About the oldest domesticates among fishes

Journal of Fish Biology (2004) 65 (Supplement A), 1–27 

doi:10.1111/j.1095-8649.2004.00563.x,availableonlineathttp://www.blackwell-synergy.com 

About the oldest domesticates among fishes 

E. K. BALON 

Department of Zoology and Institute of Ichthyology, University of Guelph, Guelph, 

Ontario N1G 2W1, Canada 

Domestication of mammals such as cattle, dogs, pigs and horses preceded that of fishes by at 

least 10 000 years. The first domesticated fish was the common carp Cyprinus carpio. Initially 

it was held as an exploited captive and did not undergo major changes in body shape or colour 

variations. About 2000 years ago, wild common carp were most abundant in the inland delta 

of the Danube River. These fish were torpedo shaped, golden-yellow in colour and had two 

pairs of barbels and a mesh-like scale pattern. Large schools of them thrived and reproduced on 

the flood plains of the Danube. The Romans kept fishes in specially built ponds at that time. 

The common carp was an ideal candidate and its rearing became more popular in medieval 

times. Common carp culture gradually became the most profitable branch of agriculture in 

central Europe and many special ponds were built. Soon common carp were being produced in 

pond systems including spawning and growing ponds. Unintentional artificial selection had 

taken place between the 12th and mid-14th century, and deep bodied and variously scaled or 

scaleless domesticated forms appeared in nearly every pond system. Some colour aberrations 

appeared in the 1950s in Japan, which, as koi, became the most expensive of fish. Common carp 

were not originally domesticated in China but wild ‘chi’ Carassius auratus occasionally appeared 

as a xanthic form that, as the goldfish, has been known since 960 A.D. By the 1200s the fish 

were used as ornamental animals in the garden pools of rich landowners. Circa 1276 to 1546, 

the Chinese began keeping golden chi in aquarium-like vessels and soon rich and poor alike 

became breeders of the fancy domesticated goldfish. The variously shaped monstrosities and 

colour aberrants were freaks but they became very fashionable at that time and still are. 

Domesticated goldfish monstrosities were first exported from China to Japan and much later 

to Europe and around the world. More recently other species have been domesticated by 

aquarists, such as the guppy Poecilia reticulata or the neon tetra Paracheirodon innesi. Other 

fishes kept as ornamentals, like swordtails Xiphophorus hellerii and platies Xiphophorus maculatus, 

the discus and angelfishes (Cichlidae), as well as those cultured for food like the rainbow trout 

Oncorhynchus mykiss, channel catfish Ictalurus punctatus or sturgeons (Acipenseridae) are merely 

exploited captives. # 2004 The Fisheries Society of the British Isles 

Key words: common carp; exploited captives; goldfish; guppy; neon tetra; rainbow trout. 

INTRODUCTION 

BACKGROUND AND CRITERIA FOR ANIMAL DOMESTICATION 

According to Clutton-Brock (1999) ‘a domestic animal can be defined as one 

that has been bred in captivity for purposes of economic profit to a human 

*The thirteenth J.W. Jones Lecture 

Tel.: þ1 519 843 2570; fax: þ1 519 767 1656; email: ebalon@uoguelph.ca 

# 2004 The Fisheries Society of the British Isles 

1

2 E. K. BALON 

community that maintains total control over its breeding, organization of 

territory, and food supply’. This definition of domestication applies also to 

fishes subject to some amendments (Diamond, 2002). Definitive answers to 

the questions of where and how organisms were first domesticated are rare 

(Zeuner, 1963; Isaac, 1970) because most of these domestications happened as 

early as the Neolithic (c. 14 000 years ago). In a truly domesticated organism, 

(a) the individual is valued and kept for a specific purpose, (b) its breeding 

is subject to human control, (c) its behaviour is different from that of the wild 

ancestor, (d) its morphology and physiology exhibit variations never seen in 

the wild and (e) some individuals at least would not survive without human 

protection. 

The best known examples of domestication are the transformations of wolves 

into breeds of dog, aurochs into cattle, guanaco into llama and alpaca, wild 

boar into domestic swine, Przewalski’s wild horse into domestic horses, and red 

jungle fowl into domestic chickens. Dependence is the most important among 

the more obvious criteria as ‘all domesticated animals depend for their day-today 

survival upon their owners’ (Livingston, 1994). 

Dependence also applies to animals in the first step of domestication, which 

Clutton-Brock (1999) calls exploited captives. Many attempts at domestication 

remain only at that level including most fishes cultured for food or ornament, 

especially where wild individuals are occasionally brought into the stock for 

breeding in captivity. For a fish to be a true domesticate criteria (d) and (e) 

above must apply. Some of these variants in colour and form would not survive 

without human protection. The original criterion of economic profit may only 

partially apply to domesticates when kept as pets. 

DOMESTICATION OF FISHES 

Fishes were domesticated much later than many other animals and plants, 

now fixed by Diamond (2002) at c. 10 500 years ago. Only the wild common 

carp Cyprinus carpio L., acquired c. 2000 years ago by the Romans in southcentral 

Europe (Balon, 1974, 1995a, b), and the goldfish Carassius auratus (L.) 

selected and released into ‘ponds of mercy’ c. 1000 years ago in China (Chen, 

1956) qualify as ‘true domesticates’. 

Common carp and goldfish both started to be changed into true domesticated 

animals in early medieval times, although in different parts of the world and for 

different purposes (Chen, 1956; Balon, 1969, 1974; Hoffmann, 1995a). In 

contrast to the extinct aurochs, ancestors of domestic cattle or the nearly extinct 

Przewalski horse, only the wild common carp may be close to extinction to be 

replaced in the wild by feral progeny of domesticated forms. 

The loss of the ancestral wild forms of any domesticated organism is a 

loss of epigenetic potential (Balon, 2002). Clutton-Brock (1999) believes ‘that 

animals bred under domestication evolve into new species as a result of reproductive 

isolation from their wild progenitors’. Domesticated animals released into 

the wild to fend for themselves usually perform poorly and, while some revert to 

resemble their wild ancestors in time, the process is nearly always incomplete. 

All such feral forms remain poor facsimiles of their progenitors (Livingston, 

1994). 

# 2004 The Fisheries Society of the British Isles, Journal of Fish Biology 2004, 65 (Supplement A), 1–27

OLDEST FISH DOMESTICATES 3 

Exploited captives are little changed from their wild ancestral form and can 

usually be returned to the wild. Animals become domesticated when they 

change form, function, colour and behaviour and often only partially resemble 

their wild ancestors, and survive poorly as feral forms if returned to the wild 

without human protection. 

A few species have been domesticated by the aquarium industry, including 

guppy Poecilia reticulata Peters, swordtails Xiphophorus hellerii Heckel, platies 

Xiphophorus maculatus (Gu¨nther), mollies Poecilia sp., neon tetra Paracheirodon 

innesi (Myers), discus and angelfishes (Cichlidae) for ornamental and zebrafish 

Danio rerio (Hamilton) for research, although some of these are of questionable 

status. Fishes reared in captivity for human food, including rainbow 

trout Oncorhynchus mykiss (Walbaum), channel catfish Ictalurus punctatus 

(Rafinesque), salmonids, Chinese and Indian pelagic spawning carps 

(Cyprinidae), tilapias (Cichlidae) and sturgeons (Acipenseridae), are exploited 

captives. Some of these do have variants, such as albino strains, that would 

qualify them as domesticated. 

