Animal Diversity Web

Ge­o­graphic Range

True clown anemone­fishes (Am­phiprion per­cula) are na­tive only to the Indo-Pa­cific Re­gion (Rosen­berg and Cruz, 1988). The species ranges from North­ern Queens­land to Melane­sia, which com­prises New Guinea, New Britain, New Ire­land, the Solomon Is­lands, and Van­u­atu (Fautin and Allen, 1992). (Fautin and Allen, 1992; Rosen­berg and Cruz, 1988)

• Biogeographic Regions
• oriental
  • native
• australian
  • native

Habi­tat

Like all anemone­fishes, A. per­cula forms sym­bi­otic re­la­tion­ships with sea anemones. It uses its host as both shel­ter and pro­tec­tion from preda­tors. Be­cause of this close re­la­tion­ship, the dis­tri­b­u­tion of suit­able host anemone species dic­tates the habi­tat of A. per­cula. As­so­ci­a­tions in­volv­ing A. per­cula and the sea anemone species Het­er­ac­tis mag­nifica, Sti­chodactyla gi­gan­tean, and Sti­chodactyla merten­sii are usu­ally found in na­ture (El­liott and Mariscal, 1996). Both sym­bionts re­side in shal­low coastal wa­ters of the trop­ics where depth rarely ex­ceeds 12 me­ters and water tem­per­a­ture ranges from 25-28 de­grees C. (Ran­dall et al., 1997; Fautin and Allen, 1992). The dis­tri­b­u­tion of sea anemones them­selves is lim­ited by the pho­to­syn­thetic ac­tiv­ity of golden-brown algae that oc­cupy the anemones’ ten­ta­cles (Fautin and Allen, 1992). The fish and anemone pair gen­er­ally oc­curs on coral reefs where the lat­ter is an­chored se­curely and the for­mer can be seen swim­ming in and out of the pro­tec­tive ten­ta­cles of its host.

When sev­eral species of anemone­fishes occur to­gether in sim­i­lar habi­tats, they tend to par­ti­tion them­selves ac­cord­ing to mi­cro­hab­i­tats and avail­able species of sea anemones. A. per­cula, for ex­am­ple will typ­i­cally oc­cupy H. mag­nifica in nearshore zones while Am­phiprion perid­eraion will oc­cupy the same species in off­shore zones. In­tense com­pe­ti­tion for lim­ited re­sources un­doubt­edly af­fects the ter­ri­to­r­ial na­ture of these fishes. Niche dif­fer­en­ti­a­tion is caused by dis­tri­b­u­tion, abun­dance, and re­cruit­ment pat­terns of com­pet­ing species (El­liott and Mariscal, 2001). (El­liott and Mariscal, 1996; El­liott and Mariscal, 2001; Fautin and Allen, 1992; Ran­dall, et al., 1997)

• Habitat Regions
• tropical
• saltwater or marine

• Aquatic Biomes
• reef
• coastal

• Range depth

  1 to 12 m

  3.28 to 39.37 ft

Phys­i­cal De­scrip­tion

A. per­cula can grow to 110 mm in length and is often dis­tin­guished by three white ver­ti­cal bars on a bright or­ange body. The an­te­rior white bar oc­curs just be­hind the eye; the mid­dle bar bi­sects the fish; the pos­te­rior bar oc­curs near the cau­dal fin. An an­te­rior pro­ject­ing bulge fur­ther char­ac­ter­izes the mid­dle bar. In ad­di­tion to the white col­or­ing, black edg­ing out­lines each fin with vary­ing thick­ness (Fautin and Allen, 1992; Grant, 1999). Al­though A. per­cula’s vi­brant col­ors are eye catch­ing, it is eas­ily con­fused with Am­phiprion ocel­laris (false clown anemone­fish). One may dis­tin­guish the two by count­ing the num­ber of dor­sal-fin spines. A. per­cula usu­ally has 10 dor­sal-fin spines, while A. ocel­laris usu­ally has 11. Also, the lat­ter never has thick black mar­gins out­lin­ing the fins (Fautin and Allen, 1992)

