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Fish reproduction

Fish reproductive organs include testes and ovaries. In 1.2


most species, gonads are paired organs of similar size,
which can be partially or totally fused.[1] There may also
be a range of secondary organs that increase reproductive
tness. The genital papilla is a small, eshy tube behind
the anus in some shes, from which the sperm or eggs are
released; the sex of a sh often can be determined by the
shape of its papilla.

1
1.1

Ovaries

Anatomy
Testes

Most male sh have two testes of similar size. In the case


of sharks, the testes on the right side is usually larger. The
primitive jawless sh have only a single testis, located in 3. Ovary of a female Atlantic cod
the midline of the body, although even this forms from
the fusion of paired structures in the embryo.[2]
Under a tough membranous shell, the tunica albuginea,
the testis of some teleost sh, contains very ne coiled
tubes called seminiferous tubules. The tubules are lined
with a layer of cells (germ cells) that from puberty into old
age, develop into sperm cells (also known as spermatozoa
or male gametes). The developing sperm travel through
the seminiferous tubules to the rete testis located in the
mediastinum testis, to the eerent ducts, and then to the
epididymis where newly created sperm cells mature (see
spermatogenesis). The sperm move into the vas deferens,
and are eventually expelled through the urethra and out
of the urethral orice through muscular contractions.

Many of the features found in ovaries are common to all


vertebrates, including the presence of follicular cells and
tunica albuginea There may be hundreds or even millions
of fertile eggs present in the ovary of a sh at any given
time. Fresh eggs may be developing from the germinal
epithelium throughout life. Corpora lutea are found only
in mammals, and in some elasmobranch sh; in other
species, the remnants of the follicle are quickly resorbed
by the ovary.[2] The ovary of teleosts is often contains a
hollow, lymph-lled space which opens into the oviduct,
and into which the eggs are shed.[2] Most normal female
sh have two ovaries. In some elasmobranchs, only the
However, most sh do not possess seminiferous tubules. right ovary develops fully. In the primitive jawless sh,
Instead, the sperm are produced in spherical structures and some teleosts, there is only one ovary, formed by the
called sperm ampullae. These are seasonal structures, fusion of the paired organs in the embryo.[2]
releasing their contents during the breeding season, and Fish ovaries may be of three types: gymnovarian, secthen being reabsorbed by the body. Before the next ondary gymnovarian or cystovarian. In the rst type,
breeding season, new sperm ampullae begin to form and the oocytes are released directly into the coelomic cavripen. The ampullae are otherwise essentially identical to ity and then enter the ostium, then through the oviduct
the seminiferous tubules in higher vertebrates, including and are eliminated. Secondary gymnovarian ovaries shed
the same range of cell types.[2]
ova into the coelom from which they go directly into
In terms of spermatogonia distribution, the structure of
teleosts testes has two types: in the most common, spermatogonia occur all along the seminiferous tubules, while
in Atherinomorph sh they are conned to the distal portion of these structures. Fish can present cystic or semicystic spermatogenesis in relation to the release phase of
germ cells in cysts to the seminiferous tubules lumen.[1]

the oviduct. In the third type, the oocytes are conveyed


to the exterior through the oviduct.[3] Gymnovaries are
the primitive condition found in lungsh, sturgeon, and
bown. Cystovaries characterize most teleosts, where
the ovary lumen has continuity with the oviduct.[1] Secondary gymnovaries are found in salmonids and a few
other teleosts.
1

1.3

1 ANATOMY

Eggs

Salmon eggs
in dierent stages of development. In some only a few
cells grow on top of the yolk, in the lower right the blood
vessels surround the yolk and in the upper left the black
eyes are visible.

Diagram of a Salmon Fry hatching. The larva has grown around the remains
sh egg: A. vitelline membrane B. chorion C. yolk D. oil of the yolk and the remains of the soft, transparent egg are discarded.
globule E. perivitelline space F. embryo
The eggs of sh and amphibians are jellylike. Cartilagenous sh (sharks, skates, rays, chimaeras) eggs are
fertilized internally and exhibit a wide variety of both
internal and external embryonic development. Most sh
species spawn eggs that are fertilized externally, typically
with the male inseminating the eggs after the female
lays them. These eggs do not have a shell and would dry
out in the air. Even air-breathing amphibians lay their
eggs in water, or in protective foam as with the Coast
foam-nest treefrog, Chiromantis xerampelina.
This young
male spinner shark has claspers, a modication to the
pelvic ns which also function as intromittent organs

1.4

Intromittent organs

Male cartilaginous shes (sharks and rays), as well as the


males of some live-bearing ray nned shes, have ns
that have been modied to function as intromittent organs, reproductive appendages which allow internal fertilization. In ray nned sh they are called gonopodiums
or andropodiums, and in cartilaginous sh they are called
claspers.

