aquaaqua-aquapress.com/pdf/AQUA21(1).pdfEugenie Clark¹, Diane R. Nelson² and Rachel Dreyer¹ 1)...

aquaInternational Journal of Ichthyology

Vol. 21 (1), 15 January 2015

AquapressISSN

0945-9871

Managing Editor:

Heiko BleherVia G. Falcone 11, 27010 Miradolo Terme (PV), ItalyTel. & Fax: +39-0382-754129E-mail: [email protected]

Scientific Editor:

Flávio C. T. LimaMuseu de Zoologia da Universidade Estadual deCampinas “Adão José Cardoso”Rua Albert Einstein s/n, Caixa Postal 6109, 13083-863 Campinas, São Paulo, Brazil Tel. +55 019-3521-6385E-mail: [email protected]

Editorial Board:

Gerald R. Allen Department of Aquatic Zoology, Western Australian Museum, Perth, Australia

Nina G. Bogutskaya Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia

Frank PezoldCollege of Science & EngineeringTexas A&M University – Corpus Christi, TX

Friedhelm KruppCurator of Fishes, Senckenberg Research Institute and Natural History Museum, Frankfurt am Main, Germany

Axel Meyer Lehrstuhl für Zoologie und Evolutions biologie, Universität Konstanz, Germany

Paolo Parenti Department of Enviromental Sciences, University of Milano-Bicocca, Milan, Italy

Mário de PinnaMuseu de Zoologia da USP, São Paulo, Brazil

John E. RandallBishop Museum, Honolulu, Hawaii, U.S.A.

Helen K. LarsonCurator Emeritus, Fishes - Museum and Art Gallery of the Northern Territory. Darwin, Australia

Scopeaqua is an international journal which publishes original sci-entific articles in the fields of systematics, taxonomy, bio -geography, ethology, ecology, and general biology of fishes.Papers on freshwater, brackish, and marine fishes will beconsidered. aqua is fully refereed and aims at publishingmanuscripts within 2-4 months of acceptance. In view ofthe importance of color patterns in species identi ficationand animal ethology, authors are encouraged to submit col-or illustrations in addition to descriptions of coloration. It isour aim to provide the international scientific communitywith an efficiently published journal meeting high scientificand technical standards.

Call for papersThe editors welcome the submission of original manu-scripts which should be sent in digital format to the scien-tific editor. Full length research papers and short notes willbe considered for publication. There are no page chargesand color illustrations will be published free of charge. Au-thors will receive one free copy of the issue in which theirpaper is published and an e-print in PDF format.

Subscription NoticeAt least one volume (4 issues) of aqua is being publishedper year, each issue comprising 38-64 pages (incl. cover). The subscription rate (for one volume = 4 issues): Personal subscription: Euro 75,00 (online edition) – Euro100,00 (online edition + print edition incl. priority mail);Institutional subscription: Euro 800,00 (online edition) –Euro 1000,00 (online edition + print edition incl. prioritymail). Subscription enquires should be sent to the publisher at the address given below or by e-mail to:[email protected] - [email protected]

Special PublicationSince 2003 Aquapress publishes a series of Special Publica-tions, which are produced at irregular intervals. All SpecialPublications have about 100 or more pages and are availableseparately from regular issues of aqua. Enquiries about sub-scriptions and prices should be sent to the publisher at theaddress given here above or by e-mail to: [email protected] - [email protected]

Claim policyOrders are regarded as firm and are not refundable. Takingout an annual subscription commits you for a minimum ofone year. Subscription will be renewed automatically. If youhave received a renewal invoice and do not wish to continuewith the subscription, please submit the cancellation in writ-ing within one month to: [email protected] or Fax: +39 0382754129

ISSN 0945-9871Publisher: Aquapress, Redazione aqua, I-27010 Miradolo Terme (Pavia), Italywww.aqua-aquapress.comPrinter: Pronto Stampa Srl – Bergamo – ItalyCopyediting and layout: Rossella Bulla© 2015 aqua, International Journal of Ichthyology

aqua - International Journal of Ichthyology

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.

b

a

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-



a



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.

b

c

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.

a

b

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.

a

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.



a

b



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.

c



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.

a

b

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



a

b

c



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.

d

e



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.

a

b



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

b

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.

c

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.

aa

b



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.

aa

b



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

b



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.

aa

b



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.

aa

b



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.

a

b

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.

a

b

(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





b

a

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.

REFERENCES ALEVIZON, W. S. 1976. Pelagic capture of young rough

triggerfish in the Caribbean. Florida Scientist 31 (1): 3-5.ALLEN, G. R. & ERDMANN, M. V. 2012. Reef Fishes of the

East Indies Vol. I-III. Tropical Reef Research, Perth, Aus-tralia. 1292 pp.

ALLEN, G. R. & ROBERTSON, D. R. 1994. Fishes of theTropical Eastern Pacific. Crawford House Press, Bathurst.332 pp.

BAILEY, K. 1985. Logs as fish aggregating devices in theequatorial Western Pacific. Regional Technical Meeting onFisheries 17: 1-8.

BARNES, A. 2005. Note on the occurrence of two rare trig-gerfishes (Balistidae) from the Gulf of Aqaba, northernRed Sea. Cybium 29(4): 407-409.

BERRY, F. H. & BALDWIN, W. J. 1966. Triggerfishes (Balis-tidae) of the eastern Pacific. Proceedings of the CaliforniaAcademy of Sciences, 4th Series 34: 429-474.

BLEEKER, P. 1865. Atlas Ichthyologique des Indes OrientalesNéêrlandaises. De Breuk & Smits, Amsterdam. 152 pp.

BLOCH, D. M. E. 1786. Naturgeschichte der ausländischenFische V1. Aus Kosten der Verfassers und in Commissionin der Buchhandlung der Realschule, Berlin, Germany.280 pp.

BÖHLKE, J. E. & CHAPLIN, C. C. G. 1993. Fishes of the Ba-hamas and Adjacent Waters, 2nd edition. University ofTexas Press, Austin, Texas. 771 pp.

BREDER, C. M., JR. & ROSEN, D. E. 1966. Modes of Repro-duction in Fishes. Natural History Press, Garden City,New Jersey. 941 pp.

BÜTTIKER, W., KRUPP, F., NADER, I., SCHNEIDER, W.(eds.). (in press). Fishes of the Red Sea In: Fauna of ArabiaVol. 26.

CARPENTER, K. E. 2002. The living marine resources of theWestern Central Atlantic. Vol. 3 Bony fishes part 2 (Opistog-nathidae to Molidae), sea turtles and marine mammals.Rome, pp 1375-2127.

CASAZZA, T. L. & ROSS, S. W. 2008. Fishes associated withpelagic Sargassum and open water lacking Sargassum inthe Gulf Stream off North Carolina. Fishery Bulletin 106(4): 348-363.

CLARK, E., KOGGE, S., NELSON, D., ALBURN, T. & POHLE,J. 2006. Burrow distribution and diel behavior of thecoral reef fish Pholidichthys leucotaenia (Pholidichthyidae).aqua International Journal of Ichthyology, 12 (2): 45-82.

CLARK, E., NELSON, D. R., STOLL, M. J. & KOBAYASHI, Y.2011. Swarming, diel movements, feeding and cleaningbehavior of juvenile venomous eeltail catfishes, Plotosuslineatus and P. japonicus (Siluriformes: Plotosidae). aquaInternational Journal of Ichthyology, 17 (4): 211-39.

CLARK, E., & POHLE, J. F. 2007. Massive colonies and be-havior of sand-diving fishes, genus Trichonotus. In 2007Joint Meeting of Ichthyologists and Herpetologists–Abstracts.

CLARK, E., POHLE, J. F. & HALSTEAD, B. 1998. Ecologyand behavior of tilefishes, Hoplolatilus starcki, H. fron-ticinctus and related species (Malacanthidae): non-mound and mound builders. Environmental Biology ofFishes, 52(4): 395-417.

DAGORN, L., HOLLAND, K. N., RESTREPO, V. & MORENO,G. 2013. Is it good or bad to fish with FADs? What arethe real impacts of the use of drifting FADs on pelagicmarine ecosystems? Fish and Fisheries, 14 (3): 391-415.

DOOLEY, J. K. 1972. Fishes associated with the pelagic Sar-gassum complex with a discussion of Sargassum commu-nity. Contributions in Marine Science 16(Mar): 1.

EMLEN, S. T. & ORING, L. W. 1977. Ecology, sexual selec-tion and the evolution of mating systems. Science 197:215-223.

ESCHMEYER, W. N. 1998. Catalog of Fishes Vol. 1-3. Cali-fornia Academy of Sciences.

FRICKE, H. W. 1980. Mating system, maternal and bi-parental care in triggerfish (Balistidae). Zeitschrift fürTierpsychologie 53: 105-122.

GARRISON, G. 2005. Isla del Coco Fishes, 2nd edition. In-stituto Nacional de Biodiversidad, Santo Domingo deHeredia, Costa Rica. 430 pp.

GILL, A. C. & RANDALL, J. E. 1997. Redescription of andlectotype designation for Balistes macrolepis Boulenger,1887, a senior synonym of Canthidermis longirostris Tor-tonese, 1954 and C. villosus Fedoryako, 1979 (Teleostei,Tetraodontiformes, Balistidae). Bulletin of the NaturalHistory Museum London (Zoology) 63 (1): 27-31.

GLADSTONE, W. 1994. Lek-like spawning, parental careand mating periodicity of the triggerfish Pseudobalistesflavimarginatus (Balistidae). Environmental Biology ofFishes 39: 249-257.

GOLDSTEIN, M. C., CARSON, H. S. & ERIKSEN, M. 2014.Relationship of diversity and habitat area in the NorthPacific plastic-associated rafting communities. MarineBiology 161: 1441-1453.