There is no dif­fer­ence in color pat­terns among sexes. Nonethe­less, di­mor­phic vari­a­tion is pre­sent, since the fe­male is larger than the male. Poly­mor­phism, al­though pre­sent in other species of anemone­fishes, does not occur in A. per­cula. Such is the case with melanis­tic (black pig­men­ta­tion) vari­a­tion in some anemone­fish species. This is gen­er­ally ab­sent in A. per­cula (Fautin and Allen, 1992). (Fautin and Allen, 1992; Grant, 1999)

• Other Physical Features
• ectothermic
• heterothermic
• bilateral symmetry

• Sexual Dimorphism
• female larger

• Range length

  110 (high) mm

  4.33 (high) in

De­vel­op­ment

After in­cu­bat­ing for 6-7 days, the eggs of A. per­cula are ready to hatch. Just be­fore then, how­ever, the em­bryo is vis­i­ble through the trans­par­ent egg mem­brane. The two no­tice­able fea­tures at this stage are the sil­very pupils con­tained within the large eyes and the red-or­ange yolk sac (Fautin and Allen, 1992). After hatch­ing, the larva is about 3-4 mm total length and trans­par­ent ex­cept for the eye, yolk sac, and a few scat­tered pig­ments. The newly hatched in­di­vid­ual ini­tially sinks to the ben­thic en­vi­ron­ment but quickly swims to the upper sur­face of the water col­umn using a process called pho­to­taxis. Es­sen­tially, the larva is able to ori­ent it­self using the shine from a moon­lit night. At this point, the larva spends a week float­ing among the plank­ton and is pas­sively trans­ported by ocean cur­rents (Fautin and Allen, 1992). The lar­val stage of A. per­cula ends when the young anemone­fish set­tles to the sea bot­tom ap­prox­i­mately 8-12 days after hatch­ing (DAH). Com­pared to other coral reef species, this is a rel­a­tively short pe­riod (Welling­ton and Vic­tor 1989).

The ju­ve­nile stage of A. per­cula is char­ac­ter­ized by a rapid de­vel­op­ment of color schemes. The white bar­ring pat­tern that is unique to this species be­gins to form around 11 DAH and may cor­re­spond to the fish’s first as­so­ci­a­tion with its host anemone (El­liott et al., 1995). Con­se­quently, con­tact with the anemone stim­u­lates A. per­cula to pro­duce its pro­tec­tive mu­cous coat (El­liott and Mariscal, 1996) (See Be­hav­ior sec­tion for a com­plete elab­o­ra­tion on ac­cli­ma­tion and pro­tec­tion from anemone ne­ma­to­cysts). The en­tire meta­mor­pho­sis from larva to ju­ve­nile is usu­ally com­pleted in a day (Fautin and Allen, 1992).

De­vel­op­ment from ju­ve­nile to adult is highly de­pen­dent on the so­cial hi­er­ar­chy of the “fam­ily group.” Each host anemone is often oc­cu­pied by a mat­ing pair plus two to four smaller fish (Fautin and Allen, 1992). Ag­gres­sion be­tween the dom­i­nant fe­male and her mate is min­i­mal, thereby caus­ing lit­tle ex­pen­di­ture in en­ergy. Each male, how­ever, bul­lies and chases the next male of smaller suc­ces­sive size until the small­est in­di­vid­ual is dri­ven away from the host anemone. As a re­sult, en­ergy that could be used for growth is in­stead ap­pro­pri­ated for com­pet­i­tive en­coun­ters. The adult pair es­sen­tially stunts the growth of ju­ve­niles (Myers, 1999).

