers prob-ably demanded to be served their customary fish, soinfluencing the importation of common carp. After all,the common carp was the largest, tastiest fish thatmight have survived the rigors of transport.

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a

There is so far no written or archeological evidenceof common carp being kept in piscinae or vivaria inRome 2000 years ago (Zeuner, 1963; Hoffmann,1995a). Since written records from this time are gen-erally scarce even regarding activities that have beenproven for that period by other means, the keeping ofcarp by Romans in those times is still highly likely.

After all, Walton (1676: 146) claimed “Tis said (byJovius [who probably was Paolo Govio, a papal physi-cian, who wrote about fishes on the Roman market in1524, Richard Hoffmann, personal communication],who hath writ of fishes) that in the lake Larius [LakeComo] in Italy carps have thriven to be more than fiftypounds weight.” These early introductions of the wild

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Fig. 6. (a) Part of the Roman Empire with roads and settlements and the Amber Road (dotted yellow). (b) Part of Pan-nonia at the Danube River frontier. Modified from Sitwell (1981).

Fig. 7. a. Present view of the excavated Roman castel-lum Celamantia near the modern village I a, with theDanube River hidden behind the flood dike on the farside. b. Common carp bones from the excavations atCelamantia; c. the strong serrated spine of the anal fin,and (d.) the maxilla, pharyngeal bone and opercle. Photoby E. K. Balon, 1993.

a

b

c

d

b

common carp west of the middle Danube were prob-ably only using the animal as an exploited captive.The history of common carp domesticationAndreska (1987: 32) states in the chapter on the com-mon carp and ponds that “old chroniclers in their writ-ings dealt not with unessential details. They have writ-ten mainly about saints, rulers, wars and wondrouscelestial signs. About who was the first in bringing (...)seeds of the carp, they left not the slightest record” (mytranslation). The earlier assumption that the keeping ofcommon carp in monastery ponds continued after thecollapse of the Roman Empire and the establishment ofChristianity is a pure conjecture regarding the periodfrom the early middle ages to the tenth century (Hoff-mann, 1995a). It is based on two logical assumptions.First, that monasteries began to be founded in the fifthand sixth centuries and soon acquired land and farms.Second, that Christianity introduced more than 180 fast-ing days per year which were mainly observed by duti-ful monks, nuns and priests, thus interest in the com-mon carp should have survived.On fasting days the only flesh that might be eaten wasthat of aquatic animals such as shellfish and fishes –and also in some regions unborn rabbits called laurices– thus these customs are directly linked to Roman tra-dition (see more in Balon, 1967c, 1995b). The only evi-

dence that the Roman taste for eating common carpcontinued is from Cassiodorus (AD 490-585;1626) whoordered delivery of the wild common carp from theDanube to Italy for the table of King Theoderic.The first to write about breeding common carp inponds was Albertus Magnus in the 1260s (1193-1280;1861), he may however compete for this privilege withthe Count of Champagne who according to Hoffmann(1994a: 142) was breeding common carp in 1258.Hoffmann states that “no remains or verbal mentionsfrom before the twelfth century suggest carp culture orartificial fish ponds. These are wild fish.” Ponds withmuddy bottoms have been built since the tenth cen-tury (Hoffmann, 1985; Andreska, 1987), and wildcommon carp became established in them in thetwelfth century at the latest, judging from the relativelyfast process of domestication as reported by Tu a(1958), Bastl (1961) and Mi ík and Tu a (1965). Soit seems that in the common carp the transition fromexploited captive to truly domesticated animal tookplace in the twelfth century.

Modern carp culture: “To build a fishpond, not justa holding tank, is to create a new aquatic habitat.Active construction of ponds for this purpose gotunder way in the eleventh century [documented Hoff-

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Fig. 8. The common carp harvest in southern Bohemia near T ebo¸ in early November 1998. Photo by E. K. Balon.

mann, 1995a: 67] and increased rapidly in the twelfthand thirteenth.” While initially these ponds werestocked with several local fishes, common carp soonproved to be the best choice for such man-made habi-tats. Hoffmann (1994a, b, 1995a: 72) compiled the evi-dence for medieval times in Europe and concludedthat: “Verbal and archeological evidence thus togethercorroborate three phases in the medieval diffusion ofcommon carp west and north from an epicentre at thenorth-west margin of its native range in Pannonia (thepiedmont section of the Danube, Balon 1967a). Thefirst phase between perhaps the seventh and theeleventh centuries principally carried the fish up theDanube and into at least some west-flowing tributariesof the middle Rhine. Some remains also indicate carpin waters north of the Danube. In a second phase(twelfth to the early fourteenth century) common carpradiated across most of the economic and culturalheartland of medieval northwestern Europe, from thelower Rhine/Maas region south to the Paris basin andBurgundy. In a final stage, probably occurring after themid-fourteenth-century shift of economic trends,extended common carp into the outer periphery of the

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Fig. 9. Harvest of common carp from a pond near Hluboka in mid-November 1998. Photo by E. K. Balon.