THE COMMON CARP 

Common carp always have two external barbels on each side of the upper 

jaw; goldfish have none. The body cavity of goldfish is lined with a dark brown 

or black pigmented peritoneum, whereas the peritoneum of the common carp is 

unpigmented. The Chinese name for carp includes several cultured genera and 

species, which is still confusing authors working on the origin of common carp 

domestication. The frequent claims that domesticated common carp came to 

Europe from China is based on past misidentification of the two species and on 

the continued repetition of this misconception (Leonhardt, 1906; Rudzin´ski, 

1961; Tamura, 1961; Hickling, 1962; Steffens, 1967, 1980; Borgese, 1980; Liu, 

2003). 

Details on the origins of the common carp have already been given by Balon 

(1974): ‘The latest, Chinese study . . . . . . on pond culture stated that ‘‘thanks to 

the creative efforts of the Chinese people for many generations, breeding of the 

carp in this country has proceeded successfully for >2000 years. From China 

the breeding of this fish spread all over the world. From Asia the rearing of the 

carp spread to Europe and later to America, Australia, and Africa’’.’ Now it is 

time to put this myth to rest. 

Kwai Sin Chak Shik, a book written during the Sung Dynasty in A.D. 1243, 

describes how ‘carp’ fry were transported in bamboo baskets in much the same 

way as they are transported and traded today. Fry were collected in rivers and 

reared in ponds as recorded in A Complete Book of Agriculture, written in A.D. 

1639. But if the fish fry were collected in rivers, it is doubtful whether they were 

C. carpio, which is not a riverine pelagic spawner. They were more likely the 

young of grass carp Ctenopharyngodon idella (Valenciennes), silver carp 

Hypophthalmichthys molitrix (Valenciennes), bighead carp Aristichthys nobilis 

(Richardson) and black carp Mylopharyngodon piceus (Richardson), which are 

true riverine pelagic spawners and cannot be bred in ponds or rice paddies. 

Furthermore, no domesticated forms of common carp, like the mirror or 

leather, or even the high-bodied forms (Pokorny´ et al., 1995), were ever reported 


4 E. K. BALON 

from China prior to known introductions from Europe (Wohlfarth, 1986). 

Furthermore, Chinese carp resemble feral carp, in most of the traits in which 

the races differ than do the European fish (Wohlfarth, 1984). Chinese common 

carp should be regarded as a semi-domesticated animal, since wild fish are 

frequently added to breeding stocks. Therefore, the view that common carp 

were domesticated in China >2000 years ago is a myth. In China the common 

carp remains an exploited captive or a feral form of the introduced European 

domesticated strains. 

Even if the common carp had been domesticated independently in China 

much earlier than in Europe, the European domestic forms were later 

introduced to eastern Asia (Basavaraju et al., 2003). It is unlikely that any 

domesticated carp remain in China or Japan that are pure-bred descendants 

of fish originally domesticated in these countries. Common carp were known 

and used in the fish culture of western Europe well before any direct European 

contact with China (Hoffmann, 1995a). The fish culture methods used in the 

west since 1200, was based on dedicated ponds kept permanently filled for 

several years during the growth of a single age class and then dried for a time 

after harvest. This is the reverse of the fish culture integrated with paddy rice 

cultivation practiced in the east. 

Taxonomy 

Cyprinus carpio is taxonomically a most confusing species. According to the 

latest review by Barusˇ et al. (2002) it remains without a holotype and was 

described ‘using specimens from pond culture, designating Europe as the terra 

typica. Presuming that all domestic European forms originated as wild carps from 

the Danube, specimens from the Danube River can be considered as typical’. 

Most of the confusion comes from giving quasi taxonomic names to feral 

specimens or populations. For example, Wu (1977) lists many, Zhou & Chu 

(1986) list 12 species from Yunan Province, China, and Barusˇ et al. (2002) cite 

>30 synonyms and over 10 sub-species, varieties and morphs from across its 

range. This compares with those given by Kottelat (1997) of c. 15 sub-species 

and eight varieties and morphs. Using the best data available, Barusˇ et al. 

(2002,) concluded that only three sub-species of C. carpio can be recognized: 

(1) the European and central Asian common wild carp, Cyprinus carpio carpio 

L., (2) the east Asian common wild carp, Cyprinus carpio haematopterus 

(Temminck & Schlegel) and (3) the south-east Asian wild carp, C. carpio 

viridiviolaceus Lacepède. None of the data are convincing, nor is the claimed 

occurrence of C. c. haematopterus throughout east Asia from the Amur River 

across China, Korea and Japan. Cyprinus haematopterus was described as a 

separate species from the area of Nagasaki, Japan by Temminck & Schlegel in 

1845. 

More than 200 years earlier, the first colonies of Portuguese and Dutch were 

established at Nagasaki. Common carp culture was by then widespread in the 

European homelands of these colonists and it is, therefore, possible that the 

common carp was introduced by them to Japan (Balon, 1995b). Tableware 

decorated with common carp images once belonging to these early Dutch 

colonists can be seen on display in a Nagasaki Museum (Balon, 1995b). While 

C. c. haematopterus became the nominal sub-species for China, it is possible that 


these were feral descendants of domesticated European fish that escaped or 

were introduced in Japan and possibly China. This taxonomic confusion may be 

resolved by molecular markers but it would be more prudent to consider the 

wild ancestor of the common carp as a single species, Cyprinus carpio, widely 

distributed from the Danube to the Amur rivers, as feral forms elsewhere and as 

naturalized wherever suitable conditions prevail (Lever, 1996). 

Wild common carp, the ancestor of the domesticated forms 

All data so far point to wild common carp from the Danube River as the 

most ancestral form that was initially kept as exploited captives and later 

domesticated (Fig. 1). The inland delta of this river has been badly degraded 

by the Gabčikovo Dam (Balon & Holcˇı´k, 1998, 1999), and common carp 

remains, found in excavations of Roman settlements in this area confirm that 

the fish was already of interest at that time (Balon, 1995b). Therefore, the 

present paper will focus entirely on describing the wild common carp from 

this area before it was extirpated or replaced by feral forms. 

Around 1955 large schools of the wild common carp were still spawning every 

year on inundated meadows along the Slovak part of the Danube and Lesser 

Danube (Misˇı´k, 1957; Be´l, 1962). Detailed studies of these fish were undertaken 

(a) 

(b) 


FIG. 1. A wild common carp (a) and its feral form (b) from the Danube delta in 1900 (from Antipa, 

1909). 


6 E. K. BALON 

on the assumption that these fish are the direct descendants of the much larger 

schools of wild common carp that the Celts and Romans exploited in the same 

area c. 2000 years earlier (Balon & Misˇı´k, 1956; Balon, 1957, 1958a, b; Misˇı´k, 

1958). These studies were already reviewed (Fig. 2) and the life history and 

ecology of the wild common carp compiled in detail by Balon (1974, 1995a, 

b) and Barusˇ et al. (2002). 

Lucullan demands, Pannonia and the first carp captives 

Two significant historical events coincided and contributed to the development 

of wild common carp from the inland delta of the Danube as a major food 

source and as an exploited captive. 