Gonopodia are found on the males of some species in the


Anablepidae and Poeciliidae families. They are anal ns
that have been modied to function as movable intromitThis
male tent organs and are used to impregnate females with milt
mosquitosh has a gonopodium, an anal n which during mating. The third, fourth and fth rays of the
functions as an intromittent organ[4][5]
males anal n are formed into a tube-like structure in

3.1

Ovuliparity

which the sperm of the sh is ejected.[6] When ready for


mating, the gonopodium becomes erect and points forward towards the female. The male shortly inserts the
organ into the sex opening of the female, with hook-like
adaptations that allow the sh to grip onto the female to
ensure impregnation. If a female remains stationary and
her partner contacts her vent with his gonopodium, she is
fertilized. The sperm is preserved in the females oviduct.
This allows females to fertilize themselves at any time
without further assistance from males. In some species,
the gonopodium may be half the total body length. Occasionally the n is too long to be used, as in the lyretail
breeds of Xiphophorus helleri. Hormone treated females
may develop gonopodia. These are useless for breeding.

3
In sh, fertilisation of eggs can be either external or internal. In many species of sh, ns have been modied
to allow Internal fertilisation. Similarly, development of
the embryo can be external or internal, although some
species show a change between the two at various stages
of embryo development. Thierry Lod described reproductive strategies in terms of the development of the zygote and the interrelationship with the parents; there are
ve classications - ovuliparity, oviparity, ovo-viviparity,
histotrophic viviparity and hemotrophic viviparity.[10]

3.1 Ovuliparity

eggs (ova),
Similar organs with similar characteristics are found Ovuliparity means the female lays unfertilised
[10]
which
must
then
be
externally
fertilised.
Examples
of
in other shes, for example the andropodium in the
ovuliparous
sh
include
salmon,
goldsh,
cichlids,
tuna
[7]
Hemirhamphodon or in the Goodeidae.
and eels. In the majority of these species, fertilisation
Claspers are found on the males of cartilaginous shes. takes place outside the mothers body, with the male and
They are the posterior part of the pelvic ns that have female sh shedding their gametes into the surrounding
also been modied to function as intromittent organs, and water.
are used to channel semen into the females cloaca during
copulation. The act of mating in sharks usually includes
raising one of the claspers to allow water into a siphon 3.2 Oviparity
through a specic orice. The clasper is then inserted
into the cloaca, where it opens like an umbrella to anchor Main article: Oviparity
its position. The siphon then begins to contract expelling
water and sperm.[8][9]
Oviparity is where fertilisation occurs internally and so
the female sheds zygotes (or newly developing embryos)
into the water,[10] often with important outer tissues
2 Physiology
added. Over 97% of all known sh are oviparous,[11] In
oviparous sh, internal fertilisation requires the male to
Oogonia development in teleosts sh varies according to use some sort of intromittent organ to deliver sperm into
the group, and the determination of oogenesis dynam- the genital opening of the female. Examples include the
ics allows the understanding of maturation and fertilisa- oviparous sharks, such as the horn shark, and oviparous
tion processes. Changes in the nucleus, ooplasm, and rays, such as skates. In these cases, the male is equipped
the surrounding layers characterize the oocyte maturation with a pair of modied pelvic ns known as claspers.
process.[1]
Marine sh can produce high numbers of eggs which are
Postovulatory follicles are structures formed after oocyte often released into the open water column. The eggs have
release; they do not have endocrine function, present a an average diameter of 1 millimetre (0.039 in). The eggs
wide irregular lumen, and are rapidly reabsorbed in a are generally surrounded by the extraembryonic memprocess involving the apoptosis of follicular cells. A de- branes but do not develop a shell, hard or soft, around
generative process called follicular atresia reabsorbs vitel- these membranes. Some sh have thick, leathery coats,
logenic oocytes not spawned. This process can also oc- especially if they must withstand physical force or descur, but less frequently, in oocytes in other development iccation. These type of eggs can also be very small and
fragile.
stages.[1]
Some sh are hermaphrodites, having both testes and
ovaries either at dierent phases in their life cycle or, as
in hamlets, have them simultaneously.