Fig. 10. Prevailing form of domesticated common carpfrom the harvest in Fig. 9. Photo by E. K. Balon.

west, an arc from southwestern France through Eng-land and southern Scandinavia into east centralEurope. Though carp are recorded in Italy at the endof the Middle Ages, the diffusion pattern at no timeaccords with radiation from there.”In time more written evidence appeared and by theearly fourteenth century culture of the common carpwas well established (Pucher, 1987; Hoffmann,1995b, 2002). Soon the entire procedure of pond carpmanagement was described in the famous Latin man-ual by Jan Dubravius (1547)1 written between 1535and 1540, and by Olbrycht Strumie˜ski (1573) whoreported on the Czech and Polish techniques of thattime (Strojnowski, 1609; Æusta, 1889; Inglot andNyrek, 1960; Szczygielski, 1967a,b; Berka, 1986). Forexample, von Hohberg (1687) mused that the rearingmethods of the common carp in his time were supe-rior to those used by the Romans who would surelyhave enjoyed the taste of their contemporary carp.Dubravius (1547) also frequently mentions theRomans in relation to the common carp, but RichardHoffmann (personal communication) spoiled the fundeclaring that “these remarks are more likely a con-temporary literary fashion to quote Latin sources thanknowledge of the Roman carp culture” (Balon, 1995b:30). What a pity!The “golden age” of pond construction for rearing thecommon carp reappeared when the wars and skir-mishes of the fifteenth century had ended. It has beenestimated that more than 25,000 carp ponds werebuilt towards the end of the fifteenth century and atthe beginning of the sixteenth century in Bohemia andMoravia alone. A special guild of mobile and roguepond builders flourished. During these 50 years,about 500 ponds per year, i.e., two ponds per day,were constructed (Andreska, 1987). Production of thecommon carp for food became the most profitablebranch of agriculture (Andreska, 1997; Guziur et al.,2003). Various wars and a decline in the price of com-mon carp later led to a dramatic reduction in the num-ber and area of these ponds from a total of 180 000ha to about 51 000 ha. At 711 ha, Ro emberg pond,which was built in the sixteenth century, one of the 34largest, still operates on the two to three year cycle ofcarp production (Kourzil and Guziur, 2004).Already Dubravius (1547) recommended a systemof several specialized ponds for the production ofmarketable common carp. The smallest of these wasthe spawning pond with a grassy bottom simulatingshallow, freshly flooded meadows into which groupsof selected parent fish were released. Soon afterspawning, the early juveniles (Balon, 1999) wereflushed into nursery ponds at a lower level, then oneyear later into growing ponds. In some years pondswere left dry in rotation, treated with lime and various

chemical and natural fertilizers in order to enhancethe carp’s food production. Some consider that T.Dubisch, the illiterate Slovak fish master (Morcinek,1909), invented the system of transferring commoncarp from one growing pond to the next, while gradu-ally lowering their densities (Billard, 1999; Guziur etal., 2003). While the production of young carp inspawning ponds was recommended for every pondsystem, it was later replaced, at least partly, by theartificial stripping of gametes under hatchery condi-tions (Woynarovich, 1962; Billard et al., 1995). Thegrowth of the common carp was enhanced by everbetter formulations of supplementary food, the systemof sequential, specially fertilized ponds remained.Production increased from 40 to more than 450 kg perha ( ítek et al., 1998), with the growth cycle of mar-ketable common carp reduced from 5-6 to 2-4 years(Fig. 8, 9, 10, 11).Sooner or later different forms of domestic commoncarp appeared in the various pond systems. When thewild common carp was introduced into a pond systemits torpedo-shaped body naturally started to change toa deep, laterally compressed, hunchbacked body.Soon individuals appeared with no regular, geometricalarrangement of scales. Instead scale arrangementswere severely irregular, and scales were reduced oreven absent. These variations soon became a sourcefor artificial selection. Ultimately, the domesticatedcommon carp are represented by a variety of forms,such as scaled carp, line carp, mirror carp, and leatheror naked carp (e.g., Bryli˜ska, 1986; Pokornø et al.,1995; Gorda et al., 1995).

