aqua Journal of and Aquatic Biology Vol. 8 (2), March 2004

Aquapress ISSN 0945-9871 aqua - Journal of Ichthyology and Aquatic Biology

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The subadult of the labrid Novaculoides macrolepidotus , a mimic of waspfishes of the

John E. Randall and Andreas Spreinat

Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA Domäneweg 1, D-37079 Göttingen, Germany

Accepted: 17.09.2003

Keywords portamento, mostrandosi riluttante a muoversi e Mimicry, marine fishes, Novaculoides , Ablabys tenendo completamente eretta la sua pinna dorsale.

Abstract Introduction A brief review of mimicry in marine fishes is followed There has long been confusion regarding the terms by the example of the subadult of the labrid fish protective resemblance and mimicry. Poulton (1898) Novaculoides macrolepidotus believed to be a mimic defined them as follows: “In the former an animal of venomous scorpaeniform fishes of the genus resembles some object which is of no interest to its Ablabys . In addition to the resemblance in form and enemy, and in so doing is concealed; in the latter an colour to species of Ablabys , the subadult of this animal resembles an object which is well-known and labrid fish mimics Ablabys behavior by showing a avoided by its enemy, and in so doing becomes con - strong reluctance to move and by holding its dorsal fin spicuous.” After their discovery of the blenny Aspi - fully erect. dontus taeniatus mimicking the cleaner Labroides dimidiatus , Randall and Randall (1960) Zusammenfassung revised Poulton’s definition of mimicry: “…an animal Nach einer kurzen Übersicht über Fälle von Mimikry resembles an object which is well-known and is bei Meeresfischen wird das Beispiel der subadulten avoided or not preyed upon by its enemy, and in so Lippfische beschrieben , die als „Nachahmer“ der gifti - doing becomes conspicuous.” gen Skorpionsfische der Gattung Ablabys aufgefasst Far from being avoided, the cleaner wrasse is werden. Neben der Ähnlichkeit in Form und Farbe zu sought by other fishes of the community for its para - den Ablabys -Arten, ahmen die subadulten Tiere des site removal services (but it also feeds heavily on Lippfischs auch das Verhalten von Ablabys nach, mucus). It serves as the model for the mimicking indem sie sich bei der Bewegung stark zurückhalten blenny to enable the latter to get close enough to nip und die Rückenflosse gänzlich aufgerichtet halten. the fins of its reef-fish prey. Cott (1957) differentiated two well-known types of Résumé mimicry, Batesian and Müllerian. Batesian mimicry is Un bref survol du mimétisme chez les poissons de when “a relatively scarce, palatable, and unprotected mer est suivi par le cas du subadulte du labridé species resembles an abundant, relatively unpalat - Novaculoides macrolepidotus susceptible d’être un able or well-protected species, and so becomes dis - imitateur des poissons-scorpions venimeux du genre guised. In Müllerian mimicry, on the other hand, a Ablabys . En plus de la ressemblance de forme et de number of different species all possessing apose - couleur avec des espèces d’ Ablabys , le subadulte de matic attributes and appearance resemble one ce labridé imite le comportement des Ablabys en another, and so become more easily recognized.” montrant une forte répugnance au mouvement et en The resemblance of the harmless black-barred érigeant au maximum sa dorsale. snake colubrinus to the venomous sea snake Platurus colubrinus has long been given as a Sommario classic example of Batesian mimicry in the sea. Una breve rivisitazione del mimetismo nei pesci Another is the black-marked pectoral fin of the com - marini è qui proposta traendo come esempio il mon sole Solea vulgaris which resembles the black subadulto del labride Novaculoides macrolepidotus, dorsal fin of the venomous weeverfishes (Trachinus ritenuto assumere le sembianze di scorpeniformi draco and T. vipera). velonosi del genere Ablabys. Oltre alle somiglianze di Remarkable examples of Batesian mimicry are colore e forma, il labride imita Ablabys anche nel com - those blennies of the genera Ecsenius , Petroscirtes ,

45 aqua vol. 8 no. 2 - 2004 The subadult of the labrid fish Novaculoides macrolepidotus , a mimic of waspfishes of the genus Ablabys and Plagiotaenia that mimic the venomous fang dorsal ray of the Asterorhombus fijiensis has blennies of the genus Meiacanthus (Springer and been shown to serve as a lure (Amaoka et al., 1994). Smith-Vaniz, 1972; Losey, 1972; Smith-Vaniz, 1976). Examples of entire fishes that are aggressive mim - Also mimicking the venomous fang blennies, though ics of benign fishes to get closer to prey are species less precisely, are one species of cardinalfish of the of the western Atlantic butter hamlets ( Hypoplectrus ) genus Cheilodipterus and two species of the that resemble damselfishes (Pomacentridae) (Ran - nemipterid genus Scolopsis (Allen et al., 1975),. dall and Randall, 1960); the juvenile of the snapper Unique is the example of McCosker (1977) depicting Lutjanus bohar that resembles the damselfishes the fright posture of the plesiopid fish Calloplesiops Chromis ternatensis and C. iomelas (Russell et al., altivelus , commonly known as the Comet, that looks 1976); the juvenile of the grouper Anyperodon like the head of the moray eel Gymnothorax meleagris . leucogrammicus that mimics labrid fishes of the Another unusual example is the mimicry of genus Halichoeres (Russell et al., 1976); and the angelfishes of the genus Centropyge by the young of juvenile of the grouper Aethalopera rogaa that resem - the surgeonfish Acanthurus pyroferus . Randall & bles certain drab damselfishes, as well as dark Randall (1960) first described this in Tahiti, with the species of the angelfish genus Centropyge (Snyder et brightly-coloured Centropyge flavissima as the model. al., 2001). They suspected that the angelfish was a cleaner and Longley (1917: 279) warned that the resemblance of therefore protected from predation. Disproving that, one fish to another does not necessarily mean that they then tested the possibility of the angelfish having mimicry is involved. Mere resemblance might be the some quality repellent to a predator. Failing to case with the cardinalfish Fowleria sp., suggested as demonstrate that, they admitted to not knowing the a mimic of scorpionfishes by Siegel and Adamson biological basis of the mimicry. Years later, after (1983), or the similar example of Fowleria abocellata experiencing the difficulty of taking underwater pho - (= F. vaiuli) advocated as mimicking Scorpaenodes tographs of the species of Centropyge , the author guamensis by Goren and Karplus (1983). Much more determined that these small angelfishes are convincing is the apparent mimicry of a scorpionfish extremely wary and quickly take cover in the reef by the percoid fish Centrogenys vaigiensis . It was when approached. Predators learn the same and try first described by Quoy and Gaimard (1824) as a for easier prey. species of Scorpaena . Cuvier in Cuvier and Valenci - Randall and McCosker (1993) adopted the term ennes (1829) realized it was not a scorpionfish and social mimicry for fishes, used previously for mixed renamed it Centropristes scorpenoides , but Quoy and flocks of birds, when two or more species of similar Gaimard’s species name prevails. The mimicry was shape and colour aggregate for the protection pro - discussed by Whitley (1935). vided by schooling. Their example in fishes was find - We present here a new example of Batesian mim - ing three anthiine fishes, a pomacentrid, and a icry, with a venomous scorpaeniform fish serving as blenniid that have a common yellow and pink colour the model. pattern and feed together in aggregations on zoo - plankton. Another example is the goatfish Mul - Novaculoides macrolepidotus , a mimic of loidichthys mimicus which is coloured yellow with blue Ablabys spp. stripes like the snapper Lutjanus kasmira and schools The fish shown in Figure 1, an estimated 8 cm total with it (Randall and Guézé, 1980). length, was photographed in March 2001 at a depth of The term aggressive mimicry has come into vogue about 8 m in the Watamu National Marine Park, to depict an animal or a part thereof that will enable a Kenya by the second author. He first thought it was a predator to get closer to its prey or attract its prey to species of the waspfish genus Ablabys (subfamily within striking range. Therefore the Batesian blenny Tetraroginae of family ), because of the mimic Aspidontus taeniatus is also an aggressive colour, the slender compressed body, and especially mimic. the higher anterior part of the dorsal fin and its origin Perhaps the best known examples of aggressive on the head. It was encountered in plant debris, so he mimicry are found in the frogfish family Antennariidae tried to get it to move it away from the debris for his (and other ceratioids). The esca (bait) of the illicium photograph. However, it could be enticed to move (fishing pole) can look very much like a small fish, only with difficulty, and then very slowly. The dorsal fin worm or shrimp, and serves to attract prey to within was held fully erect during the whole time the fish was striking distance of the mouth of the frogfish (Pietsch observed. Upon close inspection of his photographs, and Grobecker, 1978, 1987). Similarly, the slender it was decided the fish was a labrid, so he sent the lure from tissue inside the lower jaw of some of the photographs to the first author whose initial identifica - stargazers ( Uranoscopas ) is enticingly wriggled like a tion was also Ablabys . Once realizing it was a labrid, worm from the mouth of the otherwise hidden stargaz - he was able to identify it as a subadult of Novacu - ers. One snake eel has adopted a comparable oral loides macrolepidotus (Bloch). This widespread Indo- lure (McCosker, 1982). Also the elongate frilled first Pacific labrid was only recently classified in the new aqua vol. 8 no. 2 - 2004 46 John E. Randall and Andreas Spreinat

Fig. 1. Subadult of Novaculoides macrolepidotus, about 8 cm total length, Watamu, Kenya, 8 m. Photo by A. Spreinat.

Fig. 2. Large male of Novaculoides macrolepidotus in seagrass , Cebu, Philippines, 2 m. Photo by J. E. Randall.

Fig. 3. Adult of , about 9 cm total length, Mindoro, Philippines. Photo by A. Spreinat.

47 aqua vol. 8 no. 2 - 2004 The subadult of the labrid fish Novaculoides macrolepidotus , a mimic of waspfishes of the genus Ablabys genus Novaculoides (Randall, 2004). Adults are usu - McCosker, J. E. 1977. Fright posture of the plesiopid ally green and live in seagrass beds, hence the com - fish Calloplesiops altivelis : an example of Batesian mon name (Fig. 2). They do not mimicry. Science, 197 : 400–401. hold their dorsal fin erect, and they are difficult to McCosker, J. E. 1982. A new genus and two new approach closely. Any overt movement by an species of remarkable Pacific worm (Ophichthi - observer results in their quickly disappearing into the dae, subfamily Myrophinae). Proceedings of the Cal - seagrass they resemble. ifornia Academy of Sciences, 43 (5): 59–66. Three species of Ablabys are recognized, A. binota - Pietsch, T. W. & D. B. Grobecker. 1978. The com - tus (Peters), A. macracanthus (Bleeker), and A. taeni - pleat angler: aggressive mimicry in an antennariid anotus Bleeker, any of which could serve as a model anglerfish. Science, 201 : 369-370. for Novaculoides macrolepidotus . An underwater pho - Pietsch, T. W. & D. B. Grobecker. 1987. Frogfishes tos is presented here of the Waspfish, A. of the World. xxii + 420 pp. Stanford University taenianotus , the most widespread and apparently the Press, Stanford, CA. most common species of the genus. Poulton, W. B. 1898. Natural selection the cause of The photographed subadult of Novaculoides mimetic resemblance and common warning colors . macrolepidotus allowed itself to be moved to and fro Journal of the Linnaean Society (Zoology), 26 : 558- with the surge, and it resembled the brown fragments 612. of dead seagrass in which it was first encountered. In Quoy, J. R. C. & P. Gaimard. 1824-1826. Pp. 192- this sense it is an example of protective resemblance. 401 in Voyage autour du Monde…Exécuté sur les However, we believe the wrasse achieves more pro - Corvettes de S.M. “l’Uranie” et “La Physicienne” tection as a mimic of waspfishes. The mimicry is evi - pendant les Années 1817-1820 . Pillet Aîné, Paris. dent because of the constant elevation of its dorsal fin Randall, J. E. 2003. Novaculoides , a new genus for (therefore looking more like a fish than a piece of sea - the Indo-Pacific labrid fish Novaculichthys grass) and in its reluctance to move, which is charac - macrolepidotus. Aqua teristic of venomous fishes in general. Randall, J. E. & P. Guézé. 1980. The goatfish Mulloi - dicthys mimicus , n. sp. (Pisces:Mullidae) from Ocea - Acknowledgements nia, a mimic of the snapper Lutjanus kasmira We thank Ralf Boecker and Phillip C. Heemstra for (Pisces: Lutjanidae). Bulletin du Museum National reference photographs and John L. Earle, Lutz Gohr, d’Histoire Naturelle , sér. 4, 2 (2): 603-609. and Heemstra for helpful comments. Randall, J. E. & J. E. McCosker. 1993. Social mim - icry in fishes . Revue Française d’Aquariologie, 20 References (1): 5-8. Allen, G. R., Russell, B. C., Carlson, B. A. & W. A. Randall, J. E. & H. A. Randall. 1960. Examples of Starck II. 1975. Mimicry in marine fishes. Tropical mimicry and protective resemblance in tropical Fish Hobbyist, 25 (1): 47–56. marine fishes . Bulletin of Marine Science of the Gulf Amaoka, K., Senou, H. & A. Ono. 1994, Record of and Caribbean. 10 (4): 444-480. the bothid flounder Asterorhombus fijiensis from the Russell, B. C., Allen, G. R. & H. R. Lubbock. 1976. western Pacfic, with observations on the use of the New cases of mimicry in marine fishes. Journal of first dorsal-fin ray as a lure. Japanese Journal of Zoology. (London), 180 : 407-423. Ichthyology, 41 (1): 23-28. Seigel, J. A. & T. A. Adamson. 1983. Batesian mim - Cott, H. B. 1957. Adaptive Coloration in . icry between a cardinalfish () and a ven - xxxii + 508 pp. Menthuen and Co., Ltd., London. omous scorpionfish (Scorpaenidae) from the Philip - Cuvier, G. & A. Valenciennes. 1829 . Histoire pine Islands. Pacific Science, 37 (1): 75-79. Naturelle des Poissons . Vol. 3. xxviii + 500 pp. F.G. Smith–Vaniz, W. F. 1976. The saber–toothed Levrault, Paris. blennies, Tribe Nemophini (Pisces: Blenniidae). Goren, M. & I. Karplus. 1983. Preliminary observa - Monographs of the Academy of Natural Sciences of tions on the scorpion fish Scorpaenodes guamensis Philadelphia. , 19 : 1-196. and its possible mimic the cardinal fish Fowleria abo - Snyder, D. B., Randall, J. E. & S. W. Michael. 2001. cellata. Developments in Ecology and Environmen - Aggressive mimicry by the juvenile of the redmouth tal Quality, 2: 327-336. grouper, Aethaloperca rogaa . Cybium, 25 (3): 227- Longley, W. H. 1917. Studies upon the biological signif - 232. icance of animal coloration II. A revised working Springer, V. G. & W. F. Smith–Vaniz. 1972. Mimetic hypothesis of mimicry. American Naturalist, 51 : 257- relationships involving fishes of the family Blenniidae. 285. Smithsonian Contribution of Zoology , 112 : 1-36. Losey, G. S. 1972. Predator protection in the poi - Whitley, G. P. 1935. Fishes from Princess Charlotte son–fang blenny, Meiacanthus atrodorsalis , and its Bay, North Queensland. Records of the South Aus - mimics, Ecsenius bicolor and Runula laudandus tralian Museum, 5 (3): 345-363. (Blenniidae). Pacific Science, 26 (2): 129–139. aqua vol. 8 no. 2 - 2004 48 aqua, Journal of Ichthyology and Aquatic Biology

Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei)

1 2 James C. Tyler and Philip A. Hastings

1) National Museum of Natural History, Smithsonian Institution (MRC-106), Washington D.C. 20560-0106, USA. E-mail: [email protected] 2) Marine Biology Research Division, Scripps Institution of Oceanography (MC-0208), University of California San Diego, 9500 Gilman Dr., La Jolla, California 92093-0208, USA. E-mail: [email protected]

Accepted: 31.01.2004

Keywords sur base de 35 mâles. Il se distingue aisément de ses Emblemariopsis dianae, , Belize, mor - congénères par l’étendard orange distal entre les trois phological variation, cephalic sensory pores, caudal premiers rayons durs de la dorsale, sans bande skeleton blanchâtre sous l’étendard, et strictement trois pores sensoriels mandibulaires au lieu des quatre typiques Abstract pour la plupart des chaenopsidés. La nouvelle espèce Emblemariopsis dianae, new species, is described montre une différence significative quat au nombre de based on 35 males. It is easily distinguished from its pores céphaliques en rangées sélectionnées et quant congeners by the orange flag distally between the first à des détails du squelette caudal, y compris quant à three dorsal fin spines, without a whitish band below the la seule synapomorphie connue dans le genre Emble - flag, and modally three mandibular sensory pores mariopsis , la forme et la taille de l’épine neurale de instead of the four that are typical of most chaenopsids. l’avant-dernière vertèbre. La nouvelle espèce a été The new species exhibits significant variation in number trouvée uniquement dans des eaux lagunales calmes of cephalic sensory pores in selected series and in et peu profondes de la Barrière de Corail de Belize. details of the caudal skeleton, including variation in the only known synapomorphy of the genus Emblemariop - Sommario sis, the shape and size of the neural spine of the penul - Emblemariopsis dianae, nuova specie, è descritta timate vertebra. The new species has been found only sulla base di 35 individui maschi. Si distingue facilmente in low-energy, mid-shelf, lagoonal waters of the Belize dagli altri congeneri per il “flag” arancio in posizione dis - Barrier Reef. tale tra i primi tre raggi spinosi dorsali, senza la sot - tostante banda biancastra, e per un valore modale di tre Zusammenfassung dei pori sensoriali mandibolari anziché quattro come Die neue Art Emblemariopsis dianae wird auf der tipicamente riscontrato nella maggior parte dei Grundlage von 35 männlichen Exemplaren be- Chaenopsidae. La nuova specie mostra grande vari - schrieben. Ihre Vertreter lassen sich durch eine orange- abilità nel numero dei pori cefalici nelle file seriali e nei farbene Fahne zwischen den ersten drei Rücken - dettagli dello scheletro caudale, incluse variazioni del - flossenstrahlen von den nächsten Verwandten leicht l’unico carattere sinapomorfico noto per il genere unterscheiden; weiterhin durch das Fehlen eines Emblemariopsis, la forma e la grandezza della spina weißlichen Streifens unterhalb der Fahne sowie modal neurale della penultima vertebra. La nuova specie è drei Sinnesporen am Unterkiefer statt vier, wie es für die stata finora trovata solo lungo la barriera corallina del meisten Chaenopsiden üblich ist. Signifikant variierende Belize, in aree lagunari della fascia intertidale media a Merkmale sind bei dieser neuen Art die Zahl der Kopf - bassa velocità di flusso. poren sowie Einzelheiten des Schwanzskeletts - darunter die einzig bekannte Synapomorphie innerhalb Introduction der Gattung Emblemariopsis und Form und Größe der Of the 11 species of the chaenopsid tube-blenny vorletzten Wirbel. Bisher wurde die neue Art nur in den genus Emblemariopsis (sensu Hastings, 1997; the Lagunengewässern im niedrig-energetischen mittleren species summarized with keys by Williams, 2002), Riffbereich des Belize-Barriereriffs gefunden. seven are known to occur on the outer barrier reef plat - form of the Belize Barrier Reef complex. They are most Résumé frequently found in high-energy areas along the outer Emblemariopsis dianae, nouvelle espèce, est décrit edge of the barrier reef (Smith et al., 2003), such as at