Like other anemone­fishes, the unique­ness of A. per­cula de­vel­op­ment lies in adult meta­mor­pho­sis from male to fe­male (protan­drous her­maph­ro­ditism). All anemone­fishes are born as males (Wood and Aw, 2002; Fautin and Allen, 1992; Rosen­berg and Cruz, 1988), and the largest of the group re­verses sex to be­come the dom­i­nant fe­male. The sec­ond largest male sub­se­quently be­comes the dom­i­nant male. In in­stances when the fe­male dies, the dom­i­nant male re­verses sex and all other sub­or­di­nate males move up in the hi­er­ar­chi­cal lad­der. (El­liott and Mariscal, 1996; El­liott, et al., 1995; Fautin and Allen, 1992; Myers, 1999; Rosen­berg and Cruz, 1988; Welling­ton and Vic­tor, 1989; Wood and Aw, 2002)

• Development - Life Cycle
• metamorphosis

Re­pro­duc­tion

• Mating System
• monogamous

Monog­a­mous pair-bond for­ma­tions be­tween male and fe­male in­di­vid­u­als of A. per­cula are very strong and cor­re­lated with the small ter­ri­tory size that this species oc­cu­pies. De­spite being re­stricted to the im­me­di­ate vicin­ity of its host anemone, A. per­cula can breed/spawn year round due to the per­pet­u­ally warm trop­i­cal wa­ters they in­habit.

Ini­ti­a­tion of courtship is highly cor­re­lated with the lunar cycle. The moon­light serves to main­tain a high level of alert­ness in the male, which then leads to in­creased so­cial in­ter­ac­tion with the fe­male. Sev­eral days be­fore spawn­ing, the male will show mor­pho­log­i­cal and be­hav­ioral changes: fin erec­tion, chas­ing, nest prepa­ra­tion, and “sig­nal jump­ing.” This last trait is de­picted with rapid up and down swim­ming mo­tions. Fi­nally, ex­ten­sions of anal, dor­sal, and pelvic fins ac­com­pany the ag­gres­sive­ness of the male (Fautin and Allen, 1992)

The choice of nest site is im­por­tant for later sur­vival of the eggs. It is usu­ally lo­cated under the ten­ta­cles of the host anemone and se­curely po­si­tioned on a patch of cleared rock (Myers, 1999). The male has been known to nip at the bot­tom edges of the ten­ta­cles in order to cause re­trac­tion, and thus pro­vid­ing enough clear­ance to clean the area (Rosen­berg and Cruz, 1988). Ini­tially, the male clears algae and de­bris with its mouth only later to be joined by its mate (Fautin and Allen, 1992). There is clear em­pha­sis, then, on male parental care, and this will be cru­cial when the eggs be­come vul­ner­a­ble to pre­da­tion.

Ac­tual spawn­ing pro­ces­sion takes place dur­ing the morn­ing hours, and gen­er­ally lasts about 30 min­utes to more than two hours. At this stage, the con­i­cal ovipos­i­tor of the fe­male be­comes vis­i­ble. Sev­eral eggs are ex­truded through this struc­ture with each slow and de­lib­er­ate pass as the belly gen­tly brushes the nest sur­face. Fol­low­ing closely be­hind is her mate, who ex­ter­nally fer­til­izes the eggs as they are laid. The num­ber of total passes dur­ing each spawn­ing ses­sion is high, and the amount of de­posited eggs range from 100 to over 1000, de­pend­ing on fish size and pre­vi­ous ex­pe­ri­ence. Older, more ex­pe­ri­enced mat­ing pairs will pro­duce more eggs. The eggs of A. per­cula are about 3-4 mm in length (Fautin and Allen, 1992).