Reproductive strategies

Egg of lamprey
Egg of catshark (mermaids purse)
Egg of bullhead shark
Egg of chimaera

Further information: Modes of reproduction, Spawning,


Ichthyoplankton, and Pregnancy in sh
The newly hatched young of oviparous sh are called
larvae. They are usually poorly formed, carry a large yolk

3 REPRODUCTIVE STRATEGIES

sac (for nourishment) and are very dierent in appear- 3.5


ance from juvenile and adult specimens. The larval period in oviparous sh is relatively short (usually only several weeks), and larvae rapidly grow and change appearance and structure (a process termed metamorphosis) to
become juveniles. During this transition larvae must
switch from their yolk sac to feeding on zooplankton prey,
a process which depends on typically inadequate zooplankton density, starving many larvae.

3.3

Hermaphroditism

Ovoviviparity

Main article: Ovoviviparity


In ovoviviparous sh the eggs develop inside the mothers
body after internal fertilisation but receive little or no
nourishment directly from the mother, depending instead
on a food reserve inside the egg, the yolk.[10] Each embryo develops in its own egg. Familiar examples of
ovoviviparous sh include guppies, angel sharks, and
coelacanths.

3.4

Viviparity

Female
groupers change their sex to male if no male is available

Main article: Viviparity


There are two types of viviparity, dierentiated by how
the ospring gain their nutrients.
Histotrophic (tissue eating) viviparity means embryos develop in the females oviducts but obtain nutrients by consuming other tissues, such as
ova (oophagy) or zygotes.[10] This has been observed primarily among sharks such as the shortn
mako and porbeagle, but is known for a few bony
sh as well such as the halfbeak Nomorhamphus
ebrardtii.[12] An unusual mode of vivipary is
adelphophagy or intrauterine cannibalism, in which
An anemone
the largest embryos eat weaker, smaller unborn sibsh couple guarding their anemone. If the female dies,
lings. This is most commonly found among sharks
a juvenile male moves in, and the resident male changes
such as the grey nurse shark, but has also been resex.
[12]
ported for Nomorhamphus ebrardtii.
Hemotrophic (blood eating) viviparity means embryos develop in the females (or males) oviduct
and nutrients are provided directly by the parent,
typically via a structure similar to, or analogous
to the placenta seen in mammals.[10] Examples of
hemotrophic sh include the surfperches, splitns,
lemon shark, seahorses and pipesh.
Female anAquarists commonly refer to ovoviviparous and glersh, Haplophryne mollis, with atrophied males
viviparous sh as livebearers.
attached