In the last years B.C. and in the first and second century A.D., the Romans 

developed a luxurious lifestyle with an equally elaborate cuisine that led to the 

(a) 

(b) 

(c) 

1 

5 cm 

4 

3 

20 

5 

P 

12 

26 

1513 

28 

27 

V 

D 

14 

21 18 

FIG. 2. (a) Wild Cyprinus carpio carpio from the Danube River arm near Medved’ov, captured 21 May 

1954, (b) scheme for counting the meristic characters and (c) scheme for mensural characters 

(rearranged from Misˇı´k, 1958). 

# 2004 The Fisheries Society of the British Isles, Journal of Fish Biology 2004, 65 (Supplement A), 1–27 

16 

17 

19 

29 

A 

22 

24 

2 

C 

23 

25

importation of foreign foods (Friedla¨nder, 1936). Sergius Orata, Cicero’s 

teacher, devised special reservoirs where fishes for the kitchen were stored. 

These reservoirs called piscinae soon became very fashionable, for they ensured 

a permanent supply of a variety of fresh fishes independent of weather conditions 

and fishing success (Zeuner, 1963). Individuals such as Lucinius 

Muraena adopted fish rearing in piscinae, and added freshwater fishes to the 

initially stored marine ones (Pliny the Elder in translation 1958–1962). The 

patricians soon competed in building such piscinae; consul Lucullus (75 B.C.), 

whose reputation as a gourmet was well-known, dug a trench through a hill 

near Naples to bring water to his ponds that cost more than his villa. Varro 

(116–27 B.C.; 1912) and Columella (–50 A.D.; 1941–1968) claimed that the 

Roman patricians preferred seawater ponds, and that freshwater ponds (piscinae 

dulces) were considered plebeian, but the documented prejudice is proof of their 

existence. Also, the existence of plebeian owned freshwater fish ponds may 

signify that the desire to transport a wild common carp from the Danube to 

patrician piscinae in Italy was applicable also to plebeian troops and the 

accompanying tradesmen and artisans (Hoffmann, 1995a). 

In the first years AD, the Roman Empire expanded its northern boundary 

beyond the barrier of the Alps right up to the shores of the Danube River 

(Sitwell, 1981). The northernmost province, Pannonia, faced formidable forces 

of Celts and Germans across the Danube and the Empire had to establish a 

strong military presence (Fig. 3). In the second century, the comparatively short 

stretch of river between Vienna and Budapest, c. 150 miles long, required no less 

than four legions to guard it. At least 20 000 fighting legionnaires were accompanied 

by supporting troops, wives, mistresses, children, slaves and tradesmen 

to a total of >100 000 Romans. These, together with the indigenous communities 

formed a human population large enough to establish a carp-eating tradition, if 

wild carp was, at least seasonally, the most abundant and easily caught fish in 

the area. 

Archeological excavations in the region showed abundant fish remains that 

revealed that the bones of the wild common carp dominated all other fish bones 

(K. Hensel, pers. comm.). This constitutes the first direct evidence that Romans 

Alcmona 

Augusta 

Vindelicum 

Isara 

Abodiacum Pons Aoni 

Veldidena 

Naba 

Castra Regina 

Castra Batava 

Aenus 

Danuvius Lauriacum 


Vindobona 

Eburodunum 

Arlape Cetium Carnumtum 

Aquae 

Scarbantia 

Virunum 

Savaria 

Marus 

Arrabona 

Peiso Piso 

Danuvius minor 

Danuvius 

Mogentiana 

Amber Road 

Duria 

Laugaricio 

lza 

Granua 

Brigetio 

Aquincum 

Vetus Salina 

Lussonium 

Vistula 

Pathisus 

River (flumen, fluvius) 

Boundary of Empire 

Cities and towns 

Legionary fortress (castra) 

Fort (castella) 

Wall (vallum) 

Road (via) 

Amber (Road) 

0 200 km 

FIG. 3. The Roman Danube frontier of Pannonia with the Amber Road crossing the river near 

Carnuntum, and the major floodplain from Peiso Piso (modern Bratislava) to Celamantia (Izˇa). 


8 E. K. BALON 

knew the wild common carp of the Danube. It is most likely, therefore, that the 

Romans, while living on the Danube in Pannonia not only consumed common 

carp but also transported them alive to other Roman provinces, because of the 

high fashion of that time to keep fishes in piscinae. The many thousand soldiers 

after returning to Italy probably demanded their customary fish to be served 

and so influenced the importation of common carp. After all, the common carp 

was the largest, tastiest fish that would have survived the rigours of transport. 

There is nearly no written or archeological evidence of common carp being 

kept in piscinae or vivaria in Rome 2000 years ago (Zeuner, 1963; Hoffmann, 

1995a). Written records from this time are generally scarce and information is 

lacking even on activities that are known to have existed at that time. Walton 

(1676) quoting Jovius, who was probably Paolo Govio, a papal physician, who 

wrote about fishes on the Roman market in 1524 and recorded that ‘carps of 

more than fifty pounds weight’ were to be found in Lake Lurian in Italy. 

Probably these early introductions of wild common carp west of the middle 

Danube were using it only as an exploited captive fish. 

The history of common carp domestication 

The early assumption that common carp keeping was continued in monastery 

ponds after the collapse of the Roman Empire and the establishment of Christianity 

is pure conjecture for the early middle ages up to the 10th century 

(Hoffmann, 1995a). This is based on two logical assumptions; firstly, that 

monasteries began to be founded in the 5th and 6th centuries and soon gained 

land and farms; secondly, that Christianity introduced >180 fasting days per 

year that monks, nuns and priests had to respect. 

During fasting days the only animal protein that could be eaten was from 

aquatic animals such as shellfish and fishes, although unborn rabbits called 

laurices were also acceptable in some areas. This links these customs directly to 

Roman tradition (Balon, 1995b). The only evidence that the Roman taste for 

eating common carp continued, however, is from Cassiodorus (490–585 A.D.; 

1626) who ordered wild common carp to be delivered from the Danube to Italy 

for the table of King Theoderic. 

Albertus Magnus was the first to write about breeding common carp in ponds 

in the 1260s (Magnus 1193–1280; 1861), however, he may compete for this 

privilege with the Count of Champagne who was breeding common carp in 

1258 according to Hoffmann (1994). Ponds with muddy bottoms have been 

built since the 10th century (Hoffmann, 1985; Andreska, 1987), and wild 

common carp became established in them in the twelfth century at the latest, 

judging from the relatively fast process of domestication as reported by Tuča 

(1958), Bastl (1961) and Misˇı´k & Tuča (1965). So it seems that transition of 

common carp from exploited captives to truly domesticated animals took place 

in the 12th century. 

In modern common carp culture, to build a fishpond, not just a holding tank, 

is to create a new aquatic habitat. Active construction of ponds for this purpose 

got under way in the 11th century and increased rapidly in the 12th and 13th 

century (Hoffmann, 1995a). Initially these ponds were stocked with several local 

species but common carp soon proved to be the best choice for such man-made 

habitats. Hoffmann (1994, 1995a) concluded that ‘there were three phases in the 



medieval diffusion of common carp to the west and north from an epicentre at 

the north-west margin of its native range in Pannonia (the piedmont section of 

the Danube, Balon, 1967). The first phase, between the 7th and 11th centuries, 

principally carried the fish up the Danube and into some west-flowing tributaries 

of the middle Rhine. Some remains also indicate common carp in waters 

north of the Danube. In a second phase (12th to the early 14th century) 

common carp radiated across most of the economic and cultural heartland of 

medieval north-western Europe, from the lower Rhine-Maas region south to the 

Paris basin and Burgundy. A final stage, probably occurring after the economic 

shift of the mid-14th century, extended common carp into the outer periphery 

of the west, an arc from south-western France through England and southern 

Scandinavia into east central Europe. Though common carp are recorded in 

Italy at the end of the Middle Ages, the diffusion pattern at no time accords 

with radiation from there’. 