49 aqua vol. 8 no. 2 - 2004 Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei) the Smithsonian Institution’s marine laboratory at Carrie phate (1 gram per 500 ml water), and four individuals Bow Cay. To these can be added a new species from (JCT 2001-18) were taken from holes in coral heads. the lower energy, quieter waters in the lagoonal setting Subsequently, in March-April 2001 and in March 2002, of the mid-shelf cays, including the Rhomboidal Cays, special efforts were made by Diane M. Tyler and J.C. about 20 km south-west of Carrie Bow Cay, and the Tyler to collect more specimens from holes in coral cays just to the north (Wee Wee) and north-west (Blue heads and to obtain natural history data. Specimens Ground Range) of the Rhomboidals. were located by searching for their blackish coloured The Rhomboidal Cays are a group of mangrove heads protruding from holes in mounds of massive coral islands in a lagoonal mid-shelf region about 10 km west (such as Montastraea, Siderastrea, Diploria). These of the outer edge of the barrier reef. They are unusual in specimens were trapped in clear plastic tubes placed that the mangroves are anchored to the top of a live and upright over the opening of the holes immediately after lush coral reef dominated by Agaricia tenuifolia Dana, the solution of quinaldine was squirted into the holes, 1846 (a thin-leaf lettuce coral), rather than to mud (Mac - driving out the partially narcotized and disoriented spec - intyre and Rützler, 2000). Included are patch reefs with imens. The great majority of the specimens resulted well-developed mounds of coral heads between and from the use of quinaldine. around these mangrove islets (especially on the wind - If the coral containing the hole from which the fish spec - ward sides). The northern group of cays within the imen was obtained could be broken away with a hammer Rhomboidals is referred to as the Pelican Cays, and and chisel, it was often collected to examine it for eggs these structurally defined cays have been the subject of and to identify the kind of invertebrate that had produced intensive biological and geological study by Smithson - the burrow. However, most burrows were left intact, and ian scientists and their colleagues visiting the Carrie some were marked with tape to facilitate subsequent vis - Bow Cay lab. A survey of the Pelican Cays was con - its and to help in recording occupancy by other blennies ducted between 1997 and 2001 to assess this ichthy - after the original inhabitant had been collected. ofauna and to compare it with that of other regions of the The original two collections by Tyler and Davis bear Belize Barrier Reef complex (Smith et al ., 2003). During JCT 2001-__ station numbers; all collections by D.M. this survey, a new species of chaenopsid blenny was Tyler and J.C. Tyler bear JDT 2001-__ or 2002-__ stat- found in the patch reefs protecting the mangrove cays, ion numbers. The list of specimens is given chronologi - and it is described below. cally and in numerical station order, starting with the Males of the new species have a distinctively coloured JCT stations and proceeding with the JDT stations. orangish distal margin or “flag” in the anterior part of the All specimens were radiographed to aid in recording spinous dorsal fin. Unlike some species of Emblemari - meristic features. We noticed that there was variation in opsis, this region of the dorsal fin is not prominently ele - the shape and length of the neural spine of the penulti - vated. The new species also is unique in having modally mate vertebra (npu2), but the radiographs were often three mandibular sensory pores, whereas most insufficiently clear to be able to determine the morphol - chaenopsids have four. Considerable variation is pre - ogy of this neural spine. Therefore, a series of 15 spec - sent in the caudal skeleton, but most individuals have a imens was cleared and counter-stained (Dingerkus and broad truncate neural spine on the penultimate vertebra, Uhler, 1977) to better understand this variation. Cleared a feature unique among chaenopsids to Emblemariop - and stained (c&s) individuals are noted as such in the sis. list of specimens. Cephalic sensory pores of 22 specimens were Methods counted with the aid of a fine jet of compressed air. Specimens are deposited at the National Museum of Counts for bilateral pore series are reported separately Natural History, Smithsonian Institution, Washington, for each side. In some specimens, some pores were dif - D.C. (USNM, which includes collections of the former ficult to discern, accounting for the different sample United States National Museum) and the Scripps Insti - sizes for some of the pore series. Terminology follows tution of Oceanography, La Jolla, California (SIO). All Smith-Vaniz and Palacio (1974), as modified by Hast - collections were made by scuba diving in relatively shal - ings (1990). low water, about 5 m deep or less. Specimen lengths are in mm standard length (SL). The first two specimens of the new species were col - lected during the January-February 2001 segment of the Pelican Cays Fish Survey (Smith et al ., 2003) at Emblemariopsis dianae n. sp . coral heads off North Elbow Cay (northern Rhomboidal Orangeflag blenny (Figs. 1-3; Tables I, II) Cays) at a small rotenone station (JCT 2001-4) by J. C. Tyler and W. P. Davis. To date, these are the only spec - This new species previously had been mentioned in imens of the new species that have been collected with a listing of the fishes of the Pelican Cays (Smith et al ., rotenone. A few days later, in the central Pelican Cays, 2003) as Emblemariopsis sp. nov., with a photograph the same collectors tried to obtain additional specimens and the common name redflag blenny, but orangeflag of the new species specifically using quinaldine sul - blenny is more descriptive of the colour of the distinc - aqua vol. 8 no. 2 - 2004 50 James C. Tyler and Philip A. Hastings

Fig. 1. Emblemariopsis dianae. Specimens (all males and not to scale) photographed freshly preserved in formalin on same day of collection: A, USNM 365355, 16.7 mm (holotype); B, USNM 365355, 17.5 mm; C, USNM 365356, 16.4 mm; D, USNM 365356, 13.6 mm. Photos by Roy E. Clark, Jr.

51 aqua vol. 8 no. 2 - 2004 Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei)

Fig. 2. Emblemariopsis dianae. Specimens (all males and not to scale) photographed more than one year after preser - vation: A, USNM 365365, 21.1 mm (largest known specimen); B, USNM 365356, 16.4 mm; C, USNM 367251, 14.4 mm; D, USNM 367250, 12.6 mm. Photos by James F. DiLoreto. tive flag. station and all of the subsequent specimens collected Type specimens with quinaldine from holes in coral are males. Repeated A total of 35 male specimens, 12.6-21.1 mm SL (mean efforts to find female specimens (by careful observation 15.0 mm SL) . and finger combing of the algal mat) on the surface of The two specimens originally collected in a rotenone the same coral heads from which males were extracted aqua vol. 8 no. 2 - 2004 52 James C. Tyler and Philip A. Hastings

Fig. 3. Emblemariopsis dianae . Photographs of the posterior region of the vertebral columns of cleared and counter- stained specimens (arrows indicate the distal extent of the neural spine of the penultimate vertebra, npu2). Photos by Philip A. Hastings. A, USNM 365363 with a broad, truncate npu2; B, USNM 367250 with a broad, truncate npu2; C, USNM 367245 with a broad, moderately long npu2; D, SIO 03-56 with a slender, long npu2; E, USNM 365362 with a dou - ble npu2 consisting of a slender, long anterior portion and a broad, truncate posterior portion (note also the double haemal spine of the penultimate vertebra and long centrum compared with that of the antepenultimate vertebra); F, USNM 367247 with scoliosis of the posterior vertebral column.

53 aqua vol. 8 no. 2 - 2004 Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei) from their holes have been unsuccessful. Surface- April 2001, quinaldine, 16°46.467’N, 88°08.652’W; bur - dwelling females, however, are equally difficult to find rows of both specimens collected, and both of coiled and collect in the related E. ruetzleri Tyler and Tyler, Petaloconchus molluscs; one burrow relatively com - 1997, at Carrie Bow Cay, with only two females having plete, with 3.6 mm diameter aperture and more than been found after similar efforts in previous years (Tyler 100 eggs of several stages of development; one burrow and Tyler, 1997). fragmentary and only region with 3.9 mm diameter The catalogue number is followed by the number of aperture retained. specimens, size or size range, location, depth, and USNM 365365, 2, 14.6-21.1 mm, extensive coral other collecting data. heads on sandy bottom in clear water on west (lagoon) Holotype: USNM 365355, 16.7 mm, North Elbow Cay side of Billy Hawk Cay, just south of south end of Cock - (Peripheral Rhomboidal Cays), north-east side of cay, ney Cay, just south of main cut between cays in south - patch reefs on sandy bottom, coral heads from bottom ern region of Blue Ground Range, 8–15 feet depth, field almost to surface, 8 feet depth, field no. JCT 2001-4, no. JDT 2001-4, J. C. and D. M. Tyler, 7 April 2001, J. C. Tyler and W. P. Davis, 28 January 2001, rotenone, quinaldine, 16°47.371’N, 88°09.540’W; smaller speci - 16°42.338’N, 88°10.870’W. men from burrow (collected) in dead coral and larger Paratypes: USNM 365355, 17.5 mm, same data as specimen from hole in live coral, with dark head of larger holotype; this paratype and the holotype photographed specimen difficult to distinguish from roundish black freshly preserved same day of collection (Fig. 1). sponges encrusted on otherwise live parts of coral near USNM 365356, 2, 13.6-16.4 mm, coral ridge on sandy to burrow of fish; burrow of smaller specimen of coiled bottom midway between Co-Cat Cay and Little Cat Dendropoma mollusc, with aperture diameter of 3.9 Cay (Pelican Cays), opposite Avicennia Cay, many mm, eggs of several stages present but burrow too small and large coral heads (Montastraea predomi - incomplete for total egg count estimate; specimens of nating), 5-10 feet depth, field no. JCT 2001-18, J. C. aspera abundant in holes in dead Tyler and W. P. Davis, 2 February 2001, quinaldine, coral in same habitat. 16°39.802’N, 88°12.023’W; both specimens pho - USNM 365366, 1, 16.2 mm, c&s, coral ridge on sandy tographed freshly preserved on the same day of col - bottom midway between Co-Cat Cay and Little Cat Cay lection; originally four specimens; two (13.9-14.9 mm) (Pelican Cays), opposite Avicennia Cay, many small transferred to SIO 03-55. and large coral heads (same region of coral heads as USNM 365362, 2, 12.9-13.9 mm, both c&s, coral USNM 365356, JCT 2001-18), 5–10 feet depth, field no. heads on silty sand bottom just south of middle of chan - JDT 2001-5, J. C. and D. M. Tyler, 10 April 2001, quinal - nel at south end of Wee Wee Cay, just west of ( i.e., dine, 16°39.802’N, 88°12.023’W; dark head of speci - lagoon side) reef extending south of Wee Wee toward men difficult to distinguish from roundish black sponges Pelican Cays, 5-10 feet depth, field no. JDT 2001-1, encrusted on otherwise live parts of coral near to dead J. C. and D. M. Tyler, 5 April 2001, quinaldine, coral surface with burrow of fish. 16°45.795’N, 88°08.763’W; specimens of Acanthem - USNM 367243, 1, 15.5 mm, c&s, coral heads on silty blemaria aspera (Longley, 1927) abundant in holes in sand bottom just south of middle of channel at south dead coral in same habitat. end of Wee Wee Cay, just west of ( i.e., lagoon side) of USNM 365363, 4, 13.0-20.8 mm, two of these (13.0 reef extending south of Wee Wee toward Pelican Cays and 19.3 mm) c&s, same region of coral heads just (same region of coral heads as USNM 365362, JDT south of Wee Wee Cay as USNM 365362, field no. JDT 2001-1), 5–10 feet depth, field no. JDT 2002-2, J. C. 2001-2, J. C. and D. M. Tyler, 6 April 2001, quinaldine, and D. M. Tyler, 8 March 2002, quinaldine, 16°45.795’N, 88°08.763’W; fractured burrows of three 16°45.795’N, 88°08.763’W (originally two specimens; of the four specimens collected; burrow of one of the one transferred to SIO, following); also collected in two larger specimens (unknown whether 19.3 or 20.8 same habitat, Emblemariopsis signifera in algal turf of mm) of unknown worm or mollusc origin (but probably coral heads, and Acanthemblemaria aspera from holes molluscan), a relatively long and straight tube, with 12 in dead parts of coral heads. mm diameter aperture and 75 mm total length, and with SIO 03-56, 1, 13.3 mm, c&s, same data for Wee Wee more than 100 eggs of several stages of development; Cay as USNM 367243, collected along with it and also burrows of both smaller specimens (13.0 and 13.2 mm) c&s; burrows of both of these specimens from JDT collected, and both of coiled Dendropoma molluscs, 2002-2 marked with tape, and when revisited three days with aperture diameters of 3.0 and 3.4 mm, both bur - later (11 March) one burrow (original occupant 13.3 rows with eggs but burrows too incomplete for total egg mm) empty and one burrow (original occupant 15.5 count estimates. mm) occupied by 19.0 mm long Acanthemblemaria USNM 365364, 2, 15.3-15.8 mm, reef and sand spit aspera, and when revisited two days later (13 March) west north-west of Wee Wee Cay, coral heads at 4-8 one burrow (from the 15.5 mm specimen) occupied by feet depth on gently sloping silty sand bottom leading to 13.2 mm specimen of E. dianae. a deeper steep slope with much dead coral and silty USNM 367244, 1, 15.3 mm, c&s, JDT 2002-3, J. C. sediment, field no. JDT 2001-3, J. C. and D. M. Tyler, 6 and D. M. Tyler, 8 March 2002, same region of coral aqua vol. 8 no. 2 - 2004 54 James C. Tyler and Philip A. Hastings

Table I. Meristics of male Emblemariopsis dianae (* includes holotype).

Dorsal fin spines Dorsal fin rays Anal fin spines Count XVIII XIX XX 10 11 12 II III Number of 1232* 1 22* 12 33* 1 specimens Anal fin rays Pectoral fin rays Vertebrae Count 19 20 21 13-13 13-14 14-14 11+25=36 11+26=37 Number of 4 29* 1 30* 32 5 27 specimens heads at Wee Wee Cay as USNM 367362, JDT 2001- 88°12.023’W; specimen catalogued separately from 1, and USNM 367243, JDT 2002-2; Emblemariopsis three other specimens (USNM 367250, following) from signifera collected in algal turf of same coral head as same dive and location because its burrow collected; burrow in dead coral of E. dianae. burrow of coiled Dendropoma mollusc, with aperture USNM 367245, 1, 14.8 mm, c&s, extensive coral diameter of 2.8 mm and about 65 eggs at several heads on sandy bottom in clear water on west (lagoon) stages of development. side of Billy Hawk Cay, just south of south end of Cock - USNM 367250, 3, 12.6-16.3 mm, two of these (14.3 ney Cay, just south of main cut between cays in south - and 16.3 mm) c&s, JDT 2002-8, same data as USNM ern region of Blue Ground Range, 8-15 feet depth, field 367249, but no burrows collected. no. JDT 2002-4, J. C. and D. M. Tyler, 11 March 2002, USNM 367251, 6, 13.0-15.8 mm, one of these (15.8 quinaldine, 16°47.371’N, 88°09.540’W (same group of mm) c&s, North Elbow Cay (Peripheral Rhomboidal coral heads as USNM 365365, JDT 2001-4); burrow Cays), north side of cay, patch reefs on sandy bottom, marked with tape and when revisited two days later (13 coral heads from bottom almost to surface, 5–10 feet March) empty; both Acanthemblemaria aspera and A. depth, field no. JDT 2002-9, J. C. and D. M. Tyler, 15 greenfieldi Smith-Vaniz and Palacio, 1974, abundant in March 2002, quinaldine, 16°41.655’N, 88°10.250’W; holes in dead coral in same habitat. this is largest collection of individuals of E. dianae; this USNM 367246, 1, 14.7 mm, c&s, JDT 2002-5, J. C. station somewhat further to north end of cay than other and D. M. Tyler, 13 March 2002, same region of coral collection at North Elbow Cay in which E. dianae taken heads at Wee Wee Cay as USNM 367362, JDT 2001- in rotenone collection at north-east side of cay during 1, and USNM 367243, JDT 2002-2, and USNM 367243, Pelican Cays Fish Survey, USNM 365355, JCT 2001-4. JDT 2002-3; specimen from previously unseen hole in USNM 367252, 1, 15.6 mm, JDT 2002-11, J. C. and dead coral; burrow marked with tape and empty when D. M. Tyler, 17 March 2002, same region of coral heads revisited two days later (15 March). at Wee Wee Cay as USNM 367362, JDT 2001-1, and USNM 367247, 1, 13.2 mm, c&s, JDT 2002-5, J. C USNM 367243, JDT 2002-2, and USNM 367243, JDT and D. M. Tyler, 13 March 2002, same dive at same 2002-3, and USNM 367246-367247, JDT 2002-5; this locality at Wee Wee Cay as USNM 367246; specimen specimen collected from same hole (marked with tape) kept separate because replacement for 15.5 mm spec - from which 15.5 mm specimen (USNM 367243) imen, USNM 367243, of E. dianae collected five days removed on 8 March, and itself replaced by 13.2 mm previously on 8 March (see below, USNM 367252, for specimen (USNM 367247) collected on 13 March, with second replacement individual from same hole). present specimen collected from same hole on 17 USNM 367248, 1, 13.1 mm, c&s (poorly), extensive March; burrow collected, of coiled Dendropoma mollusc, coral heads on sandy bottom in clear water on west with aperture diameter of 3.5 mm and no eggs present. (lagoon) side of Billy Hawk Cay, just south of south end of Cockney Cay, just south of main cut between cays in Diagnosis southern region of Blue Ground Range, 8-15 feet depth, Males of Emblemariopsis dianae are unique among field no. JDT 2002-6, J. C. and D. M. Tyler, 13 March the species of Emblemariopsis in having a bright orange 2002, quinaldine, 16°47.371’N, 88°09.540’W (same flag on the distal one-third to one-fourth of the anterior group of coral heads as USNM 365365, JDT 2001-4, region of the spinous dorsal fin. The colour is mostly and USNM 367245, JDT 2002-4). confined to between the first and third spines (tapering USNM 367249, 1, 13.3 mm, coral ridge on sandy bot - in some larger specimens to end posteriorly between tom midway between Co-Cat Cay and Little Cat Cay the third and fourth spines, with much less orange color (Pelican Cays), opposite Avicennia Cay, many small in the tapering region) and without a chalky white band and large coral heads (same region of coral heads as along the ventral margin of the orange flag. Emblemar - USNM 365356, JCT 2001-18, and USNM 365366, JDT iopsis dianae also is unique among the species of 2001-5), 5-10 feet depth, field no. JDT 2002-8, J. C. and Emblemariopsis in modally having three mandibular D. M. Tyler, 15 March 2002, quinaldine, 16°39.802’N, sensory pores.

55 aqua vol. 8 no. 2 - 2004 Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei)

Table II. Measurements of Emblemariopsis dianae, in percent of standard length (SL). H - holotype, No - number of other specimens measured, X - mean, R - range

Character H No XR Head length 27.5 35 27.6 23.2-29.5 Head depth 18.0 31 17.2 15.2-18.6 Snout length 5.4 31 6.4 5.4- 7.4 Orbit diameter 7.2 32 8.3 6.2- 9.5 Interorbital width 3.6 27 4.0 3.2- 4.6 Body depth 17.4 29 15,7 13.9-18.1 First dorsal fin spine length 11.4 33 11.2 8.2-15.5 Second dorsal fin spine length 9.0 31 9.3 6.6-11.4 Third dorsal fin spine length 10.2 31 10.1 7.3-13.4 Fourth dorsal fin spine length – 21 10.7 7.9-14.0 Middle dorsal fin spines length 13.2 32 12.7 11.0-14.6 Rear soft dorsal fin rays length 13.2 32 14.0 12.7-16.8 First pelvic fin ray length – 23 12.3 10.4-14.4 Second pelvic fin ray length – 36 15.6 12.9-17.8 Third pelvic fin ray length – 20 9.6 7.6-11.7 Pectoral fin length 19.8 26 20.6 15.2-24.0 Caudal fin lenght 18.6 32 19.7 16.8-22.0