Comparisons of wild and domesticated commoncarp: The domesticated common carp underwentinternal and physiological changes as well as changesin its external shape, scalation and color. Detailedreviews were published elsewhere by Balon (1974,1995a,b) and Baru et al. (2002). A brief summary will

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Fig. 11. Other domesticated common carp forms, theleather and mirror, stocked into the Orava Dam Lake(Slovakia). Photo by E. K. Balon, 1958.

1 Second edition was published in 1559 probably by demand ofConrad Gessner, third edition appeared in 1596, and in 1599 DePiscinis appeared in English translation (Andreska, 1987).

suffice here: The mouth gape, the first character thatRudzi˜ski (1961) and Steffens (1964) used for cleardistinction between the wild and the domesticated com-mon carp, was much smaller in the wild than in domes-ticates (Table III). “Even more pronounced differencesare encountered among the mouth-gape indices calcu-lated by dividing 10 times the mouth gape in squarecentimeters by the length of the head in cm – 4.46 to5.57 for the wild common carp and 8.12 for domesti-cates” (Steffens, 1964). Both authors considered theenlargement of the mouth in domesticates to becaused by changes in diet, and possibly the result ofartificial selection. Domesticated common carpselected to utilize supplementary food, grew better inponds when man-made food was added (see Sibbing,1988). Studies on wild common carp in ponds (Rudz-i˜ski, 1961; Leszczy˜ska and Biniakowski, 1967)clearly proved that wild common carp progeny werebetter suited for stocking natural riverine habitats thanwere domesticates.Rudzi˜ski (1961) found and Steffens (1964) con-firmed that the intestine of the wild common carp was15 to 25% shorter than that of a domesticated carp.The index calculated by dividing the length of theintestine by standard length in centimeters was 2.11for the wild common carp and 2.64 for the domesti-cates. Longer intestines in domesticates are likely dueto utilization of vegetable food not normally consumedby wild common carp. Both authors also found, whencomparing the deeper body of the domesticates withthe cylindrical one of the wild common carp, that thecalculated muscle mass was the same in both,although it appears to be more in the domesticatedfish. It means that dressed mass of domesticates isnot larger in spite of the different body proportions.Also, both chambers of the swimbladder of the wildcommon carp are of similar size, whereas in thedomesticate the anterior chamber is always muchlarger and the posterior chamber smaller; the propor-tion in percent as given by Steffens (1980) is 61:39 forthe wild and 90:10 for the domesticate. This mayrelate to the relative greater mass of the head indomesticated common carp.

The greater strength of the wild common carp wassupported by some of the physiological attributes(Steffens, 1964). The wild common carp had 18 to19% more erythrocytes and hemoglobin than did thedomesticate; blood sugar level was 16 to 26% higherin the former. Also, the wild common carp had a muchlower water content in muscles and liver than did thedomesticate. Furthermore, the wild common carp hadgreater fat content in individual organs, more glyco-gen in the liver, and more vitamin A in the eyes, intes-tine, and liver. In addition its muscles were more vas-cularized and did not tire as quickly as those of thedomesticate. As in some exploited captive salmonids(Balon 1980, 2004), there are some indications thatcarp domesticates usually produce eggs with lessyolk and as a consequence more altricial progenythan the ancestral wild form (Kryzhanovsky et al.,1951; Balon, 1958a; Pe¸áz, 1995). This aspect willbe dealt with in the final section.As already mentioned, specific forms of the domes-ticated common carp are created and maintainedthrough constant artificial selection in nearly every indi-vidual pond culture system. Standardized breeds weredeveloped and new forms are continually tested inplaces such as the Fish Culture Research Institute atSzarvas, Hungary, the Institute of Ichthyobiology andPond Culture of the Polish Academy of Sciences atGo‚ysz, Poland, and at the Research Institute of Fish-ery and Hydrobiology of the Southern Bohemian Uni-versity at Vod¸any, Czech Republic (e.g., Gorda et al.,1995; Pokornø et al., 1995a,b; Flaj hans, 1996;Varadi et al., 2002; Guziur et al., 2003; see also Billardand Gall, 1995). The common carp is considered a del-icacy in Europe not only at Christmas, at which themain autumn harvest is aimed (Balon, 1966), but alsoin special restaurants, for which a professional cookerybook with more than 84 recipes for domesticated com-mon carp dishes was written around 1927 (Va¸ha,1993).