In time more written evidence appeared and by the early 14th century 

common carp culture was established (Pucher, 1987; Hoffmann, 1995b, 2002). 

Soon pond common carp management was described in its entirety in the 

famous Latin manual by Jan Dubravius (1547) that was written in 1535 to 

1540, and by Olbrycht Strumien´ski (1573) who reported on the Czech and 

Polish techniques of that time (Strojnowski, 1609; Sˇusta, 1889; Inglot & 

Nyrek, 1960; Szczygielski, 1967a, b; Berka, 1986). 

The ‘golden age’ of pond construction for rearing of common carp 

reappeared after the wars and skirmishes of the 15th century ended. It has 

been estimated that >25 000 common carp ponds were built in Bohemia and 

Moravia alone towards the end of the 15th and the beginning of the 16th 

century. A special guild of mobile and rogue pond builders flourished. During 

these 50 years, c. 500 ponds per year, or two ponds per day, were constructed 

(Andreska, 1987). Production of the common carp for food became the most 

profitable branch of agriculture (Andreska, 1997; Guziur et al., 2003). Various 

wars and a decline in the price of common carp later led to a dramatic reduction 

in the number and area of these ponds, which declined from a total of 

180 000 ha to c. 51 000 ha. Rozˇemberg pond, which was built in the 16th century 

and covers 711 ha is one of the 34 largest. It still operates on the 2 to 3 year 

cycle of common carp production (Kourzil & Guziur, 2004). 

Dubravius (1547) recommended a system of several specialized ponds for the 

production of marketable common carp. Smallest of all was the spawning pond 

with a grassy bottom simulating shallow, freshly flooded meadows. Selected 

parent fish were released in groups into it. Soon after spawning, the early 

juveniles (Balon, 1999) were flushed into nursery ponds, and 1 year later were 

transferred into growing ponds. Ponds were left dry in rotation, treated with 

lime and various chemical and natural fertilizers in order to enhance the natural 

food production. Some consider that T. Dubisch, an illiterate Slovak fish master 

(Morcinek, 1909), invented the system of transferring the fish from one growing 

pond to the next while gradually lowering their density (Billard, 1999; Guziur 

et al., 2003). While the production of young common carp in spawning ponds 

was recommended for every pond system, this was later, at least partly, replaced 

by artificial stripping of gametes under hatchery conditions (Woynarovich, 

1962; Billard et al., 1995). The growth of the common carp was enhanced by 


10 E. K. BALON 

ever better formulations of supplementary food, the system of sequential, 

specially fertilized ponds remained. The production increased from an initial 

40 to >450 kg ha 1 (Cˇı´tek et al., 1998), with an ever reducing duration for 

growing marketable common carp from 5–6 to 2–4 years (Fig. 4). 

Sooner or later different forms of domestic common carp appeared in the 

various pond systems. When the wild common carp was introduced into a pond 

system it naturally started to change its torpedo-shaped body into a deep, 

laterally compressed and hunchbacked body. Soon individuals appeared that 

did not have regular, geometrical arrangement of scales but showed severe 

irregularities, scale reductions or even complete scalelessness. These variations 

soon became a foundation for artificial selection. Ultimately, domesticated 

common carp are represented by a variety of forms, such as scaled carp, line 

carp, mirror carp, and leather or naked carps (Brylin´ska, 1986; Gorda et al., 

1995; Pokorny´ et al., 1995). 

The domesticated common carp not only changed its external shape, scalation 

and colour, but also underwent internal and physiological changes. Detailed 

reviews were published elsewhere by Balon (1974, 1995a, b) and Barusˇ et al. 

(2002). Briefly, the mouth gape, the first character that Rudzin´ski (1961) and 

Steffens (1964) used for clear distinction between wild and domesticated common 

carp, was much smaller in the wild than in the domesticated varieties 

(Table I). Even more pronounced differences are encountered among the 

mouth-gape indices calculated by dividing 10 times the mouth gape in cm 2 by 

length of head in cm: 4 46 to 5 57 for the wild common carp and 8 12 for 

domesticated fish (Steffens, 1964). Both Rudzin´ski (1961) and Steffens (1964) 

FIG. 4. (a), (b) Common carp harvest in southern Bohemia near Hluboka and Vodnˇany. (c), (d) harvested 

domesticated common carp (early November 1998). 


TABLE I. Mouth gape and intestine length indices for the wild common carp and 

domesticated common carp: o l 1 ,10 mouth gape in cm 2 per standard length (L S)in 

cm; o m 1 ,10 mouth gape in cm 2 per mass in g; o lc 1 ,10 mouth gape in cm 2 per 

length of head in cm; gut l 1 , length of intestine in cm per 1 cm of L S; gut m 1 , length of 

intestine in cm per 10 g of wet mass 

Index o l 1 


om 1 

olc 1 

gut l 1 

gut m 1 

Wild common carp 1 14 

With supplementary diet 1 09 0 10 4 46 2 11 3 02 

With natural diet 1 24 0 08 5 27 

Domesticated carp 1 91 

With supplementary diet 2 00 0 08 8 12 2 64 2 25 

With natural diet 2 30 0 08 8 12 

considered the enlargement of the mouth in domesticates to be caused by 

changes in diet, and possibly the result of artificial selection. Domesticated 

common carp selected to utilize supplementary food, grew better in ponds 

when man-made food was added (Sibbing, 1988). Studies on the wild common 

carp in ponds (Rudzin´ski, 1961; Leszczyn´ska & Biniakowski, 1967) proved that 

wild common carp progeny was better suited for stocking natural riverine 

habitats than were domesticated varieties. 

Rudzin´ski (1961) found and Steffens (1964) confirmed that the intestine of 

the wild common carp was 15 to 25% shorter than that of a domesticated carp. 

The index calculated by dividing the length of the intestine by standard length 

(L S) in cm was 2 11 for the wild common carp and 2 64 for the domesticated 

fish. Longer intestines in domesticated fish are probably due to utilization of 

vegetable food not normally consumed by wild common carp. Both authors 

also found, when comparing the deeper body of domesticated common carp 

with the cylindrical one of wild common carp, that the calculated muscle mass 

was the same in both, although it appears to be greater in the domesticated fish. 

This means that the dressed mass of the domesticated fish is not larger in spite 

of the different body proportions. Also, wild common carp has both chambers 

of the swimbladder of similar size, whereas the domesticated fish has the 

anterior chamber always much larger and the posterior chamber smaller; the 

proportion (%) as given by Steffens (1980) is 61 : 39 for the wild and 90 : 10 for 

the domesticate. This may be related to the relative greater mass of the head in a 

domesticated common carp. 

The greater strength of wild common carp is supported by some of its physiological 

attributes (Steffens, 1964). Wild common carp has 18 to 19% more erythrocytes 

and haemoglobin than domesticated fish and blood sugar level is 16 to 26% 

higher in the former. Also, wild common carp has a much lower water content in 

muscles and liver than domesticated fish. Furthermore, wild common carp has 

greater fat content in individual organs, more glycogen in the liver, and more 

vitamin A in the eyes, intestine and liver. In addition its muscles are better vascularized 

and do not fatigue as quickly as do those of domesticated common carp. 