3.6

Sexual parasitism

and reproduce, perhaps because it is larger.[21] Anemone


shes are sequential hermaphrodites which are born as
males, and become females only when they are mature. Anemone shes live together monogamously in a
anemone, protected by the anemone stings. The males
do not have to compete with other males, and female
anemone sh are typically larger. When a female dies
a juvenile (male) anemone sh moves in, and the resident male then turns into a female and reproductive
advantages of the large femalesmall male combination
Parthenogenesis
continue.[22] In other shes sex changes are reversible.
was rst described among vertebrates in the Amazon
For example, if some gobies are grouped by sex (male or
molly
female), some will switch sex.[14]:164[21]
Main article: Hermaphroditism
The mangrove rivulus Kryptolebias marmoratus produces
both eggs and sperm by meiosis and routinely reproduces
Hermaphroditism occurs when a given individual in a
by self-fertilization. Each individual hermaphrodite norspecies possesses both male and female reproductive ormally fertilizes itself when an egg and sperm that it
gans, or can alternate between possessing rst one, and
has produced by an internal organ unite inside the shs
then the other. Hermaphroditism is common in invertebody.[23] In nature, this mode of reproduction can yield
brates but rare in vertebrates. It can be contrasted with
highly homozygous lines composed of individuals so gegonochorism, where each individual in a species is either
netically uniform as to be, in eect, identical to one
male or female, and remains that way throughout their
another.[24][25] The capacity for selng in these shes has
lives. Most sh are gonochorists, but hermaphroditism is
apparently persisted for at least several hundred thousand
known to occur in 14 families of teleost shes.[13]
years.[26]
Usually hermaphrodites are sequential, meaning they can
Although inbreeding, especially in the extreme form
switch sex, usually from female to male (protogyny).
of self-fertilization, is ordinarily regarded as detrimenThis can happen if a dominant male is removed from
tal because it leads to expression of deleterious recesa group of females. The largest female in the harem
sive alleles, self-fertilization does provide the benet of
can switch sex over a few days and replace the domifertilization assurance (reproductive assurance) at each
nant male.[13] This is found amongst coral reef shes such
generation.[24]
as groupers, parrotshes and wrasses. It is less common
for a male to switch to a female (protandry).[14]:162 As an
example, most wrasses are protogynous hermaphrodites
within a haremic mating system.[15][16] Hermaphroditism 3.6 Sexual parasitism
allows for complex mating systems. Wrasses exhibit
three dierent mating systems: polygynous, lek-like, and Some anglersh, like those of the deep sea ceratioid
promiscuous mating systems.[17] Group spawning and group, employ an unusual mating method. Because inpair spawning occur within mating systems. The type dividuals are locally rare, encounters are also very rare.
of spawning that occurs depends on male body size.[16] Therefore, nding a mate is problematic. When scienLabroids typically exhibit broadcast spawning, releas- tists rst started capturing ceratioid anglersh, they noing high amounts of planktonic eggs, which are broad- ticed that all of the specimens were female. These incast by tidal currents; adult wrasses have no interaction dividuals were a few centimetres in size and almost all
with ospring.[18] Wrasse of a particular subgroup of the of them had what appeared to be parasites attached to
Labridae family Labrini do not exhibit broadcast spawn- them. It turned out that these parasites were highly reduced male ceratioids. This indicates the anglersh use a
ing.
polyandrous mating system.
Less commonly hermaphrodites can be synchronous,
meaning they simultaneously possess both ovaries and The methods by which the anglersh locate mates are
testicles and can function as either sex at any one time. variable. Some species have minute eyes unt for identiBlack hamlets take turns releasing sperm and eggs dur- fying females visually, while others have underdeveloped
ing spawning. Because such egg trading is advanta- nostrils, making it unlikely that they eectively nd fe[27]
geous to both individuals, hamlets are typically monog- males using olfaction. When a male nds a female, he
amous for short periods of timean unusual situation in bites into her skin, and releases an enzyme that digests the
shes.[19] The sex of many shes is not xed, but can skin of his mouth and her body, fusing the pair down to
[28]
The male becomes dependent
change with physical and social changes to the environ- the blood-vessel level.
[20]
on
the
female
host
for
survival
by receiving nutrients via
ment where the sh lives.
their shared circulatory system, and provides sperm to the
Particularly among shes, hermaphroditism can pay o female in return. After fusing, males increase in volume
in situations where one sex is more likely to survive and become much larger relative to free-living males of

4 INBREEDING

the species. They live and remain reproductively functional as long as the female lives, and can take part in
multiple spawnings.[27] This extreme sexual dimorphism
ensures, when the female is ready to spawn, she has a mate
immediately available.[29] Multiple males can be incorporated into a single individual female with up to eight males
in some species, though some taxa appear to have a one
male per female rule.[27]
One explanation for the evolution of sexual parasitism
is that the relative low density of females in deep-sea
environments leaves little opportunity for mate choice
among anglersh. Females remain large to accommodate
fecundity, as is evidenced by their large ovaries and eggs.
Males would be expected to shrink to reduce metabolic
costs in resource-poor environments and would develop
highly specialized female-nding abilities. If a male manages to nd a female parasitic attachment, then it is ultimately more likely to improve lifetime tness relative to
free living, particularly when the prospect of nding future mates is poor. An additional advantage to parasitism
is that the males sperm can be used in multiple fertilizations, as he stays always available to the female for mating. Higher densities of male-female encounters might
correlate with species that demonstrate facultative parasitism or simply use a more traditional temporary contact
mating.[30]

3.7

Parthenogenesis

to the ospring. Because gynogenetic species are all female, activation of their eggs requires mating with males
of a closely related species for the needed stimulus. The
Amazon molly, (pictured), reproduces by gynogenesis.