Description anterofrontal = 1 (44); commissural = 1 (22); frontal = 0 See Table I for meristics and Table II for measure - (39) or 1 (5); supraorbital = 2 (43); posterior infraorbital ments. Length of head about 3.6 times in SL (three = 3 (43); anterior infraorbital = 3 (44); median supratem - largest specimens, 19.3-21.1 mm, tending to have rela - poral = 1 (17), 2 (1), or 3 (2); lateral supratemporal = 2 tively smallest heads, 23.2-25.9% SL). Depth of head (25) or 3 (10); posttemporal = 4 (30); preopercular = 4 about 5.8 times in SL. Snout short, slightly less than (40); common = 1 (22); mandibular = 3 (33) or 4 (9). diameter of orbit (snout about 1.3 times in orbit). Orbit Anterior end of dorsal fin not prominently elevated; about 3.3 times in head length (three largest specimens anterior spines not forming upraised lobe, not higher tending to have relatively smallest orbits, 6.2-7.3% SL). anteriorly than posteriorly. Distal edge of anterior Bony interorbital width about 2 times in orbit. Body region of dorsal fin with gentle concavity, with mem - depth at anal fin origin about 6.4 times in SL. branes between first few spines slightly incised. First Supraorbital cirrus short, simple, speckled with dorsal fin spine about 11.2% SL, about same length melanophores like head; cirrus usually simply rounded as fifth spine, with spines gradually increasing in distally, but sometimes (especially in large specimens) length to middle of series of spines, where length is with distal edge incised into two or three small lobes, about 12.7% SL; second spine shortest, about 9.3% almost trident-like if three lobes; cirrus slender to mod - SL; third spine about 10.1% SL; fourth spine only erate in width; width 2 to 3 times in cirrus length; cirrus slightly shorter than first, 10.7% SL. length one-half to two-thirds of pupil diameter. Dorsal fin spines somewhat longer in large specimens Anterior nasal opening tubular, with a simple cirrus than in small individuals. Here and for the subsequent projecting from rear edge; cirrus of variable length, from measurements, the mean is often followed in paren - about equal to tube length to 3 times tube length. theses by the range in percent of standard length. Teeth on lower (dentary) and upper (premaxillary) First dorsal fin spine 10.7% SL (8.2-13.4%) in 20 jaws similar, each with a row of approximately 20 mod - small specimens (12.6-14.9 mm, mean 13.6 mm), and erately sized canines, extending from symphysis, 12.1% SL (10.0-15.5%) in 13 large specimens across front of jaws, and turning posteriorly along each (15.3-21.1 mm, mean 17.0 mm). Second dorsal fin side of jaws. Teeth in both jaws of moderate size ante - spine 9.0% SL (6.6-10.8%) in 19 small specimens (as riorly, largest in anterior portion of lateral row, and grad - above in size and range, here and following), and ually decreasing in size posteriorly. Anteriorly in both 9.7% SL (8.5-11.4%) in 12 large specimens. Third jaws a patch of low, pointed teeth, 2 or 3 teeth wide, dorsal fin spine 9.5% SL (7.3-11.5%) in 19 small behind outer row of canines. Anterior margin of vomer specimens, and 11.0% SL (9.2-13.4%) in 12 large with approximately 10 small pointed teeth in crescentic specimens. Fourth dorsal fin spine 9.9% SL (7.9- row, with 3 or 4 small teeth behind. Palatine with a sin - 12.3%) in 13 small specimens, and 12.0% SL (10.5- gle row of 5 small canines. 14.0%) in 8 large specimens. Posterior soft dorsal-fin Number of cephalic sensory pores in each series as rays about 14.0% SL. follows: nasal = 1 (frequency of count = 44); Pelvic fin much shorter than pectoral fin; 1,3, each ray aqua vol. 8 no. 2 - 2004 56 James C. Tyler and Philip A. Hastings extending beyond incised membrane; second ray Flag at upper front of spinous dorsal fin mostly orange, longest, third ray shortest; pelvic fin spine a very short sometimes slightly orangish red; flag most prominent splint, only visible in cleared and stained specimens; distally between first three spines, sometimes tapering first pelvic fin ray mean 12.3% SL, second ray mean to termination between third and fourth spines; from 15.6% SL, third ray mean 9.6% SL. Pectoral and caudal fourth spine posteriorly, distal edge of membranes paler fins about equal in length, pectoral fin mean 20.6% SL than below. Depth of flag usually one-third to one-fourth and caudal fin mean 19.7% SL. of first dorsal-fin spine length. No white band below Dorsal fin XX, 11 or 12 (34% with 12 rays); a longer orange flag; membranes of fin below flag same dark space between bases of third and fourth spines than grey to black as rest of dorsal fin; depth of flag mean between the others. Anal fin usually II, 20 (one speci - 28% (20–35%) of first dorsal-fin spine length or 3.6 men unusual with 3 anal fin spines). Pectoral fin usually (2.8–5.0) times in first dorsal fin spine length. with 13 rays on both sides (14% with 13 on one side and Proximal two-thirds of pectoral fin darkly speckled, but 14 on the other, 6% with 14 on both sides). Caudal fin less speckled and much paler distally. Pelvic fin with with 13 segmented unbranched principal rays and 2–4, dark interradial membranes, rays pale beyond incised usually 3, procurrent rays above and below. Vertebrae: membranes. After long preservation (Fig. 2), orangish 11 abdominal and usually 26, sometimes 25, caudal; flag in dorsal fin becoming whitish, and tan and yellow - total usually 37, sometimes 36. ish overtones on body becoming pale white. Caudal skeleton somewhat variable. All specimens examined with typical chaenopsid features of fused Etymology upper and lower hypural plates, a free minimal hypural, The species is named in honour of Diane M. Tyler of and a single epural. Neural spine of penultimate verte - the Smithsonian Institution Press, in recognition of her bra (npu2) variable in shape and length: in most speci - studies of the behavioural ecology of chaenopsids at mens (10 of 15 c&s) npu2 broad, with its distal margin Carrie Bow Cay; she is the co-collector of most of the truncate, ranging from rectilinear to gently rounded, type specimens of this new species, and her dedicated extending much less than half distance from centrum to collecting efforts over the years in and around Carrie body outline (Fig. 3A & 3B), similar to that reported for Bow Cay have procured many important materials. other species of Emblemariopsis; in two specimens (USNM 367244, 367245) npu2 broad but longer, Distribution extending a little more than half-way to body outline Emblemariopsis dianae has been collected in four (Fig. 3C); in two specimens (USNM 367243, SIO 03-56) mid-shelf lagoonal areas to the south and west of the npu2 slender and similar to more anterior neural spines, Smithsonian Institution’s marine laboratory at Carrie extending two-thirds or more to body outline (Fig. 3D); Bow Cay, Belize. Whereas the marine laboratory is on in one specimen (USNM 365362) npu2 divided into a the outer edge of the Belize Barrier Reef complex, E. slender, long anterior portion and a broad, truncate pos - dianae has been collected only in calmer lagoonal terior portion (Fig. 3E). This variation in shape and size waters. The four areas are the Elbow Cays, about 14 of npu2 not related to body size. km south-west (bearing 222°) of Carrie Bow in the Colour of males when live: as seen with only head northern Peripheral Rhomboidal Cays; the central Peli - protruding from hole in coral, uniformly black, the can Cays, about 20 km south-west of Carrie Bow, with orange flag at front of dorsal fin not visible. Specimens the Pelicans being the northern group of cays in the blanch somewhat immediately upon being driven from Rhomboidal Cays (the Rhomboidal Cays are comprised their burrow with quinaldine, with head becoming less of the Peripheral Rhomboidals and the Pelicans); Wee intensely dark, more brownish black, and body becom - Wee Cay, about 7 km south-west of Carrie Bow; and the ing lighter coloured posteriorly. Blue Ground Range, a series of north to south cays Colour when freshly preserved: (Fig. 1) speckled approximately 8 km long, some 8 km west of Carrie with brownish black melanophores, most dense on Bow. These areas are discussed in Macintyre and Rüt - head, and becoming progressively less intensely speck - zler (2000), and their ichthyofauna is detailed in Smith et led from anteriorly to posteriorly on body; body with tan al. (2003). to yellowish overtones between darker speckling; large specimens usually more darkly pigmented than small Comparisons and Variation specimens. Two to four dark spots on opercular region The morphology of E. dianae is consistent with that of and on cheek, and often present on base of pectoral fin. other species currently placed in Emblemariopsis Long - Diffuse dark blotch laterally on caudal peduncle. Vertical ley, including separate nasal bones, a single short fins generally dark grey to black, except becoming pale supraorbital cirrus (absent in some species of the at distal edges. Body along bases of dorsal and anal fins genus), and, in most specimens, a truncate neural spine with indistinct pale crescents spaced about every sec - on the penultimate vertebra (npu2), the only feature ond ray. Abdomen darker than rest of middle of body, unique to Emblemariopsis among chaenopsids (Hast - with top of abdomen tending to be bordered by indistinct ings, 1997) . Although most cleared and stained speci - slightly pale band. mens of E. dianae have a broad, truncate npu2, two

57 aqua vol. 8 no. 2 - 2004 Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei) have a broad and moderately long (half-way out to body beyond the fourth dorsal fin spine to at least the fifth or outline) npu2, two have a slender and long npu2, and sixth spine (versus to between only the third and fourth one has a divided npu2, with the anterior part slender spines in E. dianae ) and the ventral margin of the flag is and long and the posterior part broad and truncate. In bordered by a chalky white band (versus no such white the latter specimen (USNM 365362; Fig. 3E), the length band in E. dianae ). of the centrum of the penultimate vertebra is slightly Emblemariopsis dianae is similar to E. tayrona, E. greater than that of more anterior vertebrae and the ramirezi, E. occidentalis, E. leptocirrus Stephens, 1970, haemal spine is divided, much as is the neural spine, and E. signifera (Ginsburg, 1942) in having modally 13 which we interpret as indicative of incomplete fusion of pectoral fin rays and a supraoccipital cirrus. Among two posterior vertebrae. The cause of the variation in these, Emblemariopsis dianae is most similar to E. lep - these features is unknown, but the presence in one tocirrus in having a relatively low anterior region of the specimen (USNM 367247; Fig. 3F) of severe scoliosis spinous dorsal fin, with the first dorsal fin spine only and in another (USNM 367251, 13.4 mm SL) of three slightly elongate relative to more posterior dorsal fin anal fin spines may indicate disruption or instability in spines; thus, the characters of E. dianae would approx - the developmental program of some individuals. imate those of E. leptocirrus in the key to the species of Emblemariopsis dianae is unique within Emblemariop - Emblemariopsis by Williams (2002). Emblemariopsis sis in modally having three mandibular sensory pores, leptocirrus differs from E. dianae, however, in having a although some variation exists. Seventeen of 22 speci - relatively uniformly pigmented anterior dorsal fin, with - mens carefully examined for the cephalic pore system out a prominent flag distally between the more anterior have three mandibular pores on both jaws, three speci - spines; in having the first few dorsal spines of about the mens have four mandibular pores on both jaws, and same length; and in having the interradial membranes three specimens are asymmetric, with three pores on of the dorsal fin spines not prominently incised (versus one side and four on the other. Counts made prior to E. dianae having a prominent flag, the second spine dis - clearing and staining were generally confirmed for 12 tinctly shorter than the first spine, and the membranes specimens. Those specimens with four external clearly incised between the first three spines in a con - mandibular pores have four evident pores internally: cave distal edge of the fin). pores 1-3 pass through the dentary and pore 4 is in the space between the dentary and the anguloarticular. Natural History Specimens with three external pores most often have Emblemariopsis dianae males have been found only pores 1, 2 and 4 present, and pore 3 absent, but occa - in the relatively quiet, low-energy, mid-shelf, lagoonal sionally (n = 5 rami in 3 specimens) pore 3 is present as waters many kilometers inside (westward of) the outer a tiny perforation, less than one-half the diameter of the edge of the Belize Barrier Reef. This protected habitat is other mandibular pores. In these later cases, this pore in marked contrast to the high-energy waters at the is either not open through the skin or can be overlooked outer edge of the barrier reef at Carrie Bow Cay where due to its small size. two other species of Emblemariopsis, E. ruetzleri Tyler The numbers of cephalic sensory pores in most other and Tyler, 1997, and E. pricei Greenfield, 1975, are series are invariant in E. dianae except for those along common. Both E. ruetzleri and E. pricei utilize similar the posterior dorsal surface of the head (frontal, median, habitats around the reef crest, on patch reefs just inside and lateral supratemporal series), a feature common the lagoon, and in the spur and groove zone in front of among chaenopsids. However, unlike E. dianae, the the reef crest (Tyler and Tyler, 1997, 1999). This differ - number of mandibular pores is usually constant in ential distribution of E. dianae relative to E. pricei and E. species of chaenopsids ( e.g., the genus Acanthemble - ruetzleri is documented by more than 100 quinaldine maria, see Smith-Vaniz and Palacio, 1974), with only collections (deposited at the Smithsonian’s National occasional variants reported in, for example, Stath - Musum of Natural History) of these taxa of tube dwelling monotus and Ekemblemaria (Hastings, 1992; Hastings chaenopsids across the outer barrier reef platform from and Springer, 1994). Other than four of the species of the reef crest westward far into the lagoon between Car - Stathmonotus (Hastings and Springer, 1994), E. dianae rie Bow Cay and the Rhomboidal Cays. In all three of is the only chaenopsid with predominantly three rather these species, mature males are confined to holes in than four mandibular pores. Stephens’ (1970, Table I) coral, and females (not known for E. dianae ) and imma - report of five mandibular pores in all genera of chaenop - ture individuals are surface dwellers. Whereas E. pricei sids includes the “common pore,” located between the and E. ruetzleri are similar in being found only in high- jaw articulation and preopercular series (see Smith- energy habitats, versus only in low-energy habitats for Vaniz and Palacio, 1974), in the mandibular pore count E. dianae, the males of E. ruetzleri and E. dianae are and does not include data for Stathmonotus. similar in being found mostly in holes in dead coral, ver - An orangish red distal flag similar to that of E. dianae sus in holes in live coral for E. pricei. occurs in E. tayrona (Acero, 1987), E. ramirez Whereas different energy levels characterize the (Cervigon, 1999) and E. occidentalis Stephens, 1970, restricted habitats of E. dianae versus E. ruetzleri and but the flag in these three species extends posteriorly E. pricei, at least two other species of Emblemariopsis aqua vol. 8 no. 2 - 2004 58 James C. Tyler and Philip A. Hastings in this region are found in a wide variety of water condi - ity of reoccupation of suitable shelter sites for E. dianae tions: E. leptocirrus is found both in the high-energy seems to be at about the same low level as for E. ruet - waters of offshore banks such as Glovers Reef and of zleri, and it surely is not like the rapid reoccupancy of the barrier reef such as Carrie Bow Cay and English burrows typical of E. pricei. Cay, and in the low-energy waters of Cat Cay in the Pel - The holes from which specimens of E. dianae were ican Cays (Smith et al., 2003); E. signifera is common in extracted were in dead parts of the coral heads, with the the cays of the offshore banks and the barrier reef single exception of one of the two specimens in USNM (many records in Greenfield and Johnson, 1981, and in 365365 (see list of type specimens), whose hole was in Smith et al., 2003), but it has also been collected along a live part of the head. with E. dianae in the quiet waters of the mid-shelf cays Of the eight burrows that were collected of E. dianae, at Wee Wee Cay (see annotations in list of type speci - seven were the coiled shells of “worm-shell” gastropod mens). We note that both mature males and females of molluscs, family Vermetidae. Of these vermetids, five the ubiquitous E. signifera are mostly out on the surface were of species of Dendropoma and two were of of corals and associated algal mats during daylight (also species of Petaloconchus (without determination to the see long-term observations of this species in Smith and species level). The diameters of the apertures of these Tyler, 1972), rather than with at least adult males being vermetid shells ranged from 2.8 to 3.9 mm, with total mostly confined to holes in corals as in many other coiled lengths estimated at about 25 to 60 mm; the size species of Emblemariopsis. of the specimens of E. dianae occupying these burrows Previous studies at Carrie Bow Cay of E. pricei and ranged from 13.0 to 15.8 mm. Five of the vermetid bur - E. ruetzleri have demonstrated a difference between rows had fish eggs from different depositions; three of the two species in how rapidly immature males these burrows were too incomplete for estimates to be occupy vacant shelter sites. In the case of E. pricei, made of the total number of eggs, but one had 65 eggs the numerous serpulid (Spirobranchus) worm tubes and the other more than 100 eggs. One vermetid bur - from which a mature male had been extracted were row had only the region of the aperture collected, and it usually immediately occupied by an immature male is unknown whether eggs were present. One of the lurking on the nearby surface (Tyler and Tyler, 1999). seven vermetid burrows, whose entire length could be In the case of E. ruetzleri, the burrows in coiled ver - examined after being broken open, had no eggs. In metid (Dendropoma) gastropods from which the fish addition to these seven coiled vermetid shells, one of had been extracted were not so frequently or rapidly the collected burrows was of much larger size and rela - occupied, and suitable holes in coral are apparently tively straight, with an aperture diameter of 12 mm and not in short supply for E. ruetzleri. a total length of about 75 mm; more than 100 eggs were An effort in 2002 to obtain similar data for Emblemari - present, in several stages of development. The size of opsis dianae was minimally successful, with only four the specimen of E. dianae occupying this large burrow burrows tagged and subsequently revisited. One of was either 19.3 or 20.8 mm (USNM 365363), these two these burrows, from which a 14.8 mm original occupant specimens being among the three largest specimens was extracted (USNM 367245) on 11 March, was collected. It could not be determined whether this large empty when revisited on 13 March and was not exam - burrow belonged to a mollusc or a worm, but it most ined again. Another burrow, from which a 14.7 mm orig - probably belonged to a non-vermetid mollusc. Thus, at inal occupant was extracted (USNM 367246) on 13 present, we can only state that E. dianae usually utilizes March, was empty when revisited on 15 March and was the tubes formed by at least two different vermetid mol - not examined again. On 8 March at Wee Wee Cay, two luscs but is sometimes found in tubes of a presently burrows were marked from which specimens of 13.3 unknown larger invertebrate. Many of the burrows of E. mm (SIO 03-56) and 15.5 mm (USNM 367243) were dianae that were not collected seemed frequently to be extracted. When revisited on 11 March, the burrow from those of coiled vermetid gastropods. At Carrie Bow Cay, which the 13.3 mm specimen had been extracted was E. pricei is mostly found in serpulid (Spirobranchus) empty and was also empty subsequently when revisited worm tubes, whereas E. ruetzleri is mostly found in ver - on 13 and 17 March. The burrow from which the 15.5 metid (Dendropoma) shells, so E. dianae and E. ruetz - mm specimen had been extracted on 8 March had a 19 leri are similar in this respect. mm specimen of Acanthemblemaria aspera (specimen The black head of E. dianae protruding from its burrow collected, not retained) when revisited on 11 March, and is especially easy to overlook when the burrow is near when this burrow was revisited on 13 March it had a patches of the encrusting black sponge Cliona carib - 13.2 mm specimen of Emblemariopsis dianae (USNM baea Carter, 1882, that is often present on coral heads 367247); when this same burrow was revisited for the in this region (see Rützler, 2002). last time on 17 March, it had a 15.6 mm specimen of E. dianae (USNM 367252). Thus, three of the four burrows Acknowledgements that were revisited were not reoccupied, but one of them Most of the field work for this study was done in con - was, by a combination during a nine-day period of two junction with Diane M. Tyler of the Smithsonian Institu - individuals of E. dianae and one of A. aspera. The rapid - tion Press, as part of the ichthyofaunal survey of the

59 aqua vol. 8 no. 2 - 2004 Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei)

Pelican Cays undertaken by C. Lavett Smith (American ogy, and distribution (Blenniidae, Chaenopsidae, Museum of Natural History, emeritus) and colleagues Labrisomidae, Tripterygiidae). Fieldiana, Zoology, n.s. on behalf of the Caribbean Coral Reef Ecosystem 8: 1-106. (CCRE) Program of the Smithsonian’s National Hastings, P. A. 1990. Phylogenetic relationships of Museum of Natural History, under the direction of Dr. tube blennies of the genus Acanthemblemaria Klaus Ruetzler; this is contribution 671 of that program. (Blennioidei). Bulletin of Marine Science, 47 : 725-738. At the National Museum of Natural History we have Hastings, P. A. 1992. Ekemblemaria lira, a new been greatly helped with radiographic services by San - blennioid fish from Ecuador, with comments on sexual dra J. Raredon, with photography in the field by Roy E. dimorphism and relationships in Ekemblemaria Clark, Jr., and with photography at the museum by (Teleostei: Chaenopsidae). C opeia, 1992 : 769-776. James F. DiLoreto, Jeffrey T. Williams, and Sandra J. Hastings, P. A. 1997. Phylogenetic relationships of the Raredon. We thank Susan L. Jewett, Lisa F. Palmer, Coralliozetus clade of chaenopsid blennies, with Sandra J. Raredon, Cindy Klepadlo, and H.J. Walker for description of a new genus (Teleostei, Blennioidei). arranging the loan of specimens. We are grateful to Ralf Bulletin of Marine Science, 61 (3): 743-761. G. Britz and G. David Johnson, not only for the excel - Hastings, P. A. & V. G. Springer. 1994. Review of lence of their clearing and counter-staining the initial Stathmonotus, with redefinition and phylogenic analy - specimens in this study, but also for their advice on sis of the Chaenopsidae (Teleostei: Blennioidei). interpretation of the variation in the caudal skeleton. Smithsonian Contributions to Zoology, 558 : 1-48. Identification of the invertebrates that formed the bur - Macintyre, I. G. & K. Rützler (Eds.). 2000. Natural his - rows utilized by E. dianae was generously provided by tory of the Pelican Cays, Belize. Atoll Research Bul - M.G. Harasewych, in collaboration with Rüdiger Bieler letin, nos. 466-480 : 1-332. of the Field Museum of Natural History, and the Rützler, K. 2002. Impact of crustose clionid sponges on sponges often found in the vicinity of E. dianae were Caribbean reef corals. Acta Geologica Hispanica, 37 : kindly identified by Klaus Ruetzler. 61-72. For generously letting us examine their collections of Smith, C. L. & J. C. Tyler. 1972. Space resource shar - colour photographs of numerous species of Emblemar - ing in a coral reef fish community. In: Results of the iopsis in life, we thank Jeffrey T. Williams; Les Wilk of Tektite Program: Ecology of Coral Reef Fishes (Eds. Reef Net Inc., Mississauga, Ontario, Canada; Peter B.B. Collette and S.A. Earle). Natural History Museum, Wirtz of FishBase, Madeira, Portugal; and William F. Los Angeles County, Science Bulletin, 14 : 125-170. Smith-Vaniz, U.S Geological Service, Gainesville, Smith, C. L., Tyler, J. C., Davis, W. P., Jones, R. S., Florida. Smith, D. G. & C. C. Baldwin. 2003. Fishes of the Pel - For providing us with information on Emblemariopsis ican Cays, Belize. Atoll Research Bulletin, 497 : 1-111. in the southern Caribbean, we thank Fernando Smith-Vaniz, W. F. & F .J. Palacio. 1974. Atlantic Cervigon, Museo Marino de Margarita, Venezuela; fishes of the genus Acanthemblemaria, with descrip - Oscar M. Lasso-Alcala, Museo de Historia Natural La tion of three new species and comments on Pacific Salle, Venezuela; and Arturo Acero, Universidad species (Clinidae: Chaenopsidae). Proceedings of the Nacional de Colombia. Academy of Natural Sciences of Philadelphia, 125 : 197–224. References Stephens, J. S. 1970. Seven new chaenopsid blennies Acero P., A. 1987 . The chaenopsine blennies of the from the western Atlantic. Copeia, 1970 : 280-309. southwestern Caribbean (Pisces, Clinidae, Tyler, D. M. & J. C. Tyler. 1997. A new species of Chaenopsinae). III. The genera Chaenopsis and chaenopsid fish, Emblemariopsis ruetzleri, from the Coralliozetus . Boletin Ecotropica, 16 : 1-21. western Caribbean off Belize (Blennioidei), with notes Cervigon, F . 1965 . Emblemariopsis randalli nov. sp., on its life history. Proceedings of the Biological Society una nueva especie de Chaenopsidae de las costas de of Washington, 110 (1): 24-38. Venezuela. Novedades Cientificas, Contribuciones Tyler, J. C. & D. M. Tyler. 1999. Natural history of the Ocasionales del Museo de Historia Natural La Salle, sea fan blenny, Emblemariopsis pricei (Teleostei: Serie Zoologica, 33 : 1-4. Chaenopsidae), in the western Caribbean. S mithson - Dingerkus, G. & L. D. Uhler. 1977. Enzyme clearing ian Contributions to Zoology, 601 : 1-24. of alcian blue stained whole small vertebrates for Williams, J. T. 2002. Chaenopsidae. Pages 1761-1767 demonstration of cartilage. S tain Technology, 52 : in: The Living Marine Resources of the Western Cen - 229–232. tral Atlantic. Volume 3: Bony Fishes Part 2 (Opistog - Greenfield, D. W. 1975 . Emblemariopsis pricei, a new nathidae to Molidae), Sea Turtles and Marine Mam - species of chaenopsid blenny from Belize. Copeia, mals. FAO Species Identification Guide for Fishery 1975 : 713-715. Purposes (Ed. K.E. Carpenter). Food and Agriculture Greenfield, D. W. & R. K. Johnson. 1981. The Organization of the United Nations, Rome, pp. 1375- blennioid fishes of Belize and Honduras, Central 2127. America, with comments on their systematics, ecol - aqua vol. 8 no. 2 - 2004 60 aqua, Journal of Ichthyology and Aquatic Biology