The emergence of ornamental “nishikigoi”: Thecommon carp was traditionally reared for human foodin small terrace ponds alternating with rice paddies

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Table III. Mouth gape and intestine length indices for the wild common carp and domesticated common carp: o l-1 (10 xmouth gape in square centimeters per standard length in centimeters), o w-1 (10 x mouth gape in square centimeters permass in grams), o lc-1 (10 x mouth gape in square centimeters per length of head in centimeters); gut l-1 (length of intes-tine in centimeters per 1 cm of standard length), gut w-1 (length of intestine in centimeters per 10 g of wet mass).

Index ol-1 ow-1 o lc-1 gut l-1 gut w-1

Wild common carp 1.14with supplementary diet 1.09 0.10 4.46 2.11 3.02with natural diet 1.24 0.08 5.27

Domesticated carp 1.91with supplementary diet 2.00 0.08 8.12 2.64 2.25with natural diet 2.30 0.08 8.12

between the cities of Nagaoka and Ojiya in the NiigataPrefecture facing the sea of Japan, some 280 kmwest of Tokyo. The area is reputed to have one of thehighest snowfalls in the world; for about six months ofthe winter the whole area is under at least six metersof snow. These unusual conditions may have con-tributed to the frequent occurrence of color aberra-tions in the common carp cultured here. The high inci-dence of color aberrations may be caused by mela-tonin production during life in total darkness under thecover of deep snow.This mountainous area had only few roads in thepast and the local farmers were often stranded formany long winter months. It is assumed that selectionand cross-breeding of the colored aberrants of thecommon carp began here 180 years ago (Kataoka,1989; Kuroki, 1990). There is, however, little evidenceto support the appearance of these early “nishikigoi”.Plate 16 in volume one of Illustrations of JapaneseAquatic Plants and Animals, published in 1931, showssome colored carp (Fig. 12) but no modern“nishikigoi” (Ishikawa et al., 1931; Balon, 1995b).Consequently, while the appearance of the coloredcommon carp more than 70 years ago is documented(Fig. 13), selection of the true “nishikigoi” (koi for

short) probably started on a commercial scale onlyafter the Second World War, in the 1950s. By thenimproved living standards in Japan and elsewhereallowed more people to afford garden ponds. Thesegarden ponds were fully exploited for decoration with“swimming flowers” particularly in Japan, where landis more precious than in many other parts of the worldand where Buddhism encourages a love of animals.The ponds were a logical progression in the develop-ment of the miniature garden. At time when the koibegan to enjoy international popularity, Amano (1968:36) wrote that “the number produced in a year is tenmillion, amounting to 1,000,000,000 yen worth”.The koi, as they are known today (Axelrod et al.,1996) are color aberrations of the common carp,Cyprinus carpio. They probably originated from thedomesticated common carp or its feral forms, andfrom some more recent imports from Europe (doitsukoi). When the color aberrations started to appearmore frequently and demands for them increased,production and crossbreeding intensified and theirprogeny started to be severely culled, leaving only thebest colored – 1 out of 10 000 or more – for futurebreeding. The first modern “nishikigoi” come from the30 hamlets of Yamakoshi united in 1956 in the Niigata

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Fig. 12. Plate 16 from the Illustrations of Japanese aquatic plants and animals, published in 1931 attests that apart fromsome xanthic individuals, no modern koi were known at that time, contrary to popular claims that koi were at least a cen-tury older.

Prefecture (Fig. 14) where “87 per cent of the 906families [are] being fancy carp producers” (Amano,1971: 37). The beauty of the wild common carp wasreinstated here in color, not unlike that of rabbit,chicken or cattle long time ago.Today, the Japanese recognize about 15 basic coloraberrants of the common carp called koi, each ofthese aberrants with many varieties (e.g., Phipps,1989; Kuroki, 1990; and the Nishiki Goi monthly mag-azine). The standard is the carp body to remain cylin-drical and the colors to be viewed from above. Theintroduction of koi with elongated fins and scale orbody deformation is not accepted in Japan butappears frequently from producers in Israel and else-where. Even by most careful crossbreeding and par-ent selection the resultant colors of the severelyculled offspring are impossible to predict (Kataoka,1989) by genecentric ideology (Ho, 1999; Hall et al.,2004). After several cullings only the most promisingand appealing colors are retained. “Out of these[enthuses Balon, 1995b: 41-42] grow numerous beau-tiful nishikigoi, but the winners of shows are but a few

unique individuals selected from many hundreds ofthousands. In Japan the favorites are the kôhaku –the red and white koi, taisho-sanke – red, black andwhite, and the showa sanshoku – a black koi withwhite and red imposed (...). As no two specimens areever alike, the grand champions of modern showscommonly sell for US $ 150 000 or more, others oftenfor more than a million dollars each. These must bethe highest prices paid for an individual fish!” Finally,it has been found that koi grow better if allowed to lis-ten to violin music for three hours per day (Vasanthaet al., 2003). Expensive music lovers!