3.8 Others
The elkhorn sculpin (Alcichthys elongatus) is a marine
teleost with a unique reproductive mode called internal gametic association. Sperm are introduced into the
ovary by copulation and then enter the micropylar canal
of ovulated eggs in the ovarian cavity. However, actual
sperm-egg fusion does not occur until the eggs have been
released into sea water.[36]

4 Inbreeding
4.1 Inbreeding depression
The eect of inbreeding on reproductive behavior was
studied in the poeciliid sh Heterandria formosa.[37] One
generation of full-sib mating was found to decrease reproductive performance and likely reproductive success
of male progeny. Other traits that displayed inbreeding
depression were ospring viability and maturation time
of both males and females.

Exposure of zebra sh to a chemical environmental agent,


analogous to that caused by anthropogenic pollution, amMain article: Parthenogenesis
plied the eects of inbreeding on key reproductive
traits.[38] Embryo viability was signicantly reduced in
Parthenogenesis is a form of asexual reproduction in
inbred exposed sh and there was a tendency for inbred
which growth and development of embryos occur without
males to sire fewer ospring.
fertilization. In animals, parthenogenesis means development of an embryo from an unfertilized egg cell. The rst The behaviors of juvenile Coho salmon with either low or
all-female (unisexual) reproduction in vertebrates was de- medium inbreeding were compared in paired contests.[39]
scribed in the Amazon molly in 1932.[31] Since then at Fish with low inbreeding showed almost twice the aggresleast 50 species of unisexual vertebrate have been de- sive pursuit in defending territory than sh with medium
scribed, including at least 20 sh, 25 lizards, a single inbreeding, and furthermore had a higher specic growth
snake species, frogs, and salamanders.[32] As with all rate. A signicant eect of inbreeding depression on jutypes of asexual reproduction, there are both costs (low venile survival was also found, but only in high-density
genetic diversity and therefore susceptibility to adverse competitive environments, suggesting that intra-specic
mutations that might occur) and benets (reproduction competition can magnify the deleterious eects of inwithout the need for a male) associated with partheno- breeding.
genesis.
Parthenogenesis in sharks has been conrmed in the
bonnethead[33] and zebra shark.[34] Other, usually sexual
species, may occasionally reproduce parthenogenetically,
and the hammerhead and blacktip sharks[35] are recent
additions to the known list of facultative parthenogenetic
vertebrates.
A special case of parthenogenesis is gynogenesis. In this
type of reproduction, ospring are produced by the same
mechanism as in parthenogenesis, however, the egg is
stimulated to develop simply by the presence of sperm
- the sperm cells do not contribute any genetic material

4.2 Inbreeding avoidance


Inbreeding ordinarily has negative tness consequences
(inbreeding depression), and as a result species have
evolved mechanisms to avoid inbreeding. Numerous
inbreeding avoidance mechanisms operating prior to mating have been described. However, inbreeding avoidance mechanisms that operate subsequent to copulation
are less well known. In guppies, a post-copulatory mechanism of inbreeding avoidance occurs based on competition between sperm of rival males for achieving

7
fertilisation.[40] In competitions between sperm from an
unrelated male and from a full sibling male, a significant bias in paternity towards the unrelated male was
observed.[40]
Inbreeding depression is considered to be due largely
to the expression of homozygous deleterious recessive
mutations.[41] Outcrossing between unrelated individuals
results in the benecial masking of deleterious recessive
mutations in progeny.[42]

Sexual strategies

Main article: Spawn (biology) Sexual strategies

Within two or three days, the vulnerable goldsh eggs hatch into
larvae, and rapidly develop into fry

