nesting sites and behavior of the deep water triggerfish …1)_canthidermis maculata.pdf ·...

Abstract We observed the nesting behavior, eggs, and larvae of the

triggerfish Canthidermis maculata in the Solomon Islandsand in Thailand. We found groups of 3 to over 120 nests,extending from shallow water (6 m) to, more commonly,deeper water (35-40 m). Nests, approximately 1 m diame-ter, sometimes occupied every available space in the softsand bottom between reef contours, coral heads, and/orboulders. Nesting sites were usually found near drop-offsinto deep water (300+ m). We observed active nests withone or two fish near or in the nest. The guarding C. mac-ulata aerated the nest and defended it against other fishspecies, except herbivorous fish which grazed around therim of the nest. Color pattern changes on the head andpectoral region occurred when the fish were courtingand/or defending a nest. We observed courtship behaviornear dusk. Demersal eggs collected from nests hatchedduring the night in containers aboard our vessel, and thepelagic larvae immediately swam upward.

ZusammenfassungWir untersuchten das Brutverhalten, die Eier und die

Larven des Drückerfisches Canthidermis maculata um dieSalomon-Inseln und in Thailand. Wir fanden Gruppenvon 3 bis über 120 Nestern, teils im flachen Wasser (6 m),häufiger aber in tieferem Wasser (35-40 m). Die Nestermit etwa 1 m Durchmesser besetzten manchmal jeden ver-fügbaren Platz im weichen Sandgrund zwischen Riffrän-dern, Korallenköpfen und/oder Geröll. In der Regel befan-den sich die Eiablageplätze in der Nähe von Abhängen intiefes Wasser (300+ Meter). An benutzten Brutplätzenwaren ein oder zwei Fische nahe am oder im Nest zubeobachten. Das bewachende Exemplar von C. maculatabelüftete das Nest und verteidigte es gegen Vertreter ander-er Arten mit Ausnahme von pflanzenfressenden Fischen,die beim Abweiden an den Rändern der Nester geduldetwurden. Während der Balz und beim Verteidigen eines

Nestes traten am Kopf und in der Brustgegend Farbverän-derungen auf. Balzverhalten beobachteten wir in derAbendämmerung. Aus den Eiern, die wir am Boden vonNestern gesammelt hatten, schlüpften über Nacht Larvenin Behältern an Bord unseres Bootes, und die pelagischenLarven schwammen sofort nach oben.

RésuméNous avons observé le comportement de nidification, les

œufs et les larves de Canthidermis maculata aux îles Sa-lomon et en Thaïlande. Nous avons trou vé des groupes de3 à 120 nids, allant d’eaux peu profondes (6 m) et, plussouvent, d’ eaux plus pro fondes (35-40m). Les nids, d’1mde diamètre en viron, occupaient parfois tout l’espacedispo ni ble dans un fond de sable fin entre les profils de ré-cifs, les têtes de corail et/ou les grosses pierres. Les si tes denidification se trouvaient généralement près de tombants((300 + m). Nous avons observé des nids habités avec unou deux poissons dans le nid ou tout près. Le C. maculatade garde oxygénait le nid et le défendait contre d’autres es-pèces de poissons, sauf les poissons herbivores qui brou -taient au bord du nid. Des modifications du patron de col-oration intervenaient sur la tête et la région pectoralequand les poissons paradent et/ou dé fen dent un nid. Nousavons observé les comportements de parade vers le crépu -s cule. Les œufs dé mersaux collectés dans les nids éclosaientde nuit dans des contenants à bord de notre bateau, et leslarves pélagiques nageaient immédiatement vers le haut.

SommarioSono state studiate le modalità di nidificazione, le uova e

le larve del pesce balestra Canthidermis ma cu lata nelle IsoleSalomone e in Tailandia. Sono stati trovati gruppi di 3 adoltre 120 nidi, estendentesi da acque basse (6 m) a, più co-munemente, acque profonde (35-40 m). Nidi, di circa 1 mdi dia metro, a volte invadevano ogni spazio disponibile sulfondale sabbioso tra i contorni della barriera, i coralli, e/o

aqua vol. 21 no. 1 - 15 January 20151

aqua, International Journal of Ichthyology

Nesting sites and behavior of the deep water triggerfish Canthidermis maculata(Balistidae) in the Solomon Islands and Thailand

Eugenie Clark¹, Diane R. Nelson² and Rachel Dreyer¹

1) Mote Marine Laboratory, 1600 Ken Thompson Pkwy, Sarasota, FL 34236, U.S.A. E-mail: [email protected]

2) Department of Biological Sciences, East Tennessee State University, Johnson City, TN, 37614-1710, U.S.A

Received: 18 November 2014 – Accepted: 29 December 2014

i massi. Siti di nidificazione sono stati spesso trovati inacque profonde (oltre 300 m) lungo le pareti spio venti del-la barriera. Sono stati osservati nidi attivi con uno o duepesci vicino o nel nido. Il C. maculata guardiano areava ilnido e lo difendeva da altre specie di pesci, ad eccezione dipesci erbivori che pascolavano intorno al bordo del nido.Variazioni della colorazione della testa e della regione pet-torale si verificavano durante il cor teg giamento e/o la dife-sa del nido. Il corteggiamento era osservato all'approssi-marsi del tramonto. Uova demersali raccolte da nidi schi-udevano durante la notte in contenitori a bordo della nos-tra nave, e le larve pelagiche immediatamente nuotavanoverso l'alto.

INTRODUCTIONThe family Balistidae consists of 11 genera and at

least 40 species (Nelson 2006). In the deepest wa-ter genus Canthidermis, three species are recog-nized (Eschmeyer 1998): Canthidermis maculata(Bloch, 1786), Canthidermis sufflamen (Mitchill,1815), and Canthidermis macrolepis (Boulenger,1888). Canthidermis maculata, originally describedas Balistes maculatus Bloch 1786, was transferred tothe genus Canthidermis by Swainson (1839). Thethree species of Canthidermis are morphologicallysimilar and can be difficult to differentiate. Theydiffer in their geographic distributions: C. macula-ta is circumtropical, but not reported in the RedSea or the Mediterranean (Berry & Baldwin 1966;Masuda et al. 1984; Böhlke & Chaplin 1993; Gar-rison 2005; Allen & Erdmann 2012); C. sufflamen,

ocean triggerfish, is known from the Atlantic andCaribbean (Robins and Ray 1986; Böhlke &Chaplin 1993; Monteiro et al. 2008); and C.macrolepis, largescale triggerfish, is known from theRed Sea, Gulf of Oman, and western Indian Ocean(Baranes 2005; Kuiter 2014; Büttiker et al. (inpress)).Both C. maculata and C. sufflamen have been

called “ocean triggerfish.” In addition C. maculatahas been commonly named “oceanic triggerfish,”“spotted oceanic triggerfish,” “spotted triggerfish,”“white-spotted triggerfish,” and “rough trigger-fish.” “Maculata” means “spotted” and refers tothe color pattern of immature C. maculata. The of-ficial Food and Agriculture Organization andAmerican Fisheries Society common name for thisspecies is “rough triggerfish” (Carpenter 2002;Page et al. 2013). In our series of 114 diving trips (1950-2014) to

study fishes in tropical seas, mainly the Red Sea,Indo-Pacific, and Caribbean, we observed groupnesting activities of several balistid species. Thenesting behavior of shallow water triggerfish (Bal-istapus, Balistes, Melichthys, Sufflamen, Xan-thichthys) have been reported by various authors(Breder & Rosen 1966; Fricke 1980; Kawabe1984; Thresher 1984; Gladstone 1994; Kawase &Nakazono 1992; Ishihara & Kuwamura 1996;Kawase 1998, 2002, 2003a,b; Sahayak 2005; Sim-mons & Szedlmayer 2012). No detailed descrip-

aqua vol. 21 no. 1 - 15 January 2015 2

Nesting sites and behavior of the deep water triggerfish Canthidermis maculata (Balistidae) in the Solomon Islands and Thailand