The “chi” into goldfish, Carassius auratus

When preliminary studies in 1968 and 1969 indicatedthat goldfish tended to forget things when drunk, andthat Siamese Fighting Fish became more aggressiveafter a little drink or two, their attraction as experi-mental animals became irresistible.

William Hartston (1987: 7)in The Drunken Goldfish

“Of all the favourite species [informs Taylor, 1884:58] the goldfish has long been most domesticated, sothat now, like the canary among birds, it seems to bebetter adapted to confinement than even to a free rov-ing life.” It has also been an animal of choice for lab-oratory experimenting in physiology (e.g., Ostrander,2000).2 In spite of all this it is often confused with thecrucian carp, Prussian carp or even the common carp(Szczerbowski and Szczerbowski, 1996).There is no reason to confuse the chi or goldfish withcarp for goldfish, unlike common carp, have no bar-bels. The crucian carp, Carassius carassius (Lin-naeus, 1758) differs from the goldfish by (a) having 23to 33 gill rakers on the first branchial arch while wildgoldfish have 37 to 53, (b) having 28 to 29 fine denti-cles on the posterior edge of the last unbranchedspiny ray of the dorsal fin while goldfish have 10 to 11irregular denticles, (c) by having a light peritoneumwhile goldfish have a black peritoneum, (d) by havingthe upper edge of the dorsal fin raised while in gold-fish it is concave, and (e) by having a blackish spot atthe base of the caudal fin while goldfish have none.Body color of the crucian carp is usually coppery goldwhile that of the chi, the wild goldfish, is silvery-grey(Fig. 15). In spite of these differences, most FAO sta-tistics erroneously list goldfish, Carassius auratusauratus (Linnaeus, 1758), as crucian carp, but cruciancarp, C. carassius, is rarely used in aquaculture andis quite infrequent. The same mistake can be found in

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Fig. 13. Xanthic abberation of the common carp thatappeared among domesticated carp in a small privatepond in southern Bohemia. Photo by E. K. Balon, 1998.

2 I have found the most curious use of goldfish in the wonderfulautobiography by James Lovelock (2001: 406): “She inserted alive goldfish and, with a flourish and a heave of her powerfulvaginal muscles, expelled it into a bowl of water on the otherside of the room. She did this several times and missed onlyonce. (...) In the West, such an exhibition might have been criti-cized as unseemly, but in Japan, it rated as impermanent art.”

some aquarium literature (e.g., Matsui and Axelrod,1991; Wang et al., 2000), and even in the newestencyclopedia (Liu, 2003) although in the first editionall was again correct and more informative (Grzimekand Ladiges, 1973).The crucian carp, C. carassius, occurs in twomorphs: the precocial, fast growing and deep bodied

and the altricial, dwarfed morpha humilis (see fig. 15and 16 in Szczerbowski and Szczerbowski, 1996,2002). The fish occurs in most of Eurasia, except inthe northwestern and southwestern parts of Europe,and except in China, Korea and Japan. At many loca-tions it was replaced by the expanding Prussian carp,Carassius auratus gibelio (Bloch, 1783), that Kottelat

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Fig. 14. (a) The koi ponds on the terraces of Yamakoshi, Niigata, (b) selected three years old individuals at the breeder’spond, (c) price winning koi in the garden of a rich hobbyist in Kyoto, and (d) at the wholesaler in Ojiya, Niigata Prefec-ture, Japan. Photo by E. K. Balon, 1994.