After egg de­po­si­tion has fin­ished, the in­cu­ba­tion pe­riod be­gins. At this time, the male ac­tively mouths and fans the eggs, while si­mul­ta­ne­ously being on guard against any preda­tors (Rosen­berg and Cruz, 1988). Be­cause the eggs are at­tached to the bot­tom sub­strate via ad­he­sive strands, ad­di­tional pro­tec­tion is pro­vide by the over­hang­ing ten­ta­cles of the host anemone (Allen, 1997). Re­moval of dead eggs and de­bris is also im­por­tant in keep­ing a well-oxy­genated nest and is ac­com­plished by the male. The fe­male, in con­trast, is oc­cu­pied with feed­ing dur­ing this time (Fautin and Allen, 1992). (Allen, 1997; Fautin and Allen, 1992; Myers, 1999; Rosen­berg and Cruz, 1988)

• Key Reproductive Features
• iteroparous
• year-round breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sequential hermaphrodite
  • protandrous
• sexual
• fertilization
  • external
• oviparous

• Breeding season

  year round

• Range number of offspring

  100 to 1000

• Average time to hatching

  6-7 days

• Parental Investment
• no parental involvement
• pre-hatching/birth
  • protecting
    • male

Lifes­pan/Longevity

There is very lit­tle longevity data for many species of anemone­fishes. How­ever, some are recorded to have lived at least 6-10 years in na­ture. In cap­tiv­ity, the record is 18 years for Am­phiprion fre­na­tus and Am­phiprion perid­eraion. (Fautin and Allen, 1992)

• Average lifespan
  Status: wild

  6-10 years

• Range lifespan
  Status: captivity

  18 (high) years

• Typical lifespan
  Status: wild

  6 to 10 years

Be­hav­ior

In anemone­fishes par­tic­u­lar at­ten­tion has been given to be­hav­ioral com­po­nents of the sym­bi­otic re­la­tion­ship with sea anemones. Re­liance on a host has ef­fects at every par­tic­u­lar life stage. A. per­cula lays its eggs under the over­hang of an anemone’s ten­ta­cles (lee­ward side). Arved­lund et al. (2000) be­lieved that this was a preda­tor-de­ter­rence and an ol­fac­tory im­print­ing mech­a­nism. The lat­ter plays an im­por­tant role in di­rect­ing ju­ve­niles to the ap­pro­pri­ate sea anemone species later on. With a lee­ward place­ment, a max­i­mum amount of im­print­ing mu­cous can trans­fer be­tween the ten­ta­cles and eggs.

Once a ju­ve­nile, A. per­cula must lo­cate and in­habit a suit­able anemone host. Its poor swim­ming abil­ity makes it an easy tar­get for preda­tors. Cer­tain chem­i­cal cues are used, and they dif­fer among anemone­fishes; this causes pref­er­en­tial se­lec­tion for cer­tain anemone species (Fautin and Allen, 1992). El­liott et al. (1995) found that ocean cur­rents fa­cil­i­tate the lo­cat­ing process and that vi­sual cues were never used. Even when a tar­geted anemone is al­ready oc­cu­pied, the ap­proach­ing A. per­cula does not avoid it; how­ever, the ter­ri­to­r­ial na­ture of anemone­fishes causes the res­i­dent to chase away its in­truder.

Habi­ta­tion of a cho­sen anemone gen­er­ally re­quires a pe­riod of ac­cli­ma­tion (Dav­en­port and Nor­ris, 1958; Fautin and Allen, 1992, El­liott and Mariscal, 1997). The pro­tec­tive mu­cous of A. per­cula is de­vel­oped with re­peated in­ter­ac­tions with the host anemone. Two the­o­ries exist about how the mu­cous layer forms. Ei­ther the fish ac­quires it after con­tact with the ten­ta­cles (a be­hav­ioral process), or it is de­vel­oped phys­i­o­log­i­cally (a bio­chem­i­cal process). Both ex­pla­na­tions have been sup­ported, and both are be­lieved to be equally im­por­tant. Dur­ing its first en­counter with the sea anemone, A. per­cula will en­gage in a swim­ming dance, gin­gerly touch­ing ten­ta­cles first to its ven­tral fins and then to its en­tire body. It may be stung a num­ber of times be­fore full ac­cli­ma­tion oc­curs. The whole pro­ce­dure may take as lit­tle as a few min­utes to sev­eral hours. Once ac­cli­mated, though, the mu­cous pro­tec­tion may dis­ap­pear upon ex­tended sep­a­ra­tion be­tween host and fish. Con­tin­ued con­tact with the ten­ta­cles ap­pears to re­ac­ti­vate the mu­cous coat on A. per­cula. (Arved­lund, et al., 2000; Dav­en­port and Nor­ris, 1958; El­liott and Mariscal, 1997; El­liott, et al., 1995; Fautin and Allen, 1992)

