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Accepted by J. Sparks: 11 Dec. 2013; published: 22 Jan. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3755 (3): 295–300
www.mapress.com
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Article
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http://dx.doi.org/10.11646/zootaxa.3755.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:61E01A93-8578-4FC8-A342-AA8D19CCBD91
Eviota piperata, a new gobiid species from Palau (Teleostei: Gobiidae)
DAVID W. GREENFIELD¹ & RICHARD WINTERBOTTOM²
¹Research Associate, Department of Ichthyology, California Academy of Sciences, 55 Music Concourse Dr., Golden Gate Park, San
Francisco, California 94118-4503, and Professor Emeritus, University of Hawaii. Mailing address: 944 Egan Ave., Pacific Grove, CA
93950. E-mail: greenfie@hawaii.edu
²Curator Emeritus, Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, M5S 2C6; and
Department of Ecology & Evolutionary Biology, University of Toronto, Toronto, Ontario, M5S 1A1. E-mail: rickw@rom.on.ca
Abstract
A new species of dwarfgoby, Eviota piperata is described from Palau. It belongs to the cephalic sensory-pore system
Group II (lacking only the IT pore); has a dorsal/anal-fin formula of 8/8; has some pectoral-fin rays branched; no dark spot
over the ural centrum; the male genital papilla is not fimbriate; and the cheek and body are heavily peppered with chro-
matophores.
Key words: Gobiidae, Eviota piperata, Dwarfgoby, Palau
Introduction
The Indo-Pacific gobiid genus Eviota is currently represented by 83 described valid species (Eschmeyer, 2014), yet
we are aware of many more awaiting description. This situation closely parallels that found in another genus of
small Indo-Pacific gobiids, Trimma, where there are almost 75 described valid species (Winterbottom, 2011).
During synoptic surveys of Indo-Pacific reef fishes, the second author made three collecting trips to Palau (2004,
2006, 2008), each lasting about three weeks and averaging about 40 scuba collections per trip. The Palauan islands,
on the fringes of the ‘Coral Triangle’ which encompasses the world’s greatest diversity of coral reef fishes (see. e.g.
Allen and Erdmann, 2012), contain a total fish fauna (including freshwater and open sea fishes) of some 1,700
species (Winterbottom, unpub. data). Approximately 50 previously undescribed species of fishes were obtained
during these trips, mostly gobies. Here we describe one of these new species in the gobiid genus Eviota as part of
our ongoing studies of members of this genus (see Greenfield & Winterbottom, 2012 for the descriptions of two
other new Palauan species of Eviota collected during these expeditions).
The new species fits the description typical of all species of Eviota: the pelvic fins are separate and the 5th
pelvic-fin ray, if present, is unbranched; the membrane joining the 5th pelvic-fin rays is rudimentary or absent; there
are ctenoid scales on the body but no scales on the head, nape or pectoral-fin base; the breast either lacks scales or
may have a few embedded cycloid scales; the teeth in the upper jaw are in two or more rows and there are 1–3
enlarged curved canine-like teeth in the innermost row of the lower jaw just behind the jaw symphysis.
Material and methods
Counts and measurements, descriptions of fin morphology and the cephalic sensory-canal pore patterns follow
Lachner & Karnella (1980) and Jewett & Lachner (1983). Postanal midline spots, along the posteroventral midline
of the body, begin at the anal-fin origin and extend to a vertical drawn 2 to 3 scale rows anterior to the ends of the
hypurals where they articulate with the caudal-fin ray bases, the additional smaller spot posterior to this is not
counted. “The membrane joining the 5th pelvic-fin rays is always short and weakly developed and the fins lack a
frenum. The membranes joining the first four fin rays are considered to be well developed when the membranes
GREENFIELD & WINTERBOTTOM
296 · Zootaxa 3755 (3) © 2014 Magnolia Press
extend beyond the bases of the first branches; they are considered to be reduced when they are slightly developed,
not extending to the bases of the first branches” (Lachner and Karnella, 1980). Dorsal/anal fin-ray counts only
include segmented rays.