Five new shrimp gobies of the genus from islands of Oceania

John E. Randall

Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA

Accepted: 17.01.2004

Keywords Résumé Taxonomy, , Amblyeleotris , new species, On décrit ici cinq nouvelles espèces de gobies du central and south Pacific genre Ámblyleotris de l’Indo-Pacifique originaires d’île du Pacifique central et méridional: A. katherine , Abstract d’abord erronément décrite comme A. fasciata Five new species of shrimp gobies of the Indo- (Herre), sur base de 14 spécimens des îles de la Pacific genus Amblyeleotris are described from Société (localité type), des îles Cook, Marshall et Mar - islands of the central and south Pacific: A. katherine , iannes; A. biguttata , décrit à partir de 4 spécimens de formerly misidentified as A. fasciata (Herre), from 14 Nouvelle Caledonie (localité type) et des îles Salomon specimens from the Society Islands (type locality), (le nomerenvoie à deux grandes taches noires sur le Cook Islands, Marshall Islands, and Mariana Islands; menton); A. ellipse sur base de neuf spécimens de A. biguttata described from four specimens from New l’île Samoa (le nom fait allusion à la marque elliptique Caledonia (type locality) and the Solomon Islands dans la caudale); A. bellicauda , sur base d’un seul (named for two large black spots on the chin); A. spécimen de Nouvelle Calédonie (le nom se réfère ellipse described from nine specimens from American aux couleurs remarquables de la caudale ornée Samoa (named for the elliptical mark in the caudal également d’une grande marque elliptique); et A. fin); A. bellicauda described from one specimen from stenotaeniata , à partir d’un seul spécimen de Nou - New Caledonia (named for the striking colour pattern velle Calédonie (le nom désigne les fines lignes som - of the caudal fin, also featuring a large elliptical mark); bres sur le corps). Avec les descriptions ci-dessus, le and A. stenotaeniata , described from one specimen nombre total d’espèces reconnues d’ Amblyleotris est from New Caledonia (named for the narrow dark bars désormais de 33. on the body). With these descriptions, the total num - ber of recognized species of Amblyeleotris is now 33. Introduction The Indo-Pacific genus Amblyeleotris is one of 13 Zusammenfassung genera of gobiid fishes with species that live symbiot - Fünf neue Meergrundelarten der indo-pazifischen ically with snapping shrimps of the genus Alpheus . Gattung Amblyeleotris werden von den Inseln des The shrimps build and maintain a burrow in sand or zentralen und des südlichen Pazifik beschrieben: A mud, while the gobies serve as sentinels at the bur - katherinae, früher fälschlicherweise A. fasciata zuge - row entrance and take refuge in the burrow as ordnet, auf der Grundlage von 14 Exemplaren von needed. den Society-Inseln (Typenlokalität), Cook-Inseln, The genus Amblyeleotris is characterized by a mod - Marshall-Inseln und Mariana-Inseln; A. biguttata, erately elongate and compressed body; gill opening nach 4 Exemplaren von Neu-Kaledonien (Typen - extending forward to below the preopercle; transverse lokalität) und den Salomon-Inseln (benannt nach zwei cheek pattern of papillae; dorsal rays VI+I,12-19; anal großen schwarzen Flecken am Kinn); A. ellipse, nach rays I,12-19; pectoral rays 16-21; caudal fin rounded neun Exemplaren vom amerikanischen Samoa to pointed; scales in longitudinal series 58-110, (benannt nach dem elliptischen Fleck auf der ctenoid posteriorly, becoming cycloid and smaller Schwanzflosse); A. bellicauda, nach einem Exemplar anteriorly. Unique to the genus is a single pair of sen - aus Neukaledonien (benannt nach der auffälligen Far - sory papillae in a depression just behind the upper lip, bzeichnung auf der Schwanzflosse, auch hier findet instead of a pair of longitudinal rows (Winterbottom, sich ein großer Fleck); sowie A. stenotaeniata, nach 2003). The species are among the most beautifully einem Exemplar aus Neukaledonien (benannt nach coloured of fishes, typically with prominent maroon to schmalen dunklen Streifen auf dem Körper). Nach red bars and variously marked with small spots of yel - diesen Beschreibungen erhöht sich die Gesamtzahl low, blue, red, or black. der anerkannten Arten von Ambleleotris auf 33. The first species of this genus was described as

61 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Gobius fontanesii by Bleeker (1852). The following eter, and interorbital width the least bony width; upper year he named two more, Gobius gymnocephalus jaw length is taken from the front of the upper lip to the and Eleotris periophthalmus. In 1874 he created the posterior end of the maxilla; caudal peduncle depth is new genus Amblyeleotris , designating Eleotris perio - the least depth, and caudal peduncle length the hori - phthalmus as the type species. zontal distance between verticals at the rear base of It was not until 1910 that another species was the anal fin and the caudal fin base; lengths of fin described. Seale named Biat luzonica as a new spines and rays of the fins are measured to their genus and species; however, it proved to be a syn - extreme bases; caudal fin length is the horizontal dis - onym of Amblyeleotris fontanesii. tance from the base of the fin to a vertical at the tip of Fowler (1938) described the fourth valid species of the longest ray; pectoral fin length is the length of the Amblyeleotris as Pteroculiops guttatus from the longest ray; pelvic fin length is measured from the Philippines. base of the pelvic spine to the tip of the longest soft Herre (1939) named the fifth as Amblyeleotris ray. Morphometric data presented in the tables are (Andameleotris ) raoi from the Andaman Islands, and given as percentages of the standard length. Propor - in 1953 the sixth , Zebreleotris fasciata , from Bikini tional measurements in the text are rounded to the Atoll, Marshall Islands. nearest 0.05. Beginning with the description of Amblyeleotris The longitudinal scale series is the number of japonicus by Takagi (1957) from Kagoshima Prefec - oblique rows of scales between the upper end of the ture, Japan, 22 more valid species have been gill opening and the caudal fin base. The transverse described in the genus (see Eschmeyer, 1998 and scale count is made from the origin of the anal fin update of March 13, 2003 at www.calacademy.org/ obliquely upward to the base of the first dorsal fin. research/ichthyology/catalog/fishcatsearch.html). Only the lower limb gill rakers are counted because Polunin and Lubbock (1977, 1979) described seven of the smaller upper limb rakers are indistinguishable these species, but one that they named Cryptocentrus from nearby papillae. Meristic and morphometric data wheeleri is a junior synonym of Amblyeleotris fasciata given in parentheses refer to paratypes. (Herre). Wass (1984), Randall and Randall (1987), and Myers (1999) have misidentified an undescribed species from Pacific islands as A. fasciata (Herre), fol - Amblyeleotris katherine n. sp. lowing the identification of Bishop Museum speci - (Figs. 1-2; Table I) mens by D. F. Hoese. This species is described in the present paper, along with four others from islands of Amblyeleotris fasciata (non Herre) Wass, 1984: 27 Oceania to make the names available for the author’s (American Samoa); Randall and Randall, 1987: 308 book on the reef and shore fishes of the South Pacific, (Marshall Islands); Myers, 1999: 238, pl. 150 A (Mar - now in production. iana and Marshall Islands).

Materials and Methods Holotype: BPBM 14990, female, 49.7 mm, Society Type specimens of three of the new species have Islands, Tetiaroa Atoll (17.05°S, 149.30°W), outside been variously deposited at the following institutions: reef off Tiaraua Islet, 38 m, sand patch surrounded by Australian Museum, Sydney (AMS); Institute of Zool - coral and rock, spear, J. E. Randall, 23 April 1973. ogy, Academia Sinica, Taipei (ASIZP); Bernice P. Paratypes: BPBM 34757, 50.2 mm, Marshall Islands, Bishop Museum, Honolulu (BPBM); California Acad - Enewetak Atoll, Medren Island, lagoon, off power sta - emy of Sciences, San Francisco (CAS); Muséum tion, sand, 3 m, 29 July 1955 (from reference collec - National d’Histoire Naturelle, Paris (MNHN); National tion of the Mid-Pacific Research Laboratory); UGM Science Museum, Tokyo (NSMT); Royal Ontario 5622, 62.2 mm, Mariana Islands, Maug, R.S. Jones et Museum, Toronto (ROM); University of Guam, Mangi - al ., 10 February 1971 (no further collecting data); lao (UGM), and the U.S. National Museum of Natural BPBM 13916, 54.0 mm, Cook Islands, Rarotonga, off History, Washington, D.C. (USNM). The two other oil tanker buoy near harbour entrance, small pocket of species are represented only by a Bishop Museum coarse sand, 15 m, rotenone, J. E. Randall and D. B. holotype. Cannoy, 10 March 1971; BPBM 19966, 49.6 mm, Lengths of specimens are given as standard length Marshall Islands, Kwajalein Atoll, south end of lagoon, (SL), measured from the anterior end of the upper lip top of patch reef, coral rubble and coarse sand, 5 m, to the base of the caudal fin (posterior end of the spear, J. E. Randall, 8 April 1976; BPBM 39282, 2: hypural plate); head length (HL) is measured from the 44.0-48.7 mm; AMS I.42860-001, 50.2 mm; ASIZP same anterior point to the posterior end of the oper - 62617, 44.8 mm; CAS 218422, 46.4 mm; MNHN cular membrane; body depth is the greatest depth 2003-1811, 52.0 mm; NSMT–P 67416, 50.2 mm; (adjusting for any malformation from preservation); ROM 73732, 55.5 mm; USNM 374475, 51.3 mm, all body width is the maximum width just posterior to the from Mariana Islands, Guam, T. Allen, 9 August 2000 gill opening; orbit diameter is the greatest fleshy diam - (no further collecting data). aqua vol. 8 no. 2 - 2004 62 John E. Randall

Diagnosis branched lower ray of largest paratype); pelvic rays Dorsal rays VI+I,12; anal rays I,12; pectoral rays 18- I,5, the fifth soft ray branched once, the branches not 19 (usually 18); pelvic fins joined by a very short separated (ray appears simple); branched caudal membrane at base, with no frenum; fifth pelvic ray rays 14 (one paratype with 13); upper procurrent cau - longest, branched once, the branches closely parallel; dal rays 9 (8-10), the posterior two segmented; lower longitudinal scale series 67-73; predorsal scales procurrent caudal rays 9 (8-9), the most posterior seg - embedded, extending to above upper end of preoper - mented; longitudinal scale series 69 (67-73); trans - cular margin; gill opening reaching forward to below verse scale series 25 (24-27); predorsal scales middle of preopercle; body depth 4.65-5.35 in SL; embedded and not in even rows, extending to above head length 3.3-3.5 in SL; caudal fin rounded and middle of preopercle; circumpeduncular scales at moderately long, 2.7-3.2 in SL; white with numerous midlength of caudal peduncle 20; lower limb gill rak - small yellow spots on head, body, and dorsal fins; five ers 9; vertebrae 25. narrow, slightly oblique red bars, one from nape Body depth 4.65 (4.6-5.35) in SL; body width 1.7 across opercle, one under first dorsal fin, two under (1.45-1.7) in body depth; HL 3.3 (3.3-3.5) in SL; snout second dorsal fin, and the last on caudal peduncle; a short, 4.9 (4.4-5.1) in HL; profile of snout steep, form - red spot above posterior end of maxilla; a blue-edged ing an angle of about 55° to horizontal axis of body; a red arc on base of caudal fin sharp deflection in profile above eye, the angle of postorbital head and nape about 5° to horizontal axis Description of body; eye moderately large, the orbit diameter 3.8 Dorsal rays VI+I,12; anal rays I,12; all dorsal and (3.4-4.35) in HL; interorbital space very narrow, the anal soft rays branched, the last to base; pectoral rays least bony width 19 (16-20) in HL; caudal peduncle 18 (18-19; three of 13 paratypes with 19), the upper - depth 2.6 (2.3-2.5) in HL; caudal peduncle length 1.65 most and lower one or two unbranched (except for (1.55-1.7) in HL.

Fig. 1. Holotype of Amblyeleotris katherinae, BPBM 14990, 49.7 mm, Tetiaroa, Society Islands. Photo by J. E. Randall.

Fig. 2. Underwater photograph of Amblyeleotris katherinae with the snapping shrimp Alpheus ochrostriatus, Guam, Mariana Islands. Photo by J. E. Randall.

63 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Table I. Proportional measurements of type specimens of Amblyeleotris katherinae expressed as percentages of the standard length.

Holotype Paratypes BPBM BPBM CAS BPBM BPBM ROM UGM 14990 39283 218422 39282 34757 73732 5611 Sex female male female female male male male Standard length (mm) 49.7 44.0 46.4 48.7 50.2 55.5 62.2 Body depth 21.5 18.8 20.2 20.6 18.6 21.9 19.4 Body width 12.5 12.4 11.7 12.5 12.9 12.8 12.1 Head length 30.4 30.1 29.3 29.5 28.6 29.4 28.4 Snout length 6.2 6.6 5.9 5.8 6.5 6.6 6.4 Orbit diameter 8.0 8.8 8.1 8.2 7.9 7.5 6.5 Interorbital width 1.6 1.5 1.7 1.7 1.8 1.7 1.6 Upper jaw length 13.2 13.4 13.0 12.9 13.0 12.8 12.6 Caudal peduncle depth 11.8 12.0 11.7 12.1 11.8 11.8 11.4 Caudal peduncle length 18.6 18.6 18.2 17.1 18.4 18.1 18.5 Predorsal length 35.3 34.7 34.5 34.8 34.0 34.5 34.2 Preanal length 56.7 54.7 56.6 57.5 57.1 57.5 54.8 Prepelvic length 29.9 29.8 29.7 30.4 31.4 30.5 29.2 Base of dorsal fins 50.2 50.3 49.7 49.9 49.9 49.6 51.0 First dorsal spine 17.8 16.2 17.5 13.6 17.0 18.3 17.0 Longest dorsal spine 22.6 18.6 20.1 16.2 22.0 23.2 20.9 Spine of second dorsal fin 15.8 14.1 14.6 12.1 14.2 14.4 12.7 Longest dorsal soft ray 24.1 19.3 21.6 21.2 24.0 22.8 24.2 Anal fin base 26.2 26.6 26.4 27.0 26.8 26.3 27.5 Anal spine 10.0 9.8 9.7 8.3 10.3 10.6 8.4 Longest anal ray 22.5 18.1 19.8 20.5 22.8 20.1 21.5 Caudal fin length 37.3 31.8 33.0 31.0 32.9 33.4 32.5 Pectoral fin length 29.0 24.9 24.6 26.4 26.0 25.1 24.9 Pelvic spine length 8.1 9.1 8.8 8.2 8.1 9.3 8.0 Pelvic fin length 27.3 27.2 27.8 25.0 28.4 27.1 28.3

Mouth large, the maxilla reaching slightly posterior to anterior nasal opening a short membranous tube less a vertical at rear edge of orbit, the upper jaw length 2.3 than a nasal opening diameter ventroanterior to pos - (2.2-2.3) in HL; mouth oblique, forming an angle of terior nasal opening, the nasal sensory pore directly about 35° to horizontal axis of body; lower jaw slightly above the anterior. Pores of cephalic sensory system projecting; upper jaw with a band of inward-curving and sensory papillae on cheek and opercle as shown villiform teeth, those at front of jaw in about four rows, in the illustration of Amblyeleotris japonica by the teeth of inner and outer rows largest; two large well in Masuda et al . (1984: 255, fig. 95). spaced recurved canine teeth on each side of upper Scales on body progressively smaller anteriorly; jaw, the medial gap about equal to pupil diameter; side scales ctenoid posterior to below rear fourth of pec - of upper jaw with an outer row of slender incurved teeth toral fin on midside of body, farther back dorsally and (15 on holotype) about half length of anterior canines; ventrally, scales cycloid anteriorly and progressively lower jaw with a band of villiform teeth anteriorly, the more embedded; predorsal scales extending forward slender outer teeth largest and recurved (eight at front to above upper end of preopercular margin; small of jaw of holotype); typically three large, close-set, embedded scales on chest and prepectoral region; no strongly recurved, canine teeth nearly halfway back in scales on head, and none on fins except basal fourth jaw medial to a row of small teeth; side of lower jaw with to fifth of caudal fin. a single row of small slender incurved teeth; no teeth Origin of first dorsal fin over rear base of pelvic fins, on vomer or palatines. Margin of lips with a row of papil - the predorsal length 2.85 (2.9-2.95) in SL; first dorsal lae except posteriorly. Tongue rounded. spine 1.7 (1.6-1.85, except one paratype with 2.2) in Gill opening extending forward to below middle of HL; second or third dorsal spines longest (second preopercle (slightly less than orbit diameter posterior spine slightly prolonged on some specimens, including to eye). Gill rakers short, the longest on lower limb the holotype), 1.35 (1.25-1.6, except one paratype with about one-half length of longest gill filaments. 1.8) in HL; spine of second dorsal fin 1.9 (2.0-2.45) in Posterior nasal opening small, with a low anterior rim HL; penultimate dorsal soft ray longest, 1.25 (1.15- just before edge of orbit at level of lower third of eye; 1.55) in HL; origin of anal fin below or slightly posterior aqua vol. 8 no. 2 - 2004 64 John E. Randall to base of first dorsal soft ray, the preanal length 1.75 edges; fins translucent whitish, the only marking a faint (1.75-1.8) in SL; anal spine 3.05 (2.8-3.3) in HL; penul - dusky double line in lower basal part of caudal fin. timate anal soft ray longest, 1.35 (1.2-1.45) in HL; cau - dal fin rounded, 2.7 (3.0-3.2) in SL; pectoral fins Etymology pointed, the tenth or eleventh ray longest, 3.55 (3.8- This species of Amblyeleotris is named katherine at 4.05) in SL; pelvic fins broadly separated, joined only the request of Tim Allen, the collector of the Guam at extreme base by membrane, no frenum present; specimens, to honour his mother. To be treated as a fifth pelvic ray longest, reaching to or slightly posterior noun in apposition. to origin of anal fin, 3.65 (3.5-4.0) in SL. Colour of holotype when fresh: head and body Remarks white with numerous small yellow spots, faintly dark- As mentioned in the introduction, this species has edged, those on body tending to be vertically elon - long been misidentified as Amblyeleotris fasciata gate, some arranged in vertical rows; five slightly (Herre), now known to be the senior synonym of A. oblique bright red bars about half width of orbit, the wheeleri (Polunin and Lubbock). first from nape across opercle and branchiostegal Nine of the 13 paratypes were collected in Guam by membranes, edged in small blue spots on its lower the late Tim Allen in August 2000. Unfortunately, no part, the second below fifth and sixth dorsal spines, other collection data are available for his specimens. the third below third and fourth dorsal soft rays, the Amblyeleotris katherine is presently known only from fourth below tenth dorsal soft ray, and the fifth, more islands of the Pacific Plate, specifically the Mariana orange than red, on caudal peduncle; an obliquely Islands, Marshall Islands, Niue (pers. comm. Malcom P. elongate red spot above posterior end of maxilla; dor - Francis), American Samoa, Rarotonga, and Society sal fins whitish with many small dark-edged yellow Islands. The Society Islands record is based on the holo - spots; anal fin dusky yellow, the basal half with dark- type from the atoll of Tetiaroa, but the species is known edged yellow spots and blue spots and irregular blue also from Tahiti based on an underwater photograph markings, some encircling yellow spots; outer third of sent by Philppe Bacchet. The listing of this species (as fin with irregular oblique dark-edged blue streaks; A. fasciata ) from the Great Barrier Reef and Christmas caudal fin of similar colour to body basally, the mem - Island, Indian Ocean by Myers (1999: 238: pl. 150 A) branes of rest of fin pale blue with a yellow line run - appears to be an error. The species illustrated from ning parallel to rays; a boomerang-shaped, blue- Christmas Island by Allen and Steene (1988: 140, fig. edged, red mark on basal half of caudal fin containing 420) is correctly identified as the true A. fasciata . some yellow spots in its anterior part; pectoral fins Amblyeleotris katherine has been collected from translucent with whitish rays; pelvic fins white. depths of 3 to 38 m on sand, where it shares a burrow Largest specimen examined 62.2 mm SL. with the snapping shrimp Alpheus ochrostriatus . Colour of holotype in alcohol : pale tan, with five This species appears to be most similar to faint dark bars on body, visible mainly from their dark Amblyeleotris guttata (Fowler) (see Fig. 3 herein). At

Fig. 3. Underwater photograph of Amblyeleotris guttata with Alpheus ochrostriatus, Enewetak, Marshall Islands. Photo by J. E. Randall.