• Key Behaviors
• natatorial
• diurnal
• motile
• sedentary
• territorial
• dominance hierarchies

Com­mu­ni­ca­tion and Per­cep­tion

• Perception Channels
• tactile
• chemical

Food Habits

A. per­cula feeds mainly on zoo­plank­ton, such as cope­pods and lar­val tu­ni­cates. Pos­si­bly, it con­sumes algae from the sur­round­ing coral reef or even left­over food por­tions from its host anemone. The for­mer strat­egy is com­monly used by A. perid­eraion (Fautin and Allen, 1992). Fre­quently, A. per­cula will carry large pieces of food to its host anemone, pre­sum­ably to store it for later use. The anemone, how­ever, de­vours the ac­ces­si­ble food item in most cases (Grant, 1999).

Op­ti­mal ju­ve­nile growth rate was dis­cov­ered at a ra­tion of ap­prox­i­mately 6% body weight per day (John­ston et al., 2000). Ju­ve­niles are under con­sid­er­able pres­sure from the hi­er­ar­chi­cal struc­ture. The in­di­vid­ual is ha­rassed and chased by big­ger males of the “fam­ily group,” which re­sults in stunted growth. Con­se­quently, the smaller fish has a more re­stricted feed­ing area, and more en­ergy must be placed on eva­sion. Only when a larger male is re­moved (e.g. death) will the smaller ju­ve­nile ex­pe­ri­ence an ac­cel­er­a­tion in growth rate. It is be­lieved that less time being ha­rassed trans­lates into more time spent on feed­ing (Fautin and Allen, 1992).

Due to the in­creased aquar­ium trade for A. per­cula (See Eco­nomic Im­por­tance for Hu­mans) and a con­tin­ued de­ple­tion of coral reef habi­tats, there have been tremen­dous de­vel­op­ments in rear­ing of ma­rine fishes using aqua­cul­tur­ing tech­niques. One of the most chal­leng­ing ob­sta­cles is pro­vid­ing an eco­nom­i­cal, yet ef­fec­tive, feed in an ar­ti­fi­cial en­vi­ron­ment. Hoff (1996) found that A. per­cula lar­vae and ju­ve­niles could be suc­cess­fully reared on highly in­te­grated and di­verse feeds, such as ro­tifers, small par­tic­u­late dry feed, Artemia, and krill meal. Un­for­tu­nately, this proved too ex­pen­sive to be prac­ti­cal, and a regime solely based on ar­ti­fi­cial feed de­creased sur­vival and growth rates in young fishes. If, how­ever, ju­ve­niles were weaned from live Artemia 15 to 20 days after hatch­ing and fed a fish meal/ca­sein-based sub­sti­tute, sur­vival and growth rates showed no dif­fer­ence from ju­ve­niles fed en­tirely on live feed (Gor­don et al., 2000). (Fautin and Allen, 1992; Gor­don, et al., 1998; Grant, 1999; Hoff, 1996; John­ston, et al., 2000)