Goldsh

Spawning strategies

Main article: Spawning strategies

Spawning grounds

Main article: Spawning ground

Examples

Goldsh, like all cyprinids, are egg-layers. They usually


start breeding after a signicant temperature change, often in spring. Males chase females, prompting them to
release their eggs by bumping and nudging them. As the
female goldsh spawns her eggs, the male goldsh stays
close behind fertilizing them. Their eggs are adhesive and
attach to aquatic vegetation. The eggs hatch within 48 to
72 hours. Within a week or so, the fry begins to assume
its nal shape, although a year may pass before they develop a mature goldsh colour; until then they are a metallic brown like their wild ancestors. In their rst weeks of
life, the fry grow quicklyan adaptation born of the high
risk of getting devoured by the adult goldsh.
Carp
A member of the Cyprinidae family, carp spawn in times
between April and August, largely dependent upon the
climate and conditions they live in. Oxygen levels of the
water, availability of food, size of each sh, age, number
of times the sh has spawned before and water temperature are all factors known to eect when and how many
eggs each carp will spawn at any one time.[43]
Siamese ghting sh

Female goldsh spawn (discharge) eggs into the water, encouraged by male goldsh who simultaneously discharge sperm
which externally fertilizes the eggs

Prior to spawning, male Siamese ghting sh build bubble


nests of varying sizes at the surface of the water. When a
male becomes interested in a female, he will are his gills,
twist his body, and spread his ns. The female darkens
in colour and curves her body back and forth. The act
of spawning takes place in a nuptial embrace where the
male wraps his body around the female, each embrace
resulting in the release of 10-40 eggs until the female
is exhausted of eggs. The male, from his side, releases
milt into the water and fertilization takes place externally.
During and after spawning, the male uses his mouth to
retrieve sinking eggs and deposit them in the bubble nest
(during mating the female sometimes assists her partner,

10

but more often she will simply devour all the eggs that [7]
she manages to catch). Once the female has released all
of her eggs, she is chased away from the males territory,
as it is likely that she'll eat the eggs due to hunger.[44] The
eggs then remain in the males care. He keeps them in [8]
the bubble nest, making sure none fall to the bottom and
repairing the nest as needed. Incubation lasts for 2436 [9]
hours, and the newly hatched larvae remain in the nest for
the next 23 days, until their yolk sacs are fully absorbed.
Afterwards the fry leave the nest and the free-swimming
stage begins.[45]
[10]
Siamese ghting sh build bubble nests of varying
sizes.
A pair of Siamese ghting sh spawning under their
bubble nest.
One-day-old Siamese ghting sh larvae in a bubble
nest - their yolk sacs have not yet been absorbed
A 15-day-old free-swimming fry of a Siamese ghting sh

See also
Evolution of sexual reproduction

10

References

[1] Guimaraes-Cruz, Rodrigo J., Rodrigo J.; Santos, Jos E.


dos; Santos, Gilmar B. (2005). Gonadal structure and
gametogenesis of Loricaria lentiginosa Isbrcker (Pisces,
Teleostei, Siluriformes)". Rev. Bras. Zool. 22 (3): 556
564. doi:10.1590/S0101-81752005000300005. ISSN
0101-8175.
[2] Romer, Alfred Sherwood; Parsons, Thomas S. (1977).
The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 385386. ISBN 0-03-910284-X.
[3] Brito, M.F.G.; Bazzoli, N. (2003). Reproduction
of the surubim catsh (Pisces, Pimelodidae) in the
So Francisco River, Pirapora Region, Minas Gerais,
Brazil. Arquivo Brasileiro de Medicina Veterinria e
Zootecnia. 55 (5): 624633. doi:10.1590/S010209352003000500018. ISSN 0102-0935.
[4] Masterson, J. "Gambusia anis". Smithsonian Institution.
Retrieved 21 October 2011.
[5] Kuntz, Albert (1913). Notes on the Habits, Morphology of the Reproductive Organs, and Embryology of the
Viviparous Fish Gambusia anis. Bulletin of the United
States Bureau of Fisheries. Department of Commerce. 33:
181190.
[6] Kapoor BG and Khanna B (2004) Ichthyology Handbook
pp. 497498, Springer Science & Business Media. ISBN
9783540428541.

REFERENCES

Helfman G, Collette BB, Facey DH and Bowen BW


(2009) The Diversity of Fishes: Biology, Evolution, and
Ecology p. 35, Wiley-Blackwell. ISBN 978-1-40512494-2.
System glossary. FishBase. Retrieved 2013-02-15.
Heinicke, Matthew P.; Naylor, Gavin J. P.; Hedges, S.
Blair (2009). The Timetree of Life: Cartilaginous Fishes
(Chondrichthyes). Oxford University Press. p. 320. ISBN
0191560154.
Lod, T. (2012). Oviparity or viviparity? That is the
question". Reproductive Biology. 12 (3): 259264.
doi:10.1016/j.repbio.2012.09.001. Retrieved November
4, 2014.