Fig. 1. Map of Indo-Pacific. Locations of the nesting sites of Canthidermis maculata studied in Thailand and the SolomonIslands marked with red dots.

tion has been previously reported on the nestingactivity of the deep water genus Canthidermis, al-though Randall (1995) made a brief reference tonesting C. macrolepis, and Nellis (1980) publisheda short note on the reproduction of C. sufflamen.Here we report the first detailed description onnesting sites and nesting behavior of C. maculata.Canthidermis maculata is often seen in loose ag-

gregations just offshore on deep slopes and reefs(Myers 1991) or further offshore in open water(Matsuura 2001) or under FADs, where C. macu-lata have been observed in groups of hundreds orthousands (Taquet et al. 2007). They are collectedalong with other triggerfish species and sold atmarkets (Sethi et al. 2011). We first encounteredC. maculata during our expeditions to Papua NewGuinea in 1987 with Bob Halstead aboard the div-ing vessel M/V Telita while studying various sandfishes near coral reefs. Our first observations on C.maculata nesting were when we were studying thedistribution and behavior of tilefishes of the genusHoplolatilus, which inhabit the deeper slopes ofcoral reefs (Clark et al. 1998). Here we report ourconcentrated observations on C. maculata nesting

at two island groups in the Solomon Islands and atone island in Thailand.

MATERIALS AND METHODSPreliminary observations: Initially we made four

research expeditions to the Solomon Sea (1996,1997, 1998, 1999) to study Hoplolatilus (Clark etal. 1998), Pholidichthys (Clark et al. 2006), and Plo-tosus (Clark et al. 2011). During these studies wecame upon small nesting groups of Canthidermismaculata off Fonagho Island in the Russell Islands,SOL and Kicha Island in the New Georgia Islands,SOL. We made one expedition to Thailand in April2000 to study Trichonotus (Clark & Pohle 2007)and came upon a nesting area for C. maculata offone island, Ko Tachai. In June 2014 we made anexpedition to the Solomon Islands specifically tostudy the nesting behavior of C. maculata.Lisa Choquette, who operates Solomon Dive Ad-

ventures in Marovo Lagoon, New Georgia Islands,SOL, had reported to us that she observed nestingC. maculata between 2007 and 2010. From 2011to 2013 she and her skilled diving staff made spe-cial trips to Kicha Island, Male Male Island, and


Eugenie Clark, Diane R. Nelson and Rachel Dreyer

Figs 2a-b. Google Earth images of our study site for Canthidermis maculata in Russell Islands, Solomon Islands. (a) FonaghoIsland circled. (b) Inset: Enlarged view of Fonagho Island, study site off northern tip of island marked with star.

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Mbulo Island to observe and photograph C. macu-lata nesting behavior for us. We include her valu-able information and some of her many pho-tographs in this paper. In 2014 (4-16 June), one ofour experienced research divers (T. Konstantinou)stayed at the Solomon Dive Adventures to makeconcentrated observations on nesting C. maculatawith Choquette and her staff.Study sites: Our study locations in the Solomon

Islands and Thailand are shown in Fig. 1. FonaghoIsland, in the eastern Russell Islands, SOL (Fig. 2a)at 9°05’44.07”S 159°17’50.92”E, is one of severalsmall islands adjacent to a channel that separates itfrom the three easternmost islets. The longest tran-sect (1370 m) of Fonagho Island ran northeast tosouthwest. Our main study site was off the north-ern tip of the island (Fig. 2b), where a flattenedmassive rubble ridge extended northward at depthsof 6.7 to 8.2 m. We also found a few nests on thesouthwest side of Fonagho, on a rubble ridge withsand gullies extending downward to a vertical“drop-off ” wall at 37 m depth.Kicha Island is a small island (400 m W to E)

southeast of Marovo Lagoon in the New Georgia

Islands, SOL (Fig. 3a) at 8°47’21.74”S,158°19’8.56”E. Suitable nesting habitats for C.maculata were found only on the north side ofKicha (Fig. 3b, A1-A5, K1). The rest of Kicha hassheer cliffs, ridges, and drop-offs, and the shallowerareas are storm and surge swept with little sand.Mbaleva Island, SOL, is a small island (440 m

NW to SE) located at 8°28’35.64”S, 158°4’5.84”Ein north Marovo Lagoon, inside Lumalihe Passwhich opens to deep water in the New GeorgiaSound (Fig. 3c). Our study location was near aspur of coral reef off the northeast shore of Mbale-va. The substrate of the area surrounding this reefspur is a series of three coral ridges at 10 to 22 mdepth; between the ridges are sandy valleys 26 to32 m depth. One of the sand valleys leads into asand/coral rubble field at 33-35 m depth. The sur-rounding islands in north Marovo Lagoon (Hanav-isi, Matebako, Lumalihe, Sambuco) were also sur-veyed for triggerfish nests.Ko Tachai Island, Thailand, is an elongated island

(2510 m N to S) located at 9°4’25”N, 97°48’45”Ein the Similan Islands in the Andaman Sea, 47 kmoff the east coast of Thailand (Fig. 4). Approxi-



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mately 500 m from the SSE end of the island wereseven large boulders and numerous smaller ones.These boulders were approximately in a circle andwere separated from the island by a deep (42 m)channel.Underwater observations in the Solomon Is-

lands and Thailand (3328 h, 89 divers): In SOLduring 1996, 1997, 1998, 1999, and 2014, wechartered the live-aboard dive boat, M/V Bilikiki,from which our 78 research scuba divers spent a to-tal of 2399 h underwater. We studied nesting of C.maculata on 23-25 April 1996, 30 April-1 May1997, 2 & 13 April 1998, 10 April 1999, 7-10June 2014, and 16-17 June 2014. In Thailand dur-ing 2000, we chartered the live-aboard dive boat,M/Y Aqua One, from which our group of 41 scubadivers spent 929 h underwater documenting fishbehavior. We studied nesting of C. maculata at KoTachai Island, Thailand on 23-24 April 2000. Inall locations, whenever divers made pertinent observations, they recorded details on a “Kogge

Figs 3a-c. Google Earth images of study sites for Canthidermis maculata in Marovo Lagoon, New Georgia Islands, SolomonIslands. (a) Study sites near Marovo Lagoon marked with arrows: Mbaleva Island marked with red, Mbulo Island markedwith green, Kicha Island marked with yellow, and Male Male Island marked with white. (b) Enlarged view of Kicha Island.Nesting sites (A1-A5, K1) were all found on the north side of the island. (c) Enlarged view of north Marovo Lagoon. MbalevaIsland and Kahaini Island (location of Solomon Dive Adventures) circled in red; nesting site at “Mati Pangera” marked withwhite star.