HOFFMAYER, E. R., FRANKS, J. S., COMYNS, B. H., HEN-DON, J. R. & WALLER, R. S. 2005. Larval and juvenilefishes associated with pelagic Sargassum in the Northcen-tral Gulf of Mexico. Gulf and Caribbean Fisheries Institute56: 259-269.

ISHIHARA, M. & KUWAMURA, T. 1996. Bigamy ormonogamy with maternal egg care in the triggerfish, Suf-flamen chrysopterus. Ichthyological Research 43: 307-313.

KAVANAGH, K. D. & OLNEY, J. E. 2006. Ecological corre-



lates of population density and behavior in the circum-tropical black triggerfish Melichthys niger (Balistidae).Environmental Biology of Fishes 76: 387-398.

KAWABE, R. 1984. Spawning behavior of the bridled trig-gerfish, Sufflamen fraenatus, in the aquarium. JapaneseJournal of Ichthyology 31: 193-197.

KAWASE, H. 1998. Reproductive behavior and evolution oftriggerfish (Balistidae) and filefish (Monocanthidae).Japanese Journal of Ichthyology 45: 1-19 (in Japanese).

KAWASE, H. 2002. Simplicity and diversity in the repro-ductive ecology of triggerfish (Balistidae) and filefish(Monocanthidae). Proceedings of the International Com-memorative Symposium 70th Anniversary of the JapaneseSociety of Fisheries Science 68: 119-122.

KAWASE, H. 2003a. Spawning behavior and biparental eggcare of the crosshatch triggerfish, Xanthichthys mento(Balistidae). Environmental Biology of Fishes 66: 211-219.

KAWASE, H. 2003b. Maternal egg care in the bridled trig-gerfish, Sufflamen fraenatus (Balistidae) at HachijojimaIsland, Japan. Natural History Research. 7 (2): 193-197.

KAWASE, H. & NAKAZONO, A. 1992. Reproductive behav-ior of the flagtail triggerfish, Sufflamen chrysopterus. Pro-ceedings of the 7th International Coral Reef Symposium 2:905-907.

KUITER, R. H. 2014. Fishes of the Maldives: Indian Ocean.Atoll Editions, Cairns, Australia. 308 pp.

KUWAMURA, T. 1997. Evolution of female egg care inharemic triggerfish, Rhinecanthus aculeatus. Ethology 103:1015-1023.

LANELLI, Y. 2008. War and Peace in the Deep: What youneed to know about nesting triggerfish. Alert DiverMarch/April: 38-43.

LEROY, B., PHILLIPS, J. S., NICOL, S., PILLING, G. M.,HARLEY, S., BROMHEAD, D., HOYLE, S., CAILLOT, S., AL-LAIN, V. & HAMPTON, J. 2013. A critique of the ecosys-tem impacts of drifting and anchored FADs use by purse-seine tuna fisheries in the Western and Central PacificOcean. Aquatic Living Resources 26: 49-61.

LOBEL, P. S. & JOHANNES, R. 1980. Nesting, eggs, and lar-vae of triggerfishes (Balistidae). Environmental Biology ofFishes 5: 251-252.

LYCZKOWSKI-SHULTZ, J. & INGRAM, G. W., JR. 2003. Pre-liminary guide to the identification of the early lifestages of balistid fishes in the western central North At-lantic. NOAA Technical Memorandum NMFS-SEFSC-493. 13 pp.

MACKICHAN, C. A. & SZEDLMAYER, S. T. 2007. Reproduc-tive behavior of the gray triggerfish, Balistes capriscus, inthe northeastern Gulf of Mexico. Gulf and CaribbeanFisheries Institute 59: 213-218.

MASUDA, H., AAMAOKA, K., ARAGA, C., UYENO, T, &YOSHINO, T. (Eds.). 1984. Fishes of the Japanese Archipel-ago. Tokai University Press, Tokyo. 437 pp.

MATSUURA, K. 2001. FAO Guide to Fishes Tetraodontif-ormes: Balistidae Rome, pp. 3911-3917.

MONTEIRO, P., RIBEIRO, D., SILVA, J., BISPO, J. &GONÇALVES, J. M. S. 2008. Ichthyofauna assemblagesfrom two unexplored Atlantic seamounts: Northwest

Bank and João Valente Bank (Cape Verde archipelago).Scientia Marina 72 (1): 133-143.

MOORE, D. 1967. Triggerfishes (Balistidae) of the WesternAtlantic. Bulletin of Marine Science 17 (3): 689-722

MYERS, R. F. 1991. Micronesian Reef Fishes. 2nd edition.Coral Graphics, Barrigada, Guam. 298 pp.

NELSON, J. S. 2006. Fishes of the World. 4th edition. JohnWiley & Sons, Inc, New York. 601 pp.

NELLIS, D. W. 1980. Reproduction in the ocean triggerfishCanthidermis sufflamen. Caribbean Journal of Science 16(1-4): 167.

NISHIDA, T., MATSUNAGA, A., ONIKURAA, N., OIKAWA,S. & NAKAZONO, A. 2008. Fish fauna associated withdrifting sea weeds in the Chikuzen Sea, NorthernKyushu, Japan. Fisheries Science 74: 285-292.

OHTA, I., & TACHIHARA, K. 2004. Larval developmentand food habits of the marbled parrotfish, Leptoscarusvaigiensis, associated with drifting algae. Ichthyological Re-search 51: 63-69.

PAGE, L. M., ESPINOSA-PÉREZ, H., FINDLEY, L. T.,GILBERT, C. R., LEA, R. N., MANDRAK, N. E. & MAY-DEN, R. L. 2013. Common and Scientific Names of Fishesfrom the United States, Canada, and Mexico, 7th edition.American Fisheries Society, 243 pp.

PARIN, N. V. & FEDORYAKO, B. I. 1992. Pelagic fish com-munities around floating objects in the open ocean. InProceedings of the International Workshop on the Ecologyand Fisheries for Tunas Associated with Floating Objects(Feb): 447-458.

RANDALL, J. E. 1996. Caribbean Reef Fishes Revised ThirdEdition. T.H.F. Publishers, Neptune City, New Jersey.368 pp.

RANDALL, J. E. 1995. Coastal fishes of Oman. University ofHawaii Press, Honolulu, Hawaii. 439 pp.

RANDALL, J. E., ALLEN, G. R. & STEENE, R. C. 1997.Fishes of the Great Barrier Reef and Coral Sea. Universityof Hawai’i Press, Honolulu. 557 pp.

RANDALL, J. E. & MILLINGTON, J. T. 1990. Triggerfish bite– a little-known marine hazard. Journal of WildernessMedicine 1: 79-85.

ROBINS, C. R. & RAY, G. C. 1986. A Field Guide to At-lantic Coast Fishes of North America. Houghton MifflinCompany, Boston, U.S.A. 354 pp.

SAHAYAK, S. 2005. Reproductive biology of the maskedtriggerfish, Sufflamen fraenatus. Journal of the Marine Bi-ology Association of India 47 (1): 70-76.

SCHWARTZ, F. J. 2006. Balistid triggerfishes and monocan-thid filefishes (Tetraodontiformes) of North Carolina.Journal of the North Carolina Academy of Science 122 (3):118-124.

SETHI, S. N., RAJAPACKIAM, S., JAIGANESH, P. & RUDHRA-MURTHY, N. 2011. Occurrence of trigger fishes at Chen-nai. Marine Fisheries Information Service T&E Series 208:20-21.

SIMMONS, C. M. & SZEDLMAYER, S. T. 2012. Territoriali-ty, reproductive behavior, and parental care in gray trig-gerfish, Balistes capriscus, from the northern Gulf of Mex-ico. Bulletin of Marine Science 88 (2): 197-209.



SONHANA, K. S., SEETHA, P. K., KISHORE, T. G., DIVYA, D.D., DINESHKUMAR, S., NAJMUDEEN, T. M., NAIR, R. J. &ZACHARIA, P. U. 2013. Unusual landing of the spottedocean triggerfish Canthidermis maculata at Cochin Fish-eries Harbour. Marine Fisheries Information Service; Tech-nical and Extension Series 215: 35.

SWAINSON, W. 1839. On the natural history and classifica-tion of fishes, amphibians and reptiles Vol II. Longman,Orme, Brown, Green, & Longmans, London. 452 pp.

TAQUET, M., SANCHO, G., DAGORN, L., GAERTNER, J.-C.,ITANO, D., AUMEERUDDY, R., WENDLING, B. &

PEIGNON, C. 2007. Characterizing fish communities as-sociated with drifting fish aggregating devices (FADs) inthe Western Indian Ocean using underwater visual sur-veys. Aquatic Living Resources 20: 331-341.

THRESHER, R. E. 1984. Reproduction in Reef Fishes. T. F.H. Publishing, Neptune City, New Jersey. 399 pp.

VENKATARAMAN, G. & GEORGE, K. C. 1964. On the oc-currence of large concentrations of file-fish off the KeralaCoast, India. Journal of the Marine Biological Associationof India 6 (2): 321-323.



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.

AbstractPerugia (1891) described and recorded many fish species

from Argentina, Paraguay and Brazil, whose voucher speci-mens were deposited at the Museo Civico di Storia Naturaledi Genova (MSNG). Several species recorded by Perugia(1891) from Argentina were misidentified, which lead to er-roneous citations of species for the Rio de la Plata basin dur-ing more than one century, species which in fact are restrict-ed to river drainages from northern South America. The re-evaluation of the specimens identified by Perugia (1891) asPimelodus cristatus Müller & Troschel, Ramphichtys brevi-rostris Steindachner, and Myletes asterias Müller & Troschel,and the records of Serrasalmo humeralis Valenciennes, andSerrasalmo gymnogenys Günther allowed us to rectify theserecords and exclude these species from the fish fauna lists ofboth the Rio de la Plata basin and Argentina.