65 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania first glance, this would not seem to be true, because pelvic frenum present; fourth pelvic ray longest; fifth the colour pattern is so different; guttatus has faint pelvic ray branching twice; longitudinal scale series blackish bars, the first and second expanding and 91-103; no median predorsal scales; scales on side of becoming black ventrally, and its pelvic fins are nape extending to above middle of opercle; gill open - largely black. However, guttata has essentially the ing reaching forward to a vertical about two-thirds same general morphology, the same number of dor - orbit diameter behind eye; body depth 5.8-6.1 in SL; sal and anal rays, 19 pectoral rays (18-19, usually 18 head length 3.6-3.8 in SL; caudal fin pointed and long, for katherine ), 75-78 scales in longitudinal series (67- 2.2-2.6 in SL; pale brown dorsally, white ventrally, 73 for katherine ), predorsal scales extending to above with three indistinct brown bars on body, and a slightly the middle of the preopercle, and the same pelvic fin oblique brown bar from nape across opercle; pale structure. It has many bright yellow spots (though interspaces with small brown blotches, faintly inter - larger than those on katherine ) on the head, body, connected; a pair of prominent black spots on chin; and dorsal fins; the five dark bars are narrow, slightly dorsal fins yellowish grey, the first dorsal with small oblique, and in the same place as the red ones of irregular dark-edged pale blue spots and orange- katherine , and the first bar extending down from the tipped spines, the second dorsal with a broad yellow nape has blue spots in its lower part. The most salient border containing small dark-edged pale blue spots, differences of guttata are its longer, pointed caudal fin the rays tipped with orange; caudal fin with a vertically and the prolonged second to fourth dorsal spines in elongate diffuse brown spot on base. adults. Description Dorsal rays VI+I,13; anal rays I,13; all dorsal and Amblyeleotris biguttata n. sp. anal soft rays branched, the last to base; pectoral rays (Fig. 4, Table II) 19, the upper two and lowermost unbranched; pelvic rays I,5, the fifth soft ray branched twice, the branches Holotype: BPBM 38362, male, 60.7 mm, Solomon diverging; branched caudal rays 14; upper procurrent Islands, Guadalcanal, 7 miles west of Honiara, adja - caudal rays 10 (9-10), the posterior two segmented; cent to the wreck of a Japanese transport ship, silty lower procurrent caudal rays 9 (9-10), the most pos - sand at a depth of 35 m, collected with spear, J. E. terior segmented; longitudinal scale series 93 (91- Randall, 5 July 1973. 103); transverse scale series 30 (29-32); no median Paratypes: USNM 374773, 71.8 mm, same locality predorsal scales; circumpeduncular scales at as holotype, but at a depth of 50 m, collected with midlength of caudal peduncle 20; lower-limb gill rak - spear, J. E. Randall, 11 July 1973; BPBM 22549, 78.0 ers 9; vertebrae 25. mm, New Caledonia, Canala Bay, east side, point at Body depth 5.8 (5.8-6.1) in SL; body width 1.7 (1.6- south end of d’Amata Bay, muddy sand and rock with 1.7) in body depth; head length 3.6 (3.6-3.8) in SL; little live coral, at a depth of 10-12 m, collected with snout short, 4.9 (4.7-5.05) in HL; profile of snout rotenone, J. E. Randall, P. Laboute, and J.-L. Menou, steep, forming an angle of about 53° to horizontal axis 7 January 1979. MNHN 2003-2026, 63.6 mm, same of body; a sharp deflection in profile above eye, the data as for preceding specimen. postorbital head and nape nearly horizontal; eye mod - erately large, the orbit diameter 3.9 (3.9-4.0) in HL; Diagnosis interorbital space very narrow, the least bony width 20 Dorsal rays VI+I,13; anal rays I,13; pectoral rays 19; (18-20) in HL; caudal peduncle depth 2.75 (2.7-2.8) in pelvic fins joined by membrane to tips of fifth rays; HL; caudal peduncle length 1.85 (1.7-1.85) in HL.

Fig. 4. Holotype of Amblyeleotris biguttata, BPBM 38362, 60.7 mm, Guadalcanal, Solomon Islands. Photo by J. E. Randall. aqua vol. 8 no. 2 - 2004 66 John E. Randall

Table II. Proportional measurements of type specimens just in front of edge of orbit at level of lower third of of Amblyeleotris biguttata expressed as percentages of eye; anterior nasal opening a short membranous tube the standard length. about a nasal opening diameter ventroanterior to pos - terior nasal opening, the nasal sensory pore directly Holotype Paratypes above anterior nasal opening. Pores of cephalic sen - BPBM MNHN USNM BPBM sory system and sensory papillae on cheek and oper - 38362 03-2062 374773 22549 cle as shown in the illustration of Amblyeleotris japon - Sex male male male male ica by Akihito in Masuda et al . (1984: 255, fig. 95). Standard length (mm) 60.7 63.6 71.8 78.0 Scales on body progressively smaller anteriorly; Body depth 17.2 16.4 16.7 17.1 scales ctenoid posterior to tip of pectoral fin on mid - Body width 10.2 9.8 10.3 10.0 side of body, farther back dorsally and ventrally, Head length 27.9 27.2 27.3 26.7 scales cycloid anteriorly; no median predorsal scales; Snout length 5.7 5.5 5.8 5.3 scales on side of nape extending forward to above Orbit diameter 7.2 7.0 6.9 6.7 Interorbital width 1.4 1.5 1.4 1.3 middle of opercle; small scales on chest; no scales on Upper-jaw length 11.5 10.9 11.1 10.5 prepectoral region, head, or fins except basal seventh Caudal peduncle depth 10.2 9.9 9.8 10.0 of caudal fin. Caudal peduncle length 15.2 14.9 14.8 15.6 Origin of first dorsal fin slightly anterior to origin of Predorsal length 30.8 29.3 30.9 28.8 pelvic fins, the predorsal length 3.25 (3.25-3.5) in SL; Preanal length 55.5 55.0 55.1 54.5 first dorsal spine 2.1 (1.85-2.1) in HL; fifth dorsal spine Prepelvic length 30.5 29.9 31.0 28.6 longest, 1.7 (1.6-1.8) in HL; spine of second dorsal fin Base of dorsal fins 55.8 56.1 56.4 56.6 2.2 (2.05-2.2) in HL; penultimate dorsal soft ray First dorsal spine 13.3 14.9 13.0 13.6 longest, 1.5 (1.25-1.45) in HL; origin of anal fin below Longest dorsal spine 16.5 15.7 15.2 16.7 base of second dorsal soft ray, the preanal length 1.8 Spine of second dorsal fin 12.7 12.9 12.3 13.0 (1.8-1.85) in SL; anal spine 3.75 (3.3-3.75) in HL; Longest dorsal soft ray 18.3 19.6 18.9 21.2 Anal fin base 28.1 29.1 28.0 29.5 penultimate anal soft ray longest, 1.55 (1.3-1.55) in Anal spine 7.4 8.2 7.3 7.8 HL; caudal fin pointed, 2.6 (2.2-2.65) in SL; pectoral Longest anal ray 17.7 19.2 17.8 20.5 fins pointed, the eleventh ray longest, 4.65 (4.5-4.75) Caudal fin length 38.1 42.6 37.7 45.6 in SL; pelvic fins joined by membrane to tips of fifth Pectoral fin length 21.5 22.2 21.0 21.8 rays; pelvic frenum moderately developed; fourth Pelvic spine length 9.2 9.4 8.5 9.9 pelvic ray longest, reaching to or slightly posterior to Pelvic fin length 28.0 29.3 27.8 29.5 origin of anal fin, 3.6 (3.4-3.6) in SL. Largest speci - men examined, 78.0 mm SL Mouth moderately large, the maxilla reaching to or Colour of holotype when fresh: pale brown dor - beyond a vertical at rear edge of pupil (but not beyond sally, the scale edges clearly darker than centres, a vertical at rear edge of orbit), the upper-jaw length white to pale grey ventrally, with bars as described 2.4 (2.45-2.55) in HL; mouth oblique, forming an above; pale interspaces with indistinct small brown angle of about 30° to horizontal axis of body; lower blotches that are faintly interconnected; a pair of large jaw slightly projecting; upper jaw with a band of small black spots on chin; suborbital region pale blue, the inwardly curving villiform teeth, those at front of jaw in sensory papillae visible as dark lines across this zone; about four rows, narrowing to one to two rows poste - a small yellow-edged blue spot halfway between orbit riorly; two large broadly-spaced recurved canine teeth and corner of mouth; a row of small dark-edged yel - on each side at front of upper jaw, the medial gap of low spots along posterior edge and dorsally on pre - inner pair about two-thirds pupil diameter; side of opercle, preceded by a vertical row of slightly larger upper jaw with an outer row of slender inwardly curv - blue spots; dark-edged pale blue lines following bran - ing teeth (12 in holotype), averaging about half length chiostegal rays; dorsal fins yellowish grey, the first of anterior canines; front of lower jaw with an outer dorsal with small irregular dark-edged pale blue spots row of eight (four pairs) of slender recurved teeth, the and orange-tipped spines, the second dorsal with a median four about half length of upper canines, with a broad yellow border containing small dark-edged pale band of villiform teeth behind; a large recurved canine blue spots, the rays tipped with orange; caudal fin with tooth about one-third way back in lower jaw, followed pale yellowish membranes, blue rays, and orangish by a single row of small slender inwardly curving brown streaks in outer part of fin (fin damaged so teeth; no teeth on vomer or palatines. Margin of lips position of streaks is not well-defined); anal fin dusky with a row of prominent papillae except posteriorly. with a narrow white band at base, edged distally with Tongue rounded. Gill opening extending forward to a a narrow greyish blue band and an orange line; pec - vertical about two-thirds orbit diameter posterior to toral fins pale yellowish grey; pelvic fins light grey with eye. Gill rakers short, the longest at angle about one- a dusky orange line on each ray. half length of longest gill filaments. Colour of holotype in alcohol: light brown with Posterior nasal opening small, with a low anterior rim three diffuse dark brown bars on body, one centred

67 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania below fifth dorsal spine, one below fourth and fifth dor - (orange for biguttata ), and the faint brown blotches in sal soft rays, and the third below posterior part of sec - the pale intermediate zones of the body arranged in ond dorsal fin; a slightly oblique dark brown bar from vertical rather than randomly. A decision on the taxo - nape across opercle; two very dark brown spots on nomic status of the Papua New Guinea fish should chin about half orbit diameter in size, partly hidden in await the collection of additional material, especially in groove medial to lower jaw; a large, diffuse, vertically the area between Madang and Guadalcanal. elongate brown spot on caudal fin base; fins light Amblyeleotris biguttata is most similar in colour to A. brown, the first dark bar of body extending into base fontanesii (Bleeker), which also has only three brown of first dorsal fin, and the second and third bars to a bars on the body, one across the nape and opercle, a lesser extent into second dorsal fin. vertically elongate brown spot on the caudal fin base, and small yellow dots on the head. In addition, Etymology fontanesii has the pelvic fins fully joined, a pelvic Named biguttata from Latin meaning two spots, in ref - frenum, the first dorsal fin of near uniform height, and erence to the pair of prominent back spots on the chin. a pointed caudal fin. On presently available data it can be separated from A. biguttata as it has 15 dorsal soft Remarks rays, 16 or 17 anal soft rays, and a greater maximum The description of this species is based on two lots size (A. fontanesii exceeds 200 mm total length). of specimens from Guadalcanal, Solomon Islands and one from New Caledonia. The depth range at which the fish were collected was 11 to 50 m, and the Amblyeleotris ellipse n. sp. bottom was silty sand. The fish were associated with (Fig. 6; Table III) an unidentified alpheid shrimp and took refuge in the shrimp’s burrow. Amblyeleotris sp. 20 Wass, 1984: 27 (Tutuila, Ameri - The author collected six specimens of what is either can Samoa). a very closely related species or a geographical vari - ant of Amblyeleotris biguttata in the lagoon of the Holotype: BPBM 11332, female, 37.6 mm, American Madang area of Papua New Guinea in November, Samoa, Tutuila, Fagasa Bay, south-west side, sand 1984. The bottom was silty sand, and the depth range near edge of reef, at depth of 23 m, collected with was 26-30 m. The specimens are deposited at the spear, J. E. Randall, 5 September 1971. Bishop Museum under numbers 32554, 32579, Paratypes: AMS I.21994-001, 2: 43.9-44.0 mm, 32604, and 32606; they vary from 37-76 mm SL. American Samoa, Tutuila, Taema Bank, 32 m, Their meristic data are the same except that one rotenone, R. C. Wass and I. Swan, 11 June 1976; specimen has 20 pectoral rays on one side, and the ASIZP 62618, 32.4 mm; CAS 218423, 32.4 mm; longitudinal scale count varies from 89-96 (compared MNHN 2003-1812, 39.5 mm; NSMT-P 67417, 36.3 to 91-103 for biguttata). The only clear difference in mm; ROM 73733, 43.4 mm; USNM 374476, 42.9 mm, proportional measurements is the length of the pelvic all with same data as AMS I.21994-001. fins of comparable size to biguttata , 30.0-33.2% SL (compared to 28.0-29.5 in biguttata). The colour is Diagnosis very similar (Fig. 5), including the two black chin Dorsal rays VI-I,13; anal rays I,13; pectoral rays 19- spots. The only obvious colour differences are the 20 (usually 19); pelvic fins joined by a short mem - black-tipped spines and rays of the dorsal fins brane at base, with no frenum; fifth pelvic ray longest,

Fig. 5. Amblyeleotris cf biguttata, BPBM 32554, 73.0 mm, Madang Province, Papua New Guinea. Photo by J. E. Randall. aqua vol. 8 no. 2 - 2004 68 John E. Randall branched once, the branches not separated; longitu - orbit diameter 3.6 (3.4-4.15) in HL; interorbital space dinal scale series 80-89; no median predorsal or very narrow, the least bony width 19.5 (18.5-20) in prepectoral scales; scales on side of nape extending HL; caudal peduncle depth 2.8 (2.85-2.9 ) in HL; cau - to above middle of opercle; body depth 5.55-6.15 in dal peduncle length 1.75 (1.85-1.95) in HL. SL; head length 3.25-3.45 in SL; gill opening extend - Mouth large, the maxilla reaching slightly posterior to ing forward to below middle of preopercle; caudal fin a vertical at rear edge of orbit, the upper jaw length rounded and moderately long, 2.8-3.1 in SL; pale yel - 2.7 (2.6-2.9) in HL; mouth oblique, forming an angle of low dorsally, white ventrally, densely spotted with about 35° to horizontal axis of body; upper jaw with a small pale blue spots; three broad dusky red bars on band of villiform teeth, with two large, well-spaced, body, and a narrower darker one from nape across recurved canine teeth on each side, the medial gap opercle and throat, broadly bordered by yellow con - about equal to pupil diameter; side of upper jaw with taining small blue spots; a faint brown bar across pos - an outer row of slender inwardly-curving teeth (16 in terior caudal peduncle, merging with brownish red of holotype) about half length of anterior canines; lower caudal fin base; second dorsal and anal fins with a jaw with a broader band of villiform teeth anteriorly, broad longitudinal dusky band, the second dorsal yel - the slender outer row of about 10 teeth notably largest low at base with dark-edged blue markings, the anal and recurved; typically three (varying from one to white at base with a broad yellow margin containing four), large, close-set, strongly-recurved, canine teeth dark-edged blue markings; caudal fin with a large nearly halfway back in jaw medial to a row of small blue-edged elliptical mark that varies in zones from teeth; side of lower jaw with a row of small slender brownish orange to orange red, yellow, and black inwardly-curving teeth; no teeth on vomer or palatines. Edge of lips with a row of papillae except Description posteriorly. Tongue rounded. Dorsal rays VI-I,13; anal rays I,13; all dorsal and anal Gill opening extending forward to below middle of soft rays branched, the last to base; pectoral rays 19 preopercle (slightly less than orbit diameter posterior (19-20; two of 10 paratypes with 20), the uppermost to eye). Gill rakers short, the longest on lower limb and lowermost unbranched; pelvic rays I,5, the fifth soft about one-half length of longest gill filaments. ray branched once, the branches contiguous; branched Posterior nasal opening small, with a low anterior rim caudal rays 14; upper procurrent caudal rays 10 (9-10), just in front of edge of orbit at level of lower third of the posterior two segmented; lower procurrent caudal eye; anterior nasal opening a short membranous tube rays 9 (8-9), the most posterior segmented; longitudinal less than a nasal opening ventroanterior to posterior scale series 86 (80-89); transverse scale series 28 (28- nasal opening, the nasal sensory pore directly above 30); no median predorsal scales; circumpeduncular the anterior nasal opening. Pores of cephalic sensory scales in midlength of peduncle 20; lower-limb gill rak - system and sensory papillae on cheek and opercle as ers 9 (9-11); vertebrae 25. shown in the illustration of Amblyeleotris japonica by Body depth 5.75 (5.55-6.15) in SL; body width 1.7 Akihito in Masuda et al . (1984: 255, fig. 95). (1.5-1.8) in body depth; head length 3.45 (3.25-3.45) Scales on body progressively smaller anteriorly; in SL; snout short, 4.5 (4.45-4.95) in HL; profile of scales ctenoid posterior to rear fourth of pectoral fin snout steep, forming an angle of about 55° to hori - on midside of body, farther back dorsally and ven - zontal axis of body; a sharp deflection in profile above trally; scales cycloid anteriorly; no median predorsal eye, the angle of postorbital head and nape about 5° scales; small scales extending forward on side of to horizontal axis of body; eye moderately large, the nape to above middle of opercle; small scales on

Fig. 6. Holotype of Amblyeleotris ellipse, BPBM 11332, 37.6 mm, Tutuila, American Samoa. Photo by J. E. Randall.

69 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Table III. Proportional measurements of type specimens of Amblyeleotris ellipse expressed as percentages of the stan - dard length.

Holotype Paratypes BPBM CAS NSMT ROM AMS AMS 11332 218423 67417 73733 I.21994 I.21994 Sex female male male male female male Standard length (mm) 37.6 32.4 36.3 43.4 43.9 44.0 Body depth 17.4 18.0 17.9 17.7 16.5 16.3 Body width 10.1 10.8 10.0 11.2 10.7 10.9 Head length 29.1 30.7 30.5 29.8 30.0 29.2 Snout length 6.5 6.2 6.6 6.3 6.7 6.6 Orbit diameter 8.0 9.1 8.2 8.1 8.0 7.0 Interorbital width 1.5 1.5 1.6 1.6 1.6 1.6 Upper aw length 10.9 11.9 11.1 11.3 11.4 10.9 Caudal peduncle depth 10.3 10.5 10.7 10.4 10.3 10.2 Caudal peduncle length 16.6 15.8 16.5 16.0 15.9 15.8 Predorsal length 34.4 33.7 33.5 33.8 33.0 34.5 Preanal length 55.5 55.7 55.1 54.5 55.0 54.3 Prepelvic length 31.1 32.0 30.5 30.4 31.1 31.6 Base of dorsal fins 52.3 52.4 52.5 54.2 52.3 53.2 First dorsal spine 15.2 15.4 15.7 15.0 16.1 15.9 Longest dorsal spine 19.2 18.4 20.1 19.6 18.7 20.2 Spine of second dorsal fin broken 15.5 18.3 16.3 15.9 14.6 Longest dorsal soft ray 19.4 20.4 20.8 22.9 22.8 22.0 Anal fin base 28.3 29.4 28.5 29.4 29.7 29.7 Anal spine 9.1 10.2 10.4 9.4 10.3 9.4 Longest anal ray 18.8 19.2 19.3 20.4 20.1 19.5 Caudal fin length 32.5 33.8 32.5 35.8 34.3 33.9 Pectoral fin length 24.7 25.4 26.5 26.9 25.9 25.5 Pelvic spine length 10.3 9.5 9.8 9.2 9.1 9.3 Pelvic fin length 25.8 27.9 29.4 29.3 27.3 25.6 chest; no scales on head, and none on fins except with dark bluish brown spots, whitish on snout and basal fourth to fifth of caudal fin. cheek with small dark-edged bluish spots; an oblique Origin of first dorsal fin over rear base of pelvic fins, dark orangish brown bar from nape across posterior the predorsal length 2.9 (2.9-3.0) in SL; first dorsal opercle and throat, broadly edged in yellow containing spine 1.9 (1.85-2.0) in HL; third or fourth dorsal spines blue spots; posterior edge of branchiostegal mem - longest (none prolonged), 1.5 (1.45-1.65) in HL; spine branes blue, preceded by a bright orange line; a hemi - of second dorsal fin broken in holotype, 1.65-2.0 in HL spherical brown spot above posterior end of maxilla; of paratypes; penultimate dorsal soft ray longest, 1.5 edge of maxilla and mandible yellow posteriorly; ven - (1.3-1.5) in HL; origin of anal fin below base of first troposterior edge of orbit yellowish with an oblique dorsal soft ray, the preanal length 1.8 (1.8-1.85) in SL; dark-edged blue line; first dorsal fin dusky anteriorly, anal spine 3.2 (2.9-3.2) in HL; penultimate anal soft yellow posteriorly with dark-edged blue spots that are ray longest, 1.55 (1.45-1.6) in HL; caudal fin rounded, horizontally elongate basally in fin, and form a broad 3.1 (2.85-3.2) in SL; pectoral fins pointed, the ninth to band in each membrane in outer part; basal third to eleventh rays longest, 4.05 (3.7-3.95) in SL; pelvic fourth of second dorsal fin yellow with horizontally- fins broadly separated, joined only at extreme base by elongate dark-edged blue spots, some crossing two to membrane, no frenum present; fifth pelvic ray longest, three membranes; outer part of fin dusky, shading to reaching to or slightly posterior to origin of anal fin, 3.9 grey distally, with a dark-edged yellow line on each (3.4-3.9) in SL. A small species, the largest specimen membrane parallel to rays; basal third of anal fin measured 44 mm SL. white, grading outwardly to grey; rest of fin dusky with Colour of holotype when fresh : body pale yellow a broad outer dusky yellow margin containing irregu - dorsally, shading to white ventrally, densely marked lar dark-edged blue spots; caudal fin with blue rays with small irregular pale blue spots; body with three and a large elliptical mark, much of which is edged in dusky red bars a little broader than pale interspaces, blue, that joins the brownish red basal part of fin and the first beneath middle of first dorsal fin, the next two extends to margin of fin posteriorly; basal part of mark below second dorsal fin; a faint fourth brown bar pos - purplish, followed by zones (in clockwise direction) of teriorly on caudal peduncle, merging with brownish yellow, black, yellow, white, yellow and bright orange; red base of caudal fin; head orangish brown dorsally centre of fin enclosed by elliptical mark white with an aqua vol. 8 no. 2 - 2004 70 John E. Randall indistinct streak of grey in each membrane; upper and cauda from New Caledonia, described below. In fact, lower margins of caudal fin broadly yellowish, the bellicauda was first regarded as a colour variant of upper margin with a few blue spots basally, narrowing ellipse . The two have the same fin ray and scale posteriorly and ending at tip of sixth branched ray; counts, both lack median predorsal and prepectoral lower margin ending at tip of eleventh branched ray; scales, and there is much similarity in colour pattern. pectoral fins with transparent membranes and faint It was determined as a distinct species when it was pink rays; pelvic fins grey with a white leading edge. noticed that the gill opening did not extend as far for - Colour of holotype in alcohol : head and body ward as that of ellipse , the small scales on the nape pale tan; opercular membrane and posterior edge of reach to above the middle of the opercle, and the branchiostegal membranes reddish brown; fins with ctenoid scales are more posterior. For more compar - translucent membranes and whitish rays; caudal fin isons, see Remarks for bellicauda . with a submarginal reddish brown streak from the third to sixth branched rays. Some paratypes with these markings more pigmented; in addition, the Amblyeleotris bellicauda n. sp. lower part of the life colour elliptical mark in the cau - (Figs. 7, 8; Table IV) dal fin is visible from dark edges, the base of the dor - sal fin is dusky, and the outer part of the anal fin has Holotype: BPBM 34318, 49.7 mm, New Caledonia, wavy dark bands. outside barrier reef off Tenia Island (22°1’1”S, 165°55’5”E), at depth of 3 m over coarse sand with Etymology large ripple marks, collected with spear, J. E. Ran - This species is named ellipse from the English and dall, 25 March 1990. French for a geometric figure, in reference to the large elliptical mark in the caudal fin. The word is derived Diagnosis from the Greek elleipsis meaning to fall short (i.e. to fall Dorsal rays VI-I,13; anal rays I,13; pectoral rays 19; short of a circle). It is treated as a noun in apposition. pelvic fins joined by a short membrane at base, with no frenum; fifth pelvic ray branched three times; lon - Remarks gitudinal scale series 89; no median predorsal scales Amblyeleotris ellipse is presently known only from and no prepectoral scales; body depth 5.35 in SL; Tutuila, American Samoa. The first specimen was col - head length 3.55 in SL; gill opening extending forward lected by the author in 1971 in Fagasa Bay at a depth to below posterior margin of preopercle; mouth not of 23 m on sand next to a reef. The goby was living very large, the maxilla reaching to below centre of symbiotically in a burrow with an alpheid shrimp. R.C. eye, the upper-jaw length 2.65 in HL; caudal fin Wass and I. Swan collected eight specimens with slightly pointed and moderately long, 2.9 in SL; fourth rotenone in 1976 at Taema Bank in 32 m. Wass pelvic ray longest; pale yellow dorsally, shading to (1984: 27) reported the species in his annotated white ventrally, densely marked with small pale blue checklist of the fishes of Samoa as Amblyeleotris sp. spots; four red bars on body, the first slightly oblique 20 and provided a colour description. He deposited beneath first dorsal fin, the next two broader ventrally, the specimens in the Australian Museum, Sydney. Six below second dorsal fin, and the fourth posteriorly on of these have been transferred as paratypes to other caudal peduncle and on caudal fin base; head brown - museums. ish yellow dorsally, white ventrally, with numerous This species is very similar to Amblyeleotris belli - small pale blue spots and a red bar from nape across