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sheet” including depth, time, and fish observa-tions, and mapped diagrams on graph paper on theback-side. We’ve been using Kogge sheets (de-signed and supplied by S. Kogge) to describe sig-nificant dive observations since 1988. Additional photographs: Additional photograph-

ic evidence of C. maculata nesting in other parts ofthe Indo-Pacific was provided to us by the experi-enced divers: Lisa Choquette, Bob Halstead,Martha Kiser, and Douglas Seifert. Video recordings: Our videographers (Cho-

quette, Culter, Kiser, Moltzer, J. Nelson, Rubin,Rumiser, Stoll) used several types of video camerasin underwater housings to record C. maculataswimming in large aggregations off the reef and in-dividuals or pairs over their nesting sites insand/coral rubble and on reef ledges. In June 2014,two of our divers (Culter, Rumiser) left their Go-Pro cameras at an active nest on separate days totake photos at set time intervals (1 photo every 5

sec for appx. 30 min) to document nesting behav-ior without divers present.Mapping of nesting sites: Divers measured nests

at study sites using compass readings, surroundingsubstrate, and a measuring tape or a dive slatemarked with centimeters, and they mapped nest-ing sites on Kogge sheets. Occasionally divers esti-mated nest size visually (Fig. 5a). At nest sites atKicha Island, SOL, divers measured nests andmapped them in relation to adjacent coral heads(“bommies”) and coral reefs. At Fonagho Island,SOL, divers used a nylon cord and set up an X andY axis that divided the rubble field into 4 quad-rants (A, B, C, D). The X-axis was laid NNE-SSWwith a 45 m cord; the Y-axis was laid ESE toWNW with a 36.5 m cord. Divers mapped andmeasured the nests in each quadrant (Fig. 5b) us-ing small tags to mark each nest. The nylon cordwas removed from the site, and the tags wereburied for reuse the following year. Nesting sites at



Fig. 4. Google Earth image of study site for Canthidermis maculata in Thailand. Ko Tachai Island, 47 km off Thai coast,marked with white arrow.



Figs 5a-b. Measuring nest area of Canthidermis maculata. (a) Diver R. Amos showing relative size of nest, ~1 meter diameter,K1, Kicha Island, Solomon Islands. Photo by L. Choquette, 2011. (b) Research divers M. Lovejoy, M. Dougherty, and B. A.Johnson measuring distance between nests, Fonagho Island, Solomon Islands. Photo by D. Nelson, 1997.

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Kicha Island and Fonagho Island were re-visitedduring subsequent trips to check for C. maculatanesting activity. At Mbaleva Island, SOL, diversused a surface buoy marker (red float) to mark thedive location and mapped nests in relation to buoylocation and coral reef contours. Kogge sheets werecompiled to map the Mbaleva nesting area. At KoTachai Island, Thailand, divers measured C. macu-lata nests and mapped them in relation to large,submerged boulders.

Collection and observation of eggs: When one ortwo C. maculata were observed within a nest,divers looked for eggs. If egg clusters were visibleon close examination, a small portion (<10%) wascollected from the nest, brought to the surface in aplastic bag, put into a bowl on the vessel, and usuallystirred occasionally. The eggs were examined in seawater under a magnifying glass. A few mL of surfacesea water were added to the bowl in the evening, andthe eggs were monitored during the night.Phases of the moon: We determined moon phase

in relation to dates and locations of nesting C. mac-ulata at: http://www.timeanddate.com/world-clock/moonrise.html.Abbreviations: cm: centimeter; diam: diameter;

FAD: fish aggregating device; h: hours; mL: milli-liters; PNG: Papua New Guinea; s: seconds; SL:standard length; SOL: Solomon Islands; TL: totallength.

RESULTS Underwater observations in Solomon Islands:

At Kicha Island (A1-A5, K1), SOL, 36 diversaboard the M/V Bilikiki in 1996, 1997, and 2014spent 378 h underwater surveying the area for C.



Table I. Measurements of seven ovoid nests of Canthider-mis maculata, K1, Kicha Island, Solomon Islands, 23 April1996.

Water depth Rim to Rim Nest Height Nest # (m) Diameter (cm) (cm)

1 8,8 84 x 97 10

2 9,1 70 x 98 11

3 9,4 77 x 103 9

4 10,6 98 x 104 14

5 11,2 90 x 110 14

6 9,1 10 x 109 _ a

7 8,8 _ a _ a

Fig. 6. Location of 10 Canthidermis maculata nests at K1, Kicha Island, Solomon Islands, 23 April 1996. Divers L to R: R.Petzold, E. Clark, M. J. Stoll.



Fig. 7. Location of 24 Canthidermis maculata nests at northern tip of Fonagho Island, Solomon Islands, 30 April 1997.

maculata and their nests. On 23 April 1996, wefound seven C. maculata nests on a coral/sand/rub-ble ledge on the northwest side of Kicha Island(K1), SOL, with a pair of C. maculata on a nest ata depth of 8.8 m. The measurements of these nestsare shown in Table I and the corresponding loca-tion of these nests are shown in Fig. 6 (nests 1-7).At nests 6 and 7, fish were present and not dis-turbed for data. Nest 7 had one triggerfish (~65-70cm TL) that stayed in the nest and one triggerfish(~60 cm TL) that came into the nest, but a thirdtriggerfish (~55 cm TL) repeatedly swam past thenest approximately 3 m above it. That afternoonwe found three more nests for a total of 10 nests(Fig. 6). Over 35 C. maculata were observed at 12m depth along the drop-off with other groups offish (Naso hexacanthus, Caesio sp., Caranx sp.). On the following day at the same site (24 April

1996, K1), one diver videoed 30+ C. maculataswimming along the drop-off at 12+ m depth.During a dusk dive at K1, three divers looked forbut did not observe courtship or egg laying.In 1997 we returned to K1, Kicha Island. On 25

April we observed no adult C. maculata and nonests were visible. On 4 May, we dived at K1 to ob-serve the tilefish Hoplolatilus fronticinctus and ob-served 29 C. maculata swimming in the water col-umn at ~12 m depth with schools of Naso hexacan-thus, Caranx latus, and Caesio sp. There were nosigns of Canthidermis maculata nesting. In June 2014, our divers surveyed Kicha Island

and the nearby islands, Mbulo and Male Male. Noactive C. maculata nests were reported during anydives. On 18 June 2014 we dived at K1 (morning)and A1-A5 (afternoon) and observed 40+ inactiveC. maculata nests throughout this area at depthsover 10 m, but no triggerfish were observed. Astorm prevented diving at these islands 19-20 June2014. On 21 June 2014, we dived the north side ofKicha Island (A1-A5, K1) and observed 25+ C.maculata at 14 m depth over 10 m away from thereef at K1. We also observed 40+ Naso hexacanthusover the reef at A4, A5, and K1, and 200+ Caesio

sp. in the water column around the reef. That af-ternoon (21 June 2014) we observed five C. macu-lata at about 30 m depth but no nests at Mbulo Is-land and observed no C. maculata or their nests atMale Male Island.On 17 May 2014, Daniela Tombion, dive master

aboard M/V Bilikiki, observed 5 active C. maculatanests at Karanjou Island, SOL in the northeasternpart of Marovo Lagoon (pers. comm.). On 17 June2014, we observed these inactive C. maculata nests,which were structured with large coral rubblearound the perimeter of the nest at depths from 13to 21 m. No C. maculata were visible in the area.At Fonagho Island, SOL aboard the M/V Bilikiki

in 1997, 1998, 1999, and 2014, 43 divers spent129.2 h observing and recording behavior of nest-ing C. maculata. On 30 April 1997 we observed 15nests, four with a C. maculata over each nest. Nestswere in a massive coral rubble field in shallow wa-ter (6.7-8.2 m) off the northern tip of the island.On 1 May 1997 we mapped the locations of 24nests (Fig. 7). The number of C. maculata nestscounted in quadrants A, B, C, and D were 6, 3, 8,and 7, respectively. All nests were approximately 1m diam. Only one nest (B1) was active with a sin-gle fish over the nest. No eggs were observed inany nests at this site. Divers did not find any C.maculata nests off the rubble shelf down to 70 mdepth. On 7 May 1997 we checked the northerntip of Fonagho and found that the entire nest siteappeared abandoned. No C. maculata were seenover the nest area, however, on the same dive, over50 C. maculata were observed southwest of Quad-rant C in relatively shallow water (6-9 m).We checked the north side of Fonagho for Can-

thidermis maculata activity thrice in April 1998,once in April 1999, and once in June 2014. On 2April 1998, the nests contained rubble and no C.maculata were observed over any nests. We saw sixC. maculata swimming along the drop-off awayfrom the reef. No tags buried the previous year(1997) were found. On 13 April 1998 we notedthe remains of one C. maculata nest on the rubble



Table II. Measurements of three nests of Canthidermis maculata, Mbaleva Island, Solomon Islands, 15 June 2014.