ResumenPerugia (1891) describió y registró muchas especies de

peces de Argentina, Paraguay y Brasil, cuyos ejemplares es-tán depositados en el Museo Civico di Storia Naturale diGenova (MSNG). Varias especies registradas por Perugia(1891) de Argentina, fueron erróneamente identificadas,hecho que condujo a citaciones incorrectas de especies parala cuenca del Río de la Plata durante mas de un siglo; algu-nas especies están restringidas a drenajes de ríos del nortede Sudamérica. La re-evaluación de los ejemplares identifi-cados por Perugia (1891) como Pimelodus cristatus Müller& Troschel, Ramphichtys brevirostris Steindachner y Myletesasterias Müller & Troschel, y los registros de Serrasalmohumeralis Valenciennes y Serrasalmo gymnogenys Günther,nos permitió rectificarlos y excluirlos de las listas de ictio-fauna de ambos, cuenca del Rio de la Plata y Argentina.

ZusammenfassungPerugia (1891) hat viele Fischarten aus Argentinien,

Paraguay und Brasilien beschrieben und aufgezeichnet,deren Belegexemplare im Museo Civico di Storia Naturaledi Genova (MSNG) hinterlegt wurden. Einige Arten, die

Perugia (1891) aus Argentinien beschrieben hat, warenfehlbestimmt, was über ein Jahrhundert lang zur irrtüm-lichen Anführung von Arten für das Rio-de-la-Plata-Beckenführte, die in Wirklichkeit auf Einzugsgebiete im nördlichenSüdamerika beschränkt sind. Die Neubewertung der Exem-plare, die Perugia (1891) als Pimelodus cristatus Müller &Troschel, Ramphichtys brevirostris Steindachner und Myletesasterias Müller & Troschel bestimmt hatte, und die Nach-weise von Serrasalmo humeralis Valenciennes und Serrasalmogymnogenys Günther ermöglichten uns die damaligen Aufze-ichnungen zu berichtigen und die Arten von den Fischfau-na-Listen des Rio-de-la-Plata-Beckens sowie Argentiniens zustreichen.

RésuméPérugia (1891) a décrit et enregistré beaucoup d’espèces

de poissons d’Argentine, du Paraguay et du Brésil dont lesspé cimens conservés ont été déposés au Museo Civico diStoria Na turale di Genova (MSNG). Plusieurs espèces col-lectées en Argentine par Pérugia (1891) ont été mal iden-tifiées, ce qui a conduit à l’affectation erronée d’espècesdans le bassin du Rio de la Plata pendant plus d’un siècle,des espèces qui, en fait, sont limitées à des bassins de coursd’eau du nord de l’Amérique du Sud. La réévaluation desespèces identifiées par Pérugia (1891) comme Pimeloduscristatus Müller et Tro schel, Rhamphichtys brévirostris Stein-dachner et Myletes aste rias Müller et Troschel, et les men-tions de Serrasalmo hu me ra lis Valenciennes et Serrasalmogymnogenys Günther nous ont permis de corriger ces men-tions et d’exclure ces espèces des listes de la faune piscicoledu Rio de la Plata et d’Argentine.

SommarioPerugia (1891) ha descritto e registrato molte specie di

pe sci provenienti da Argentina, Paraguay e Brasile, i cui esemplari sono stati depositati presso il Museo Civico diStoria Naturale di Genova (MSNG). Diverse specie regi -strate dall’Argentina sono state erroneamente identificatedal Perugia (1891), causando per oltre un secolo citazioni



On some freshwater fish species reported by Perugia (1891) from Argentina

María de las Mercedes Azpelicueta1 and Stefan Koerber2

1) Conicet-División Zoología Vertebrados, Museo de La Plata, 1900 La Plata, Argentina. E-mail: [email protected]

2) Friesenstr. 11, 45476 Muelheim, Germany. E-mail: [email protected]

Received 23 June 2014 – Accepted 02 January 2015

erronee di presenza nel bacino del Rio de la Plata di specieche in realtà sono confinate in bacini idrogeografici dellaparte settentrionale del Sudamerica. La rivalutazione degliesemplari identificati dal Perugia (1891) come Pimeloduscristatus Müller & Troschel, Ramphichtys brevirostris Stein-dachner, e Myletes asterias Müller & Troschel, e le regis-trazioni di Serrasalmo humeralis Valenciennes, e Serrasalmogymnogenys Günther ci ha permesso di correggere questesegnalazioni e di escludere queste specie dalla lista dei pescipresenti nel bacino del Rio de la Plata e in Argentina

INTRODUCTIONThe Rio de la Plata basin drains large portions of

Argentina, Bolivia, Brazil, Paraguay, and Uruguay,being the second largest drainage system in SouthAmerica and the fourth largest in the world. TheParaná River flows 4.000 km southwards from itssources in the Precambrian Brazilian Shield to itsmouth in the Pampa Plain (Quirós, 2003).Perugia (1891) published a list of South Ameri-

can fishes housed at the Museo Civico di StoriaNaturale di Genova (MSNG), based on specimensfrom Argentina, Paraguay and Brazil. Severalspecies described as new in that paper are still con-sidered valid, among others, Trigonectes balzanii,Pseudocorynopoma doriae, and Gymnogeophagusbalzanii, and he provided the first record from Ar-gentina for several other species.Although dating back from 120 years ago, most of

the specimens reported by Perugia are still extantand available at the ichthyological collection ofMSNG, which enabled us to re-examine it andamend some identifications when necessary. Re-cently, Koerber & Azpelicueta (2009) presented ev-idence that Weyenbergh’s Xiphophorus obscurus is asenior synonym of Jenynsia pygogramma and Koer-ber & Casciotta (2012) identified Perugia’s Crenici-chla lacustris as C. scottii, finally excluding the firstfrom the list of Argentinean freshwater fishes.This contribution attempts to clarify the system-

atic status of five additional species of freshwaterfishes, which have been erroneously included inthe lists of Argentinean ichthyofauna and from theRio de la Plata basin through the last century.

RESULTSPimelodus cristatus Müller & Troschel, 1849Weyenbergh sent two specimens from Tucumán

(Argentina) to Perugia who recorded them asPimelodus cristatus Müller & Troschel 1849. Thespecies was cited by Ringuelet et al. (1967) (asPimelodella cristata), citing Perugia (1891) as thesource for the sole record from Argentina. In the last

available list on freshwater fishes from Argentina,López et al. (2003) listed the species based on the in-formation provided by Ringuelet et al. (1967). Sub-sequently, Liotta (2005) provided Ringuelet et al.(1967) as the only known Argentinean record.The fish fauna of the rivers of Tucumán has been

recently investigated by several authors who didnot confirmed the presence of this species in thatarea. Mirande & Aguilera (2010) only reportedPimelodella gracilis (Valenciennes 1835), P. griffiniEigenmann, 1917, and P. laticeps Eigenmann,1917 in the Northwest of Argentina (NOA). Bock-mann & Guazzelli (2003) considered P. cristata asa valid species, but restricted its distribution toGuyana and French Guiana.Perugia’s specimens are still extant and belong to

two lots (MSNG 8133, MSNG 14543, Figs 1, 2),both examined by the authors. Both specimens(87.19-101.69 mm SL) correspond to Pimelodellagracilis (Valenciennes, 1835) which is indeed simi-lar to P. cristata. Pimelodella gracilis has the uppercaudal-fin lobe much longer and with similarwidth than the lower one, whereas P. cristata has anarrower upper caudal-fin lobe, which is scarcelylonger than the lower lobe. In addition, some mea-surements taken from six specimens of P. gracilisfrom Rio Paraná in Corrientes (Argentina) and sixspecimens of P. cristata from the Corantijn River,Nickerie District (Suriname), differ respectively inthe following measurement ratios: eye diameter20.08-23.25 % of head length (vs. 23.07-26.31 %of head length), eye diameter in percentage ofsnout 77.77-93.33 % (vs. 100.0-125.0 %) and arelatively shorter length of the adipose fin 35.63-40.68 % of standard length (vs. 38.92-48.78 % ofstandard length).As no other record for the species other than the

one from Perugia (1891) is known in Argentina,we consider that P. cristata does not occur either inthe Rio de la Plata basin and the Argentinean fresh-waters.

Ramphichtys brevirostris Steindachner, 1868The specimens examined by Perugia (1891) and

listed as Ramphichtys brevirostris were collected bySpegazzini in Resistencia, capital of the Argentineanprovince of Chaco. Ringuelet et al. (1967) listed Pe-rugia s record (as Hypopomus brevirostris) as the firstrecord for Argentina. López et al. (2003) repeatedthe information from Ringuelet et al. (1967), list-ing the species as Brachyhypopomus brevirostris (acombination proposed by Mago-Leccia, 1994).



Albert & Crampton (2003) ambiguously citedthe species as distributed at ’South America: East-ern South America from Orinoco to la Plata River’but occurring exclusively in the countries ‘Bolivia,Brazil, Suriname, Venezuela’. Albert (2001) as-signed lots from Guyana, Venezuela, and the Ama-zonian basin in Bolivia and Brazil to Brachyhypopo-mus brevirostris, but listed those from Argentinaand the Río San Francisco basin in Brazil as B. cf.brevirostris, showing consequently his doubts aboutthe distribution of the species southward from theAmazon basin.Liotta (2005) listed Argentinean records for

Brachyhypopomus brevirostris from 11 different lit-erature references. All citations of the species fromArgentina predate the descriptions of B. bombillaLoureiro & Silva, 2006, B. draco Giora, Malabarba& Crampton, 2008, and B. gauderio Giora & Mal-abarba, 2009. Almirón et al. (2010) published thefirst Argentinean records of these three species.They reexamined several lots previously listed as B.brevirostris. Referring to B. brevirostris, they statedthat “it is possible that some other Argentineanrecords of this species could correspond to B. bom-billa, B. draco or B. gauderio”.We have examined Perugia’s specimens (MSNG

27458), which included one specimen of Eigenman-

nia trilineata López & Castello 1966 (128.66 mmSL, Fig. 3). As for the remaining specimens, one be-long to Brachyhypopomus bombilla (97.8 from tip ofsnout to last anal fin ray insertion), while the otheris a damaged Brachyhypopomus specimen, probablybelonging to the same species (102.0 mm fromsnout tip to last anal fin ray insertion). The smallerspecimen is in good state of preservation, with thescattered dark brownish pigmentation over brownbackground clearly visible; also, it has the upper jawslightly longer than lower jaw, and a very slendershort caudal filament (Fig. 4).To avoid future confusions, the specimen of

Eigenmannia trilineata has been separated in an in-dividual lot (MSNG 56988). Besides B. brevi-rostris, we also take this opportunity to remove Hy-popomus artedi (Kaup 1856) (listed by López et al.2003) from the list of fishes from Argentina; H.artedi is present in Atlantic drainages of Guyana,Surinam, French Guiana, and northern Brazil (Al-bert & Crampton 2003).