As in some exploited captive salmonids (Balon, 1980, 2004) there are some 

indications that domesticated common carp usually produce eggs with less yolk 



and as a consequence more altricial progeny than the ancestral wild form 

(Kryzhanovsky et al., 1951; Balon, 1958a; Penˇáz, 1995). 

Specific forms of the domesticated common carp are created and maintained 

through constant artificial selection in nearly every pond culture system. Standardized 

breeds were developed and new forms continue to be constantly tested 

in some institutes, such as the Fish Culture Research Institute at Szarvas, 

Hungary, the Institute of Ichthyobiology and Pond Culture of the Polish 

Academy of Sciences at Gołysz, Poland, and the Research Institute of Fishery 

and Hydrobiology of the Southern Bohemian University at Vodnˇany, Czech 

Republic (Billard & Gall, 1995; Gorda et al., 1995; Pokorny´ et al., 1995; 

Flajsˇhans, 1996; Varadi et al., 2002; Guziur et al., 2003). The common carp is 

considered a delicacy in Europe, and is served particularly at Christmas (Balon, 

1966). The fish, however, is appreciated throughout the year and is served in 

special restaurants. A dedicated cookbook containing >84 recipes for domesticated 

common carp dishes was written around 1927 (Vanˇha, 1993). 

The emergence of koi – the most expensive fish in the world 

Common carp was traditionally reared for human food in small terrace ponds 

alternating with rice paddies between Nagaoka and Ojiya cities of Niigata 

Prefecture, c. 280 km west of Tokyo, Japan. The area is known to have one of 

the highest snowfalls in the world and the whole area is under at least 6 m of 

snow for c. 6 months. These unusual conditions may have contributed to the 

more frequent occurrence of colour aberrations in common carp cultured here. 

The high incidence of colour aberrations may be caused by melatonin production 

during the prolonged life in total darkness under the cover of deep snow. 

In the past, this mountainous area had only few roads and the local farmers 

were often isolated during winter months. It is assumed that, c. 180 years ago, 

coloured aberrants of the common carp began to be selected and crossbred here 

(Kataoka, 1989; Kuroki, 1990). There is, however, little evidence to support 

such early appearance of ‘nishikigoi’. Plate 16 in volume one of Illustrations of 

Japanese Aquatic Plants and Animals, published in 1931, shows some coloured 

carps but no modern ‘nishikigoi’ (Ishikawa et al., 1931; Balon, 1995b). Consequently, 

although coloured common carp is recorded from >70 years ago, 

commercial scale selection of true ‘nishikigoi’ (koi) probably started only in 

the 1950s. By then, increased living standards in Japan and elsewhere allowed 

more people to afford garden ponds. Especially in Japan, where land is more 

precious than in most other parts of the world and Buddhism favours a love of 

animals, garden pools were used fully for decoration and ‘swimming flowers’ 

were a logical progression in the development of the miniature garden. Even in 

the early days of koi popularity, Amano (1968) recorded an annual production 

of some 10 million fish, amounting to ‘1 000 000 000 yen worth’. 

Koi, as they are known today (Axelrod et al., 1996), are colour aberrations 

C. carpio. They probably originated from domesticated common carp or its feral 

forms, and from some more recent imports from Europe (doitsu koi). When 

colour aberrations started to appear more frequently and demands for them 

increased, production and crossbreeding intensified and their progeny were 

severely culled, leaving only the best coloured individuals one out of 10 000 or 

more, for further breeding. The first modern ‘nishikigoi’ come from 30 hamlets 



FIG. 5. (a) The koi ponds at Yamakoshi, Japan, (b) selected 3 year-old individuals at the breeder’s pond, 

(c) in the garden of a rich hobbyist in Kyoto and (d) at the wholesaler in Ojiya, Niigata Prefecture, 

Japan. 

of Yamakoshi in the Niigata Prefecture (Fig. 5) where 87% of the 906 families 

became ‘fancy carp’ producers (Amano, 1971). 

Today, the Japanese recognize c. 15 basic colour aberrants of the common 

carp that are called koi. Each of these aberrants has many varieties (Phipps, 

1989; Kuroki, 1990). The standard is for the carp body to remain cylindrical 

and the colours to be viewed from above. Koi with elongated fins and scale or 

body deformation is not accepted in Japan but appears frequently from producers 

in Israel and elsewhere. The colours resulting from crossing the severely 

culled offspring are impossible to predict, even with the most careful parent 

selection (Kataoka, 1989). After several cullings only the most promising and 



appealing colours are retained and from these grow numerous beautiful 

‘nishikigoi’ (Balon, 1995b). The winners of shows, however, are but a few 

unique individuals selected from many hundreds of thousands. In Japan the 

favourites are ‘koˆhaku’ (red and white koi), ‘taisho-sanke’ (red, black and white) 

and ‘showa sanshoku’ (a black koi with white and red imposed). As no two 

specimens are ever alike, the grand champions of modern shows commonly sell 

for considerable amounts of money. 

THE GOLDFISH 

The goldfish has long been the most domesticated fish and now seems to be 

better adapted to confinement than to a ‘free life’ (Taylor, 1884). It has been an 

animal of choice for laboratory experiments in physiology (Ostrander, 2000) 

and is particularly susceptible to the effects of alcohol (Hartston, 1987) and so is 

a good model for research on the effects of alcohol. 

Goldfish are often confused with the crucian carp Carassius carassius (L.), 

Prussian carp Carassius auratus gibelio (Bloch) or even common carp 

(Szczerbowski & Szczerbowski, 1996). There is no reason for this confusion as 

goldfish have no barbels while common carp do. The crucian carp differs from 

the goldfish by (a) having 23 to 33 gill rakers on the first branchial arch while 

goldfish have 37 to 53, (b) having 28 to 29 fine denticles on the posterior edge of 

the last unbranched spiny ray of the dorsal fin while goldfish have 10 to 11 

irregular denticles, (c) by having a light peritoneum while goldfish has a black 

peritoneum and (d) by having a blackish spot at the base of the caudal fin while 

goldfish have none. Body colour of the crucian carp is usually coppery gold 

while that of the wild goldfish is silvery-grey. In spite of these differences, most 

FAO statistics erroneously list goldfish as crucian carp, but crucian carp is 

rarely used in aquaculture and is quite infrequent. The same mistake can be 

found in some aquarium literature (Matsui & Axelrod, 1991), and even in the 

newest encyclopedia (Liu, 2003) although in the first edition all was correct 

(Grzimek & Ladiges, 1973). 

The crucian carp occurs in two morphs: the precocial, fast growing and deep 

bodied and the altricial, dwarfed morpha humilis (Szczerbowski & Szczerbowski, 

1996, 2002). The fish occurs in most of Eurasia, except in the north-western 

and south-western parts of Europe, China, Korea and Japan. Expanding populations 

of Prussian carp replaced C. carassius in many locations. Kottelat (1997) 

recognized C. gibelio (Bloch) as a separate species. Prussian carp is silvery in 

colour, and is a gynogenetic, all female, triploid sub-species (Pelz, 1987; 

Halačka & Lusková, 2000) that lives in the peripheral areas of the distribution 

of the nominal wild goldfish. The Prussian carp with both sexes present, occurs 

only close to the distribution of the goldfish. The wild goldfish is native to 

China, the southern Amur River basin and Korea. It was introduced to Japan 

from China between 1502 and 1620 (Matsui, 1934). Available data show that 

the systematics of the genus Carassius in East Asia is confusing (Hensel, 1971; 

Kawanabe & Mizuno, 1989), and Kottelat (1997) recognizes at least five genetically 

and morphologically distinct stocks in Japan, which are considered distinct 

species or sub-species. 