Outer rim to Top of rim to Inner rim to Rim height Nest height Nest # Depth (m) outer rim (cm) top of rim (cm) inner rim (cm) (cm) (cm) 1 23 168 132 86 8 15

2 28 229 168 102 15 23

3 29 183 132 66 8 18

shelf but the rest of the nests had become filledwith coral rubble; no C. maculata were observed inarea. On 18 April 1998, 10 April 1999, and 6 June2014 divers surveyed the north side of Fonagho Is-land for evidence of C. maculata or nesting activity,but we found none. In June 2014, the three giantclams shown in Fig. 7 were no longer present,probably removed for their meat and shells.On 22 April 1997 we dived off the southwest tip

of Fonagho Island and found six concavities at 21m depth near a coral reef ledge that appeared to beinactive C. maculata nests; six C. maculata were ob-served in the water column nearby. On 6 June2014 we dived off the southwest tip of Fonagho Is-land but observed no C. maculata or remains oftheir nests.At Mbaleva Island, SOL in 2014, 32 divers

aboard the M/V Bilikiki spent 132 h underwatersurveying the area for C. maculata and their nests.On 5 June 2014 Lisa Choquette reported that atthe “Mati Pangera” dive site one of her divers (H.Amos) counted 93 C. maculata nests, with one ortwo fish over each nest. On 7 June 2014 one of us(RD) found a relatively deep (31 m) sand valleywith eight C. maculata nests, three of which hadone fish over each nest. This sand valley led into aflat sand/coral rubble field at 34 m depth with 10+C. maculata nests; only one nest was active with asingle fish over it. This open area was not observedby Choquette’s dive team, and these nests were inaddition to the 93 active nests observed on 5 June2014. On 8, 9, 16, and 17 June 2014, we returnedto this coral/rubble field and mapped this areawhen no fish were present and located 27 C. macu -



Fig. 8. Location of 90 (out of 120+) Canthidermis maculata nests at “Mati Pangera” dive site, Mbaleva Island, Solomon Is-lands, June 2014. Depths indicated in meters; location of cleaning station observed 9 June 2014 marked with yellow star.

lata nests (Fig. 8). The nests of C. maculata werenot found shallower than 13 m. Nests of the titantriggerfish, Balistoides viridescens, were observed onthe coral reef spur at 3.7 to 15.2 m depth. On 8 June 2014, Choquette led our divers to the

“Mati Pangera” dive site. Nests were observed with-in three sand valleys (Fig. 8), and approximatelyone-third of the original 93 nests were still active.When one of our divers (Culter) recorded behavior(1 photo/5 s for 34 min) of one C. maculata overits nest with no divers present, the fish circledwithin the perimeter of the nest clockwise andcounter-clockwise, approximately one rotationevery 10 s. The fish would occasionally leave the nest but quickly return to the nest and contin-ue circling. No courtship or mating was docu-mented. On 9 June 2014, approximately 100 C. maculata

were observed swimming in the water column

away from the reef at a depth of 28 m. On 15 June2014, one of our divers (T. Konstantinou) took de-tailed measurements of three C. maculata nests,shown in Table II. Rim height is the height fromthe surrounding substrate to the top of the rim;nest height is the height from the bottom of thenest to the top of the rim. Bottom time, current,and weather conditions prevented further detailedmeasurements of this area. On 8 June 2014 we surveyed Hanavisi Island,

southeast of Mbaleva, and on 9-10 June 2014 wesurveyed the east and west side of Lumalihe Pas-sage, Lumalihe Island and Sambuco Island (Fig.3c). No C. maculata fish, nesting activity, or suit-able nesting habitats were observed around theseislands. Matebako Island, a tiny island northwest of

Mbaleva Island (Fig. 3c), was surveyed by T. Kon-stantinou, L. Choquette, and her divers. This is-



Figs 9a-b. Study site for Canthidermis maculata off Ko Tachai Island, Thailand. (a) Location of 9 nests around submergedboulders, 24 April 2000. (b) Inset: Ko Tachai Island with depth contours; nesting site at southern tip of island marked withpurple star.

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land has habitat suitable for nesting triggerfish, butno C. maculata adults or nests were observed.On 16 June 2014 we returned to Mbaleva Island

and observed three C. maculata near cabbage coral,Turbinaria sp.: one fish was over a nest at 16 mdepth and two fish were in the water column at 14- 15 m depth; a second nest was visible ~2 m away(Fig. 8). That afternoon, a 177 m logging freighterfrom China, Qi Sheng, navigated through Lumalihe

Passage into north Marovo Lagoon and anchoredapproximately 200 m from our surface buoy mark-er and prevented our further diving. We made onedive that afternoon and one dive the next morningto refine our map of the large nesting area (Fig. 8),but after only two dives we were forced to leave thearea for divers’ safety as tugboats were bringing car-go to the Qi Sheng at irregular intervals. Underwater observations in Thailand: At Ko



Table III. Measurements of nine circular or ovoid nests ofCanthidermis maculata, Ko Tachai Island, Thailand, 23April 2000.

Water depth Outside Inside Nest # (m) diameter (cm) diameter (cm)

1 29,3 105 x 120 46 x 54

2 29,6 165 x 168 75 x 78

3 30,2 183 120

4 30,2 169 120

5 31,1 150 x 165 75 x 75

6 32,3 150 x 165 69 x 69

7 33,5 172 105

8 33,5 172 134

9 33,5 166 133

Fig. 10. Active Canthidermis maculata nest in sand rubblesubstrate, ~1 m diameter at top of rim, Milne Bay Province,Papua New Guinea. Photo © B. Halstead, 1998.

Fig. 11. Inactive, abandoned Canthidermis maculata nest, C2, Fonagho Island, Solomon Islands. Photo by D. Nelson, 1997.



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Tachai Island, Thailand in April 2000, 24 diversaboard the M/Y Aqua One spent 40.4 h underwaterto observe C. maculata. On 23 April 2000, wefound a group of nine C. maculata nests at the baseof the boulders only on the open ocean side neardeep water (Fig. 9a) off the southern end of KoTachai Island (Fig. 9b). Measurements of thesenests are shown in Table III. One of the crew divers(Nikom) reported that he had regularly observedC. maculata nesting at this site for the last sevenyears (1993-1999).Canthidermis maculata nesting observations:

On 7 April 1997, one of our divers (L. Benveniste)observed 5 C. maculata nests and 9 individualsnear an unchartered reef (“Canyon Reef” at 9°58.578’S, 150° 49.827’E) located near Duchess Is-land in Papua New Guinea. On 24 May 2012 inPNG, Bob Halstead and Martha Kiser (aboard the

M/V Golden Dawn) observed 100+ C. maculata atMeehan’s Bommie, an isolated offshore reef ~ 200m long in the Dampier Strait between West NewBritain and Umboi Island (5° 51.873’S, 148°23.287’E). Over 20 active C. maculata nests wereobserved at depths from 12 to 16 m.On 23 Sept 2014 off Cocos Island, Costa Rica (5°

33’29”N, 87° 2’53”W), Douglas Seifert and EmilyIrving observed 60+ C. maculata near a channelseparating Cocos Island from Manuelita Island.Observed nesting activity included 12+ depressionsin the sand at 21 m depth, and one C. maculatawas above one nest. Underwater observation summary: Year-round

nesting activity (Table IV) is shown by our data anddata collected by L. Choquette. The majority of theseevents occurred at Kicha Island, SOL (Fig. 3b). Canthidermis maculata nests: Nests of C. ma -

Figs 12a-c. Group nesting of Canthidermis maculata. (a) Coral reef with 6 nests, K1, Kicha Island, Solomon Islands. Photoby L. Choquette, 2011. (b) One of us (RD) over coral rubble field off Mbaleva Island, 34 m depth. Six of 27 nests visible.Photo by A. Konstantinou, 2014. (c) Two “shelf nests” visible; nest at right on higher level, A2, Kicha Island, Solomon Is-lands. Photo by L. Choquette, 2011.