Myletes asterias Müller & Troschel, 1844Perugia (1891) recorded the occurrence of Myletes

asterias from Misiones (Argentina), based on alarge specimen of 55 cm TL received from the ves-sel commander Bove. Perugia’s specimen is still


María de las Mercedes Azpelicueta and Stefan Koerber

Fig. 1. MSNG 8133, Pimelodella gracilis (Valenciennes 1835), 87.19 mm SL, Argentina, Tucumán.

Fig. 2. MSNG 14543, Pimelodella gracilis (Valenciennes, 1835), 101.69 mm SL, Argentina, Tucumán.

housed at MSNG (MSNG 14870), but due to itssize could not be shipped for loan and we were on-ly able to examine photographs (Figs 5-6). Bothphotographs clearly show that the specimen be-longs to Piaractus mesopotamicus (Holmberg,1887) for having ovoid body shape (Figs 5-6). Pi-aractus mesopotamicus is a widely distributedspecies in the Paraguay and Parana river basins. Pe-rugia s Myletes asterias or Myloplus asterias as isknown today (Jégu et al. 2004), has an orbicularbody shape and also, a maximum body length ofabout 250 mm SL.The record of Perugia is the first and sole record of

Myletes asterias from Argentina; Ringuelet et al.(1967) and López et al. (2003) considered thespecies as valid for the Argentinean ichthyofauna asMyloplus asterias; however, in both papers, the au-

thors did not mentioned any examined material andpresumably listed the species based on the record byPerugia (1891). Similarly, Liotta (2005) cited Roa &Permingueat (1999), who merely followed Ringueletet. al (1967). Jégu et al. (2004) provided the distrib-ution of Myloplus asterias (Müller & Troschel 1844)from the Amazon River basin and the Northern andEastern Guiana Shield rivers in Brazil and Guyana.Thus, Myloplus asterias is not recorded in the Rio dela Plata basin and should not be considered as beingpart of the Argentinean fauna.

Serrasalmo humeralis Valenciennes, 1850Perugia (1891) reported this species also based on

material collected by Spegazini, in Resistencia. Un-fortunately this specimen could not be found atthe MSNG.



Fig. 3. MSNG 56988, Eigenmannia trilineata López & Castello, 1966, 128.63 mm SL, Argentina, Resistencia.

Fig. 4. MSNG 27458, Brachyhypopomus bombilla Loureiro & Silva, 2006, 102 mm from snout tip to last anal-fin ray (at top)and 97.8 mm from snout tip to last anal-fin ray (at bottom), Argentina, Resistencia.

Nonetheless, Perugia (1891) described the speci-mens as possessing “la caudale com intensa fascianera ed uno stretto margine esterno bianco“. Thiscondition in the caudal fin indicates that the spec-imen examined by Perugia correspond to S. macu-latus Kner 1858, the sole species with that pigmen-tation pattern.The species S. humeralis was treated as a synonym

of S. rhombeus (Linnaeus 1766) by Norman(1929). Ringuelet et al. (1967) followed Norman(1929) synonym, and considered S. rhombeus withPerugia s material as the first record from Argenti-na, an action which was followed by subsequentauthors. Recently, Jégu (2003) treated S. rhombeusand S. humeralis as valid species and indicated the



Fig. 5. MSNG 14870, Piaractus mesopotamicus (Holmberg, 1887), 550 mm TL, Argentina, Misiones. At top, left lateral view;at bottom, right lateral view. Scale bar= 10 cm.

Fig. 6. MSNG 14870, Piaractus mesopotamicus (Holm-berg, 1887), 550 mm TL, Argentina, Misiones. Detail ofhead showing upper jaw teeth.



Table I. Freshwaters species described or cited by Perugia (1891) from Argentina. N indicates the number which assigns theorder in which the species is cited in the paper; # indicates species currently known not to occur in Argentina, which stillneed reexamination.

N Name of species in Perugia (1891) Present combination 5 Percichthys laevis Percichthys trucha 6 Percichthys vinciguerrae Percichthys vinciguerrae 12 Sciaena amazonica Plagioscion ternetzi 35 Atherinichthys bonairiensis Odontesthes bonariensis 39 Acara vittata Bujurquina vittata 40 Crenicichla johanna Crenicichla johanna# 41 Crenicichla saxatilis Crenicichla saxatilis# 42 Crenicichla lacustris Crenicichla scottii 54 Hypophthalmus marginatus Hypophthalmus oremaculatus 55 Platystoma lima Sorubim lima 56 Piramutana albicans Pimelodus albicans 57 Pimelodus maculatus Pimelodus maculatus 58 Pimelodus pati Luciopimelodus pati 59 Pimelodus cristatus Pimelodella gracilis 60 Pimelodus sapo Rhamdia quelen 61 Pimelodus argenteus Pimelodus argenteus 62 Pimelodus spegazzinii Parapimelodus valenciennis 63 Pimelodus elongatus Pimelodella elongata# 66 Ageniosus brevifilis Ageneiosus inermis 67 Ageniosus militaris Ageniosus militaris 70 Corydoras punctatus Corydoras punctatus# 71 Rhinodoras knerii Oxydoras knerii 72 Auchenipterus ceratophysus Trachelyopterus ceratophysus# 74 Callichthys laevigatus Hoplosternum littorale 75 Plecostomus barbatus Pseudancistrus barbatus# 76 Plecostomus bicirrhosus Hypostomus plecostomus# 77 Plecostomus punctatus Hypostomus punctatus# 79 Loricaria maculata Loricariichthys maculatus 80 Trychomicterus cordovensis Trichomycterus corduvensis 81 Heptapterus mustelinus Heptapterus mustelinus 82 Macrodon trahira Hoplias malabaricus 83 Erythrinus unitaeniatus Hoplerythrinus unitaeniatus 84 Pyrrhulina brevis Pyrrhulina australe 85 Curimatus ciliatus Psectrogaster curviventris 86 Prochilodus reticulatus Prochilodus lineatus 87 Prochilodus argenteus Prochilodus lineatus 88 Parodon nasus Parodon nasus 89 Anastomus fasciatus Schizodon borellii 91 Curimatus frederici Leporinus acutidens 92 Leporinus affinis Leporinus octofasciatus 94 Tetragonopterus rufipes Tetragonopterus argenteus 95 Tetragonopterus cordovae Astyanax cordovae 96 Tetragonopterus nigripinnis Markiana nigripinnis 98 Tetragonopterus rutilus Astyanax rutilus 99 Tetragonopterus fasciatus Astyanax fasciatus# 101 Tetragonopterus dichrourus Moenkhausia dichroura 102 Chirodon interruptus Cheirodon interruptus 104 Pseudocorynopoma doriae Pseudocorynopoma doriae 105 Anacyrtus bonariensis Roeboides microlepis 107 Anacyrtus humeralis Galeocharax humeralis 108 Cynodon vulpinus Raphiodon vulpinus 109 Salminus maxilloxus Salminnus brasiliensis 110 Xiphoramphus hepsetus Oligosarcus jenynsii 112 Serrasalmo humeralis Serrasalmus maculatus 113 Serrasalmo gymnogenis ? 115 Myletes asterias Piaractus mesopotamicus 116 Cynolebias porosus Cynolebias porosus# 118 Cynolebias bellotti Austrolebias bellottii 119 Cynolebias robustus Austrolebias robustus 120 Jenynsia lineata Jenynsia multidentata 121 Gambusia gracilis Jenynsia obscura 122 Girardinus decemmaculatus Cnesterodon decemmaculatus 128 Engraulis poey Lycengraulis poeyi# 136 Ramphichthys brevirostris Brachypopomus bombilla in part Eigenmannia trilineata in part 138 Carapus fasciatus Gymnotus sp.# 139 Symbranchus marmoratus Synbranchus marmoratus

following geographical distribution: for S.rhombeus, Amazon and Orinoco River basins,north and eastern Guiana Shield rivers, and North-eastern Brazilian coastal rivers, whereas for Ser-rasalmus humeralis, Amazon River basin in Bolivia,Brazil, and Peru. Thus, we consider that therecords of S. humeralis and S. rhombeus should beboth excluded from lists of Argentinean ichthyo -fauna.

Serrasalmo gymnogenis Günther, 1864The specimen examined by Perugia (1891) was

not found in the collection of MSNG. An analysisof the literature related with that species shows dif-ferent synonyms for the species S. gymnogenis, suchas Pristobrycon aureus Spix & Agassiz 1829 follow-ing Eigenmann (1915) or S. serrulatus Valenci-ennes 1850 sensu Norman (1929). Ringuelet et al.(1967) and López et al. (2003) followed Norman(1929) with Perugia’s material as the first recordfrom Argentina. Although Jégu (2003) confirmedthe synonymy of Serrasalmus gymnogenys with Pri -stobrycon aureus and considered S. serrulatus asvalid, none of them are present in the Rio de la Pla-ta basin. Both species names should be removedfrom ichthyofaunal lists in the area but it is impos-sible to confirm the identification of material ex-amined by Perugia (1891).