Fig. 7. Holotype of Amblyeleotris bellicauda, BPBM 34318, 49.7 mm, New Caledonia. Photo by J. E. Randall.

71 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Fig. 8. Underwater photograph of Amblyeleotris bellicauda, Papua New Guinea. Photo by P. L. Colin. posterior opercle; an irregular yellow-edged blue line with a medial gap of about one-fourth orbit diameter; extending ventrally from opercular membrane to bran - side of upper jaw with an outer row of 19 slender chiostegal membranes; first dorsal fin grey, shading to inwardly curving teeth about half length of anterior greyish blue posteriorly, with dark-edged yellow mark - canines; lower jaw with three large, close-set, ings; second dorsal fin grey with a dark-edged yellow strongly-recurved, canine teeth nearly halfway back in line parallel to rays on each membrane, the base yel - jaw medial to a row of small teeth; side of lower jaw low with dark-edged blue markings; anal fin yellow with a row of small slender incurved teeth; no teeth on with irregular dark-edged blue markings and a broad vomer or palatines. Edge of lips with a row of papillae median blackish red band; caudal fin with a large, except posteriorly. Tongue rounded. irregular, elliptical, blue-edged mark enclosing a Gill opening extending forward to below posterior dusky orange area; upper and lower edges of fin pale margin of preopercle. Gill rakers short, the longest on yellow with blue spots. lower limb about one-half length of longest gill fila - ments. Description Posterior nasal opening small, just anterior to edge Dorsal rays VI-I,13; anal rays I,13; all dorsal and of orbit near level of middle of eye, with a low rim on anal soft rays branched, the last to base; pectoral rays front edge; anterior nasal opening larger, a short 19, the uppermost and lower two rays unbranched; membranous tube immediately ventroanterior to pos - pelvic rays I,5, the fifth ray branched three times, the terior nasal opening; nasal sensory pore dorsoante - branches diverging; branched caudal rays 14; upper rior to posterior nasal opening by a distance equal to procurrent caudal rays 10, the posterior two seg - nasal opening. Pores of cephalic sensory system and mented; lower procurrent caudal rays 9, the most pos - sensory papillae on cheek and opercle as shown in terior segmented; longitudinal scale series 89; trans - the illustration of Amblyeleotris japonica by Akihito in verse scale series 29; no median predorsal scales; Masuda et al . (1984: 255, fig. 95). circumpeduncular scales in midlength of caudal Scales on body progressively smaller anteriorly; peduncle 20; lower limb gill rakers 9; vertebrae 25. ctenoid posteriorly and cycloid anteriorly, the most ante - Body depth 5.35 in SL; body width 1.85 in body rior ctenoid scales on midside of body below the third depth; head length 3.55 in SL; snout short, 4.55 in HL; and fourth dorsal spines; no median predorsal scales; profile of snout steep, forming an angle of about 50° small scales on side of nape extending only to above to horizontal axis of body; a sharp deflection in profile upper end of gill opening; no scales on head, prepec - above eye, the remaining profile of head and nape toral region or fins except basal fifth of caudal fin. horizontal; eye moderately large, the orbit diameter Origin of first dorsal fin over rear base of pelvic fins, 3.8 in head length; interorbital space very narrow, the the predorsal length 3.0 in SL; first dorsal spine 2.0 in least bony width 16.5 in HL; caudal peduncle depth HL; fourth dorsal spine longest (but not prolonged), 2.7 in HL; caudal-peduncle length 1.95 in HL. 1.65 in HL; spine of second dorsal fin 1.65 in HL; Mouth not very large, the maxilla reaching to below eleventh and twelfth dorsal soft rays longest, 1.2 in centre of eye, the upper jaw length 2.65 in HL; mouth HL; origin of anal fin below base of second dorsal soft oblique, forming an angle of about 35° to horizontal ray, the preanal length 1.75 in SL; anal spine 2.55 in axis of body; lower jaw slightly projecting; upper jaw HL; penultimate anal soft ray longest, 1.5 in HL; cau - with a band of villiform teeth, the front of jaw with two dal fin slightly pointed, 2.9 in SL; pectoral fins pointed, large well-spaced recurved canine teeth on each side, the ninth and tenth rays longest, 4.1 in SL; pelvic fins aqua vol. 8 no. 2 - 2004 72 John E. Randall

Table IV. Proportional measurements of the holotype of irregular black-edged yellow markings, except basally Amblyeleotris bellicauda expressed as percentages of where pale blue with larger yellow markings, this pat - the standard length. tern continuing onto rest of fin; basal fifth of second dorsal fin yellow with dark-edged pale blue spots, Holotype including a blue arc at upper end of third and fourth BPBM red bars that barely extend into fin; rest of fin pale 34316 grey with a long yellow line parallel to rays in each Sex female membrane; anal fin yellow with irregular dark-edged Standard length (mm) 49.7 blue markings and a broad middle blackish red band; Body depth 18.7 anal fin with a broad middle dark brownish red band, Body width 10.1 narrowly edged above and below by orange, with a Head length 28.2 dorsal extension to base of third red bar on body and Snout length 6.2 a lesser one to middle of second red bar; basal part of Orbit diameter 7.4 Interorbital width 1.7 fin yellow with dark, edged blue spots and lines; outer Upper jaw length 8.4 fifth of fin dusky yellow with dark-edged blue spots Caudal peduncle depth 10.4 and lines; caudal fin with a large, irregular, elliptical, Caudal peduncle length 14.5 brownish red mark, edged in bright blue except ante - Predorsal length 33.1 riorly where it merges with red base of fin; central part Preanal length 56.5 of fin enclosed by brownish orange elliptical mark with Prepelvic length 31.4 a row of small blue spots on membrane between Base of dorsal fins 52.2 eighth and ninth branched rays; broad upper margin First dorsal spine 14.1 of fin yellow with dark, edged blue spots, ending at Longest dorsal spine 17.3 fifth branched ray (where elliptical mark becomes Spine of second dorsal fin 17.2 marginal); lower margin of fin to tip of tenth branched Longest dorsal soft ray 23.9 Anal fin base 28.0 ray dusky orange-yellow with small, elongate, dark- Anal spine 11.0 edged blue spots; pectoral fins with translucent mem - Longest anal ray 18.5 branes and pale salmon pink rays; pelvic fins with Caudal fin length 34.2 pale purplish grey membranes and white rays. Pectoral fin length 24.5 Colour of holotype in alcohol : pale yellowish with Pelvic spine length 11.1 traces of four dark bars on body; a light brown bar on Pelvic fin length 25.7 nape extending down over posterior half of opercle to throat; a submarginal pale-edged dark brown line on broadly separated, joined only at extreme base by opercular membrane extending to posterior edge of membrane, no frenum present; fourth pelvic ray branchiostegal membranes; branchiostegal mem - longest, nearly reaching origin of anal fin, 3.9 in SL; branes brown with a curved pale, edged dark brown fifth pectoral ray branched, each branch divided, and line across middle part; dorsal fins pale with a brown each of these branches divided. line forming an arc in basal part of second dorsal fin Colour of holotype when fresh : pale yellow dor - from tenth to last ray; anal fin brown dusky with irreg - sally, shading to white ventrally, densely marked with ular, longitudinal, pale, edged dark brown lines small pale blue spots; four red bars on body, the first basally and at margin; caudal fin brown except for a slightly oblique beneath first dorsal fin, the next two broad pale upper margin from base to tip of fourth below second dorsal fin, becoming broader ventrally, branched ray and lower margin to tip of fourteenth and the fourth posteriorly on caudal peduncle and on branched ray; irregular lengthwise dark brown lines caudal fin base; head brownish yellow dorsally, white on lower third of ventral brown part of fin; pectoral fins ventrally, with numerous small dark-edged pale blue pale; pelvic fins with membranes and fifth soft ray spots and a brownish red bar from nape across pos - dusky. terior opercle and adjacent body to throat, and extending forward medially on branchiostegal mem - Etymology branes; an irregular yellow-edged blue line submar - Named bellicauda , from the Latin bellus meaning ginal on opercular membrane continuing ventrally on beautiful and cauda meaning tail, in reference to the branchiostegal membranes; a vertical row of small strikingly colourful caudal fin. Treated as a noun in blue spots at anterior edge of brownish red bar on apposition. opercle, continuing as a yellow-edged blue line in an arc on branchiostegal membranes to below end of Remarks jaws; area of branchiostegal membranes enclosed by Amblyeleotris bellicauda is described from a single pale yellow arc with blue spots; a faint narrow dusky specimen collected by the author in 1990 outside the bar from eye to posterior end of maxilla; first four barrier reef of Tenia Island, New Caledonia at a depth membranes of first dorsal fin purplish grey with small of 3 m where the coarse sand showed large ripple

73 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Fig. 9. Amblyeleotris ogasawarensis, BPBM 35098, 51.0 mm, Chichi-jima, Ogasawara Islands. marks. The fish was at the entrance to a burrow hence coexisting at the last locality with A. bellicauda . shared with an alpheid shrimp. The species also Amblyeleotris ogasawarensis shares with ellipse and occurs at Papua New Guinea, as shown by the under - bellicauda the same fin ray counts, a similar count of water photograph of Fig. 8 taken by Patrick L. Colin; scales in longitudinal series (Yanagisawa, 1978, gave therefore it should also be expected in the Solomon the range as 80-94), no median predorsal or prepec - Islands and Vanuatu. toral scales, and some features of colour. The head As mentioned in the account of A. ellipse , the holo - colour pattern is almost the same (note especially the type of A. bellicauda was first believed to be a geo - details of the opercle and branchiostegal membrane graphic colour variant of A. ellipse because there is so coloration by comparing Figs. 6 and 7 with Fig. 9 in much similarity in the life colour to that of ellipse , the this paper), the dorsal fin coloration is also very simi - fin ray and scale counts are the same, and there are lar (the black areas in the fins of figures 4 and 5 are no median predorsal or prepectoral scales. A more from the black background colour passing through the detailed comparison revealed many differences: the transparent fins). The caudal fin of ogasawarensis gill opening does not extend as far forward on belli - lacks the enclosed ellipse, but if one were to connect cauda (to below edge of preopercle, compared to the blue-edged red streaks, one per membrane, in the below middle of preopercle in ellipse ); the small outer part of the fin, there would be an ellipse. The scales on the side of the nape extend only to the most obvious differences in colour of ogasawaresis upper end of the gill opening (to above the middle of from ellipse and bellicauda are the narrower brownish the opercle on ellipse ); the ctenoid scales begin more red bars on the body, each pale interspace bisected posteriorly in bellicauda ; the caudal fin is more by a brown streak dorsally, the fifth bar entirely on the pointed; the fourth pelvic ray is longest in bellicauda caudal peduncle instead of posteriorly on the pedun - (the fifth longest in ellipse ); and the fifth pelvic ray cle and on the caudal fin base, and the extension of branches three times in bellicauda , but only once in the first bar on the body into the base of the first dor - ellipse where the branches are closely adjacent (so sal fin. The most significant morphological difference the ray appears to be unbranched). A comparison of is the complete joining of the pelvic fins by membrane Tables 3 and 4 reveals that bellicauda has a deeper to the tips of the fifth rays in ogasawarensis . body, shorter head; shorter caudal peduncle, much shorter upper jaw, shorter spines of the first dorsal fin, and longer anal and pelvic spines. Amblyeleotris stenotaeniata n. sp. These two species are most similar to Amblyeleotris (Figs. 10, 11; Table V) ogasawarensis Yanagisawa (see Fig. 9 herein) described from the Ogasawara Islands and Ryukyu Amblyeleotris sp. 4, Laboute and Grandperrin, 2000: Islands. Randall et al . (1997: 525, lower fig.) recorded 406, upper left fig. (New Caledonia). it from the Capricorn Group of the southern Great Bar - rier Reef. The Bishop Museum has specimens from Holotype: BPBM 27088, male, 66.0 mm SL, New the Ogasawara Islands; Ryukyu Islands; Negros, Caledonia, north-east coast, Canala Bay, east side, Philippines; and Lizard Island, northern Great Barrier point at south end of d’Amata Bay, , collected by J.E. Reef. In addition, the author has photographed what Randall, P. Laboute, and J.-L. Menou, 7 January appears to be this species at Kiritimati Atoll (Christ - 1979, with rotenone at depth of 10-12 m over muddy mas Island), Line Islands and at New Caledonia, sand and rock with little live coral. aqua vol. 8 no. 2 - 2004 74 John E. Randall

brown spots, an oblique dark brown line behind upper Diagnosis half of eye; a vertically elongate dark brown spot Dorsal rays VI+I,13; anal rays I,14; pectoral rays 19; extending upward from posterior end of maxilla; dor - pelvic fins joined at base by a very short membrane, sal fins dusky yellow with irregular dark-edged blue without a frenum; fifth pelvic ray longest, branched spots; a bright yellow spot distally on each soft dorsal once, the branches not separated; longitudinal scale membrane, rimmed below by a black-edged blue arc; series 81; median predorsal scales extending to anal fin with a white band at base; caudal fin with blue above middle of preopercle; body depth about 5.5 in rays, yellow membranes with brownish red lines, and SL; head length 3.75 in standard length; gill opening a brownish red arc at base. extending forward to a vertical about one-half orbit diameter behind eye; mouth moderately large, the Description maxilla reaching to between verticals at posterior Dorsal rays VI-I,13; anal rays I,14; all dorsal and anal edge of pupil and rear edge of orbit, the upper jaw soft rays branched to base; pectoral rays 19, the upper - length 2.5 in HL; third dorsal spine longest, 2.25 in HL; most and lowermost unbranched; pelvic rays I,5, the caudal fin pointed and long, 2.4 in SL; body pale tan fifth soft ray branched once, the branches contiguous; (under magnification yellow with small pale blue branched caudal rays 14; upper procurrent caudal rays spots), shading to white ventrally, with a narrow 9, the most posterior segmented; lower procurrent cau - brownish red bar from nape onto opercle and four dal rays 9, the posterior two segmented; longitudinal more on body, each on body in centre of a diffuse light scale series 81; transverse scale series 30; median brown bar the edges of which are darker yellowish predorsal scales 21, extending to above middle of pre - brown on about ventral half of each bar; upper half of opercle; circumpeduncular scales in midlength of each pale interspace with four or five small dark caudal peduncle 22; gill rakers 9; vertebrae 25.

Fig. 10. Holotype of Amblyeleotris stenotaeniata, BPBM 27088, 66.0 mm, New Caledonia. Photo by J. E. Randall.

Fig. 11. Underwater photograph of Amblyeleotris stenotaeniata, New Caledonia. Photo by T. Allen.

75 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Table V . Proportional measurements of the holotype of der papillae except posteriorly. Tongue rounded. Amblyeleotris stenotaeniata expressed as percentages Gill opening extending forward to below preopercle of the standard length. on a vertical about one-half orbit diameter posterior to eye. Gill rakers short, the longest on lower limb about Holotype one-half length of longest gill filaments. BPBM Posterior nasal opening large, with a low dorsal rim 27088 just anterior to edge of orbit at level of lower part of Sex male pupil; anterior nasal opening larger, a short membra - Standard length (mm) 66.0 nous tube immediately ventroanterior to posterior Body depth 18.1 nasal opening; nasal sensory pore dorsoanterior to Body width 10.2 posterior nasal opening by a distance equal to nasal Head length 26.7 Snout length 5.9 opening. Pores of cephalic sensory system and sen - Orbit diameter 6.5 sory papillae on cheek and opercle as shown in the Interorbital width 1.5 illustration of Amblyeleotris japonica by Akihito in Upper jaw length 10.6 Masuda et al. (1984: 255, fig. 95). Caudal peduncle depth 10.6 Scales on body progressively smaller anteriorly; Caudal peduncle length 14.8 scales ctenoid posterior to rear half of pectoral fin on Predorsal length 31.7 midside of body, farther back dorsally and ventrally; Preanal length 52.7 Prepelvic length 29.7 scales cycloid anteriorly; scales on nape somewhat Base of dorsal fins 55.3 embedded, the median predorsal scales extending First dorsal spine 20.6 forward to a point three-fourths orbit diameter poste - Longest dorsal spine 21.4 rior to a vertical at rear edge of orbit; no scales on Spine of second dorsal fin 11.8 head, and none on fins except basal fifth of caudal fin. Longest dorsal soft ray 22.8 Origin of first dorsal fin over rear base of pelvic fins, Anal fin base 31.8 the predorsal length 3.15 in SL; first dorsal spine 1.3 Anal spine 8.3 Longest anal ray 21.2 in HL; third dorsal spine longest, 1.25 in HL; no dorsal Caudal fin length 41.2 spines prolonged as filaments; spine of second dorsal Pectoral fin length 23.5 fin 2.25 in HL; twelfth dorsal soft ray longest, only Pelvic spine length 10.5 slightly longer than penultimate ray, 1.15 in HL; origin Pelvic fin length 27.6 of anal fin below base of first dorsal soft ray, the pre - anal length 1.9 in SL; anal spine 3.2 in HL; twelfth anal Body depth 5.5 in SL; body width 1.75 in body depth; soft ray longest, only slightly longer than penultimate head length 3.75 in SL; snout short, 4.55 in HL; profile ray, 1.25 in HL; caudal fin pointed, 2.4 in SL; pectoral of snout steep, forming an angle of about 70° to hori - fins pointed, the tenth ray longest, 4.25 in SL; pelvic zontal axis of body; a sharp deflection in profile above fins broadly separated, joined only at extreme base by eye, the angle of postorbital head and nape about 5° membrane, no frenum present; fifth pelvic ray longest, to horizontal axis; eye moderately large, the orbit reaching to base of second anal soft ray, 3.65 in SL. diameter 4.1 in HL; interorbital space very narrow, the Colour of holotype when fresh: body tan dorsally least bony width about 18 in HL; caudal peduncle (on magnification yellow with small pale blue spots), depth 2.5 in HL caudal peduncle length 1.8 in HL. shading to white ventrally, with four diffuse oblique Mouth moderately large, the maxilla nearly reaching brown bars, the centre of each a narrow brownish red a vertical at rear edge of orbit on left side, and just to bar, and the edges on lower part darker yellowish below rear edge of pupil on right side, the upper jaw brown; first bar beneath first dorsal fin, the third and length 2.5 in HL; mouth oblique, forming an angle of fourth below second dorsal fin, and the last on caudal about 25° to horizontal axis of body; lower jaw peduncle; upper half of each pale interspace with four strongly projecting; front of upper jaw with a band of or five small dark brown spots; a narrow dark brown - villiform teeth, two large, well-spaced, recurved ish red bar on nape curving downward across oper - canine teeth on one side and three on the other, the cle; head light brown, pale bluish adjacent and ven - medial gap nearly equal to pupil diameter; side of troposterior to eye, with an oblique dark brown line upper jaw with an outer row of 14 slender incurved faintly edged in yellow and blue behind upper part of teeth about half length of anterior canines; lower jaw eye; a small irregular dark brown spot adjacent to with a broader band of villiform teeth anteriorly, the upper part of orbit; a vertically-elongate dark brown outer row of about eight slender teeth notably largest spot, partly edged anteriorly with a yellow line, and recurved; three large, close-set, strongly- extending upward from posterior end of maxilla; eight recurved, canine teeth nearly halfway back in jaw faint small pale blue spots on cheek and opercle; medial to a row of small teeth; side of lower jaw with opercular membrane yellow with a blue submarginal a row of small slender incurved teeth; no teeth on line that continues onto branchiostegal membranes, vomer or palatines. Margin of lips with a row of slen - where edged posteriorly in a bright red and dark aqua vol. 8 no. 2 - 2004 76 John E. Randall brown line; first dorsal fin dusky yellow with irregular fourth dorsal soft rays, the third below base of tenth dark-edged blue spots; second dorsal fin similar, but and eleventh rays, and the fourth on caudal peduncle; a broad outer zone of blackish pigment obscures the spaces between bars with four or five small brown yellow and blue spotted pattern; a bright yellow spot spots; a narrow dark brown bar on nape curving distally on each soft dorsal membrane, rimmed below downward across opercle; two small middorsal dark by a black-edged blue arc; about basal fifth of anal fin brown spots on nape anteriorly in zone of predorsal white with a blue line separating it from outer dusky scales; an oblique dark brown line on nape behind part of fin; caudal fin with blue rays, yellow mem - upper part of eye; a small irregular dark brown spot branes with brownish red longitudinal lines, a brown - adjacent to upper part of orbit; a vertically-elongate ish red arc at base, and a submarginal row of small dark brown spot above posterior end of maxilla; a blue spots along outer upper edge of fin; pectoral fins dark brown streak on outer branchiostegal mem - with dusky rays and transparent membranes; pelvic branes; dorsal fins faintly dotted with small brown fins dusky yellow with blue lines, the fourth ray and specks, the last two rays brown; anal fin more densely adjacent membrane brownish red. flecked with brown except basally; caudal fin with a An underwater photograph of this species is shown vertically-elongate dark brown spot in outer scaled in the book on New Caledonia fishes by Laboute and basal portion of fin, with traces of dark pigment curv - Grandperrin (2000: 406, upper left fig.) as ing posteriorly from upper and lower ends; pectoral Amblyeleotris sp. 4. It differs in colour from the holo - fins pale; pelvic fins finely flecked with brown, darkest type in having some of the small dark brown spots in over long medial part of fin. the pale interspaces on the body connected by brown lines. Also it has an oblique blue-edged yellow line Etymology just below the ventroposterior edge of the orbit. This Named stenotaeniata from the Greek stenos for nar - area is now pale bluish on the holotype, but it may row and taeniatus meaning banded, in reference to have had a yellow line in life. the narrow oblique dark bars on the body. Colour of holotype in alcohol: pale yellowish brown with a series of 11 small dark brown spots Remarks along back, four of which have a narrow dark brown This species is presently known only from a single bar extending obliquely downward and forward onto specimen from Canala Bay on the north-east coast of upper half of body, the first just behind base of fifth New Caledonia, the underwater photograph men - dorsal spine, the second below base of third and tioned above, and an underwater photograph by Tim