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Figs 13a-b. Canthidermis maculata cleaning nest, Ko Tachai Island, Thailand. (a) Individual with dark region over eye and atpectoral fin base picking up coral rubble with mouth. (b) Individual in Fig. 13a with dark saddle mark across its snout spit-ting coral rubble out of its nest. Photos by A. Snowhite, 2000.

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culata appeared as white, shallow concavities insand and small rubble, with coarser rubble aroundthe rim. They were either highly structured with asand/fine rubble rim (Fig. 10) or unstructuredwithin course coral rubble. Once nests were aban-doned by C. maculata, their structure was difficultto determine (Fig. 11). Nests were sometimes lo-cated in close groups utilizing every available patchof sand (Figs 12a-b) and constructed along thecontours of the reef (Fig. 12c). The circular orovoid structure of a C. maculata nest was createdand maintained by the fish blowing water at sandparticles or picking up coral rubble with its mouthand spitting it outside the nest (Figs 13a-b). LisaChoquette observed C. maculata using nests re-peatedly throughout the year at Kicha Island (Fig.3b. A1-A5, K1) (pers. comm.). At the Islands of Kicha, Mbulo, and Male Male,

SOL, the structure of the nest was related to depth.Shallow (<15 m) nests surrounded by sand and/orfine coral rubble had definite form and structure.Deeper (> 15 m) nests on sand slopes or ledgeslacked structure and were confirmed as a C. macu-lata nest when one or two fish were observed overthe sandy area. In August 2014, Lisa Choquette returned to our

study site off Mbaleva Island, SOL and observedthat in areas subject to current, C. maculata nestsdid not have a wall but were discernable by a slightdepression in the sand (pers. comm.). The few C.maculata nests in areas protected from the currentretained their nest walls. Color changes of adult Canthidermis maculata:

Temporary, variable dark facial and pectoral mark-ings were visible on nesting C. maculata. In some,prominent facial markings appeared with pale pec-



Table IV. Nesting activity of Canthidermis maculata observed in the Pacific, arranged by time of year, 1996-2014.

Month Day Days Year Number of Number Depth of Location to Full Moon C. maculata of Nests Nests (m)

Feb 23 -13 2012 100+ 111 4 - 30 A2-A5, Kicha, SOL 25 -11 2012 3+ 3 11, 27, 32 A2-A5, Kicha, SOL

April 7 +13 1997 10+ 6 - 9 32 Canyon Reef, PNG 21 +3 2011 2 43 7 - 30 Kicha, SOL 23 -10 1996 35+ 10 8 - 11 K1, Kicha, SOL 24 +5 2000 10+ 9 29 - 34 Ko Tachai, Thailand 30 +7 1997 25+ 24 6 - 10 Fonagho, SOL

May 6 -11 2011 4 1 15 Kicha, SOL 17 +2 2014 20+ 5 13 - 21 Karanjou, SOL 21 -4 2013 20+ 1 not recorded K1, Kicha, SOL

24 -12 2012 100+ 20+ 12 - 16 Meehan's Bommie, PNG

June 5 +1 2012 100+ 100+ 15 - 40+ NW Male Male, SOL

5 -8 2014 100+ 93 12 - 37 Mbaleva, SOL

7 -6 2014 50+ 30+ 21 - 35 Mbaleva, SOL

12 +8 2012 20+ 2 not recorded K1, Kicha, SOL

16 +3 2014 3 2 16 Mbaleva, SOL

July 21 +6 2011 8 10+ not recorded NW Male Male, SOL

22 +7 2011 25+ 25 10 - 30 Kicha, SOL

Aug 6 +4 2012 20+ 17 9+ A4-A5, Kicha, SOL

7 +5 2012 not recorded 5 33 - 40 Male Male, SOL

7 +5 2012 5+ 5 24 - 30+ A5-K1, Kicha, SOL

Sept 23 +15 2014 60+ 12 21 Cocos Island, Costa Rica

Oct 2 +2 2012 200+ 100+ 10 - 28 A3-A5, K1, Kicha, SOL

Nov 2 -9 2011 100+ 50+ 20 - 45 A1, Kicha, SOL

4 -7 2011 1000+ 500+ 7 - 45+ A2-A5, K1, Kicha, SOL

6 -5 2011 100+ 51 12 - 30+ A1-A5, K1, Kicha, SOL

14 +3 2011 not recorded 5 21 - 37 Kicha, SOL



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Figs 14a-e. Facial and pectoral markings of Canthidermis maculata associated with nesting. (a) Individual over its nesting areawith prominent facial and pale pectoral fin markings, A5, Kicha Island, Solomon Islands. Photo by L. Choquette, 2011. (b)Individual over its nest with dark pectoral and pale facial markings, Ko Tachai Island, Thailand. Photo by N. Sumanate,2000. (c) Individual over its nest with prominent facial and pectoral markings, Ko Tachai Island, Thailand. Photo by A.Snowhite, 2000. (d) Overall view of nest in coral rubble, individual on nest with prominent facial and pectoral markings;second nest visible in background, K1, Kicha Island, Solomon Islands. Photo by R. Petzold, 1996. (e) Individual over its nestwith moderate facial and prominent pectoral markings; second nest visible in background, Male Male Island, Solomon Is-lands. Photo by R. Amos, 2012.

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Figs 15a-b. Close-ups of facial and pectoral markings on Canthidermis maculata in the water column. Note variation in facial(solid vs. patchy) and pectoral (shape of spot) markings between individuals in (a) and (b), Meehan’s Bommie, Papua NewGuinea. Photos © B. Halstead, 2012.

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toral markings (Fig. 14a). Others had prominentpectoral markings without facial markings (Fig.14b) or both facial and pectoral markings wereprominent and highly visible on the fish over a nest(Figs 14c-d). On some individuals a white line ap-peared in front of the eye (Fig. 14e), but this area al-so appeared black in other individuals. Facial mark-ings appeared either solid (Fig. 15a) or patchy (Fig.15b). Pectoral markings typically appeared as a sol-id circle or square at the base of each pectoral fin. Over a nest, C. maculata typically displayed pec-

toral markings (Fig. 16a). When two C. maculatawere observed on a nest, both fish displayed mod-erate to prominent facial and pectoral markings(Fig. 16b). In the water column above the nest, C.maculata were typically uniformly gray, while thoseover the nest displayed some variance of facial andpectoral markings (Fig. 16a). Video of nesting C.maculata taken by M. Kiser on 24 May 2012 inPNG demonstrated that markings appear and fadein as little as a few seconds. Color changes were also noted on C. maculata be-

ing cleaned by other fish. On 9 June 2014 at Mbal-eva Island, SOL (Fig. 8), one of our divers (M.Kiser) videoed three C. maculata being cleaned bywrasses Labroides dimidiatus and a juvenile Corisbatuensis. These triggerfish became uniformly darkgray and positioned themselves head up in the wa-ter column; two C. maculata opened their opercu-lum and allowed L. dimidiatus to clean their gills.On 23 Sept 2014 at Cocos Island, Costa Rica,Douglas Seifert observed varying numbers (30 to60) of C. maculata being cleaned by butterflyfish,angelfish, and wrasse. All C. maculata observed atthis cleaning station became dark gray (Fig. 17). Defensive behavior and egg predation: When an

