I.
Lifetime spawning frequency
Reproduction in Fishes Semelparity — spawn once
Why?
I. Lifetime spawning frequency Iteroparity — spawn multiple times
II. Spawning cycles
semelparous iteroparous
III. Modes of spawning
IV. Sex change and mating systems
II. Spawning cycles III. Modes of reproduction
• semelparous — annual (or multiples thereof) • oviparity — egg laying
• iteroparous — annual, lunar, daily; & cycles within cycles • ovoviviparity — livebearing, no maternal nourishment
daily • viviparity — livebearing with maternal nourishment
monthly
yearly
Tradeoff: number vs. size of eggs/offspring Oviparity
• external fertilization — gametes shed into water
Number of offspring (eggs or young) and degree of parental care
- most common mode
Species Relative Egg or Mean # of Eggs Degree of
Offspring Size or Offspring Parental Care - ancestral condition
Mola mola very small 28,000,000 none, free floating
pelagic eggs - simple
cod very small 9,000,000
haddock very small 2,000,000 - small, but numerous eggs
sturgeon medium 200,000 eggs buried, no • internal fertilization — eggs fertilized internally, but
other care
salmon medium 5,000 later released into water
sculpin medium 140
stickleback medium 60 high
- uncommon
medaka medium 30 - many elasmobranchs
sand shark very large 1-2 extreme: uterus
w/ cannibalism - often very large eggs in cases
1
Internal Fertilization in Oviparous species Ovoviviparity
• internal fertilization — eggs fertilized internally & hatch
internally, but no direct maternal nourishment
advantage
- young are generally larger
and more advanced
disadvantages
- low dispersal of offspring
- few offspring produced
- if mother dies while pregnant, offspring die, too
examples:
- most sharks and rays
- coelacanth
- rockfishes
copulation: white tip reef sharks
Viviparity
• internal fertilization & direct maternal nourishment
- ovarian fluids
- placenta (some sharks)
advantage
- young are very large and advanced
disadvantages
- same as for ovoviviparity
- energetic cost to mother
2
Internal Fertilization: males have an intromittent organ
1. modifications of pelvic or anal fins:
- claspers: elasmobranchs, chimeras
- gonopodia: guppies, mollies
2. genital papilla: non-bony tube
- rockfishes, surfperches, cottids
viviparity - lemon shark birth
Viviparity
viviparity - shiner surfperch birth
examples:
- requiem sharks (family Carcharhinidae) & hammerheads
- guppies & other freshwater livebearers
- marine surfperches
Embiotocid embryos Return to Oviparity:
with trophotaenia
Modes of Oviparity
1) broadcast spawning
2) demersal, non-guarding
3) demersal, guarding
4) brooders
3
1) Broadcast Spawning
1) Broadcast Spawning
- most marine fishes
• advantages
• eggs & sperm shed into water column & drift away
• larval & juvenile/adult habitat completely different (bipartite life cycle) - zero energy expenditure after gametes released
- best chance of long distance dispersal
• disadvantages
- very low survival of offspring
- caused by:
* lack of parental protection
* small larval size
* inability to find or return to appropriate adult habitat
1) Broadcast Spawning: Timing & Location
• maximize transport of eggs & larvae away from the reef
- spawn during outgoing tide
- downcurrent side of reef
- upward spawning rush to place gametes high in water column
• two hypotheses:
- minimize exposure to predators
- maximize dispersal (“spread the risk”)
2) Demersal Non-Guarding
• hiders and non-hiders
• eggs usually placed or glued on bottom Brook trout eggs
• common in freshwater fishes (minnows), but not in marine
fishes
advantages
- zero energy expenditure after gametes released
- eggs may be better protected
- greater control over where offspring end up
disadvantages
- still quite low survival of offspring
- high risk of loosing entire batch of eggs
- lower dispersal
- nest sites can become limited
4
Grunion spawning Grunion spawning
Skate egg cases Swell shark egg case
3) Demersal Guarding 3) Demersal Guarding
• common among reef fishes (e.g., gobies, damselfish, advantages
triggerfish) - increased survival due to parental protection
- increased survival due to parental care
• males usually guard