Measurements were made to the nearest 0.1 mm using an ocular micrometer and dial calipers, and are
presented as percentage of Standard Length (SL). All specimen lengths are SL in mm. Cyanine Blue 5R (acid blue
113) stain was used to make pores more obvious (Akihito et al. 1993; Saruwatari et al. 1997; Nakabo 2002) and an
airjet used to observe them. For measurements, values for the holotype are given first, followed by the range for all
types and the mean in parentheses.
Specimens have been deposited in the following museums: AMS – Australian Museum, Sydney; CAS –
California Academy of Sciences, San Francisco; ROM – Royal Ontario Museum, Toronto, and USNM – United
States National Museum (Smithsonian), Washington D.C.
Eviota piperata n. sp.
Peppered Dwarfgoby
(Figs. 1–3)
Holotype. ROM 84490, 20.1 male, Palau, Sonsoral State, about middle of E coast of Fana Island, 05°21’17”N,
132°13’27”E, reef slope (30–44°) of coral rock with some rubble and coarse sand patches, moderate amounts of
hard coral (Acropora, Pocillopora, Millepora), many dark colored sponges, 6–9 m, rotenone, field number RW08-
06, Winterbottom, R., Westneat, M., Williams, J., Holleman, W., Hubley, B., Winterbottom, M., McCord, C., Rall,
H., 12 September 2008.
Paratypes: ROM 94753 (ex-ROM 84490), 7 males, 17.4–20.1, 18 females, 8.1–18.8, taken with holotype,
photographed specimen in vial; AMS I.146290-001, 18.2 male, 18.8 female, taken with holotype; CAS 236164,
20.5 male, 18.5 female, taken with holotype; USNM 410661 18.3 male, 18.1 female, taken with holotype, ROM
94754 (ex-ROM 84501), 4 males, 15.5–19.6, 5 females 12.1–16.1.
Other material examined: ROM 84501 (123), 10.7–18.6 mm, single specimen in vial photographed. Palau,
Sonsoral State, off SE tip of Pulo Anna, about 10 m from surf line, 04°39’09”N, 131°57’16”E, Halimeda, heavy
cover of hard corals (Acropora, Pocillopora, Millepora), plate corals, lettuce corals, U-shaped channel about 10 m
wide at seaward side and 3 m wide at shore side, floor 17 m at outside shelving to 9 m at tip, numerous caves at
base of tip, floor mostly of live coral, 7–15 m, rotenone, field number RW 08-09, Winterbottom, R., Westneat, M.,
Williams, J., Holleman, W., Hubley, B., Winterbottom, M., McCord, C., 13 September 2008. ROM 76386 (9),
10.4–14.7 mm, single specimen in vial photographed, Palau, Ngatopang, W coast Babeldaob Island, lagoon close
to outer barrier reef, 07°29’32.1”N, 134°25’28.6”E, bommies rising from sand/silt adjacent to coral rubble, brown
algae, 0–7.3 m, rotenone, field number RW 04-23, Winterbottom, R., Holleman, W., Hubley, B., Winterbottom, D.,
30 May 2004.
Diagnosis. The following combination of characters distinguish E. piperata from congeners: cephalic sensory-
pore system Group II (lacking only the IT pore), or lacking the IT and various combinations of the POP, from both
absent to only the upper or lower pore absent; dorsal/anal–fin formula 8/8; some pectoral-fin rays branched; no
dark spot over ural centrum; male genital papilla not fimbriate; cheek and body heavily peppered with
chromatophores.