REMARKSPerugia s (1891) contribution was historically im-

portant as it dealt with a great number of fishspecies (144), from Mato Grosso (Brazil) to PuntaArenas (Chile) in southern South America. Perugia(1891) listed and in many cases discussed 91 fresh-water species and 53 marine species (including 12euryhaline species). He also described nine newspecies, six of which are still considered to be valid,and one genus which is still valid, Pseudocorynopo-ma. The six freshwater species described by Perugiaand still recognized as valid at present (presentcombinations indicated with*) are: Pseudocorynopo-ma doriae, Pimelodus argenteus, Geophagus balzanii(*Gymnogeophagus balzanii), Tetrago nop terus nigri -pinnis (*Markiana nigripinnis), Tetrago nopterus li -ne atus (*Astyanax lineatus), and Haplo chilusbalzanii (*Trigonectes balzanii). Perugia (1891) de-scribed a single new marine species, Salilota bovei,currently considered a synonym of Salilota au -stralis. About 65 % of the freshwater and 20 % ofmarine species listed were correctly identified byPerugia (1891). There are 14 synonyms of marine

species. All freshwater species cited by Perugia(1891) from Argentina are listed in the table 1, in-cluding the name given by that author, and thepresent combination. It is still necessary to exam-ine the species marked with (#), which are notknown to occur in Argentinean waters.In this contribution we dealt with five freshwater

species listed by Perugia (1891) which have beenincluded in lists of the Argentinean ichthyofaunaduring more than a century. All related fields of en-quiry as phylogeny, ecology, biogeography, phylo-geography, among others, depend on a soundknowledge of taxonomy. Biogeographic studies orfaunal lists, either of regional areas or within a na-tional perspective, are important tools not only forefforts in biodiversity conservation in general butalso for evaluations of large environmental im-pacts, such as, for example, construction of hydro-electric dams. Naturally any list, whether nationalor local, is always simply a snapshot of the momentwhen it was prepared and it might be quickly out-dated by the advance on the taxonomical and bio-geographical knowledge of a given taxon. To avoidthe ongoing use of erroneous records these listsshould be periodically ‘cleaned up’ when newknowledge makes prior errors obvious. Thus, theexamination of the fishes recorded by Perugia(1891) and the historical revisions are contribu-tions to the better knowledge of the ichthyofaunafrom the Rio de la Plata basin, especially the onefrom Argentinean freshwaters.

ACKNOWLEDGEMENTSThanks are due to Giuliano Doria (MSNG) for

the loan of material under his care and pho-tographs of Piaractus mesopotamicus; two anony-mous reviewers for pertinent corrections and sug-gestions that improved the original manuscript,and F. C. T. Lima for valuable suggestions.

REFERENCESALBERT, J. S. 2001. Species diversity and phylogenetic sys-

tematics of American knifefishes (Gymnotiformes,Teleostei). Miscellaneous Publications, Museum of Zoology,University of Michigan 190: 1-127.

ALBERT, J. S. & CRAMPTON, W. G. R. 2003. Family Hy-popomidae (Bluntnose knifefishes). In: Checklist of theFreshwater Fishes of South and Central America. (eds. R.E.Reis, S. O. Kullander & C. J. Ferraris): 495-497.EDIPUCRS, Porto Alegre, Brazil.

ALMIRÓN, A. E., CASCIOTTA, J. R., CIOTEK, L., GIORGIS,P., SONEIRA, P. & RUÍZ DÍAZ, F. 2010. Pisces, Gymnoti-formes, Hypopomidae, Brachyhypopomus Mago-Leccia,



1994: First country record of three species of the genus,Argentina. Check List 6 (4): 572-575.

BOCKMANN, F. A. & GUAZZELLI, G M. 2003. Family Hep-tapteridae (Heptapterids). In: Checklist of the FreshwaterFishes of South and Central America. (eds. R.E. Reis, S.O.Kullander & C.J. Ferraris): 406-431. EDIPUCRS, PortoAlegre, Brazil.

EIGENMANN, C. H. 1915. The Serrasalminae and Mylinae.Annals of the Carnegie Museum 9 (3-4): 226-272.

JÉGU, M. 2003. Subfamily Serrasalminae (Pacus and pira-nhas). In: Checklist of the Freshwater Fishes of South andCentral America. (eds. R. E. Reis, S. O. Kullander & C.J. Ferraris): 182-196. EDIPUCRS, Porto Alegre, Brazil.

JÉGU, M., HUBERT, N. & BELMONT-JÉGU, E. 2004. Réha-bilitation de Myloplus asterias (Müller & Troschel, 1844),espèce-type de Myloplus Gill, 1896 et validation du genreMyloplus Gill (Characidae: Serrasalminae). Cybium 28(2): 119-157.

KOERBER, S. & AZPELICUETA, M. 2009. On two Jenynsiataxa (Cyprinodontiformes: Anablepidae) from Argenti-na, originally placed in the poeciliid genus Xiphophorus.Zootaxa 2028: 51-58.

KOERBER, S. & CASCIOTTA, J. 2012. On the erroneousrecords of Crenicichla lacustris (Castelnau, 1855) fromfreshwaters of Argentina. Ichthyological Contributions ofPecesCriollos 25: 1-4.

LIOTTA, J. 2005. Distribución geográfica de los peces deaguas continentales de la República Argentina. ProBiota -Serie Documentos 3: 1-701.

LÓPEZ, H. L., MIQUELARENA, A. M., & MENNI, R. C.

2003. Lista comentada de los peces continentales de laArgentina. ProBiota - Serie Técnica y Didáctica 5: 1-85.

MAGO-LECCIA, F. 1994. Electric fishes of the continentalwaters of America. Caracas, Fundación para el Desarrollode las Ciencias Físicas, Matemáticas y Naturales. 206 pp.

MIRANDE, J. M. & AGUILERA, G. 2010. Los peces de la sel-va pedemontana del noroeste argentino. In: Selva Pede-montana de las Yungas. (eds. A. Brown, P. G. Blendinger,T. Lomáscolo, & P. Garcia Bes): 169-211. FundaciónProYungas, Ediciones del Subtrópico.

NORMAN, J. R. 1929. The South American characid fishesof the subfamily Serrasalmoninae, with a revision of thegenus Serrasalmus, Lacépède. Proceedings of the ZoologicalSociety of London 4 (30): 781-829.

PERUGIA, A. 1891. Appunti sopra alcuni pesci sud-ameri-cani conservati nel Museo Civico di Storia Naturale diGenova. Annali del Museo Civico di Storia Naturale ‘Gia-como Doria’ 10: 605-657.

QUIRÓS, R. 2003. The Plata River Basin: InternationalBasin Development and Riverine Fisheries. FAO-Pro-ceedings of the second international symposium on themanagement of large rivers for fisheries 1: 253-272.

RINGUELET, R. A., ARÁMBURU, R. H. & ALONSO DE

ARÁMBURU, A. 1967. Los peces argentinos de agua dulce.Gobernación de la provincia de Buenos Aires, Comisiónde Investigación Científica. La Plata. 602 pp.

ROA, B. H. & PERMINGEAT, E. D. 1999. Composición yabundancia de la fauna íctica en dos estaciones demuestreo del embalse de Yacyretá, Argentina. Revista deIctiología 7 (Número especial): 49-57.



Abstract A new species of syngnathid pipefish, Festucalex rufus is de-

scribed from Milne Bay Province, Papua New Guinea on thebasis of four specimens, 26.5-37.8 mm SL collected fromcoral reef habitat in 10-20 m depth. It is distinguished fromother members of the genus on the basis of a combination offeatures including a short snout (2.8-3.2 in head length), rel-atively low pectoral-ray count (10-11), and a lateral trunkridge that terminates on the penultimate trunk ring. It is asmall species with observed and collected individuals gener-ally less than 40 mm SL and the single brood-pouch malecollected measures 36.3 mm SL.

ZusammenfassungBeschrieben wird eine neue Art der Seenadeln, Festucalex

rufus, auf der Grundlage von vier Exemplaren mit 26,5 bis37,8 mm SL, die in der Provinz Milne-Bucht, Papua-Neuguinea, über einem Korallenriff in 10 bis 20 m Tiefegefangen wurden. Sie unterscheiden sich von anderen Artender Gattung durch eine Kombination von Merkmalen,darunter eine kurze Schnauze (2,8-3,2 der Kopflänge), rela-tiv wenige Brustflossenstrahlen (10-11) sowie einenseitlichen Grat auf dem Rumpf, der auf dem vorletztenRumpfring endet. Die Vertreter der neuen Art sind klein, diebeobachteten und gesammelten Individuen maßen grund-sätzlich weniger als 40 mm SL, und das einzelne Männchenmit Bruttasche unter den gefangenen Tieren hatte eineLänge von 36,6 SL.

RésuméUne nouvelle espèce de Syngnathidé, Festucalex rufus, est

décrite provenant de Milne Bay Province, Papouasie-Nou-velle-Guinée, sur base de quatre spécimens, de 26,5 - 37,8mm LS, collectés dans un récif corallien, à une profondeurde 10 - 20 m. Elle se distingue des autres membres du genrepar une combinaison de caractéristiques comprenant un ro -stre court (2,8 - 3,2 de la longueur de la tête), un nombre re -

la tivement réduit de rayons pectoraux (10 - 11) et une crêtede tronc latérale qui s’achève sur l’avant-dernier anneau detronc. C’est une petite espèce dont les individus collectés etobservés font généralement moins de 40 mm de LS et le seulmâle en incubation collecté faisait 36,3 mm de LS.