All three most common Carassius species, the crucian carp, goldfish and 

Prussian carp, reach an average size of c. 35 cm and a body mass of 1 kg. 

Crucian carp and goldfish were distinguished during the Sung Dynasty after 

968 A.D., claims Szczerbowski (2002), but since these two species do not 

naturally occur in sympatry this claim may be without foundation. The 

goldfish, was introduced to Europe and elsewhere from China as an already 

domesticated monstrosity. ‘Valenciennes (1842: 108) summarized the history of 

the introduction of Carassius auratus to Europe [writes Kottelat, 1997, p. 52]. 

He states that some authors (but does not say which ones) consider that it was 

introduced to Europe as early as 1611 or 1691 and that Yarrell reports that it 

had been introduced by the Portuguese from Java (. . .) to South Africa and 

from there to Lisboa. The first introduction to England dates to 1691 (Pennant, 

1812: 490) and to France to 1755 (Hervey & Hems, 1968. . .). It was bred in 

northern Europe for the first time in Holland in 1728 according to Sterba (1987: 

272).’ Before this, domesticated goldfish had to come by boat from China and 

must have been very expensive. As such their owners would guard them from 

release into natural waters. Invasions from aquaria and garden pools into all 

suitable habitats around the world came later. 

Purnell’s Encyclopedia of Animal Life (Burton, 1969) makes the following 

statement with regard to goldfish: ‘The more fancy breeds of goldfish are freaks, 

no matter how attractive some of them may look. To recite their names is 

enough to make this point: veiltail, eggfish, telescope, calico, celestial, lionhead, 

tumbler, comet or meteor and pearl scale. There are also the water bubble eye, 

blue fish, brown fish, brocade, pompon and fantail and many others. Some 

breeds are monstrosities rather than freaks’. 

Wild goldfish, the ancestor of the domesticated monstrosities 

The wild goldfish has a deep body, silver-grey or olive-green colouration and 

thick caudal peduncle. The body is covered by large cycloid scales, and the 

upper edge of the dorsal fin is slightly concave. Its feral forms are mostly 

slimmer and smaller, much like the altricial morpha humilis common in the 

crucian carp. The low and round body form, however, is also common in the 

wild goldfish and in the silver-greyfish caught for food by Chinese fishermen. 

The domesticated goldfish ancestor, ‘chi’, has a relatively short snout, 25 to 

37% of head length, small, protrusible mouth, and 0 4–4 0 pharyngeal teeth. 

The morphometric and meristic characters differ little between the wild goldfish 

from China and the Prussian carp from the Amur River (Table II), although the 

data from various operators and times may be imprecise. 

Gynogenetic forms are rarely found in the goldfish. Originally, the number of 

chromosomes was reported to be 94 (2n) in the goldfish (Makino, 1939), and 

bisexual populations of the Prussian carp (Cherfas, 1966). In Vasilyev’s (1985) 

review, however, goldfish had 100 (2n) chromosomes and in Chiarelli et al. 

(1969) 104 (2n) chromosomes. The triploid forms that reproduce by gynogenesis 

have 141 to 160 (3n) chromosomes (Penˇa´z et al., 1979). The goldfish is the only 

cyprinid that has morphologically differentiated sex chromosomes (Szczerbowski, 

2002). 

The goldfish is a portional spawner and releases two to three batches of eggs at 

each mating at 8 to 10 day intervals. It deposits adhesive eggs on grass and fine 



TABLE II. Morphometric and meristic characters of 73 individuals from China of wild 

Carassius auratus, compared to wild Carassius gibelio from the Amur River (after 

Szczerbowski, 2002) 

Carassius auratus Carassius gibelio 

Per cent of L S 57 mm 118 mm 110 mm 280 mm 

Head length 33 0 29 3 26 8 30 0 

Predorsal distance 52 3 29 3 51 3 52 9 

Body depth 38 8 39 5 41 7 48 7 

Caudal peduncle length 16 7 16 8 17 2 17 9 

Minimum body depth 15 0 15 8 16 1 17 0 

Pectoral origin to pelvic base 21 1 22 1 20 5 21 5 

Pelvic base to anal base 23 2 31 4 – – 

Base length of dorsal fin 35 5 38 1 34 1 39 4 

Base length of anal fin 12 5 12 7 10 5 12 0 

Longest pectoral ray length 19 1 21 6 18 9 20 6 

Longest dorsal fin ray length 18 5 22 7 17 8 19 8 

Longest anal fin ray length 16 2 19 6 15 4 17 6 

Longest pelvic ray length 20 9 23 2 20 7 21 5 

Branched rays of dorsal fin 16–18 15–19 

Branched rays of anal fin 5–6 5–6 

Lateral line (pored scales) 28–30 28–33 

Gill rakers 37–46 39–50 

Vertebrae count 26–31 39–50 

willow-like roots at water temperatures 18–19 C, very much like the common 

carp. Activated eggs of the goldfish are spherical or slightly oblong, 1 05 by 

1 14 mm, transparent with a pale yellow or greyish-green tint. The embryos 

hatch 3 to 8 mm long, still with a large yolk sac and hang on plants shortly before 

swimming and seeking exogenous food (Battle, 1940; Mansueti & Hardy, 1967). 

The wild goldfish is native to lateral waters of rivers and lakes of East and 

Central Asia, with the centre of distribution in China (Chen & Fang, 1999). The 

goldfish has acquired global distribution in temperate and subtropical areas 

from escaped domesticated forms and deliberate releases. It has naturalized, 

feral populations in >20 European countries. Outside of Europe, the goldfish 

has been introduced to South Africa, Madagascar and Mauritius, and is widespread 

in Asia. It was introduced in 1874 to North America and Oceania, and 

from 1856 to 1930, to various countries in South and Central America and the 

Caribbean. A slightly earlier date for North American introduction is given by 

Fuller et al. (1999). In some localities, it occurs in sufficient abundance to be 

marketed as a food fish (Szczerbowski, 2002). 

The goldfish is one of the most resistant fishes to natural environmental 

perturbations as well as to man-made pollutants. It tolerates a wide temperature 

range and high water turbidity. It is the only fish that survives in shallow lakes 

with total oxygen depletion during some years or seasons (as low as 

0 6mgl 1 O2). The goldfish survives within a pH range of 4 5 to 10 5, but 

prefers the optimal range of pH 7 2to84. It can survive c. 2 weeks in water 



at pH 4 5. The goldfish is more tolerant than any other fish to heavy metals or 

organochlorine insecticides (Szczerbowski, 2002). 

The goldfish has been less successful than the common carp in establishing 

itself in larger water bodies, but does best in small shallow lakes heavily overgrown 

by aquatic plants. It grows to much smaller sizes than a common carp. 

