96
J.H. Choat
tactics and spawning is also reported for the Caribbean resident spawner, the bluehead
wrasse (Warner 1995 ) . Kuwamura et al. ( 2009 ) provided details of spawning behaviour
in 14 species of Indo-Pacifi c parrotfi sh including members of the genera Calotomus,
Chlorurus and Scarus at a single site in Okinawa. Pair-spawning was observed in all
14 species and invariably involved a single female and a terminal phase male;
streaking behaviour by competing males was observed in seven species and groupspawning in seven species. However the frequency of group spawning was represented disproportionately by individuals of the two most abundant species daisy
parrotfi sh, Chlorurus sordidus , and rivulated parrotfi sh, Scarus rivulatus .
Similar information on locality-specifi c population and reproductive biology is
not available for the larger transient spawning species. There are fewer aggregations
compared with resident spawning species, spawning may be at dusk or nocturnal
(Heyman and Kjerfve 2008 ; Colin 2010a ) and there is often a lack of age-specifi c
life history information, especially for many serranid species, including the well
studied Nassau grouper, Epinephelus striatus . A feature of the comparison between
resident and transient spawning is the sensitivity of resident spawning species to
variations in density (Warner 1995 ; Petersen and Warner 2002 ) . Depending on the
local environment surgeonfi shes show a range of behaviours from localized pairspawning to very large resident aggregations on the same reef. The present information suggests that species that display classical transient spawning aggregations are
more conservative with respect to reproductive behaviour. However this aspect of
transient spawning needs to be further examined through a comparison of tropical
West Atlantic and Indo-west Pacifi c snappers and groupers as reproductive behaviour in these groups may have a strong biogeographical element.
4.4 Ecological Features Associated with Resident
and Transient Spawning Modes
4.4.1 The Infl uence of Size and Trophic Ecology
on the Classifi cation of Resident and Transient
Spawning Modes
This section examines the ecological variables that might predict spawning modes
in reef fi shes and is divided into two sections. The fi rst considers 20 families of reef
fi shes and examines two basic features, the size of fi shes in the different families
and their trophic biology. Large size (here expressed as fork length in cm) is an
important characteristic of transient spawning migrations (Domeier and Colin 1997 ;
Nemeth 2008 , Chaps. 1 and 9 ), as aquatic transport costs scale with size (Denny
1993 ) and specifi c metabolic demand decreases with increasing size (Shuter and
Post 1990 , Chap. 2 ). Predation on reef fi shes may also infl uence the type of spawning
mode. Recent studies have emphasized the importance of predators on reef environments arguing that on undisturbed reef habitats predators will have dramatic impacts
J.H. Choat
tactics and spawning is also reported for the Caribbean resident spawner, the bluehead
wrasse (Warner 1995 ) . Kuwamura et al. ( 2009 ) provided details of spawning behaviour
in 14 species of Indo-Pacifi c parrotfi sh including members of the genera Calotomus,
Chlorurus and Scarus at a single site in Okinawa. Pair-spawning was observed in all
14 species and invariably involved a single female and a terminal phase male;
streaking behaviour by competing males was observed in seven species and groupspawning in seven species. However the frequency of group spawning was represented disproportionately by individuals of the two most abundant species daisy
parrotfi sh, Chlorurus sordidus , and rivulated parrotfi sh, Scarus rivulatus .
Similar information on locality-specifi c population and reproductive biology is
not available for the larger transient spawning species. There are fewer aggregations
compared with resident spawning species, spawning may be at dusk or nocturnal
(Heyman and Kjerfve 2008 ; Colin 2010a ) and there is often a lack of age-specifi c
life history information, especially for many serranid species, including the well
studied Nassau grouper, Epinephelus striatus . A feature of the comparison between
resident and transient spawning is the sensitivity of resident spawning species to
variations in density (Warner 1995 ; Petersen and Warner 2002 ) . Depending on the
local environment surgeonfi shes show a range of behaviours from localized pairspawning to very large resident aggregations on the same reef. The present information suggests that species that display classical transient spawning aggregations are
more conservative with respect to reproductive behaviour. However this aspect of
transient spawning needs to be further examined through a comparison of tropical
West Atlantic and Indo-west Pacifi c snappers and groupers as reproductive behaviour in these groups may have a strong biogeographical element.
4.4 Ecological Features Associated with Resident
and Transient Spawning Modes
4.4.1 The Infl uence of Size and Trophic Ecology
on the Classifi cation of Resident and Transient
Spawning Modes
This section examines the ecological variables that might predict spawning modes
in reef fi shes and is divided into two sections. The fi rst considers 20 families of reef
fi shes and examines two basic features, the size of fi shes in the different families
and their trophic biology. Large size (here expressed as fork length in cm) is an
important characteristic of transient spawning migrations (Domeier and Colin 1997 ;
Nemeth 2008 , Chaps. 1 and 9 ), as aquatic transport costs scale with size (Denny
1993 ) and specifi c metabolic demand decreases with increasing size (Shuter and
Post 1990 , Chap. 2 ). Predation on reef fi shes may also infl uence the type of spawning
mode. Recent studies have emphasized the importance of predators on reef environments arguing that on undisturbed reef habitats predators will have dramatic impacts