Fig. 12. Underwater photograph of with Alpheus bellulus, Redang, Malaysia. Photo by J. E. Randall.

77 aqua vol. 8 no. 2 - 2004 Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania

Allen reproduced here as Fig. 11. National Museum, 85 : 31-135. The holotype was collected in the same rotenone Herre, A. W. C. T. 1939. On a collection of littoral station as two of the paratypes of Amblyeleotris bigut - and freshwater fishes from the Andaman Islands. tata . Laboute and Grandperrin described the habitat Records of the Indian Museum, 41 (4): 327-372. as detrital sand in the lagoon in a zone of current at Herre, A. W. C. T. 1953. The tropical Pacific Eleotridae 10-20 m. Robert F. Myers (pers. comm.) pho - with vomerine teeth with descriptions of two new gen - tographed the species underwater in an area of silty era and two new species from the Marshall Islands . sand in 3 m in New Caledonia. The most similar The Philippine Journal of Science, 82 (2): 189-192. species is Amblyeleotris diagonalis Polunin and Lub - Laboute, P. & R. Grandperrin. 2000. Poissons de bock (see Fig. 12 herein), which has the same four Nouvelle-Calédonie . 520 pp. Editions Catherine narrow oblique bars on the body and one from the Ledru, Nouméa. nape across the opercle, with small dark spots in the Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & T. upper part of the pale interspaces. It differs in colour Yoshino (Eds.). 1984. The Fishes of the Japanese in having an oblique yellow-edged dark brown bar Archipelago. Vol. 1 (text: xxii + 437 pp.) and vol. 2 from behind the lower part of the eye to beneath the (plates). Tokai University Press, Tokyo. posterior end of the maxilla and an oblique dark Myers, R. F. 1999. Micronesian Reef Fishes , ed. 3. vi brown line from the eye to the upper lip. There are two + 330 pp. Coral Graphics, Guam. meristic differences; diagonalis has 13 anal soft rays Polunin, N. V. C. & R. Lubbock. 1977. Prawn-asso - (14 in stenotaeniata ) and 67-75 scales in longitudinal ciated gobies (Teleostei: Gobiidae) from the Sey - series (81 in stenotaeniata ). chelles, Western Indian Ocean: systematics and ecology. Journal of Zoology, London, 183 : 63-101. Acknowledgements Polunin, N. V. C. & R. Lubbock. 1979. Five new I am grateful to Mark McGrouther of the Australian prawn-associated gobies (Teleostei: Gobiidae) of Museum for authorizing the transfer of six specimens the genus Amblyeleotris . Bulletin of the British of Amblyeleotris ellipse as paratypes to other muse - Museum of Natural History , 36 (4): 239-249. ums. Thanks are also due Malcolm P. Francis for his Randall, J. E., Allen, G. R. & R. C. Steene. 1997. record of A. katherinae from Niue, Philippe Bacchet Fishes of the Great Barrier Reef and Coral Sea . for his underwater photograph of the species from Second edition, xx + 557 pp. University of Hawaii Tahiti, Patrick L. Colin for his underwater photograph Press, Honolulu. of A. bellicauda , Robert F. Myers for sending his and Randall, J. E. & H. A. Randall. 1987. Annotated Tim Allen’s underwater photographs of A. stenotaeni - checklist of the fishes of Enewetak Atoll and other ata , and Loreen R. O’Hara for X-rays. The manu - Marshall Islands. Chapter 28, pp. 289-324, 3 pls. in script was commented on by Gerald R. Allen, William D.M. Devaney, E.S. Reese, B.L. Burch, and P. Hel - N. Eschmeyer, and Richard Winterbottom and frich (eds.), The Natural History of Enewetak Atoll, reviewed by two anonymous referees. Vol. II, Biogeography and Systematics . DOE/EV/ 00703. T1-Vol. 2 (DEB7006111). U.S. Dept. of References Energy, Office of Scientific and Technical Informa - Allen, G. R. & R. C. Steene. 1988. Fishes of Christ - tion, Oak Ridge, Tennessee. mas Island Indian Ocean . 197 pp. Christmas Island Seale, A. 1910. New species of Philippine fishes. Natural History Association, Australia. The Philippine Journal of Science , 4 (6): 481-543. Bleeker, P. 1852. Derde bijdrage tot de kennis der Takagi, K. 1957. Descriptions of some new gobioid ichthyologische fauna van Celebes. Nagtuurkundig fishes of Japan, with a proposition on the sensory Tijdschrift voor Nederlandsch Indië , 3: 739-782. line system as a taxonomic appliance. Journal of Bleeker, P. 1853. Diagnostische beschrijvingen van the Tokyo University of Fisheries , 43 (1): 97-126. nieuwe of weinig bekende vischsoorten van Batavia. Wass, R. C. 1984. An annotated checklist of the fishes Tiental I-VI. Natuurkundig Tijdschrift voor Neder - of Samoa. NOAA Tech. Rep. NMFS SSRF , 781 : 1-43. landsch Indië , 4: 451-516 Winterbottom, R. 2003. A redescription of Crypto - Bleeker, P. 1874. Notice sur les genre Amblyeleotris, centrus crocetus Wongratana, a redefinition of Valenciennesia et Brachyeleotris. Verslagen en Myersina Herre (Acanthopterygii; Gobiidae), a key to Mededeelingen der Koninklijke Akademie van the species, and comments on relationships. Ichthy - Wetenschappen , 8: 372-376. ological Research , 49 : 69-75. Eschmeyer, W. N. 1998. , 3 vol - Yanagisawa, Y. 1978. Studies on the interspecific umes: 1-2905, California Academy of Sciences, San relationship between gobiid fish and snapping Francisco. shrimp. I. Gobiid fishes associated with snapping Fowler, H. W. 1938. Descriptions of new fishes shrimps in Japan. Publications of the Seto Marine obtained by the United States Bureau of Fisheries Biological Laboratory , 24 (4/6): 269-325. steamer “Albatross”, chiefly in Philippine Seas and adjacent waters. Proceedings of the United States aqua vol. 8 no. 2 - 2004 78 aqua, Journal of Ichthyology and Aquatic Biology

Kiunga bleheri , a new Blue-Eye (Pisces: Pseudomugilidae) from fresh waters of Papua New Guinea

Gerald R. Allen

Department of Aquatic Zoology, Western Australian Museum, Francis Street, Perth, WA 6000, Australia and Conservation International, 1999 M Street, Suite 600, Washington, D.C. 20036, USA

Accepted: 26.02.2004

Keywords Sommario Taxonomy, freshwater fishes, Kiunga , new species, Kiunga bleheri, nuova specie, è descritta sulla base Pseudomugilidae, Papua New Guinea di 9 esemplari, di 16.7-28.4 mm SL, raccolti da H. Ble - her nel 1994 e nel 2003 lungo il Fly River nelle vici - Abstract nanze di Kiunga, Papua Nuova Guinea. Si distingue Kiunga bleheri , new species, is described from 9 dall’unica altra specie dello stesso genere, K. ballochi, specimens, 16.7-28.4 mm SL, collected by H. Bleher in anch’essa nota dalla stessa località, per la seconda 1994 and 2003 from the Fly River system in the vicinity dorsale e l’anale molto più brevi, per una differenza of Kiunga, Papua New Guinea. It differs from its only modale significativa del numero dei raggi della sec - known congener, K. ballochi, also from the Kiunga onda pinna dorsale, 6 invece di 5 file di scaglie area, on the basis of its much shorter second dorsal trasversali sul corpo e per avere di solito la maggior and anal fins, significant modal difference in the num - parte dei raggi della seconda dorsale e dell’anale non ber of second dorsal fin rays, 6 versus 5 transverse ramificati. scale rows on the body, and in usually having most of the second dorsal and anal fin rays unbranched. Introduction Blue-eyes of the family Pseudomugilidae are tiny, Zusammenfassung delicate fishes that inhabit fresh, brackish, and marine Die neue Art Kiunga bleheri wird auf der Grundlage waters of New Guinea and Australia. Their common von neun Exemplaren mit einer Länge von 16,7 bis name is derived from the neon blue colour of the iris 28,4 mm SL beschrieben, die von H. Bleher in den that is typical of most species. They were formerly Jahren 1994 und 2003 im Fly-River-System in der included in the family Melanotaeniidae, Nähe von Kiunga in Papua-Neuguinea gesammelt but a detailed study by Saeed et al. (1989) provided wurden. Die Vertreter dieser neuen Art unterscheiden convincing evidence for separate family recognition. sich von denen der nahe verwandten Art K. ballochi, The family was most recently reviewed by Allen die ebenfalls im Kiunga-Gebiet vorkommt, durch die (1995), who recognised 15 species in 3 genera. Two deutlich kürzere zweite Rückenflosse und Afterflosse, additional species from New Guinea have been eine signifikant andere Anzahl der Flossenstrahlen in described since that time (Allen et al., 1998; Allen and der zweiten Rückenflosse, eine Schuppenzahl von 6 Renyaan, 1999). statt 5 in der Querreihe, sowie dadurch, dass die meis - The present paper describes a second species in ten Strahlen der zweiten Rückenflosse und der After - the previously monotypic genus Kiunga Allen (1983). flosse ungegliedert sind. The members of this genus are distinguished from other blue-eyes on the basis of their tiny first dorsal Résumé fin, keel-like structure between the pelvic and anal Kiunga bleheri , espèce nouvelle, est décrit d’après 9 fins, and external villiform teeth on the jaws. The spécimens de 16,7-28,4 LS, collectés par H. Bleher genus was placed in a separate subfamily, Kiunginae, en 1994 et 2003 dans le bassin de la rivière Fly, prés on the basis of these characters as well as various de Kiunga, Papouasie-Nouvelle-Guinée. Il se distin- osteological features (Saeed et al., 1989). gue de son seul congénère connu, K. ballochi, égale - Both species of Kiunga were collected near the vil - ment de la région de Kiunga, par une seconde dor - lage of Kiunga in the Fly River system of south-west - sale et une anale nettement plus courtes, par une dif - ern Papua New Guinea. There are no records of the férence spécifique affirmée des rayons de la seconde genus from outside this area. The previously dorsale, par 6 rangées d’écailles transversales au lieu described species, K. ballochi , was collected about 40 de 5 sur le corps et par les rayons de la seconde dor - km north of Kiunga in 1982. The new species, K. ble - sale et de l’anale habituellement non ramifiés. heri , was discovered by H. Bleher in 1991 and two

79 aqua vol. 8 no. 2 - 2004 Kiunga bleheri , a new Blue-Eye (Pisces: Pseudomugilidae) from fresh waters of Papua New Guinea specimens were collected in 1994. However, most of pectoral and pelvic fin lengths – measured from the the specimens that form the basis of the present base to the tip of the longest ray; caudal fin length – description were collected by Bleher in 2003. measured from the base of the fin (hypural junction) to the tip of the longest ray. Materials and Methods Data in parentheses in the descriptions refer to Counts of fin spines are given in Roman numerals range of values for paratypes if differing from the holo - and soft (segmented) rays in Arabic. The last ray of type. Type specimens are deposited at the Australian the second dorsal and anal fins is divided near the Museum, Sydney (AMS) and the Western Australian base and counted as a single element. Gill raker Museum, Perth (WAM). counts refer to the first brachial arch and include 1-2 rudimentary elements on the upper limb. Other counts and measurements were taken as follows: lateral Kiunga bleheri n. sp. scales - number of scales in horizontal row from upper (Figs 1-4; Tables I-II) corner of gill cover to caudal fin base (hypural junc - tion); transverse scales - number of scales in vertical Holotype: WAM 32455-001, male, 28.4 mm SL, Tare row between anal fin origin and base of dorsal fin; Creek, 11.5 km from central market of Kiunga on predorsal scales - number of scales along midline of Konkonda Road (approximately 6°04’S, 141°13’E), nape in front of first dorsal fin; cheek scales - total Western Province, Papua New Guinea, collected by number of scales covering the suborbital and preop - H. Bleher with dipnets, 7 November 2003. erculum ; scales on opercular series – total number of Paratypes: AMS I.42990-001, 2 specimens, 24.1- scales covering the opercle bones; standard length 24.4 mm SL, collected with holotype; WAM 32455- (SL) - measured from the tip of the upper lip to the 002, 4 specimens, 16.7-25.8 mm SL, collected with caudal fin base (hypural junction); head length – mea - holotype; WAM P.31572-001, 2 specimens, 20.0-25.4 sured from the tip of the upper lip to the upper rear mm SL, 26 km north-east of Kiunga (approximately edge of the gill opening; body depth – the greatest 5°58’S, 141°10’E), Western Province, Papua New depth to the base of the dorsal fin; body width – mea - Guinea, collected by H. Bleher with dipnets, 27 Octo - sured just posterior to the opercular flap; snout length ber 1994. – measured between the front of the upper lip and the anterior edge of the bony orbit; eye diameter – hori - Diagnosis zontal measurement across middle of eye; interorbital Dorsal rays II to IV-12 or 13; anal rays I, 15 to 17; width – the least bony width; caudal peduncle depth – soft (segmented) dorsal and anal rays usually is the least depth; caudal peduncle length - measured unbranched, except last 2-3 rays; transverse scales 6; between two vertical lines, one passing through the second dorsal and anal fins much shorter than those base of the last anal ray and the other through the of K. ballochi , 1.4-2.1 and 1.5-2.1 in head lengh caudal fin base; predorsal, preanal, and prepelvic dis - respectively. tances – measured from tip of upper lip to respective origins of first dorsal fin, anal fin, and pelvic fins;

Fig. 1. Holotype of Kiunga bleheri, male, 28.4 mm SL, Kiunga area, Papua New Guinea. Photo by G. R. Allen. aqua vol. 8 no. 2 - 2004 80 Gerald R. Allen

Fig. 2. Aquarium photo of live holotype of Kiunga bleheri. Photo by H. Bleher.

Fig. 3. Aquarium photo of live paratype of Kiunga bleheri, 25.8 mm SL. Photo by H. Bleher. Description ter 2.2 (2.2 to 2.5); interorbital width 2.3 (2.2 to 2.6); Dorsal rays III-12 (II to IV-12 or 13); anal rays I,15 depth of caudal peduncle 2.0 (1.9 to 2.1); length of (I,15 to 17); pectoral rays 11; pelvic rays I,5; branched caudal peduncle 1.1 (1.2 to 1.4), all in head length. caudal rays 10; principal caudal rays 14; accessory First dorsal fin originates about opposite anal fin ori - caudal rays 6 + 6; lateral scales 30 (29 or 30); trans - gin; longest element of first dorsal fin 3.4 (3.9-4.6), of verse scales 6; scales between dorsal fins 3; predor - second dorsal fin 1.4 (1.7-2.1), of anal fin 1.7 (1.5- sal scales 14 (14 or 15); preopercle scales 2 (addi - 2.1), all in head length. Most of soft (segmented) rays tional scale possibly missing on all specimens); of second dorsal fin unbranched except last 1-2 rays scales on opercular series 5 (5 or 6); gill rakers on first frequently branched. Last 8 soft (segmented) rays of arch 2 + 8 = 10 (1-2 + 8 = 10 or 11). anal fin branched in holotype and last 1-2 rays fre - Body depth 3.1 (2.7 to 3.0); head length 4.2 (4.1- quently branched in paratypes. Pelvic fins short, the 4.5), predorsal distance 2.1 (1.9-2.1), preanal dis - tips not reaching origin of anal fin when depressed; tance 1.9 (1.8-1.9), prepelvic distance 2.7 (2.6-2.7), innermost pelvic fin ray connected to abdomen along all in SL. Greatest width of body 2.3 (2.3 to 2.5) in most of length by delicate membrane (usually dam - body depth. Snout length 3.4 (3.4 to 4.1); eye diame - aged during collecting or examination); urogenital

81 aqua vol. 8 no. 2 - 2004 Kiunga bleheri , a new Blue-Eye (Pisces: Pseudomugilidae) from fresh waters of Papua New Guinea

Fig. 4. Aquarium-bred Kiunga bleheri, approximately 20 mm SL. Photo by G. R. Allen.

Fig. 5. Aquarium photo of Kiunga ballochi, approximately 24 mm SL, Kiunga area, Papua New Guinea. Photo by G. R. Allen. openings positioned on midventral line about one- “wing” of premaxilla mainly enlarged, thorn-like, with third to one-half distance between pelvic fin origin and conical shapes; a pair of prominent patches of similar posterior tip of depressed pelvic fins; length of pelvic thorn-like teeth at front of premaxilla on either side of fin 1.7 (1.6 to 2.0) in head length. Pectoral fins median symphysis; innermost teeth of upper jaw pointed, longest rays 1.2 (1.3 to 1.5) in head length. small and villiform. Lower jaw with less conspicuous Caudal fin emarginate, its length 0.9 (1.0 to 1.2). teeth confined mainly to anterior portion, about 15 to Jaws oblique, mouth terminal; maxilla extends pos - 20 on each side, those at front of jaws largest and teriorly to level between anterior edge of eye and arranged in two large patches on each side of median anterior edge of pupil; premaxilla without accessory symphysis. ascending process (see Fig. 3A in Allen, 1983). Upper Scales cycloid, relatively large and elongated jaw (premaxilla) covered with about 46 to 60 teeth anteroventrally, arranged in regular horizontal rows; arranged in 2 to 4 irregular rows on each side that are predorsal scales extending to middle of interorbital, exposed when the mouth is closed, those on lateral including two relatively enlarged scales on frontal- aqua vol. 8 no. 2 - 2004 82 Gerald R. Allen

Table I. Proportional measurements of selected type specimens of Kiunga bleheri expressed as percentages of the stan - dard length.