• Primary Diet
• planktivore

• Animal Foods
• aquatic crustaceans
• other marine invertebrates
• zooplankton

• Plant Foods
• algae

• Foraging Behavior
• stores or caches food

Pre­da­tion

The sym­bio­sis be­tween A. per­cula and its host anemone serves as an ef­fec­tive anti-pre­da­tion mea­sure. Pro­tected within the ten­ta­cles of the sea anemone, A. per­cula be­longs to a unique group of fishes that are not stung by the ne­ma­to­cysts. It is be­lieved that a thick mu­cous layer cloaks the fish from de­tec­tion and re­sponse by anemone ten­ta­cles (Rosen­berg and Cruz, 1988). Fish species lack­ing in this phys­i­o­log­i­cal adap­ta­tion are cap­tured and de­voured by the sea anemone. It is no sur­prise, then, that A. per­cula has very few preda­tory foes as adults. Pres­ence of dan­ger im­me­di­ately elic­its a re­sponse to seek shel­ter deep within its host. Al­though adults are rel­a­tively safe from pre­da­tion, the eggs of A. per­cula are sus­cep­ti­ble and must be guarded by the dom­i­nant male. The most com­mon day preda­tors are wrasses (fam­ily Labri­dae) and other dam­selfishes (fam­ily Po­ma­cen­tri­dae). Night preda­tors of eggs are gen­er­ally not fishes but in­ver­te­brates like brit­tle stars (Ophiotrichi­dae, Ophiochim­i­dae, and Ophio­der­mati­dae) (Arved­lund et al., 2000). (Arved­lund, et al., 2000; Rosen­berg and Cruz, 1988)

• Known Predators

  • wrasses (Labridae)
  • damselfish (Pomacentridae)
  • "brittle stars (Ophiotrichidae, Ophiochimidae, and Ophiodermatidae)"

Ecosys­tem Roles

A. per­cula in­ter­acts with its sea anemone host and other anemone­fish species. The sym­bi­otic re­la­tion­ship is well doc­u­mented to ben­e­fit the fish, but equal re­wards exist for the anemone. In ex­change for pro­tec­tion, A. per­cula may feed, oxy­genate, and re­move waste ma­te­r­ial from its host (Rosen­berg and Cruz, 1988). In ad­di­tion, it may pre­vent cer­tain coe­len­ter­ate feed­ers, such as but­ter­fly fishes, from prey­ing on the anemone (Allen, 1997). Be­cause anemone­fishes are highly ter­ri­to­r­ial, A. per­cula dri­ves away in­trud­ers, in­clud­ing those that harm its sym­bi­otic host. Whether these ac­tions are self-serv­ing or al­tru­is­tic is not known, but both species gain ad­van­tage. (Allen, 1997; Rosen­berg and Cruz, 1988)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

A. per­cula and other anemone­fishes are some of the most col­or­ful fish species avail­able for the aquar­ium trade. They also demon­strate in­ter­est­ing be­hav­iors and are eas­ily adapt­able to cap­tiv­ity (Fautin and Allen, 1992). Con­se­quently, these char­ac­ter­is­tics make them good ref­er­ence fishes for sci­en­tific re­search, es­pe­cially when con­duct­ing nu­tri­tional stud­ies and de­ter­min­ing egg and lar­val qual­ity (Gor­don et al., 2000). (Fautin and Allen, 1992; Gor­don, et al., 1998)

• Positive Impacts
• pet trade
• ecotourism
• research and education

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

None known

Con­ser­va­tion Sta­tus

The de­ple­tion of coral reef habi­tats and ma­rine aquar­ium fishes has pre­sented a rel­a­tively new mar­ket in aqua­cul­ture. It is pos­si­ble to rear A. per­cula in con­trolled con­di­tions (Gor­don et al., 2000), and it may even­tu­ally play a sig­nif­i­cant role in main­tain­ing sta­ble pop­u­la­tions. At pre­sent, this species is not threat­ened or en­dan­gered. (Gor­don, et al., 1998)

• IUCN Red List

  No special status

• US Federal List

  No special status

• CITES

  No special status

Con­trib­u­tors

William Fink (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Jeff Lee (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor.

Ref­er­ences

Allen, G. 1997. Ma­rine Fishes of Trop­i­cal Aus­tralia and South-East Asia. Perth: West­ern Aus­tralian Mu­seum.