(1997) recognized as a separate species, C. gibelio.The Prussian carp, silverish in color, is mostly a gyno-genetic, all female, triploid subspecies (Pelz, 1987;Hala ka and Lusková, 2000) that lives in the periph-eral areas of the distribution of the nominal chi or wildgoldfish, C. auratus auratus. The latter subspeciesoccurs with both sexes present. The wild goldfish isnative to China, and some adjacent areas. It wasintroduced to Japan from China between 1502 and1620 probably already as monstrositas (Matsui,1934). “Available data show that at least five geneti-cally and morphologically distinct stocks are known inJapan which are considered as distinct species orsubspecies [writes Kottelat, 1997: 52, and after sayingthat] the systematics of the genus Carassius in EastAsia is confusing” (see also Hensel, 1971; Kawanabeand Mixuno, 1989).All three most common Carassius species – the cru-cian carp, chi or goldfish and Prussian carp reach onlyan average size of about 35 cm and a body mass of1 kg, usually much less. The crucian carp and goldfishwere already distinguished during the Sung Dynastyafter 968 AD, claims Szczerbowski (2002: 14), butsince these two species do not naturally occur in sym-patry this claim may be without foundation. Anyway,the goldfish, C. auratus auratus, was introduced toEurope and elsewhere from China already as adomesticated monstrosity. “Valenciennes (1842: 108)summarised the history of the introduction of Caras-sius auratus to Europe [writes Kottelat, 1997: 52]. Hestates that some authors (but does not say which)consider that it was introduced to Europe as early as1611 or 1691 and that Yarrell reports that it had beenintroduced by the Portuguese from Java (...) to SouthAfrica and from there to Lisboa. The first introductionto England dates to 1691 (Pennant, 1812: 490) and toFrance to 1755 (Hervey & Hems, 1968 ...). It was bredin northern Europe for the first time in Holland in 1728according to Sterba (1987: 272).” Before this last datedomesticated goldfish had to come by boat fromChina and, therefore, must have been very expen-sive. As such their owners would guard them fromrelease into natural waters. The invasions fromaquaria and garden pools into all suitable habitatsaround the world came later.In the Purnel’s Encyclopedia of Animal Life (1968:918) the following statement reveals all: “The morefancy breeds of goldfish are freaks, no matter howattractive some of them may look. To recite theirnames is enough to make this point: veiltail, eggfish,telescope, calico, celestial, lionhead, tumbler, cometor meteor and pearl scale. There are also the waterbubble eye, blue fish, brown fish, brocade, pomponand fantail and many others. Some breeds are mon-strosities rather than freaks.” There is a Japaneseversion of calico oranda called azuma nishiki, themodern usage and depiction of most monstrosities tobe found especially in Li (1988), Wang et al. (2000) or

in Johnson and Hess (2004) and within the Internet,e.g. www.goldfishconnection.com.

Wild “chi”, the ancestor of the domesticated coloraberrations and monstrositiesThe wild “chi”, Carassius auratus auratus, has a deepbody, silver-grey or olive-green coloration and thickcaudal peduncle (Fig. 15). Large cycloid scales coverthe body, and the upper edge of the dorsal fin is slightlyconcave. Its feral forms are mostly slimmer andsmaller, much like the altricial morpha humilis commonin the crucian carp.However, the low, round body formis also common in the wild ancestor of goldfish, in thesilver-grey catches for food by Chinese fishermen.Szczerbowski (2002: 5) gives as the sole distinguishingmeristic character between C. auratus auratus and C.auratus gibelio the lateral line pore counts: 21 to 36 forthe former and 27 to 35 for C. a. gibelio. Out of the 25wild chi captured in the River Li waters at Yangshuo Ihandled on 15 April 2005, lateral line pores rangedfrom 27 to 31 (mean 40.3).Gynogenetic forms are not found in the goldfish.Originally, the number of chromosomes was reportedto be 94 (2n) in the goldfish (Makino, 1939), andbisexual populations of the Prussian carp (Cherfas,1966). In Vasilyev’s (1985) review, however, goldfishended up with 100 (2n) chromosomes and in Chiarelliet al. (1969) with 104 (2n) chromosomes. The triploidforms that reproduce by gynogenesis have 141 to 160(3n) chromosomes (Pe¸áz et al., 1979). The goldfishis the only cyprinid that has morphologically differen-tiated sex chromosomes (Szczerbowski, 2002).The goldfish is a portional spawner and releases twoto three batches of eggs in each season at 8 to 10 dayintervals. It deposits the adhesive eggs on grassblades and fine willow-like roots at water tempera-tures 18-19° C, very much like the common carp. Acti-vated eggs of the goldfish are spherical or slightlyoblong, 1.05 by 1.14 mm, transparent with a paleyellow or greyish-green tint. The embryos hatch 3to 8 mm long, still with a large yolksac and hangonto plants shortly before swimming and seekingexogenous food (Battle, 1940; Mansueti and Hardy,1967).The chi, wild ancestor of goldfish, is native to lateral-level waters of rivers and lakes of East and CentralAsia, with the center of distribution in China (e.g.,Chen and Fang, 1999). From escaped domesticatedforms and deliberate releases the goldfish hasacquired global distribution in temperate and subtrop-ical areas, naturalized, feral “populations developingin more than 20 European countries (...). Outside ofEurope, the goldfish has been introduced to SouthAfrica, and to Madagascar and Mauritius, and is wide-spread in Asia. It was introduced in 1874 to NorthAmerica3 (...), to Oceania, and from 1856 to 1930, to