[11] Peter Scott: Livebearing Fishes, p. 13. Tetra Press 1997.


ISBN 1-56465-193-2
[12] Meisner, A; Burns, J (1997).
Viviparity in the
Halfbeak Genera Dermogenys and Nomorhamphus
(Teleostei: Hemiramphidae)".
Journal of Mordoi:10.1002/(sici)1097phology.
234: 295317.
4687(199712)234:3<295::aid-jmor7>3.3.co;2-p.
[13] Shapiro DY (1984) Sex reversal and sociodemographics
processes in coral reef shes Pages 103116 in GW Potts
and RK Wootoon, eds., Fish reproduction: Strategies and
tactics, Academic Press.
[14] Moyle PB and Cech JJ (2004) Fishes, An Introduction to
Ichthyology. 5th Ed, Benjamin Cummings. ISBN 978-013-100847-2
[15] Robertson, D.R.; R.R. Warner. Sexual patterns in the
labroid shes of the Western Caribbean II: the parrotshes
(Scaridae)". Smithsonian Contributions to Zoology. 255:
126. doi:10.5479/si.00810282.255.
[16] Kazancioglu, E.; S.H. Alonzo (August 2010). A comparative analysis of sex change in Labridae supports the
size advantage hypothesis. Evolution. 64 (8): 2254
2264. doi:10.1111/j.1558-5646.2010.01016.x. PMID
20394662.
[17] Colin, P.L.; L. J. Bell. Aspects of the spawning of labrid
and scarid shes (Pisces, Labroidei) at Enewetak Atoll,
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10

12

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11

Further references

Agarwal NK (2008) Fish Reproduction APH Publishing. ISBN 9788131303573.


Babin PJ, Cerd J and Lubzens E (Eds) (2007) The
Fish Oocyte: From Basic Studies to Biotechnological
Applications Springer. ISBN 9781402062339.
Bone Q and Moore R (2008) Biology of Fishes
Chapter 8: Reproduction and Life Histories,
pp.
217255.
Taylor & Francis.
ISBN
9781134186310.
Cabrita E, Robles V and Paz Herraez P (Eds)
(2008) Methods in Reproductive Aquaculture: Marine and Freshwater Species CRC Press. ISBN
9780849380549.
Cole, Kathleen Sabina (Ed) (2010) Reproduction
and Sexuality in Marine Fishes: Patterns and
Processes University of California Press. ISBN
9780520264335.
Hoar WS, Randall DJ and Donaldson EM (Eds)
(1983) Fish Physiology: Volume 9: Reproduction
Part A: Endocrine tissues and hormones. Academic
Press. ISBN 9780080585291.
Hoar WS, Randall DJ and Donaldson EM (Eds)
(1983) Fish Physiology: Volume 9: Reproduction
Part B: Behavior and fertility control. Academic
Press. ISBN 9780080585307.
Jakobsen T, Fogarty MJ, Megrey BA and Moksness
E (Eds) (2009) Fish Reproductive Biology: Implications for Assessment and Management John Wiley &
Sons. ISBN 9781444312126.
Melamed P and Sherwood N (Eds) (2005)
Hormones and Their Receptors in Fish Reproduction
World Scientic. ISBN 9789812569189.
Potts GW, Wootton RJ and Wootton RJ (Eds)
(1984) Fish reproduction: strategies and tactics Academic Press. ISBN 9780125636605.
Rocha MJ, Arukwe A and Kapoor BG (Eds)
(2008) Fish Reproduction CRC Press.
ISBN
9781578083312.

12

External links

EXTERNAL LINKS

11

13
13.1

Text and image sources, contributors, and licenses


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Fish reproduction Source: https://en.wikipedia.org/wiki/Fish_reproduction?oldid=743956075 Contributors: Marshman, Zxcvbnm,


Rjwilmsi, Wavelength, RussBot, Epipelagic, Magioladitis, Chiswick Chap, Oshwah, Flyer22 Reborn, ImageRemovalBot, AnomieBOT,
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