individual C. maculata guarded a nest with eggs,this triggerfish displayed defensive behavior, whichincluded raising its first dorsal fin trigger (Fig. 18a)and chasing threatening fish away from the nest. AtK1, Kicha Island, SOL on 23 April 1996 from16:46 to 17:55, one of us (EC) observed C. macu-lata guarding its nest and chasing away other fishthat came into the nest: Halichoeres hortulanus,Balistapus undulatus, Pseudobalistes flavimarginatus,Balistoides viridescens, Chaetodon auriga, Lutjanusgibbus, Mulloidichthys vanicolensis, and Heniochussp. The defending triggerfish did not chase acan-thurids (including Zebrasoma velifer) or 20+ sur-geonfishes; these herbivorous fish fed around therim of the nest. However, Canthidermis maculatawas observed being driven away from its nest by a

much smaller orange-lined triggerfish, Balistapusundulatus. At Kicha Island, SOL, Lisa Choquetteobserved C. maculata defending the eggs inside itsnest against the titan triggerfish, Balistoides viri-descens (Fig. 18b) (pers. comm.). When C. maculata abandoned its nest, we ob-

served predation on the eggs by various fish. AtKicha Island, SOL, the orange-lined triggerfish,Balistapus undulatus, and the aggressor (Fig. 18b),Balistoides viridescens, predated on the C. maculataeggs (Fig. 19a). At Fonagho Island, SOL, divers in-advertently drove a guarding C. maculata off itsnest, and the goatfish Parupeneus multifasciatus andthe checkerboard wrasse Halichoeres hortulanus atethe C. maculata eggs (Fig. 19b).Observation of Canthidermis maculata embryos

and larvae: Egg masses were collected at K1, KichaIsland on 25 April 1996 at 12:30, at Ko Tachai Is-land on 23 April 2000 at 16:30, at Mbaleva Islandon 7 June 2014 at 11:00, and at Mbaleva Island on8 June 2014 at 09:10. Once these eggs were put in-to a glass or bowl, over 500 were easily seen on thebottom of the container. The eggs measured 0.7mm in diameter (Fig. 20), with no oil droplet visi-ble but developing embryos were present.Three of the egg collections were stirred occasion-

ally in the evening. These eggs began hatching ear-ly the next morning (03:35-03:50, Fig. 21a) andcontinued hatching for about two hours. Larvaewere almost 2 mm TL (Fig. 21b). Eggs collected on8 June 2014 were not stirred and no eggs hatched.Aggregations of Canthidermis maculata: Large

(500+) aggregations of C. maculata have been ob-served in open water throughout its range (Fig.22a). Lisa Choquette observed over 40 aggrega-tions of C. maculata (20-300 individuals) duringApril 2011-September 2014 around the islands ofKicha, Male Male, Mbulo, and Mbaleva, but nonesting behavior was observed (pers. comm.). Anaggregation of over 75 C. maculata was photo -graphed in Raja Ampat, Indonesia, at depths from18 to 34 m (Fig. 22b), but no nesting behavior wasobserved (D. Seifert, pers. comm.). Thousands of C. maculata were observed in

Kimbe Bay, Lolobau Island, PNG on 9 Aug 2012.Nests were in the area, but no obvious nesting ac-tivity was observed (B. Halstead, pers. comm.).This was the largest aggregation of C. maculata onmultiple reefs that Halstead had ever observed.

DISCUSSION AND CONCLUSIONSTriggerfishes of the family Balistidae have short



a

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but strong jaws with eight long protruding incisi-form teeth in the outer row of both jaws and an in-ner row of six smaller teeth which buttress thesmaller ones in front (Randall 1996). Large trigger-fishes can be dangerous to divers, especially whenthey are nesting (Randall & Millington 1990;Lanelli 2008). There are no records of any Canthi-dermis species attacking or biting a diver, and ourdivers have never been attacked by C. maculata. Wefound C. maculata to be shy and easily driven offits nest by approaching divers, even when we tooksamples of their eggs. Nesting of Canthidermis maculata: Throughout

the year, C. maculata occupy the same nesting sites,but it is unknown if the same individuals return tothe site for each nesting event. Due to our limitedcourtship data, we were unable to classify this trig-gerfish into one of the mating systems described byEmlen & Oring (1977). Nesting in C. maculata has

been observed during all phases of the moon (TableIV). The largest observed nesting events were inFebruary (2012, Kicha), June (2014, Mbaleva) andNovember (2011, Kicha). At the southern end ofKo Tachai Island, Thailand, C. maculata are proba-bly nesting all year-round but the M/Y Aqua Oneonly dives at this location once each year before themonsoon (Nat Sumanate, pers. comm.).We have observed balistids cleaning their nests by

blowing water into the sand and moving coral rub-ble with their mouths (unpublished data). Similarnest-building behavior has been reported in thegray triggerfish, Balistes capriscus (MacKichan &Szedlmayer 2007). The orange-lined triggerfish,Balistapus undulatus, and the yellowmargin trigger-fish, Pseudobalistes flavimarginatus, have been ob-served excavating nests by the fish turning on itsside and flexing its body laterally while flapping itsdorsal and caudal fins (Lobel & Johannes 1980).



Figs 16a-c. Two individuals of Canthidermis maculata over nest. (a) Canthidermis maculata with dark pectoral marking overits nest in foreground; second C. maculata with moderate pectoral marking associated with nest, Mbaleva Island, SolomonIslands. Photo © D. Seifert/www.douglasunderwater.com, 2014. (b) Both individuals with facial and pectoral markings overnest, Male Male Island, Solomon Islands. Photo by L. Choquette, 2012. (c) Individual with facial and pectoral markings overnest, unmarked individual above nest, other individuals and nests visible, Meehan’s Bommie, Papua New Guinea. Photo ©B. Halstead, 2012.

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Canthidermis maculata does not exhibit sexual di-morphism (Moore 1967), and we were unable todifferentiate male and female C. maculata when div-ing. Based on the few courtship behaviors observedby our divers and L. Choquette and her divers, islikely that only females guarded the eggs. Maternal-only egg care is reported in the white-banded trig-gerfish, Rhinecanthus aculeatus (Kuwamura 1997),the halfmoon triggerfish, Sufflamen chrysopterus(Ishihara & Kuwamura 1996), and the masked trig-gerfish, Sufflamen fraenatus (Kawase 2003b).At Kicha Island and Mbaleva Island, SOL, there

appears to be a boundary between the shallowernests of the titan triggerfish, Balistoides viridescens,and the relatively deeper nests of Canthidermismaculata. The guarding behavior of C. maculataappears to be ineffective for defending its nest/eggsagainst B. viridescens. Therefore the minimumnesting depth of C. maculata may be limited bynesting B. viridescens, which become territorialduring nesting (Randall & Millington 1990; Ran-dall et al. 1997; Lanelli 2008). Nesting of other Canthidermis sp.: Nesting Can-

thidermis sufflamen have been observed in theCaribbean. Nellis (1980) observed one C. suffla-men over a ~60cm nest in a sandy plain at approx-

imately 18 m depth, within a half mile of a steepdrop-off which surrounded St. Croix, US VirginIslands. Although Nellis dived this area for 30years, he did not witness Canthidermis nesting be-havior again (pers. comm.). Michael Feeley (NPS)has occasionally noted nesting C. sufflamen at 30-34 m depth in the Dry Tortugas (pers. comm.).Michael Burton (NOAA) has observed large(100+) groups of C. sufflamen at Riley’s Hump,Tortugas Ecological Reserve, Dry Tortugas andnesting events were documented on 20 May 2003,9 June 2009, and 10 July 2009 (pers. comm.). There are few reports of nesting C. macrolepis.

This triggerfish species “comes to the shallows tolay demersal eggs in a nest in sand” (Randall 1995).A pair of C. macrolepis was photographed over anest at 7 m depth at Fahl Island, Oman (Gill &Randall 1997). One of us (EC) observed one Can-thidermis nest in the north Red Sea in the 1970s;based on species distribution this observation wasprobably C. macrolepis.Nesting activity compared to lunar cycles: The

nesting of triggerfish and other reef fish has oftenbeen thought to be associated with the phase of themoon. Thresher (1984) suggested a correlation be-tween the lunar cycle and spawning of balistids.