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3 Innes (1949: 47) states that “marvelous specimens brought toAmerica by Admiral Annon in 1874.”

various countries in South and Central America andthe Caribbean (...). A slightly earlier date for NorthAmerican introduction is given by Fuller et al. (1999).In some localities, it occurs in sufficient abundance tobe marketed as a food fish” (Szczerbowski, 2002: 20).The goldfish is one of the most resistant fish to nat-ural environmental perturbations and man-made pol-lutants. It tolerates a wide temperature range and highwater turbidity. It is the only fish that survives in shal-low lakes with total oxygen depletion during someyears or seasons (e.g., 0.6 mg l-1 O2). The goldfishsurvives within a pH range of 4.5 to 10.5, but prefersthe optimal range of pH 7.2 to 8.4. In water at pH 4.5it can survive about two weeks. The goldfish is moretolerant than any other fishes to heavy metals ororganochlorine insecticides (Szczerbowski, 2002).The goldfish has been less successful than the com-mon carp in establishing itself in larger water bodies, butdoes best in small shallow lakes heavily overgrown byaquatic plants. It also grows to much smaller sizes thana common carp. “Females generally grow larger and livelonger than males [writes Moyle, 2002: 171 about thisfish in California]. As a result the male: female sex ratiochanges from 1 : 1 in small fish to 13-16 : 100 in fishmeasuring more than 15 cm total length. Although fishin the wild rarely live longer than 6-8 years, maximumages of 30 years have been recorded in aquaria.” For

example, in 1967 or 1968, goldfish were introduced intoKillarbey Lake in New Brunswick, Canada. Only sevenyears later, in 1974, they yielded 43% of the feral bronzeand 57% of variously colored fish. The largest of themmeasured 34 cm, weighed 1 kg and was 8 years old,having matured at age 2 (Hooper and Gilbert, 1978).

The history of goldfish domesticationWe may consider ourselves lucky that some 30years after his detailed studies on goldfish ShisanChen (1956)4 of the Academia Sinica and Beijing Uni-versity decided to review the history of domesticationof the chi once more, in spite of a changed political cli-mate. Since Ting-pong Koh’s (1934) study on thattopic, Chen had amassed additional evidence on thehistory of domestication of the goldfish. Now “forced”to insert a page where among other “timely” claims hewrites: “In the light of Michurin principles the writerreexamines his past labours and discovers manyerrors and shortcomings, which may be groupedunder the following three headings (...)” (Chen, 1956:288). None of these are relevant. He managed to pub-

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Fig. 15. Chi, the wild ancestor of goldfish, Carassius auratus, from a side water of the Li River near Yangshuo, China.Photo by E. K. Balon, 2005.

4 Chinese biologist Chen Zhen (Shisan C. Chen, 1894-1957)began in the 1920s to study the heredity, evolution and varia-tions of goldfish. He published very valuable writings in thisrealm that have been highly regarded by Chinese and foreignscientists. Under his influence scientists have paid ever moreattention to Chinese goldfish” (Li, 1988: 17-18).

lish the English version of his study (first published inChinese 1954) ten years before the rampages of theRed Guards (see e.g. Wong, 1996; Chang, 2003).The goldfish was created as ornamental fish domes-ticate in China. Sometimes a xanthic aberration wouldappear, as is the case in many other fishes (e.g.,Balon and Frank, 1953; Balon, 1964), among the sil-ver-grey chi, the most common food fish in China (Fig.16). These red or yellow aberrations are common butonly a few reach fishermen because most of themwould be eliminated in nature because of their con-spicuous appearance. “People in the old societyattached a supernatural significance to the appear-ance of the wild golden chi and regarded them assacrosanct (...). But there were places, like the NinghaiDistrict (H.4), where they served as food” (Chen, 1956:292)5.Buddhism teaches abstinence from taking the life ofany creature and urges one to do at least one gooddeed per day. Setting free a rare golden chi musthave been thought a better deed than releasing acommon animal. Hence, ever “since the beginning ofthe Sung Dynasty when Governor Ting Yen-tsan of theDuchy of Wu-yueh discovered golden and yellow chi ina pond outside the city of Kiahsing, that pond came tobe called ‘Goldfish Pond’. Subsequently, the pondbecame a pond of mercy. In the goldfish pond wereother varieties of fish and turtles besides the golden chi,which were forbidden to catch” (Chen, 1956: 294).Other such ponds followed at the pagodas inHangzhow and Nanking where the goldfish were caredfor by the local monks. The records thus document thebeginning of goldfish accumulation in ponds of mercyat the outset of Sung Dynasty, between 968 and 975