Description. Dorsal-fin rays VI+I,8 (all); anal-fin rays I,8 (all); pectoral-fin rays 17 (16[11], 17[12]), rays 7–
16 branched in holotype; fifth pelvic-fin ray 7.8% (7.2–16.1, 10.4%) of 4th ray; 6 relatively long branches on 4th ray
(4[6], 5[3], 6[1]); 3 (2[6], 3[4]) segments between consecutive branches of 4th pelvic-fin ray; pelvic-fin membrane
reduced, about ¼ the distance to the first branch; 12 (12[4], 13[4]) branched and 17 (all) segmented caudal-fin rays;
24 (24[3], 25[7]) lateral scale rows; transverse scale rows 6 (all); vertebrae 24; midline of abdomen with cycloid
scales; first dorsal fin triangular in shape, first two spines of first dorsal fin may be filamentous in males as small as
12.5 mm, extending back past end of the second dorsal fin in holotype; genital papilla in male smooth, not
fimbriate, fringed at tip, extending to anal-fin spine; female genital papilla short and rounded with smooth sides,
several short finger-like projections on the end; body moderately slender, front of head rounded with an angle of
about 70° from the horizontal axis; mouth oblique, forming an angle of about 50° to horizontal axis of body, lower
jaw not projecting; maxilla extending to posterior third of pupil; anterior tubular nares short, just reaching to edge
Zootaxa 3755 (3) © 2014 Magnolia Press · 297
EVIOTA PIPERATA
of upper lip; gill opening extending forward to posteroventral edge of preoperculum; cephalic sensory-pore system
pattern Group II (only IT pore missing), or lacking the IT and various combinations of the POP, from both absent to
only the upper or lower pore absent.
Measurements (based on holotype and 11 paratypes, 12.1–20.1). Head length 31.4 (30.3–34.2, 32.2); origin of
first dorsal fin 33.7 (33.7–37.9, 36.4), lying behind posterior margin of pectoral-fin base; origin of second dorsal
fin 56.6 (54.9–60.3, 57.6), slightly in advance of anal-fin origin; origin of anal fin 60.5 (58.7–62.7, 60.3); caudal-
peduncle length 22.7 (22.4–24.8, 23.9); caudal-peduncle of moderate depth 13.5 (10.9–15.3, 13.3); body relatively
slender, its depth 21.4 (19.6–22.2, 21.1); eye diameter 8.7 (8.7–11.7, 11.1); snout length 5.1 (4.2–6.0, 5.1);
pectoral-fin length 28.8 (25.5–37.5, 31.4); pelvic-fin length 55.3 (30.1–55.3, 36.6).
Color in preservative of holotype (Fig. 1). Background color of head and body pale yellowish. Head with a
heavy peppering of chromatophores on the top and nape, on the cheeks, and on the operculum. A concentration of
chromatophores along the posterior edge of the preoperculum that extends down to a heavy peppering of
chromatophores on the branchiostegal membranes. A cluster of chromatophores below the eye extending down
behind the jaws. Interorbital area, snout, nares and jaws with a lighter peppering. Pupil of the eye clear, iris black.
A heavy band of chromatophores on the pectoral-fin base. Peppering of chromatophores less intense on the body
than on the head, scale pockets outlined with chromatophores, more intense on the dorsal half of the body.
Abdomen peppered. A thin dark line running along the middle of the sides of the body along the caudal peduncle.
Five obvious, dark, postanal midline spots followed by a smaller spot, matched by a similar spot at the top of the
caudal-fin base. Caudal, anal and second dorsal fins peppered with chromatophores and melanophores.
Filamentous spine of first dorsal fin with a series of spaced brown spots. Remainder of fin peppered with small
brown spots, its distal margin darker. Pectoral and pelvic fins lighter than other fins.
FIGURE 1. Eviota piperata, holotype preserved, ROM 84490. Photograph by D.W. Greenfield.
FIGURE 2. Eviota piperata, holotype fresh, ROM 84490. Photograph by R. Winterbottom.
GREENFIELD & WINTERBOTTOM
298 · Zootaxa 3755 (3) © 2014 Magnolia Press
FIGURE 3. Eviota piperata, paratype fresh, ROM 94753. Photograph by R. Winterbottom.