SommarioUna nuova specie di pesci ago, Festucalex rufus è descritta

dalla provincia di Milne Bay, Papua Nuova Guinea, sullabase di quattro esemplari di dimensioni 26,5-37,8 mm SLraccolti lungo la barriera corallina a 10-20 m di profondità.Si distingue dagli altri membri del genere sulla base di unacombinazione di caratteri, tra cui: un muso breve (2,8-3,2 inlunghezza della testa), numero di raggi pettorali relativa-mente basso (10-11) e una cresta laterale sul tronco che ter-mina sul penultimo anello del tronco stesso. Si tratta di unapiccola specie: gli individui osservati e raccolti in genere era-no inferiori a 40 mm SL e l’unico maschio con tasca ventraleraccolto misurava 36,3 mm SL.

INTRODUCTIONThe syngnathid genus Festucalex Whitley 1931 was

reviewed by Dawson (1985) and further discussedby Fricke (2004). The group is distinguished fromother family members by a combination of featuresthat include the presence of a tiny caudal fin (con-taining 10 rays), both superior and inferior trunkand tail ridges continuous, anal fin present (with 4rays), lateral trunk ridge more or less strait (not de-flected ventrally near anal ring), dorsal-fin origin onthe trunk, principal tail ridges lacking prominenthook-like projections on the posterior angles of eachtail ring, and the median dorsal snout ridge lowwithout projections, more or less concave in lateralprofile. The group was previously known to contain



Festucalex rufus, a new species of pipefish (Syngnathidae) from Milne Bay Province, Papua New Guinea

Gerald R. Allen1 and Mark V. Erdmann2

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: [email protected]

2) Conservation International Indonesia Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar80235 Indonesia, and California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118,

USA. E-mail: [email protected]

Received 4 October 2014 – Accepted 12 November 2014



seven species, which are restricted to the tropical andtemperate Indo-Pacific region: F. cinctus (Ramsay1882) from eastern and northern Australia, F. eryth -ra eus (Gilbert 1905), which is widely distributedfrom Mozambique to Hawaii, F. gibbsi Dawson1977 from Indonesia and the Great Barrier Reef, F.kul bickii Fricke 2004 from New Caledonia, F. proli -xus Dawson 1984 from Indonesia and the Philip-pines, F. scalaris (Günther 1870) from Western Aus -tra lia, and F. wassi Dawson 1977 from Fiji andSamoa. The present paper describes the eighth known

member of the genus. The species was initially pho-tographed in situ by Ned DeLoach in 2008 at MilneBay Province, Papua New Guinea. Additional obser-vations at Indonesia (Bali and Ceram) and the So lo -mon Islands indicate that it is relatively widespread,although inconspicuous due to its small size andcryptic habits. Underwater photographs appeared inseveral recent publications (Kuiter 2009; Allen et al.2010; Allen & Erdmann 2012), invariably misiden-tified as a variant of Micrognathus pygmaeus Fritzsche1981. Four specimens were finally obtained by theauthors during visits to Milne Bay in 2013-2014 andform the basis of the present description.

MATERIALS AND METHODSMethods of counting and measuring, and termi-

nology used for trunk and tail ridges follow those ofDawson (1985). Trunk rings are counted from thering bearing the pectoral fins to the ring bearing theanus. Tail rings begin with the first ring behind theanus to the penultimate ring, excluding the terminalelement bearing the caudal fin. Standard length isthe straight line distance from the tip of the lowerjaw to the base of the caudal fin. Head length is thedistance from the tip of the lower jaw to the rearmargin of the opercle. Snout length is measuredfrom the tip of the lower jaw to the anterior marginof the eye. Snout depth is the least vertical dimensionof the snout. Body depth is the maximum depth,measured at the first trunk ring between the outermargins of the superior and median ventral trunkridges, and body width is measured between the cor-responding lateral trunk ridges of the first trunk ring.Eye diameter is measured horizontally. Predorsallength is measured from the tip of the lower jaw tothe dorsal-fin origin. Preanal length is measuredfrom the tip of the lower jaw to the anal opening.Prepectoral length is measured from the tip of thelower jaw to the base of the upper base of the pec-toral fin.

Type specimens are deposited at the Western Aus-tralian Museum, Perth (WAM) and the UnitedStates National Museum of Natural History, Wash-ington, D.C. (USNM). The description of the newspecies follows the format of Fricke (2004). Countsand proportional measurements include the valuefor the holotype followed in parentheses by the rangefor the paratypes if different. Percentage of SL valuesare also included for each proportional measurementand include the range for all four type specimens.

Festucalex rufus, n. sp. Pink Pipefish(Figs 1-5)

Holotype: WAM P. 33884-001, female, 37.8 mmSL, Cheri’s Reef dive site, 10° 18.023’ S, 150°57.177’ E, about 5.5 km west of Nuakata Island,Milne Bay Province, Papua New Guinea, 18-20 m,hand net, M. V. Erdmann, 16 June 2013. Paratypes: WAM P. 33884-002, male, 36.3 mm SL,collected with holotype; USNM 432234, 2 speci-mens, 26.5-37.6 mm SL, Barracuda Point, 10°15.458’ S, 150° 43.458’ E, about 16 km west of EastCape, Milne Bay Province, Papua New Guinea, 10m, hand net, G. R. Allen, 12 September 2014.Diagnosis: A species of Festucalex with 10-11 pec-

toral fin-rays, 18-19 dorsal-fin rays, 15-16 + 31 bodyrings (total 46-47), snout length 2.8-3.2 in headlength; lateral trunk ridge ending on penultimatetrunk ring, no spine-like projections on principle tailridges, and colour in life generally pinkish red.Description: Dorsal rays 18 (2 paratypes with 19);

anal rays 4; pectoral rays 10 (2 paratypes with 11);caudal rays 10. Trunk rings 15 (16); tail rings 31; to-tal rings 46 (47); subdorsal rings 2 + 4; total subdor-sal rings 6.Head 8.8 (7.5-9.1) in SL (11.0-13.3 % of SL). Me-

dian dorsal snout ridge entire, not strongly elevated,without bony knobs or spines, not confluent withorbital ridges. A low, indistinct opercular ridge; op-erculum with striae. Other head ridges low and in-distinct. Snout length 3.0 (2.8-3.2) in head length(3.8-4.2 % of SL). Snout depth 6.2 (6.3-6.8) in headlength (1.8-2.0 % of SL). Eye diameter 4.8 (4.4-4.9)in head length (2.4-3.1 % of SL). Body relatively smooth, without conspicuous

ridges and rings. Superior and inferior trunk ridgescontinuous with their respective tail ridges. Lateraltrunk ridge not deflected near anal fin, ends mid-lat-erally on penultimate trunk ring. Margins of head

and body ridges entire, neither denticulate or serrate.Dermal flaps absent. Body depth 23.6 (20.4-24.1) inSL (4.2-4.9 % of SL). Body width 26.8 (26.3-30.1)in SL (3.3-3.8 % of SL). Preanal length 2.5 (2.4-2.6)in SL (38.3-41.9 % of SL). Principal tail ridges be-hind dorsal fin without small posterior spine-likeprojections on each ring. Brood pouch under tailrings 1-12, pouch folds present, semi pouch closure.

Dorsal-fin origin on trunk; dorsal-fin base not el-evated. Dorsal-fin base length 10.6 (10.4-12.1) inSL (8.2-9.6 % of SL). Predorsal length 2.6 (2.5-2.6)in SL (37.8-39.3 % of SL). Pectoral-fin length 41.1(29.5-40.3) in SL (2.4-3.4 % of SL). Prepectorallength 7.9 (7.4-8.5) in SL (11.8-13.6 % of SL).Caudal-fin length 34.4 (25.5-33.2) in SL (2.9-3.9% of SL).


Gerald R. Allen and Mark V. Erdmann

Fig. 1. Festucalex rufus, preserved holotype, 37.8 mm SL, near Nuakata Island, Milne Bay Province, Papua New Guinea. Pho-to by G. R. Allen.

Fig. 2. Festucalex rufus, preserved holotype, 37.8 mm SL, showing close-up view of head. Photo by G. R. Allen.

Fig. 3. Underwater photograph of Festucalex rufus, female holotype 37.8 mm SL, Milne Bay Province, Papua New Guinea.Photo by G. R. Allen.



Fig. 4. Underwater photograph of Festucalex rufus, female, approximately 38 mm SL, East Cape area, Milne Bay Province,Papua New Guinea. Photo by N. DeLoach.

Fig. 5. Underwater photograph of Festucalex rufus, approximately 38 mm SL, East Cape area, Milne Bay Province, PapuaNew Guinea. Photo by N. DeLoach.

C o l o u r i n a l c o h o l (Figs 1-2): Holotype uni-formly pale pink. Paratypes ranging from pale pinkto pale grey, nearly whitish. Microscopic examina-tion of the type series reveals a pattern of fine, inter-laced greyish striae on the head and body.C o l o u r i n l i f e (Figs 3-5): Overall pinkish red,

covered with matrix of thin white striae on head andbody, which closely matches the pattern of spiculesevident on the tubular sponge with which it associ-ates.Sexual dimorphism: The two sexes are similar in

general appearance with the exception of the female

brood pouch situated on the tail between rings 1-12,which is present in the 36.3 mm SL paratype (WAMP. 33884-002). The presence of a fully developedpouch with ripe eggs, indicates a small maximumsize for this species. All observed specimens havebeen under about 40 mm SL. Distribution and habitat: The new species is cur-

rently known from several sites in the East Capearea of Milne Bay Province, Papua New Guinea. Ithas also been observed at Indonesia (northern Baliand Ceram) and the Solomon Islands. It frequentsexposed coral reef slopes and is generally encoun-tered solitarily or in pairs at depths ranging fromabout 10-25 m. The pipefish exhibits highly effec-tive camouflage with regards to its shape andcolour, closely matching an unidentified, shorttubular sponge among which the fish is invariablyfound (Fig. 4).Remarks: Festucalex rufus is distinguished from

other members of the genus on the basis of a combi-nation of features including a very short snout (2.8-3.2 in head length), relatively low pectoral-ray count(10-11) and a lateral trunk ridge that terminates onthe penultimate trunk ring (ends on tail rings in allother species except F. erythraeus). It is most similarto F. erythraeus and F. kulbickii, differing from bothspecies in having a shorter snout (2.8-3.2 in F. rufusvs. 2.1-2.6). It further differs from F. erythraeus inhaving fewer tail rings (31 vs. 32-37) and from F.kulbickii in having the lateral trunk ridge ending onthe penultimate trunk ring (vs. fifth to ninth tailring) and lacking small posterior spine-like projec-tions on the principle tail ridges of each ring. A keyto the species of Festucalex is provided below.