AD. However, until at least the year 1089 the goldfishwere not different from the wild chi except for color,were afraid of humans and did not take food thrown tothem. They were captives exploited for religious pur-poses.The documents reviewed by Chen (1956: 299), the

Bedside Companion (1214), On Names (1241) andDreams of the Past (1247) “explain the fact that the fadof the ruling class of the time brought a special, fish-breeding trade called yu-er-huo. These breederslearned to feed the goldfish with a kind of red animalcula(...) found in putrid water [tubifex worms?] and discov-ered the secrets of propagating goldfish so as to enablethem to offer a selection of rare varieties of goldfish togods on festival days.” By the year 1241 the goldfishhad become tamed and used to the food offered byhumans. While in the ponds of mercy, goldfish livedamong the common chi, other fishes and turtles, the pri-vate ponds constructed by the rich now contained onlyxanthic goldfish. It was, therefore, easier to breed them,cross the preferred individuals, and for the first time pro-duce the golden yellow, silver white, and the variegationof black and white (tortoiseshell) goldfish. This initialdomestication began “around the year 1163 in the gold-fish pond of the Te Shou Palace, east of Hsin KungChiao in the city of Hangchow” (Chen, 1956: 299).In general, ponds and garden pools were affordableonly to landowners. The goldfish, now in several colorforms, were first kept only by royalty and rich mandarinswho constructed more ponds, or rather garden pools,for their keeping and breeding. In 1330 during the YuanDynasty, the goldfish were brought to Beijing and later(1506-1521) in large numbers outside the ForbiddenCity. At that time they were also introduced to Japan.Chen (1956) marks the transition from pools toaquarium-like vessels from 1276 to 1546. Besides thejade vessels and fancy bowls for keeping goldfish,earthen vessels became common and affordable toeveryone (Fig. 17). After 1548, rich and poor alikebecame keepers and breeders of fancy domesticatedgoldfish, and the beginning of the aquarium hobby forthe masses started.“Kwei Yu-kuang (1506-1571), a noted man of lettersin the Ming Dynasty, wrote a long poem on ‘fire fish’:

Kept in water, of no ancient line,The fire fish does in new-styled fashion shine.A perfect marvel of few inches’ size,It has the brilliance of the rainbow’s dyes, . . .The narrow compass of the vat they takeWith equanimity as stream and lake . . .The children shout at them in unison,

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Fig. 16.Wild chi being sold at the sidewalk of Yangshuo.Photo by E. K. Balon, 2005.

5 Just this once, the evidence discovered by Shisan Chen (H.4):Chen, Chi-ching (122) in The Topography of Chihcheng, wood-cut edition of the Sung Family of Linhai in Chia Ching Reign, vol.25, Ninghai District: “Chuching Pond is forty li west of the city; itwinds about the neighbouring hills and has an area of hundredsof acres... It produces gold and silver chi of the most delicioustaste. But they are so mysterious that fishing boats approachingthem often turn upside down.”

And all day play about the vats for fun . . .This fish in every coastal home being bred,Often has to inland Kiangsu spread.”

(Chen, op.cit.: 302).

Later the “fire fish” was renamed the “cinnabar fish”.In the woodcut edition of Sketches of Scenes in theImperial Capital by Liu and Yu (1634) many goldfishmonstrosities are named and described.So it seems obvious that the creation of monstrosi-ties in shape (Fig. 18) in addition to already bred coloraberrations was closely related to the keeping ofindoor aquaria – a very popular hobby after the year1548. The goldfish became the pet of the masses, apopular pastime in every home and everywhere(claims Li, 1596 in another woodcut edition of MateriaMedica). The aquarium culture enabled close obser-vation and selection of the individual fish, somethinghardly possible in garden pools. In addition to pre-senting lists of many color varieties in A Handbook onthe Cinnabar Fish Chien-te Chang (1596) says: “Thebeauty of the cinnabar fish lies not only in their extra-ordinary brilliance of color, but also in their tails, pat-

terns, and bodily form, which distinguish them fromthe common fish. Whether long or short, the bodymust have the beauty of plumpness as a primaryqualification” (Chen, 1956: 305).Selected pairs of the domesticated goldfish three tofour years old start

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