Arved­lund, M., I. Bundgaard, L. Nielsen. 2000. Host im­print­ing in anemone­fishes (Pisces: Po­ma­cen­tri­dae): does it dic­tate spawn­ing site pref­er­ences?. En­vi­ron­men­tal Bi­ol­ogy of Fishes, 58: 203-213.

Dav­en­port, D., K. Nor­ris. 1958. Ob­ser­va­tions on the sym­bio­sis of the sea anemone *Sto­ichac­tis* and the po­ma­cen­trid fish, *Am­phiprion per­cula*. Bi­o­log­i­cal Bul­letin, 115(3): 397-410.

El­liott, J., J. El­liott, R. Mariscal. 1995. Host se­lec­tion, lo­ca­tion, and as­so­ci­a­tion be­hav­iors of anemone­fishes in field set­tle­ment ex­per­i­ments. Ma­rine Bi­ol­ogy, 122: 377-389.

El­liott, J., R. Mariscal. 1997. Ac­cli­ma­tion or in­nate pro­tec­tion of anemone­fishes from sea anemones?. Copeia, 2: 284-289.

El­liott, J., R. Mariscal. 2001. Co­ex­is­tence of nine anemone­fish species: dif­fer­en­tial host and habi­tat uti­liza­tion, size and re­cruit­ment. Ma­rine Bi­ol­ogy, 138: 23-36.

El­liott, J., R. Mariscal. 1996. On­to­ge­netic and in­ter­spe­cific vari­a­tion in the pro­tec­tion of anemone­fishes from sea anemones. Jour­nal of Ex­per­i­men­tal Ma­rine Bi­ol­ogy and Ecol­ogy, 208: 57-72.

Ern, G. 1999. Grant's Guide to Fishes. Scar­bor­ough: E.M. Grant Pty Ltd..

Fautin, D., G. Allen. 1992. Field Guide to Anemone­fishes and their Host Sea Anemones. Perth: West­ern Aus­tralian Mu­seum.

Gor­don, A., H. Kaiser, P. Britz, T. Hecht. 1998. Ef­fect of feed type and age-at-ween­ing on growth and sur­vival of clown­fish *Am­phiprion per­cula* (Po­ma­cen­tri­dae). Aquar­ium Sci­ences and Con­ser­va­tion, 2: 215-226.

Grant, E. 1999. Grant's Guide to Fishes. Scar­bor­ough: E.M. Grant Pty Ltd..

Hoff, F. 1996. Con­di­tion­ing, Spawn­ing and Rear­ing of Fish with Em­pha­sis on Ma­rine Clown­fish. Dade City: Aqua­cul­ture Con­sul­tants, Inc..

John­ston, G., T. Hetcht, L. Oeller­mann, H. Kaiser. 2000. Ef­fect of feed­ing fre­quency and ra­tion on the growth of ju­ve­nile clown­fish (*Am­phiprion per­cula*). 10th South­ern African Ma­rine Sci­ence Sym­po­sium (SAMSS 2002): Land, Sea and Peo­ple in the New Mil­len­nium--Ab­stracts.

Myers, R. 1999. Mi­crone­sian Reef Fishes. Guam: Coral Graph­ics.

Ran­dall, J., G. Allen, R. Steene. 1997. Fishes of the Great Bar­rier Reef and Coral Sea. Bathurst: Craw­ford House Pub­lish­ing.

Rosen­berg, S., G. Cruz. 1988. The anemone­fishes of the Indo-Pa­cific. Sea Fron­tiers, 34: 16-21.

Welling­ton, G., B. Vic­tor. 1989. Plank­tonic lar­val du­ra­tion of one hun­dred species of Pa­cific and At­lantic dam­selfishes (Po­ma­cen­tri­dae). Ma­rine Bi­ol­ogy, 101: 557-567.

Wood, E., M. Aw. 2002. Reef Fishes: Corals and In­ver­te­brates of The South China Sea. United King­dom: New Hol­land Pub­lish­ers.