Key to the species of Festucalex (modified from Dawson 1985)

1a. Pectoral rays 10-14, snout length 2.0-3.2 in headlength ............................................................. 2

1b. Pectoral rays 16-17, snout length 1.7-1.8 inhead length ........................................... F. wassi

2a. Lateral trunk ridge ends on penultimate trunkring to fifth tail ring ....................................... 3

2b. Lateral trunk ridge ends on tenth to 13th tailring ................................................... F. prolixus

3a. Total rings 53-59 (usually more than 54) ...... 43b. Total rings 46-53............................................ 64a. Trunk rings modally 16-17; principal ridges of

tail rings with either 1-6 knobs or projectionson last 1-2 tail rings or with spine-like projec-tions on each ring........................................... 5

4b. Trunk rings modally 19; principal ridges of tailrings entire, without knobs or spiny projections........................................................... F. scalaris

5a. Trunk rings modally 17; principal ridges of last1-2 tail rings with 1-6 knobs or projections; typ-ically with narrow pale bar on postorbital ................................................................... F. cinctus

5b. Trunk rings 15-16; principal tail ridges bearinga small, posterior spine-like projection on eachring ................................................. F. kulbickii

6a. Trunk rings modally 15-16; without distinctridges on pectoral-fin base; lateral trunk ridgeends between penultimate trunk ring and firsttail ring ......................................................... 7

6b. Trunk rings modally 18; 2 distinct ridges onpectoral-fin base, lateral trunk ridge ends onsecond to third tail rings .................... F. gibbsi

7a. Snout length 2.1-2.6 in head length; snoutdepth 2.5-3.9 in snout length....... F. erythraeus

7b. Snout length 2.8-3.2 in head length; snoutdepth 2.0-2.2 in snout length ............. F. rufus

Etymology: The new species is named rufus (Latin:red or reddish) with reference to its distinctivecolouration.

ACKNOWLEDGEMENTSWe are extremely grateful to Rob Vanderloos, own-

er and operator of MV Chertan, who provided theopportunity to collect the type specimens. Underwa-ter photographs of the new species were generouslycontributed by Ned DeLoach. Lynne and Roger VanDok reported the first sighting of this fish fromMilne Bay Province and also informed us of theirsightings at Bali, Ceram, and the Solomon Islands.

REFERENCESALLEN, G. R. & ERDMANN, M. V. 2012. Reef Fishes of the

East Indies. Volumes I-III. Tropical Reef Research, Perth,Australia, 1292 pp.

ALLEN, G. R., STEENE, R., HUMANN, P. & DELOACH, N.2010. Reef Fish Identification Tropical Pacific. Fourth edi-tion. New World Publications, Jacksonville, Florida, 457pp.

DAWSON, C. E. 1985. Indo-Pacific Pipefishes (Red Sea to theAmericas). Gulf Coast Research Laboratory, Ocean Springs,Mississippi, 230 pp.

FRICKE, R. 2004. Review of the pipefishes and seahorses(Teleostei: Syngnathidae) of New Caledonia, with descrip-tions of five new species. Stuttgarter Beiträge zurNaturkunde. Serie A (Biologie) 668: 1-66.

KUITER, R. H. 2009. Seahorses and their relatives. AquaticPhotographics, Seaford, Australia, 333 pp.


Gerald R. Allen and Mark V. Erdmann


Index of aqua Vol. 20 (1-4)(Index by: 1. Author(s); 2. New Taxa; 3. Biology/Ecology/Biography/Reviews)

Author(s):Allen, Gerald. R., Hadiaty, Renny K. and Unmack, Peter J.: Melanotaenia flavipinnis, a new species of Rainbowfish (Melanotaeniidae) from Misool Island, West Papua Province,Indonesia. aqua 20 (1): 35-52, 15 January 2014Allen, Gerald R., Peristiwady, Teguh and Erdmann, Mark V.: Vanderhorstia lepidobucca, a new species of shrimpgoby from Sulawesi, Indonesia. aqua 20 (2): 81-86, 15 April2014Allen, Gerald R. and Hadiaty, Renny K.: Two new species of freshwater gudgeons (Eleotridae: Mogurnda) from the Arguni Bay Region of West Papua, Indonesia. aqua 20 (2):97-110, 15 April 2014Allen, Gerald R., Unmack, Peter J. and Hadiaty, Renny K.: Three new species of Rainbowfishes (Melanotaeniidae) from the Birds Head Peninsula, West Papua Province, In-donesia. aqua 20 (3): 139-158, 29 July Anderson, William D. Jr.: The plates in Holbrook’s Ichthyology of South Carolina, 1860. aqua 20 (4): 173-218, 15 October 2014Dalton Tavares Bressane Nielsen, Mayler Martins and Ricardo Britzke: Description of a new species of annual fish, Maratecoara gesmonei (Cyprinodontiformes: Rivulidae)from the rio Xingu system, Amazon basin, Brazil. aqua 20 (2): 87-96, 15 April 2014 Darshan, Achom, Dutta, Rashmi, Kachari, Akash, Gogoi, Budhin, Aran, Kamhun and Das, Debangshu Narayans: Creteuchiloglanis payjab, a new species of glyptosterninecatfish (Siluriformes: Sisoridae) from Yomgo River, Arunachal Pradesh, India. aqua 20 (2): 73-80, 15 April 2014 Keivany, Yazdan and Esmaeili, Hamid Reza: Threatened fishes of the world: Aphanius pluristriatus (Jenkins 1910) (Cyprinodontidae). aqua 20 (2): 67-72, 15 April 2014 Nielsen, Dalton Tavares Bressane, Martins, Mayler, Medeiros de Araujo, Luciano and dos Reis Suzart, Rogério: Hypsolebias shibattai, a new species of annual fish (Cyprin-odontiformes: Rivulidae) from the rio São Francisco basin, northeastern Brazil. aqua 20 (1): 27-34, 15 January 2014Nielsen, Dalton Tavares Bressane and Brousseau, Roger: Description of a new annual fish, Papiliolebias ashleyae (Cyprinodontiformes: Rivulidae) from the upper Rio Mamorébasin, Bolivia. aqua 20 (1): 53-59, 15 January 2014Nielsen, Dalton Tavares Bressane, Martins, Mayler, Medeiros de Araujo, Luciano, Origuela de Lira, Fabio and Faour, Amer: Hypsolebias trifasciatus, a new species of annualfish (Cyprinodontiformes: Rivulidae) from the rio Preto, rio São Francisco basin, northeastern Brazil. aqua 20 (3): 131-138, 29 July 2014 Randall, John E.: The goatfishes Parupeneus cyclostomus, P. macronemus and freeloaders. aqua 20 (2): 61-66, 15 April 2014Suzuki, Toshiyuki and Randall, John E.: Four new gobiid fishes of the genus Bryaninops from the East Indies. aqua 20 (1): 11-264, 15 January 2014Vagelli, Alejandro A.: Ephemeral Sexual Dichromatism in Quinca mirifica (Teleostei, Apogonidae), a Black Apogonid with Solitary Behavior. aqua 20 (1): 1-10, 15 January2014Valdesalici, Stefano and Eberl, Wolfgang: Aphyosemion mengilai, a new killifish species from the northern Massif du Chaillu, central Gabon (Cyprinodontiformes: Notho-branchiidae). aqua 20 (2): 111-116, 15 April 2014 Valdesalici, Stefano and Brousseau, Roger: A new Papiliolebias species (Teleostei: Cyprinodontiformes: Rivulidae) from Bolivian Amazon. aqua 20 (3): 117-122, 29 July 2014Vieira Volcan, Matheus, Krause Lanés, Luis Esteban and Cardozo Gonçalves, Ândrio: Austrolebias bagual, a new species of annual fish (Cyprinodontiformes: Rivulidae) fromsouthern Brazil. aqua 20 (4): 161-172, 15 October 2014Walsh, Fenton: Cirrhilabrus squirei, a new wrasse (Perciformes; Labridae) from the Great Barrier Reef and Coral Sea, Australia. aqua 20 (3): 123-130, 29 July 2014