Fig. 17. Canthidermis maculata in “dark phase” near cleaning station at Cocos Island, Costa Rica. Photo © D. Seifertwww.douglasunderwater.com, 2014.



Figs 18a-b. Defensive behaviors of Canthidermis maculata, A2, Kicha Island, Solomon Islands. (a) Individual displayingmodified first dorsal fin “trigger.” (b) C. maculata (on left) defending its nest against Balistoides viridescens (titan triggerfish).Photos by L. Choquette, 2011.

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Figs 19a-b. Predation on Canthidermis maculata eggs in nest. (a) Balistapus undulatus (orange-lined triggerfish) and Balis-toides viridescens (titan triggerfish) ate eggs after C. maculata abandoned its nest following encounter with titan triggerfishshown in Fig. 18b, A2, Kicha Island, Solomon Islands. Photo by L. Choquette, 2011. (b) Parupeneus multifasciatus (manybargoatfish) and Halichoeres hortulanus (checkerboard wrasse) ate eggs after divers inadvertently drove C. maculata away fromits nest, Fonagho Island, Solomon Islands. Photo by D. Nelson, 1997.

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Fig. 20. Canthidermis maculata eggs (~0.7mm) collected from nest on 7 June 2014 off Mbaleva Island, Solomon Islands,photo by J. R.-Queralt.

Figs 21a-b. Larvae of Canthidermis maculata. (a) One of us(RD) observing C. maculata eggs hatching at 4:30am on 8June 2014. Hatched larvae circled in yellow. Photo by W.Hall. (b) C. maculata larva (~2mm) that hatched 8 June 2014.Photo © J. Ares.a

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Figs 22a-b. Aggregations of Canthidermis maculata. (a) Large aggregation of over 300 C. maculata over deep water, Konacoast, Hawai’i. Photo © M. Ushioda/www.coolwaterphoto.com, 2012. (b) Aggregation of over 75 C. maculata near reefslope, Misool, Raja Ampat, Indonesia. Photo © D. Seifert/www.douglasunderwater.com, 2011.

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Figs 23a-b. Aggregations of Canthidermis macrolepis in the South Red Sea. (a) Aggregation of over 30 C. macrolepis in watercolumn near a drop-off into deep (200+ m) water. (b) Loose aggregation of over 20 C. macrolepis and over 20 Acanthurus ni-gricans (black surgeonfish) intermingled over sand/coral rubble substrate. Photos by D. Neslon, 1995.

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Figs 24a-b. Early illustrations of Balistes maculatus. (a) Bloch 1786, illustration of immature type specimen with dark spots;specimen size and collection locality not recorded. (b) Bleeker 1865, specimen with white spots, 34 cm SL, collected off Ok-inawa, Japan, appears to be immature.

We found no correlation between the lunar cycleand the nesting periods of Canthidermis maculatawe report in this paper (Table IV).Eggs: Eggs of triggerfish in the family Balistidae

can incubate up to 55 hours (Lyczkowski-Shultz &Ingram 2003). Eggs of balistids have been observedhatching at night (Lobel & Johannes 1980; Ishi-

hara & Kuwamura 1996). The C. maculata eggs wecollected at K1, Kicha Island, SOL were probablylaid at 6-7 pm the day before, and we estimate thatthe incubation time of C. maculata eggs is 33-36 h.It is likely that C. maculata larvae hatch and rise tothe surface during the middle of the night to es-cape detection by potential predators.

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Larvae and Juveniles: Larvae and juveniles of C.maculata, size range 11-24 mm SL, have beenfound in floating Sargassum in the Gulf Stream(Casazza & Ross 2008) and the northern Gulf ofMexico (Hoffmayer et al. 2005), however this trig-gerfish is not always associated with the Sargassumcommunity (Dooley 1972). Juvenile (~30 mm TL)C. maculata collected in Caribbean waters (78ºW14°N) were placed in a small aquarium and for-aged on small shrimp present in Sargassum once itwas added to the aquarium (Alevizon 1976). Larvaland juvenile C. maculata and C. sufflamen havebeen found in the North Atlantic (Lyczkowski-Shultz & Ingram 2003) and a juvenile of bothspecies has been collected in separate trawls in wa-ters over 2300 m deep (Schwartz 2006). “Small,inedible” C. maculata accounted for 2551 of 2741lbs and 2293 of 2493 lbs caught in two trawlinghauls in December 1928 in the Arabian Sea(Venkataraman & George 1964). Juvenile C. ma -cu lata (17.1-82.9 mm SL) were found in Naka-gusuku Bay, Okinawa Island, Japan (Ohta &Tachihara 2004). Nishida et al. (2008) collecteddrifting seaweed during 32 non-consecutive

months and found one C. maculata (40 mm SL)and Goldstein et al. (2014) examined 242 debrisobjects and found one “juvenile” C. maculata, sug-gesting these triggerfish are not evenly distributedthroughout their range and may be selective intheir association with types of rafting community.Once C. maculata larvae hatch and swim upward,they are likely taken by wind and currents (Alevi-zon 1976) into the open ocean to mature, formingaggregations when they are large juveniles andadults.Aggregations: Canthidermis maculata have been

found in the open ocean and around floating de-bris (Allen & Robertson 1994). Large aggregationsof C. maculata are well known. Taquet et al. (2007)observed aggregations up to 100 around FADs intropical waters near Reunion Island and massiveaggregations up to 5000 around FADs in equator-ial waters in the Seychelles.Bycatch in the tuna seine purse fisheries at FADs

in the Atlantic, Indian, and Pacific Oceans oftencontains C. maculata (Dagorn et al. 2013). In atrawl operated at 60-80 m depth in the Bay of Ben-gal, 17% of the triggerfish caught were C. maculata



Fig. 25. Immature Canthidermis maculata lectotype ~24 cm TL, ZMB 5904 in Museum für Naturkunde, Berlin, Germany.White spots visible on body and bases of dorsal, anal, and caudal fins. Photo by A. Konstantinou, 2013.



Figs 26a-b. White spotted patterns on freshly caught immature specimens of Canthidermis maculata. (a) Specimen 9 cm SL,collected 13 March 1988 in the Maldives. (b) Specimen 26.3 cm SL, collected 3 June 1975 off Okinawa, Japan. Photos byJ. E. Randall.

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(Sethi et al. 2011). Over 250 C. maculata (290-375 mm TL) were caught in the Arabian Sea in agill-net at Cochin Fisheries Harbour, Cochin, In-dia (Sobhana et al. 2013). Canthidermis maculata(30-50 cm SL) are regularly found around floatingobjects in all the oceans (Parin & Fedoryako1992). Large aggregations of C. maculata formaround FADs in Indian Ocean (Taquet et al. 2007)and the Western and Central Pacific Ocean (Bailey1985; Leroy et al. 2012). Adult C. maculata mayremain in the same aggregation in the open ocean,move to relatively shallow water to nest, and returnto open water as a group. Canthidermis macrolepis also forms large aggrega-

tions. On 14 June 1990, one of us (EC) encoun-tered a large (150+) aggregation of C. macrolepis offthe Hanish Islands, Yemen. These triggerfish weremixed with a group of 100+ whitecheek surgeon-fish, Acanthurus nigricans, and both species contin-uously encircled three divers like a merry-go-roundfor at least ten minutes. In 1995 in the South RedSea, one of us (DRN) observed C. macrolepis in ag-gregations near a drop-off into deep water (Fig.23a) and in loose, mixed aggregations with A. ni-gricans over the sandy substrate (Fig. 23b). Nonesting behavior of C. macrolepis was observed.Canthidermis maculata has been observed in large

aggregations with other species. In the Solomon Is-lands, the majority of C. maculata aggregationswere observed with the sleek unicorn surgeonfish,Naso hexacanthus. The benefit for either species inthese aggregations is not known. Kavanaugh & Ol-ney (2006) suggest that large populations of trig-gerfish could benefit from a higher tolerance ofconspecifics to reduce aggressive encounters andtherefore energy expenditure.Patterns of Canthidermis maculata: Juveniles of