(cleaning and oxygenation)
disadvantages
- high cost to parent(s): energy and time
- increased risk of mortality in parent(s)
- nest sites may become limited
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4) Brooding
• young brooded in:
– mouths (males of cardinalfishes, jawfishes,
catfishes; female cichlids)
– on body (some male pipefishes)
– pouches (male seahorses & some pipefishes)
• offspring usually (but not always — e.g.,
cardinalfishes) well developed when released
Haplochromis spawning
Haplochromis mouthbrooding female
seahorse oviposition
& male giving “birth”
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4) Brooding
advantages
• increased survival due to parental protection
• increased survival due to parental care (cleaning and IV. Sex change
oxygenation)
- among vertebrates, found only in fishes
• no need to find or maintain a nest
disadvantages
• high cost to parent(s): energy and time
• if parent dies, so do offspring
• low dispersal
Sexual Strategies
• gonochorism — separate sexes, no sex change
- advantage: allows sexual specialization
- disadvantage: may not maximize fitness Why are fish able to change sex?
• hermaphroditism
- simultaneous • simple gonads and reproductive tracts
- sequential • sex chromosomes are similar or identical
- protogynous = female then male
- protandrous = male then female
Why do fish change sex?
Why do fish change sex?
Size advantage model - predicts sex change if your current
reproductive value (RV) is less than what it would be if you
were the opposite sex
• to maximize fitness by maximizing lifetime
reproductive output RV = (expected fecundity at a given size)
×
(probability of surviving to that size)
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Why would RV differ between the sexes? RV changes with body size and mating system:
1) relative cost of gametes 1) female — increases linearly with size
• sperm are cheap, eggs expensive 2) male — depends on mating system
• therefore, females usually limited by ability to produce • competition for mates or sperm competition
eggs or hold them or embryos in the body
(69 human offspring; Russian woman, 1700’s)
Gonochorism
• males usually limited by access to females
(850+ children: Emperor of Morocco 1600-1700’s) male
# eggs produced
female
2) mating system...
or fertilized
• monogamy - one partner
• polygamy - multiple partners
- polygyny - many females, one male
- polyandry - many males, one female
Size (body mass)
RV changes with body size and mating system: immature
Protogyny -- California sheephead
1) female — increases linearly with size
2) male — depends on mating system
• competition for spawning sites or limited access to females
Protogyny female
male
# eggs produced
female
male
or fertilized
Size (body mass)
RV changes with body size and mating system:
Protandry -- anemonefish
1) female — increases linearly with size
2) male — depends on mating system
• no defensible resources
• limited # of sperm needed
• no direct competition between males
• can occur if matings are random or there is stable monogamy
Protandry
# eggs produced
female
or fertilized
male
Size (body mass)
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Trimma okinawae Paragobiodon echinocephalus
Reversible Sex Change
or
Simultaneous Hermaphroditism
Reversible sex change is very rare
female
(a handful of gobies)
# eggs produced
male Why?
or fertilized
• energetically expensive?
• lost mating opportunities? Lythrypnus zebra
Size (body mass)
Gobiodon histrio
Simultaneous hermaphroditism is common in two
groups of fishes:
• deep sea fishes
Simultaneous hermaphroditism seems like best strategy • small serranids
• Why is it so rare?
Cheating
Small serranids (Hypoplectrus, Serranus)
barracudina barracudina • egg trading allows cheaters to be detected and punished
Deep sea fishes • cheaters abandoned
• population densities very low • abandoned cheaters lose mating opportunities while
searching for new mate
• must be able to act as either sex when you encounter a
Serranus tigrinis
conspecific
• selection so strong that cheating species went extinct?
Hypoplectrus indigo
Serranus tabacarius
lancetfish