New Taxa:Aphyosemion mengilai n. sp. A new killifish species from the northern Massif du Chaillu, central Gabon (Cyprinodontiformes: Nothobranchiidae). aqua 20 (2): 111-116, 15April 2014Austrolebias bagual n. sp. A new species of annual fish (Cyprinodontiformes: Rivulidae) from southern Brazil. aqua 20 (4): 161-172, 15 October 2014Bryaninops annella n. sp. Four new gobiid fishes of the genus Bryaninops from the East Indies. aqua 20 (1): 11-264, 15 January 2014Bryaninops earlei n. sp. Four new gobiid fishes of the genus Bryaninops from the East Indies. aqua 20 (1): 11-264, 15 January 2014Bryaninops tectus n. sp. Four new gobiid fishes of the genus Bryaninops from the East Indies. aqua 20 (1): 11-264, 15 January 2014Bryaninops translucens n. sp. Four new gobiid fishes of the genus Bryaninops from the East Indies. aqua 20 (1): 11-264, 15 January 2014Cirrhilabrus squirei n. sp. A new wrasse (Perciformes; Labridae) from the Great Barrier Reef and Coral Sea, Australia. aqua 20 (3): 123-130, 29 July 2014Creteuchiloglanis payjab n. sp. A new species of glyptosternine catfish (Siluriformes: Sisoridae) from Yomgo River, Arunachal Pradesh, India. aqua 20 (2): 73-80, 15 April2014 Hypsolebias shibattai n. sp. A new species of annual fish (Cyprinodontiformes: Rivulidae) from the rio São Francisco basin, northeastern Brazil. aqua 20 (1): 27-34, 15 Janu-ary 2014Hypsolebias trifasciatus n. sp. A new species of annual fish (Cyprinodontiformes: Rivulidae) from the rio Preto, rio São Francisco basin, northeastern Brazil. aqua 20 (3): 131-138, 29 July 2014 Maratecoara gesmonei n. sp. Description of a new species of annual fish, Maratecoara gesmonei (Cyprinodontiformes: Rivulidae) from the rio Xingu system, Amazon basin,Brazil. aqua 20 (2): 87-96, 15 April 2014Melanotaenia ericrobertsi n. sp. Three new species of Rainbowfishes (Melanotaeniidae) from the Birds Head Peninsula, West Papua Province, Indonesia. aqua 20 (3): 139-158,29 JulyMelanotaenia flavipinnis n. sp. A new species of Rainbowfish (Melanotaeniidae) from Misool Island, West Papua Province, Indonesia. aqua 20 (1): 35-52, 15 January 2014Melanotaenia laticlavia n. sp. Three new species of Rainbowfishes (Melanotaeniidae) from the Birds Head Peninsula, West Papua Province, Indonesia. aqua 20 (3): 139-158, 29JulyMelanotaenia multiradiata n. sp. Three new species of Rainbowfishes (Melanotaeniidae) from the Birds Head Peninsula, West Papua Province, Indonesia. aqua 20 (3): 139-158,29 JulyMogurnda arguni n. sp. Two new species of freshwater gudgeons (Eleotridae: Mogurnda) from the Arguni Bay Region of West Papua, Indonesia. aqua 20 (2): 97-110, 15 April2014Mogurnda kaimana n. sp. Two new species of freshwater gudgeons (Eleotridae: Mogurnda) from the Arguni Bay Region of West Papua, Indonesia. aqua 20 (2): 97-110, 15 April2014Papiliolebias ashleyae n. sp. Description of a new annual fish, Papiliolebias ashleyae (Cyprinodontiformes: Rivulidae) from the upper Rio Mamoré basin, Bolivia. aqua 20 (1):53-59, 15 January 2014Papiliolebias francescae n. sp. A new Papiliolebias species (Teleostei: Cyprinodontiformes: Rivulidae) from Bolivian Amazon. aqua 20 (3): 117-122, 29 July 2014Vanderhorstia lepidobucca n. sp. A new species of shrimpgoby from Sulawesi, Indonesia. aqua 20 (2): 81-86, 15 April 2014

Biology/Ecology/Biography/Reviews:Ephemeral Sexual Dichromatism in Quinca mirifica (Teleostei, Apogonidae), a Black Apogonid with Solitary Behavior. aqua 20 (1): 1-10, 15 January 2014The goatfishes Parupeneus cyclostomus, P. macronemus and freeloaders. aqua 20 (2): 61-66, 15 April 2014The plates in Holbrook’s Ichthyology of South Carolina, 1860. aqua 20 (4): 173-218, 15 October 2014Threatened fishes of the world: Aphanius pluristriatus (Jenkins 1910) (Cyprinodontidae). aqua 20 (2): 67-72, 15 April 2014

Examples of correct reference formats:

BLABER, S. J. M. 1980. Fish of the Trinity inlet system ofNorth Queensland, with notes on the ecology of fishfaunas of tropical Indo-Pacific estuaries. AustralianJournal of Marine and Freshwater Research 31:137-46.

DAY, J. H., BLABER, S. J. M., & WALLACE, J. H. 1981.Estuarine fishes. In: Estuarine Ecology with ParticularReference to Southern Africa. (Ed. J.H. Day.): 197-221. A. A. Balkema, Rotterdam.

DIMMICH, W. W. 1988. Ultrastructure of North Ameri-can cyprinid maxillary barbels. Copeia 1988 (1): 72-79.

TREWAVAS, E. 1983. Tilapiine Fishes of the GeneraSarotherodon, Oreochromis and Danakilia. BritishMuseum (Natural History), London, 583 pp.

6. Submission of manuscript and illustrations: The man-uscript and illustrations must be submitted digitally to theScientific Editor:

Flávio C. T. LimaMuseu de Zoologia da Universidade Estadual

de Campinas “Adão José Cardoso”Rua Albert Einstein s/n, Caixa Postal 6109,

13083-863 Campinas, São Paulo, Brazil E-mail: [email protected]

to whom all subsequent correspondence shall be ad-dressed. Texts, tables, and graphs should be in Microsoft-compatible electronic form.

After the paper has been accepted for publication, illustra-tions as high-resolution TIF files or original photographs(ideally transparencies; otherwise glossy prints, preferablyin the size in which they will appear - the type area of aquais 158 x 224 mm, one column is 76 mm wide) must besent to:

Aquapress, The Managing Editor Via G. Falcone 11,

27010 Miradolo Terme (Pavia), ItalyE-mail: [email protected]

Authors should retain copies of all materials for reference.

Proofs of accepted papers will be sent as PDF files by e-mail attachment to the corresponding author.

7. Evaluation of manuscripts: manuscripts will be evaluat-ed by the editors and referees. Papers are accepted on theunderstanding that they have not and will not be pub-lished elsewhere.

8. Reprints: Authors will receive one free copy of the issuein which their paper appears and an e-print in PDF for-mat. Additional copies may be ordered from Aquapress.

Caution: To place e-print in PDF format into GoogleScholar or elsewhere for free access infringes copyrights.

1. Manuscript preparation: manuscripts must be submit-ted in English. In exceptional cases aqua may providetranslations of manuscripts written in French, German,Italian, or Spanish.

Manuscripts must be word-processed in MicrosoftWORD and submitted in an electronic form. Generic,specific, and sub-specific names must be italicised. All pa-pers must conform to the International Code of Zoologi-cal Nomenclature. Authors are strongly advised to followthe format set out in previous publications of aqua.

2. Title: the title must be short and should precisely iden-tify the main topic of the article. Names of genera orspecies are followed by the systematic group to which theybelong. Author name(s) are given in full beneath the title,followed by the complete mailing and e-mail address(es).

3. Abstract: the abstract should not exceed 250 words anddraw attention to the principal conclusions. It should notcontain any uncommon abbreviations or literature cita-tions. The inclusion of abstracts in other languages is op-tional.

4. Subject matter: the text of the manuscript is usuallyarranged in four main sections: Introduction, Materialsand methods (including a key to abbreviations), Results,and Discussion. Other subdivisions may be chosen de-pending on the material presented. Acknowledgementsshould be placed between the text and references.

Scientific names of genera, species, and subspecies shouldbe followed by the name(s) of author(s) and the year ofpublication on first mention. A description of a new taxonmust contain the following sections: Material, Diagnosis,Description, and Affinities. Synonyms must be arranged in chronological order. Identification keys must be di-chotomous.

Holotype and paratypes must be clearly identified, the in-stitution in which they have been deposited named, andthe catalogue numbers given. Private collections are not ac-ceptable as repositories for holotypes.

DNA sequences must be archived in GenBank, DNADatabank of Japan or European Molecular Biology Labora-tory. Voucher specimens associated with DNA sequencesmust be deposited in a recognized research collection acces-sible to the professional community. Accession numbers forsequences and catalog number for vouchers must be includ-ed as a table in an appendix to the final manuscript.

The metric system and SI units must be used. Tempera-tures are given in °C. Fractions should not be used.

5. References to literature: the name-year system must beused. The list of references should be placed at the end ofthe paper, alphabetically arranged according to authorname. Only those publications cited in the paper may beincluded. Titles of journals must be given in full.

Guidelines for Authors

New rainbowfishes of the Aru Archipelago in a upcoming issue of aqua.

aquaInternational Journal of Ichthyology

Vol. 21 (1), 15 January 2015

Contents:

Eugenie Clark, Diane R. Nelson and Rachel Dreyer: Nesting sites and behavior of the deep water triggerfish Canthidermis maculata (Balistidae) in the Solomon Islands and Thailand ............ 1-38

María de las Mercedes Azpelicueta and Stefan Koerber: On some freshwater fish species reported by Perugia (1891) from Argentina .................................................................................... 39-46

Gerald R. Allen and Mark V. Erdmann: Festucalex rufus, a new species of pipefish (Syngnathidae) from Milne Bay Province, Papua New Guinea .................................................................................... 47-51

Index of aqua Vol. 20 (1-4) ............................................................................................................ 52

Papers appearing in this journal are indexed in: Zoological Record; BioLIS – Biologische Literatur Information Senckenberg;

www.aqua-aquapress.com; www.aquapress-bleher.com; www.Joachim-Frische.comhttp://scholar.google.it/scholar?q=aqua+international+journal+of+ichthyology&btnG=&hl=it&as_sdt=0%2C5

Cover photo: Canthidermis maculata with dark pectoral marking over its nest in foreground; second C. maculata with moderatepectoral marking associated with nest, Mbaleva Island, Solomon Islands. Photo © D. Seifert/www.douglasunderwater.com, 2014.

aquaaqua-aquapress.com/pdf/AQUA21(1).pdfEugenie Clark¹, Diane R. Nelson² and Rachel Dreyer¹ 1)...

Documents

Transcript of aquaaqua-aquapress.com/pdf/AQUA21(1).pdfEugenie Clark¹, Diane R. Nelson² and Rachel Dreyer¹ 1)...