Holotype Paratype Paratype Paratype Paratype Paratype WAM WAM WAM WAM AMS AMS P.32455 P.32455 P.31572 P.31572 I.42990 I.42990 -001 -002 -001 -001 -001 -001

Standard length (mm) 28.4 25.8 25.4 24.5 24.4 24.1 Body depth 32.7 33.3 36.6 36.3 35.7 36.1 Body width 14.4 13.2 14.6 15.1 15.6 15.8 Head length 23.6 22.1 23.6 24.1 24.6 24.1 Snout length 7.0 6.6 6.7 6.1 6.1 5.8 Eye diameter 10.9 10.1 10.2 9.8 10.7 10.8 Interorbital width 10.2 8.9 10.6 9.4 9.4 9.5 Depth caudal peduncle 11.6 10.5 12.2 11.4 11.9 12.0 Length caudal peduncle 22.2 19.0 16.5 19.2 18.4 18.7 Predorsal distance 48.2 48.4 52.8 50.2 50.4 51.9 Preanal distance 51.8 54.7 53.1 52.2 52.0 53.5 Prepelvic distance 37.0 37.2 38.2 38.0 38.9 38.2 2nd dorsal fin base 22.9 19.0 22.0 20.4 22.1 22.0 Anal fin base 33.5 32.6 34.3 31.8 34.4 34.0 Pectoral fin length 19.4 15.9 17.3 18.0 17.2 16.6 Pelvic fin length 13.7 11.2 14.2 14.7 13.5 13.7 Longest 1st dorsal ray 7.0 5.4 5.1 5.3 12.7 6.2 Longest 2nd dorsal ray 16.9 10.9 13.4 14.3 11.9 11.6 Longest anal ray 13.7 12.0 15.4 13.5 23.4 12.9 Caudal fin length 26.4 20.9 19.7 24.5 23.4 23.7

Table II. Frequency of counts for second dorsal fin rays blackish marginal band on second dorsal fin, relatively of Kiunga. pronounced in holotype, but narrow in 25.8 mm SL paratype; second dorsal fin of holotype with yellow Fin ray count submarginal band, remainder of holotype’s fin and most of fin of 25.8 mm SL paratype translucent; cau - Species 11 12 13 14 15 16 dal fin translucent; anal fin of holotype translucent K. ballochi 14 961 with pale yellowish hue and narrow blackish margin, K. bleheri 252 that of 25.8 mm SL paratype mainly translucent with nearly imperceptible blackish edge; pelvic fins slightly interorbital; 2 scales below eye covering rear portion yellowish to translucent whitish, often with thin black of cheek; pores of cephalic sensory canals relatively anterior margin; pectoral fins translucent. large and conspicuous, about 9-10 in dentary-preop - Colour in alcohol: overall whitish with scale mar - ercle series and similar number in supraorbital series; gins of anterodorsal portion of body thinly outlined anterior nostril opening relatively small with low rim, with black; thin black line along middle of side from placed on side of snout next to premaxillary; posterior pectoral region to base of caudal fin; concentration of nostril opening enlarged, next to upper anterior corner melanophores on middorsal line between origin of first of eye and similar in appearance to other sensory dorsal fin and interorbital; weaker concentration of pores in supraorbital region. dark pigment on ventral mid-line of caudal peduncle, Colour in life: body mainly semitransparent with breast and abdomen; fins mainly translucent except internal swim bladder clearly evident; lining of stom - outer edge of second dorsal fin of holotype black. ach silvery; iris golden to bluish; a prominent dark brown stripe along middle of side and thinner black Remarks strip on mid-dorsal line between origin of first dorsal Kiunga bleheri clearly differs from its only known fin and interorbital; uppermost portion of back and congener, K. ballochi (Fig. 5), on the basis of its much upper edge of caudal peduncle of holotype with shorter second dorsal and anal fins (respective values orange hue; concentrations of pepper-like for K. bleheri of 1.4-2.1 and 1.5-2.1 in SL versus 0.8- melanophores along bases of dorsal and anal fins; 1.3 and 0.9-1.4 for K. ballochi) . There is also a tiny first dorsal fin of holotype (Fig. 2) pale yellowish, significant difference in the modal number of rays in that of 25.8 mm SL paratype (Fig. 3) colourless; the second dorsal fin (Table II), as well as the pattern

83 aqua vol. 8 no. 2 - 2004 Kiunga bleheri , a new Blue-Eye (Pisces: Pseudomugilidae) from fresh waters of Papua New Guinea

6

7

Fig. 6 (upper). Tare Creek, Papua New Guinea, about 1 km above Konkonda road crossing. Kiunga bleheri occurs upstream of this point where the environment remains undisturbed. The habitat contains dense streamside foilage and a true aquatic plant, the reddish, round-leaved Barclaya morleti. Photo by H. Bleher. Fig. 7 (lower). Type locality of Kiunga bleheri. Photo by H. Bleher. aqua vol. 8 no. 2 - 2004 84 Gerald R. Allen of branching. In K. ballochi all soft (segmented) rays Etymology of the second dorsal and anal fins are usually The new species is named after Heiko Bleher of branched, whereas in K. bleheri most of the rays Milan, Italy who collected the type specimens and except the last 2-3 are unbranched. The anal fin of the was first to observe it in the wild. holotype appears to be aberrant in this respect, with the last 8 rays being branched. Finally, there is also a Acknowledgements difference in the number of transverse scale rows on Thanks are due Heiko Bleher for collecting the type the middle of the side: 6 rows in K. bleheri compared specimens and dispatching them to the Western Aus - with 5 rows in K. ballochi. tralian Museum. Mr. Bleher also provided pho - Although the type specimens were not well fixed and tographs of live specimens shortly after capture. I am sex determination is therefore difficult, the more also indebted to Charles Nishihira of Honolulu, colourful holotype appears to be a male. Hawaii, for allowing me to photograph live aquarium The genus Kiunga is presently known only from a specimens that he bred in captivity. small area near Kiunga, Papua New Guinea. Bleher (personal communication) reports that K. ballochi is References no longer present at the type locality and only known Allen, G. R. 1983. Kiunga ballochi , a new genus and collection site, which lies approximately 40 km north species of rainbowfish (Melanotaeniidae) from of Kiunga next to the heavily-travelled road to the min - Papua New Guinea. Tropical Fish Hobbyist, 32 (2): ing town of Tabubil. The area is situated in the upper 72-77. Fly River system, about 900 km upstream from the Allen, G. R. 1995. in nature and the sea. Further details of the type locality were provided aquarium . Tetra, Melle, Germany, 180 pp. by Allen (1987). Allen, G. R., Ivantsoff, W., Shepard, M. A. & S. J. The new species was collected by Bleher from Tare Renyaan. 1998. Pseudomugil pellucidus (Pisces: Creek, located 11.5 km from the central market place Pseudomugilidae), a newly discovered blue-eye from in Kiunga along the Konkonda Road, which termi - Timika-Tembagapura region, Irian Jaya. Aqua Jour - nates at the Alice (Ok Tedi) River. According to Ble - nal of Ichthyology and Aquatic Biology, 3 (1): 1-8. her’s field notes, the upper portion of Tare Creek Allen, G. R. & S. J. Renyaan. 1999. Pseudomugil (about 1.5 km upstream of road crossing, Fig. 7) is ivantsoffi (Pseudomugilidae) a new species of blue- crystal clear with a mixed sand and rock bottom, but eye from the Timika region, Irian Jaya. aqua Journal further downstream in the vicinity of Konkonda Road of Ichthyology and Aquatic Biology, 3 (4): 163-168. the habitat has been severely degraded due to cas - Saeed, B., Ivantsoff, W. & G. R. Allen, 1989. Taxo - sava and banana cultivation. Consequently the fish is nomic revision of the family Pseudomugilidae (Order now rarer in this part of the creek than during his pre - ). Australian Journal of Marine and vious visits in 1991 and 1994. Freshwater Research, 40 : 719-787 The type specimens from Tare Creek were collected about 1.5 km upstream from the road crossing. At this point the stream is about 2.5 m in width and during low water periods is only 50-80 cm deep. A group of approx - imately 80 K. bleheri were observed at the type locality, swimming in the middle of the stream where the current was relatively fast. Temperature (26.2°C), pH (6.0), and a conductivity value of 10 µS/cm were recorded. There was no aquatic vegetation, although the Pandanus – lined stream was littered with a large amount of drift - wood and fallen trees. The only other fish inhabitants included gudgeons (Mogurnda and Ophieleotris sp.) and the rainbowfish Melanotaenia goldiei . The only known locations for the genus Kiunga are situated near vehicular roads. It seems highly unlikely that the two species are restricted to these locations. A study of topographical maps and satellite imagery of southern New Guinea indicates there is a strong possibility the fish will be found in adjacent areas with similar elevation (about 125 m), vegetation, and gen - eral water characteristics. Unfortunately the adjacent areas are located in dense jungle and are inaccessi - ble except on foot. Therefore the conservation status of the genus remains uncertain.

85 aqua vol. 8 no. 2 - 2004 Obituaries Three distinguished ichthyologists and collectors who died in 2003 It has been a sad year for science and for fish-lovers around the world. Three great personalities have passed away. One of them, R. R. Miller, was a member of the aqua editorial board from the very beginning. I would like to add my tribute to those of many others. These men will be remembered not simply because their names are now borne by a number of fish species – far fewer, in fact, than the number they discovered. Their names should also be remembered for an incredible record of fieldwork, ichthyological and natural history research, for their enthusiasm, and for a formidable contribution to the inventory of the species on our planet.

Robert Rush Miller (1916 - 2003) I will always remember our meet - ment in 1987. He began to work ing at the International Cichlid Con - with fishes in 1940 as a graduate ference, 1991, in Orlando, Florida. student, collaborating with Carl L. Bob and I talked for hours about Hubbs, and marrying Frances the genus "Cichlasoma" and about Hubbs in that year. Frances made the changes that took place when important contributions to his Sven Kullander separated some research program until her death in South American species from the 1987. The Miller children - Gifford, Central American species of this Francis, Roger, Ben, and Lawrence genus. Bob had studied the Cichli - - were also important contributors to dae for decades and I considered the field work, as were many gradu - him the world's foremost expert on ate students. The latter were all wel - the American members of this comed into the ichthyological family group. I still hold that opinion. over the years; all were made to feel Bob was a person who put friend - part of a tradition of lasting impor - ship before all else. He gave me tance to natural history. invaluable help with species identi - Bob published over 300 articles fication, as with the compilation of a contributing to ichthyology and con - book I have been working on for servation. These ranged from sys - more than a decade. He also tematic and experimental studies of worked on a large book on Mexican The late R.R. Miller with H. Bleher at the pupfishes, Cyprinodon, in Death Freshwater fishes for many years the ICC 91. Valley, to continuing studies of the (see below). We had great times at American Cichlid fishes of the western United States and Mexico. He Association meetings, and conferences of the Ameri - worked on trout, minnows, suckers, live-bearers, and can Society of Ichthyologists and Herpetologists, and sculpins, and also pioneered work in karyology, elsewhere. (At the 2001 ASIH conference Bruce hybridization, fossil fishes, and in experimental studies. Turner organized a symposium in his honour.) Bob When we first met he was Emeritus Professor at the and I had a great deal in common. He loved fishes at University Of Michigan Museum Of Zoology, Ann Arbor, least as much as I do. He used to say: “I like fish in all presiding over one of the largest preserved freshwater forms, I even love to eat them”. collections in the world. He was a brilliant man, the We shared a desire to help build up the world inven - owner of a lively mind, with a prodigious memory for tory of freshwater species. Our views were also facts, dates and details. shared on careless dam construction on the Lacantun Bob's field work and laboratory studies were affected river, and on other ecological disasters brought about by declining health in the 1990s. His work on the by human activity, causing damage to the environ - Freshwater Fishes of Mexico was carried forward by ment and breeding sites of millions of fishes. As early W. L. Minckley, Steve Norris, and Martha Gach, with as 1961, Bob's call for conservation: "Man and the Bob participating when health permitted. The final changing fish fauna of the American South-West," manuscript was sent the University of Chicago Press and his list of "Extinct, rare, and endangered Ameri - by Steve Norris in July, 2001. The last fishes Bob can freshwater fishes,” laid the foundations for the described were written up in the fall of 2002. American fish conservation movement. Goodbye friend, colleague and teacher. Thank you Robert Rush Miller was a faculty member in the for being Robert Rush Miller. Department of Zoology (later Biology) and a Curator in the Museum of Zoology from 1948 until his retire - Heiko Bleher – Managing Editor aqua vol. 8 no. 2 - 2004 86 Guy Teugels (1954 - 2003) The late Professor Guy Teugels was Senior Sci- More recently, he extended his research interests to entist and Curator of the Ichthyology Collection of the the freshwater fishes of Indonesia and Vietnam. Guy Royal Museum for Central Africa (Musée Royal de was a longstanding collaborator of the WorId Fish I’Afrique Centrale, MRAC) in Tervuren, Belgium, and Center, involved in the development of FishBase, a Professor at the Catholic University of Louvain. partner in research projects, and a resource person in Guy was a leading international expert on African training workshops in Africa. freshwater and brackishwater fishes, especially cat - I knew Guy very well. He and I and the late Jean- fishes, and on aquatic biodiversity. He led many pro - Pierre Gosse ( aqua vol. 4(4)) enjoyed a close work - jects, interacted with numerous institutions in Africa, ing relationship, not only in Tervuren, but also in and published widely in a number of languages, on Africa, as well as at conventions of the European taxonomy, species revisions and descriptions, genet - Society of Ichthyology and during private meetings. ics, biogeography, ecology, aquaculture, and the He was always extremely busy, at the same time effects of deforestation, dam construction and indus - always extremely helpful. He worked continuously trial pollution on fish populations. In particular, he con - and with intensity, made numerous field trips and, fol - tributed significantly to the development of a number lowing the compilation of the West African species, of definitive and pioneering works on African fishes, finally completed a manuscript on the freshwater notably the Check-list of the Freshwater Fishes of fishes of Cameroon and Gabon. It is to be hoped this Africa (CLOFFA) in 1991, and Faune des poissons can be printed soon. Guy died far too young, unable d’eaux douces et saumâtres de I’ Afrique de I’Ouest. to achieve his final ambition of covering the Central Tome I and II. Collection Faune Tropicale, in 1990 African fishes as well. It would take quite an effort to and 1992. fill the gap he leaves behind.

Jean-Claude Nourissat (1942 - 2003) Cichlasoma nourissati from the their environment; to find and Rio de la Pasion, Guatemala, have new species described and Paretroplus nourissati , from from remote areas. That night, the Mangarahara River, near over an excellent dinner of Ripp- Mandritsara in north-central chen mit Kraut in a restaurant in Madagascar, are two of the most Frankfurt’s famous Sach- beautiful cichlids in the world. senhausen district, we talked of Today, thanks to the late Jean- nature, expeditions, collecting Claude, they are kept and and discoveries. I provided him admired by aquarium hobbyists with some contacts in Brazil and worldwide. The world of aquaria offered him some advice. This and tropical fish may have lost last included a few words about one of its most interesting mem - malaria, an illness I have had bers, but the scientific commu - since 1967. I was not to know nity will miss him too. Jean- then that he would to die of it Claude discovered many new some 25 years later... cichlids from Central America Subsequently Jean-Claude and Madagascar. met many of my friends and col - I recall the day in 1978 when leagues such as Willy Schwartz, we first met. He walked into my Jacques Géry and Patrick de greenhouse outside Frankfurt- Rham. His first expedition, to am-Main and said: ”You must be the Amazon, took place that Heiko Bleher, the collector. It’s a same year. Later he concen - pleasure to meet you”. But it must be said, the plea - trated on Central American countries and Madagas - sure was mine. car. Jean-Claude must have gone on as many as 30 He had come from southern France to meet me, and such expeditions, more than 20 of them with Patrick. to see if I could help him. He wanted to collect fish We met many more times after that. He would come species as well, to propagate them and thus ensure to see me in Germany, and every encounter was a that some individuals survive; to keep live fishes and pleasure – I hope for both of us. He was the only learn as much as possible about their biology and really fearless person I ever met; prepared to cross

87 aqua vol. 8 no. 2 - 2004 any border and go anywhere to find a fish. We had aquariums of 15 and 28 cubic metres, as well as a much in common. good number of very large and medium-sized tanks. During his long presidency of the French Cichlid It must have been one of the most extensive private Association (A.F.C), of which he was a co-founder, “fish rooms” ever built. He kept his very large cichlids this gentleman repeatedly invited me to hold lectures in natural conditions in a gigantic pool outside. He at their annual conventions. Regrettably, Jean-Claude also had a wild duck collection, and a very large vine - was not a prolific writer, but he did publish a few arti - yard. He had the largest collection of Central Ameri - cles between 1992 and 2002 in the Revue Française can and Madagascar cichlids, superior to those of all d’Aquariologie and in the American magazine Cichlid the public aquariums I know of. Some of the species News, as well as in the magazine of the Association he had collected, such as Ptychocromoides katria , France Cichlid. In December 2002, together with were not available anywhere else at that time. Patrick de Rham, he brought out the very first mono - Widely respected, Jean Claude was the heart and graph on the endemic cichlids of Madagascar – “Les soul of the French Cichlid Association, His impressive Cichlides Endemiques de Madagascar”. contributions to this worldwide fraternity of ichthyolo - Jean-Claude was a dentist by profession but his gists and aquarium hobbyists are permanent. My passion was fish. In his Toulon house he had an enor - deepest sympathies go to his dear wife Nicole and to mous “fish room” an imposing construction almost his family, the cichlid world and the French Cichlid larger than his own living quarters, containing two Association.

Book review Rham, P. de & J.-C. Nourissat, 2002. Les Cichlides The distinguishing features of unidentified species are Endemiques de Madagascar, Association France given in great detail, as well as information on their life Cichlid. French edition. ISBN: 2-9513502-0-2 Price: in nature and in captivity. The habitat and collecting € 35.00 (including shipping). May be purchased sites of almost every species or variety, are described, directly from A.F.C.; also there is also an account of by credit card by contacting the life and diet of the local Roland Staub: fishermen and other inhabi - roland.staub@evc. tants. Their collecting meth - ods and the holding facilities Patrick de Rham and the late used are also mentioned. Jean-Claude Nourissant There follows an excellent undertook a series of field trips description of the biodiver - to almost all of the 38 river sys - sity of Madagascar. Many tems in Madagascar. Through other fresh- and brackishwa - collection and research they ter species of the island are built up a substantial knowl - touched on and photographs edge of the endemic cichlids of of some are included. The that island. authors examine the diver - Following an informative sity of the endemic cichlids introduction, the book gives in some detail, and discuss extensive information on their conservation; sugges - each of the 5 endemic gen - tions are made for protecting era: Paratilapia; Pty - their habitats and for captive chochromis; Ptychochro - breeding in Madagascar and moides; Oxytylapia and elsewhere. Their own expe - Paretroplus and all their riences with cichlid repro - known species. The genus duction are also described. Lamena Allgayer, 1998 is At the end of this admirable considered to be synony - work there is a checklist of mous with Paretroplus the 48 forms discovered by Bleeker, 1868. The varieties the authors, plus a new vari - of each discovered species ety discovered on their last are described, and detailed field trip. An English lan - descriptions of their locations guage version is soon to be and habitats are provided. published. aqua vol. 8 no. 2 - 2004 88 Instructions to Authors al Indo-Pacific Estuaries. Australian Journal of Marine and Freshwater Research, 31 :137-46. aqua is an international journal which publishes original sci- Day, J. H., Blaber, S. J. M., & J. H. Wallace. 1981. Estuarine entific articles in the fields of systematics, taxonomy, ethology, Fishes. In: Estuarine Ecology with Particular Reference to South - ecology, biogeography, and general biology of fishes, amphibians, ern Africa. (Ed. J.H. Day.): 197-221. A. A. Balkema, Rotterdam. aquatic invertebrates and plants. Papers dealing with freshwater, Dimmich, W. W. 1988. Ultrastructure of North American cyprinid brackish, and marine organisms will be considered for publication. maxillary barbels. Copeia , 1988 (1): 72-79. Scientific articles of interest to a wide readership are especially Trewavas, E. 1983. Tilapiine Fishes of the Genera Sarotherodon, welcome. Full length research papers and short notes will also be Oreochromis and Danakilia . British Museum (Natural History), considered for publication. London, 583 pp. 1. Manuscript preparation : manuscripts must be submitted in 7. Author details : these should be placed at the beginning of the English. In exceptional cases aqua may provide translations of manuscript, immediately after the title, and contain: full name(s) of manuscripts written in French, German, Italian, or Spanish. author(s), place(s) of work, and address(es) (including e-mail) for Manuscripts must be word-processed in Microsoft WORD and correspondence. submitted in an electronic form to the scientific editor. Hard copies 8. 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Additional reprints may be ordered from Aquapress. aqua Journal of Ichthyology and Aquatic Biology Vol. 8 (2), March 2004 Contents: John E. Randall and Andreas Spreinat: The subadult of the labrid fish Novaculoides macrolepidotus , a mimic of waspfishes of the genus Ablabys ...... 45-48

James C. Tyler and Philip A. Hastings: Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei) ...... 49- 60

John E. Randall: Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania ...... 61-79

Gerald R. Allen: Kiunga bleheri , a new Blue-Eye (Pisces: Pseudomugilidae) from fresh waters of Papua New Guinea ...... 79-85

Obituaries - three distinguished ichthyologists and collectors who died in 2003. Robert. R. Miller 1916-2003; Guy Teugels 1954-2003 and Jean-Claude Nourissat 1942-2003 ...... 86-88

Book review ...... 88

Papers appearing in this journal are indexed in: Zoological Record; Biolis - Biologische Literatur Information Senckenberg; www.aquageo.com; www.Joachim-Frische.com

Cover photo: Underwater photograph of Amblyeleotris katherinae with the snapping shrimp Alpheus ochrostriatus, Guam, Mariana Islands. Photo by J. E. Randall.

A different colour form of Kiunga ballochi G. R. Allen 1983. A beautiful species now extinct in nature. (See pages 79-85 in this issue and in forthcoming aqua Vol. 8 (3)). Photo by N. Armstrong.