Females generally grow larger and live longer than males (Moyle, 2002). As a 

result the male: female sex ratio changes from 1 : 1 in small fish to 0 13–0 16 : 1 

in fish measuring >15 cm total length (L T). Although fish in the wild rarely live 

longer than 6–8 years, maximum ages of 30 years have been recorded in 

aquaria. For example, goldfish were introduced into Killarbey Lake in New 

Brunswick, Canada in 1967 or 1968. Only 7 years later, in 1974, they yielded 

43% of the feral bronze and 57% of variously coloured fish. The largest of them 

measured 34 cm, weighed 1 kg and was 8 years old, having matured at age 2 

years (Hooper & Gilbert, 1978). 

The history of goldfish domestication 

Shisan Chen (1956) has reviewed the history of domestication of the ‘chi’ 

for a second time 30 years after his detailed studies on the fish. He has amassed 

additional evidence on the history of domestication of the goldfish since 

Ting-pong Koh’s (1934) study on that topic. 

The goldfish was created as a domesticated ornamental fish in China. As is 

the case in many other fishes (Balon & Frank, 1953; Balon, 1964), a xanthic 

aberration sometimes appears among the silver-grey ‘chi’, the most common 

food fish in China. These red or yellow aberrations are usual but rare and only 

few reach fishermen because most of them would be eliminated in nature due to 

their conspicuousness. People used to attach a supernatural significance to the 

appearance of the wild golden ‘chi’ and regarded them as sacrosanct but there 

were places, like the Ninghai District, where they were served as food (Chen, 1956). 

Buddhism teaches abstinence from taking the life of any creature and urges 

to do at least one good deed per day. To set free a rare golden ‘chi’ must have 

been a better good deed than the release of a common animal. Hence, ever since 

the beginning of the Sung Dynasty when Governor Ting Yen-tsan of the Duchy 

of Wu-yueh discovered golden and yellow ‘chi’ in a pond outside the city of 

Kiahsing, that pond came to be called ‘Goldfish Pond’. Subsequently, the pond 

became a ‘pond of mercy’. In the goldfish pond were other varieties of fishes 

and turtles besides the golden ‘chi’, which it was forbidden to catch (Chen, 

1956). Other such ponds followed at the pagodas in Hangchow and Nanking, 

where the local monks cared for the goldfish. The records thus document the 

beginning of goldfish accumulation in ponds of mercy at the outset of the Sung 

Dynasty, between 968 and 975 A.D. Until at least the year 1089, however, 

goldfish were not different from wild ‘chi’, were afraid of humans and did 

not take food thrown to them. They were captives exploited for religious 

purposes. 

The documents reviewed by Chen (1956), the Bedside Companion (1214), On 

Names (1241) and Dreams of the Past (1247) explain the fact that the ‘fad’ of the 

ruling class of the time brought into existence a special, fish-breeding trade 

called ‘yu-er-huo’. These breeders learned to feed the goldfish with a kind of red 

animalcula found in putrid water (tubifex worms?) and discovered the secrets of 



propagating goldfish so as to enable them to offer a selection of rare varieties to 

gods on festival days. By 1241 the goldfish had become tamed and used to the 

food offered by humans. In the ponds of mercy, goldfish lived among the 

common ‘chi’, other fishes and turtles, but the private ponds constructed by 

the rich now contained only xanthic goldfish. It was, therefore, easier to breed 

them, cross the preferred individuals, and for the first time produce the golden 

yellow, silver white, and variegated of black and white (tortoiseshell) goldfish. 

This initial domestication began c. 1163 in the goldfish pond of the Te Shou 

Palace, east of Hsin Kung Chiao in the city of Hangchow (Chen, 1956). 

In general, ponds and garden pools were affordable only to landowners. 

The goldfish, now in several colour forms, were first kept only by royalty and 

rich mandarins who constructed more ponds, or rather garden pools, for 

their keeping and breeding. Goldfish were brought to Beijing in 1330, during 

the Yuan Dynasty, and in 1506–1521 appeared in large numbers outside the 

Forbidden City. At that time they were also introduced to Japan. 

Chen (1956) notes the transition from pools to aquarium-like vessels 

from 1276 to 1546. Besides the jade vessels and fancy bowls for keeping 

goldfish, earthen vessels became common and affordable to everyone. After 

1548, rich and poor alike became keepers and breeders of fancy domesticated 

goldfish. 

The creation of ‘monstrosities’ in shape, in addition to colour aberrations, was 

closely related to the keeping of indoor aquaria, which became a very popular hobby 

after 1548. The goldfish became the pet of the masses, a popular pastime in many 

homes everywhere. Aquarium culture enabled close observation and selection of 

individual fish, something hardly possible in garden pools. In A Handbook on the 

Cinnabar Fish Chien-te Chang (1596, after Chen, 1956), beside presenting lists of 

many colour varieties, says: ‘The beauty of the cinnabar fish lies not only in their 

extraordinary brilliance of colour, but also in their tails, patterns, and bodily form, 

which distinguish them from common fish. Whether long or short, the body must 

have the beauty of plumpness as a primary qualification’. 

Selected pairs of the domesticated goldfish 3 to 4 years old started to 

reproduce in aquaria on aquatic plants. After mating and deposition of 

gametes, the parent fish had to be removed from the tank as fast as possible, 

otherwise they would have eaten the eggs. Eggs, embryos and larvae incubated 

in such aquaria were exposed to numerous and various stresses of biological, 

physical and chemical nature. These in turn caused epigenetic alterations in 

their ontogenies some of which led to the development of new ‘monsters’ that 

survived longer thanks to human care. Ultimately, through artificial selection 

some of these ‘monstrosities’ underwent genetic assimilation, and became more 

or less inheritable (Balon, 1990, 2002). 

Unintentional artificial selection and crossbreeding played an enormous role 

in the production of more and more bizarre individuals. For example, most of 

the future red goldfish appear black when young and so severe culling had to be 

applied several times in succession to raise a few ornamental individuals. So, in 

the short space of 97 years, between 1547 and 1643, at the end of the Ming 

Dynasty, new varieties were produced with an increasing degree of variation 

(Chen, 1956). Varieties such as golden helmet, golden saddle, stork’s pearl, 

brocaded back, seven stars, red head-and-tail, purple eyes, snow eyes, the 



three tails, four tail, nine tail, bulging eyes, the bicaudal and the dragon eye 

(Chen, 1956) have became quite common. The double tail was first mentioned 

in 1579 (Linnaeus later described this variety of goldfish as typical Cyprinus 

auratus) and the dragon eye in 1592; by 1596 the short body was already in 

existence, the dragon back without the dorsal fin appeared by 1726, and the 

duck’s-egg fish by 1780 (Chen, 1956). By 1848 goldfish breeders in China were 

already using deliberate artificial selection and obtained the lion-head, goosehead 

and the narial bouquet goldfish. 

ARE THERE ANY OTHER DOMESTICATED FISHES? 

There is no good reason to treat fishes differently from other animals with 

respect to domestication, although ‘fish culturists’ are often tempted to do so 

(Kim et al., 2004). The number of fish species in aquaculture has increased 

rapidly during the last few decades. Some as food fishes, for stocking in the 

wild, to be used as bait, or for the aquarium hobbyist. Commercial fish farming 

is for profit, subsistence fish farming maybe less so, but in both instances the 

enhanced production of fishes is the goal. The vast majority of cultured freshwater 

fishes are raised in ponds, though various other types of culture systems 

are also employed (Stickney, 1986). Both exploited captives or ‘domesticates’ 

can be produced in each such culture system. 