C. maculata show a distinct spotted pattern. Bloch(1786) illustrated the spots as dark (Fig. 24a) andBleeker (1865) illustrated the spots as white (Fig.24b). Bloch’s type specimen is lost. The dried lec-totype specimen for this species (ZMB 5904), de-posited at Museum für Naturkunde, Berlin, Ger-many, shows faint white spots (Fig. 25). Specimensof young and juvenile C. maculata exhibit whitespots (Figs 26 a-b); however, the collector (J. Ran-dall) did not note the presence or absence of spotsduring collection (pers. comm.). Juvenile andyoung Canthidermis maculata have been observedwith conspicuous white spots (Moore 1967; Alevi-zon 1976; Lanelli 2008) (Fig. 27a). Moore (1967)observed the spotted pattern was not always visible

on C. maculata specimens. Alevizon (1976) noteda coloration change in the juvenile (30 mm SL) C.maculata when Sargassum was added to the aquar-ium. Photo enlargement of Fig. 27a shows smallerblack spots interspersed with white spots (Fig.27b), possibly suggesting that conspicuous whiteand inconspicuous black spots occur on the samefish but in different locations. On 26 December2011 one C. maculata individual with black spotswas photographed by L. Choquette at a FAD nearKicha Island, SOL (Fig. 27c). Only one of the250+ photographs taken by L. Choquette in SOLshowed this dark spotted pattern. We have never observed adult C. maculata with a

conspicuous spot pattern over the body. They areuniformly gray and can display moderate to promi-nent facial and pectoral markings, usually duringmating. The spotted pattern is apparently a col-oration phase for young and juvenile C. maculata.Bloch’s original description, which does not give aSL or TL, is probably an immature specimen. Summary and future research: Based on our ob-

servations of nesting Canthidermis maculata andthe limited observations of nesting C. macrolepisand C. sufflamen, we suggest that the three speciesof Canthidermis have similar nesting behavior: lay-ing demersal eggs in shallow concavities in sandand/or coral rubble near deep water. Nesting of C.maculata occurs year-round and does not seem tobe related to lunar cycle (Table IV). Parental care ofthe eggs provides aeration and guards eggs againstpotential predators, although the latter is not al-ways successful (Figs 18b & 19a).The spawning frequency of an individual C. ma -

culata has yet to be determined. These triggerfishhave no identifying markers, and tagging was notattempted. How often does an individual return tonest? Does an individual utilize nesting habitatsthroughout its range or does it always return to onenesting site? If a fish returns to the same nestingarea, does it have a preference for the same nest? Unlike other balistids, massive aggregations of

Canthidermis maculata congregate around FADs inthe open oceans. Adaptation to the pelagic envi-ronment may contribute to the large numbers ofthis triggerfish found circumtropically. Large nest-ing areas could be necessary to populate this speciesthroughout its range. It would be interesting to dis-cover areas in the Indian and Atlantic Oceans withhabitat suitable for large numbers of nesting C.maculata.The Solomon Islands and eastern Papua New





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Guinea could be part of a series of important nest-ing areas for C. maculata in the Pacific. The prolificnesting areas in Marovo Lagoon may be only an ex-ample of extensive nesting areas in the Solomon Is-lands that produce massive amounts of C. maculatalarvae. Oceanic currents around the Solomon Is-lands flow into the East Australian Current andprobably provide a continuous supply of Canthi-dermis maculata larvae and young to the nearbylarge, open waters of the western South Pacific.

ACKNOWLEDGEMENTSWe are grateful to our volunteer research divers

whose observations, photographs, and video pro-vided data during our concentrated studies inThailand and the Solomon Islands: Tom Alburn,John Ares, Don Blair, Patrice Boeke, Jim Culter,Mike Dougherty, Amy Fleischer, Anita George,Patty Gergen, Wendy Hall, Carol Hermann, Bar-ney Holland, Rich and Laura Howard, BruceHunter, Emily Irving, Beth Ann Johnson, MalloriJohnson, Ben and Ginny Kendall, Richard Kiegler,Ivi Kimmel, Martha Kiser, Steve Kogge, Aya, Tak,and Niki Konstantinou, Mopsy Lovejoy, CathyMarine, Alice McNulty, Maya Moltzer, Krishnan

Natarajan, Jack Nelson, Ruth Petzold, John Robin-son, Bev and Tom Rodgerson, Pete and FawnRogers, John F. Pohle, Jann Rosen-Queralt, Dou-glas Seifert, Ken Spence, Mary Jane Stoll, Joan Ra-bin, Judith Rubin, David Rumiser, AndrewSnowhite, Patty Sturtevant, Brad Tanner, MichelleTilghman, Madi Verbeek, and Eli Weiss. BevRodgerson made the logistical arrangements forthese field trips supported through the Universityof Maryland Foundation and Mote Marine Labo-ratory. East Tennessee State University providedpartial travel support for DRN.Lisa Choquette and her diving team at Solomon

Dive Adventures: Hite Amos, Rino Amos, RonaldAmos, Dellington Bare, Bryan Palmer, NicolSchilling, and Barry Watts provided invaluable da-ta and hundreds of photographs from the nestingsites around Kicha Island, greatly increasing theamount of data to analyze for year-round activityof this deep water triggerfish. Bob Halstead andJohn E. Randall provided valuable photographs ofCanthidermis maculata and observational notes onPNG and other parts of the Indo-Pacific.We appreciate the good care of the crews on our

research vessels: the M/Y AQUA ONE in Thailand



Figs 27a-c. Canthidermis maculata spot variation. (a) Loose aggregation of white-spotted individuals, Big Island, Hawai’i.Photo © J. D. Watt, 2006. (b) Enlargement of head portion of individual in Fig. 27a showing faint, tiny black spots betweenlarger white spots. (c) Black-spotted individual at Fish Aggregating Device (FAD) near Kicha Island, Solomon Islands. Photoby L. Choquette, 26 Dec 2011; photo contrast enhanced by L. Mitchell.

c

(2000), especially dive master, Nat Sumanatemeya;and the M/V Bilikiki in the Solomons (1996,1997, 1998, 2014), especially dive masters DanielaTombion and Csaba Erdos.The Mote Marine Laboratory and its staff provid-

ed basic support for the preparation of this manu-script. Lawson Mitchell and Alexis Balinski,Graphic Designers, produced the maps and thelayout of the figures. Ruthanne Mitchell preparedthe final figures for nest sites (Figs 7-8). CarolMiller and Rick Magee made translations of perti-nent articles from German and French. SusanStover and Alyson Gamble, Mote librarians, gaveus much help in tracking down obscure references.

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ERRATA:

aqua Vol 20 (4), 15 October 2014: Austrolebias bagual, a new species of annual fish (Cyprinodontiformes: Rivulidae) fromsouthern Brazil by Matheus Vieira Volcan Luis Esteban Krause Lanés and Ândrio Cardozo Gonçalves, pp.161-172Should be: Received 10 July 2014 - Accepted 22 September 2014

aqua Vol 20 (4), 15 October 2014: The plates in Holbrook’s Ichthyology of South Carolina, 1860 by William D. Anderson, Jr.,pp. 173-2181) Should be: Received 9 June 2014 - Accepted 21 August 2014

2) p. 217, right column, line 6, close up “& c” to “&c.” should be:HOLBROOK, J. E. 1855a. An account of several species of fish observed in Florida, Georgia, &c. Journal of the Academy of Natural Sciences of Philadelphia, 2nd series, 3 (1): 47-58, plates 5 & 6.

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