Most fishes in culture are still only exploited captives but a few are on 

the threshold of becoming domesticated, most for the aquarium. Clutton- 

Brock (1999) distinguished between domesticated animals (dogs, cattle, sheep, 

swine and goats) and exploited captives (cats, elephants, camels, llamas, reindeer, 

yaks and water buffaloes). 

The best-known example is the small viviparous guppy whose males attain 3 cm 

standard length (LS) and females c. 5cmLS. Guppies originate from Trinidad and 

Venezuela but were released worldwide for mosquito control. Populations now 

live in Africa, India, North America, Italy and even in the warm springs of 

Hungary and Slovakia. Wild forms are smaller than the domesticated varieties. 

The colouration of the wild guppies varies according to habitat and many 

different coloured populations were described (Frank, 2002). Endler (1980) 

explained the reason for males being much smaller than females and the 

colour polymorphism in P. reticulata. Natural populations in Trinidad and 

north-eastern Venezuela are so polymorphic that no two males are alike. The 

colour patterns in a particular population represent a balance between selection 

for crypsis by predators and selection for conspicuousness by sexual selection 

(Endler, 1983). Black melanic spots are voluntarily increased in size during 

courtship, but reduced in size and intensity at other times. Carotenoid 

spots (red, orange and yellow) genetically decrease in size and frequency with 

increased predation and may be direct indicators of male physical fitness 

(Endler, 1980, 1983). 

This predisposition to colour variations facilitated artificial selection in captivity 

where the multitude of forms and colours far exceeds that found in nature. Guppies 

were bred with short fins, long fins and into those that have the natural olive-grey 

colour replaced by another colour. Because of inbreeding the short-fin morphs had 

to be severely culled to avoid vertebral deformities or stunted growth. In guppies 



form has been sacrificed for colour. The long-fin fish have problems with mating; 

males have difficulty inserting the gonopodium and to transfer spermatophores. 

Some breeders anaesthetize the females to give the males a chance to inseminate 

them (Wischmath, 1993). The various colour forms are then crossbred, but in some, 

where both sexes have elongated fins only artificial insemination enables reproduction. 

There are standard breeds, like triangle (the caudal fin is triangular), fantail, 

veiltail, bannertail, lyretail, topsword, bottomsword, doublesword, speartail, 

roundtail and spadetail. None of the long-tailed and most colourful fish would 

survive in the wild and, therefore, can be considered truly domesticated. 

Other livebearers common in the aquarium hobby are the swordtails and platies 

of the genus Xiphophorus, thatincludec. 17 species from Central America. Swordtail 

males have elongated lower caudal-fin rays; platies have none. Most of the 

morphs in aquaria have resulted from hybridization and artificial selection of three 

species: X. helleri, X. maculatus and Xiphophorus variatus (Meek). Natural hybrids 

are observed only rarely because the different species rarely live in sympatry or are 

separated in the wild by behaviour. The same applies to fancy mollies of the genus 

Poecilia (formerly Mollienesia). The popular black molly was obtained by crossing 

Poecilia sphenops Valenciennes and Poecilia latipinna (Lesueur). The black Yucatan 

molly Poecilia petenensis Günther was introduced into the cross. Crossing further 

with Poecilia velifera (Regan) produced the high-finned molly. ‘The progenies of 

wild specimens bred in captivity greatly resemble their parents. The first apparent 

differences are usually in size. The bred captives often remain like their ancestors in 

appearance and behaviour for years. (. . .) The genera Xiphophorus and Poecilia are 

among these very variable fishes, even in the wild, that develop many regional and 

colour variants’ (Wischmath, 1993). 

The most popular fish in the aquarium trade, the neon tetra was discovered by 

chance in one of the Amazon tributaries near the border of Peru and Colombia. 

Individuals of this small fish were shipped to Paris and Hamburg. The Shedd 

Aquarium in Chicago acquired five of them for its new exhibition. Initially, it 

seemed impossible to breed these fish in captivity; but once this was mastered, in 

the 1950s, the species became the most popular aquarium fish of all. Not long after, 

neon tetra became domesticated by selection for a longfin (Fig. 6) as well as for 

golden, diamond-head, albino and xanthic morphs and aberrations (Eliásˇ, 2003). 

The same reasoning applies to the discus, a cichlid from Amazonia, in which 

selection of some colour aberrations formed the basis for many aquarium 

forms. The majority of these were achieved by hybridization of two known 

species, Symphysodon discus Heckel, and Symphysodon aequifasciatus Pellegrin, 

with some contribution of the questionable sub-species Symphysodon discus 

willischwartzi Burgess, Symphysodon aequifasciatus axelrodi Schultz and 

Symphysodon aequifasciatus haraldi Schultz. Each occurs in the wild in many 

colour varieties, especially the green discus, S. aequifasciatus (Degen, 1995). 

These cichlids are unique in having a discoidal shape strongly laterally 

compressed, similar to the marine butterflyfishes of the genus Chaetodon. 

Approximately the size of a human hand, the S. aequifasciatus, which is widespread 

in the Amazon from Bele´m to Peru, has 50 to 61 vertical scale rows from 

the head to the caudal fin, while S. discus, endemic to the Rio Negro, has 44 to 

48. This classification, however, may change anytime (Burgess, 1991). The question 

now remains if the colour variations of discus, achieved mainly by crossing 



FIG. 6. Domesticated female neon tetras, Paracheirodon innesi: (a) long-fin, (b) albino and (c) xanthic 

forms (from J. Eliásˇ, drawings by S. Frank, reproduced with permission). 

specimens from various taxa and localities are to be designated simply as hybrids 

rather than domesticated fishes. The same problem applies to the other cichlid 

genus of Amazonia, the angelfishes Pterophyllum with two valid species 

Pterophyllum dumerilii (Castelnau) and Pterophyllum scalare (Lichtenstein). 

So far the answer to the question posed at the beginning of this section is that 

probably only guppies and neon tetra in modern aquarium culture can be 

considered as domesticated. The zebra fish, which has been developed for 

research, may be soon a true ‘domesticate’. Other taxa are exploited captives 

as hybrids and colour aberrations in unstable breeding systems with a constant 

addition of wild conspecifics. 

In other species such as the rainbow trout (Behnke, 2002), or the channel 

catfish, used in aquaculture for human food, there are separate hatchery strains 

but no true domesticated forms are known. The same is true for the other 



artificially propagated and cultured fishes including the Chinese and Indian 

carps, grass carp, silver carp, bighead carp, catla, rohu, mrigal and colbasu, 

as well as the salmonids, tilapias, basses, zanders or walleyes (Stizostedion sp.), 

tench Tinca tinca (L.), wels Silurus glanis L., walking catfish Clarias batrachus 

(L.), or sturgeons. At this time all are exploited captives but have potential to 

become domesticated in the future. 

First, my thanks go to K. Lorenzen for inviting me to present this Jack Jones 

Memorial Lecture. Without this invitation it would have never been written. Next, 

I thank R. Hoffmann for advice of a historian and some archeological literature, the 

anonymous referee for constructive comments, and C. Flegler-Balon for repeated reading 

of the final drafts and constant support. The J.W. Jones Lecture, entitled ‘About the 

oldest domesticates among fishes and the epigenetic dichotomy of fish culture’, was 

shortened by the editors to adapt it to the published proceedings. 

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About the oldest domesticates among fishes

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