Color of fresh holotype (Fig.2). Background color of head and body light brown. Scale pocket margins
outlined with chromatophores, more intense on dorsal surface. Head darker than body, heavily peppered with large
chromatophores, more concentrated on preopercular margin, under eye, behind jaws, and under head. Two dark
bars crossing top of head behind eyes. Nares dark with black tips. Pupil of eye black, iris yellow with orange
spokes radiating out from pupil. Pectoral-fin base dark, slightly darker on top and bottom. Body with six
subcutaneous bars: the first at anterior half of first dorsal fin; the second at posterior half of first dorsal fin; third
from origin of second dorsal fin to anal-fin origin; fourth at end of anal fin; fifth and sixth on caudal peduncle. Five
postanal midline spots on ventral surface of body: the first at anal-fin origin, second at end of anal fin; third, fourth,
and fifth on caudal peduncle, followed by a smaller spot at ventral caudal-fin margin next to procurrent spines, this
last spot matched at top of caudal fin. Caudal and anal fins brown, heavily peppered with melanophores.
Filamentous spine of first dorsal fin with a series of spaced black spots with orange halos. Remainder of fin brown
with an orange tinge, its distal margin darker. A narrow light area running from third black spot of first dorsal spine
down across fin to base of fourth spine. A single black spot on membrane between spines five and six. Second
dorsal fin brown, heavily peppered with chromatophores, lighter on distal half. Spine with four spaced black spots
similar to those on the first spine of the first dorsal fin, similar dark spots on the soft rays. Pectoral and pelvic fins
lighter than other fins.
Fresh color of paratype. In some specimens, the peppering of chromatophores on the head is lighter revealing
the concentration of chromatophores on various portions of the head (Fig. 3).
Distribution. Only known from Palau. Eviota piperata was collected throughout the Palauan Islands, from the
north tip of Babeldaop to Helen Reef, the southernmost of the South West Islands. It was locally abundant in the
South West Islands (except at Helen Reef, where only two specimens were collected despite nearly two weeks of
sampling), with a single collection from Pulo Anna containing 131 specimens, and a total of over 330 specimens
collected in limited collecting effort (7 days). Two three-week long expeditions to the main Palauan Islands (in
2004 and 2006) resulted in the collection of only 34 specimens of this species. The five South West Islands
(excluding Helen Reef) are all small coral limestone islands perched on submerged volcanic peaks, with small
shallow lagoons and steep outer slopes to deepwater.
Etymology. The specific epithet is from the Latin piper (pepper) referring to the heavy peppering of
melanophores over the head and rest of the body.
Comparisons. Of the 30 described species of Eviota in cephalic sensory-pore system Group II (lacking only
the IT pore) (Table 1), only four others have branched pectoral-fin rays and a dorsal/anal-fin formula of 8/8 (E.
dorsimaculata, E. indica, E. latifasciata, and E. rubra). Eviota dorsimaculata, E. indica and E. latifasciata all have
a dark spot on the caudal peduncle over the ural centrum that is lacking in E. piperata, and neither E. indica nor E.
latifasciata have an elongation of the first dorsal fin, which rarely is slightly elongated in E. dorsimaculata (the
first spines in males of E. piperata may be filamentous). Eviota rubra also has branched pectoral-fin rays and no
spot over the ural centrum; however, E. rubra has a distinctive fimbriate genital papilla in both the male and
female, whereas the papillae are non fimbriate in E. piperata. Eviota rubra also lacks the heavy peppering of
melanophores present in E. piperata as well as other color differences (Greenfield & Randall, 1999, 2004-Fig. 62).
Zootaxa 3755 (3) © 2014 Magnolia Press · 299
EVIOTA PIPERATA
TABLE 1. Species of Eviota included in the cephalic sensory-canal pore system Group II (IT missing), dorsal/anal-fin
formulas (D/A) and pectoral-fin branching (S=simple, BR=branched).
Because the number of preopercular pores (POP) can be variable in E. piperata it also is necessary to compare it to
five other species of Eviota that lack both the IT and POP: E. lacrimae, E. ocellifer, E. pinocchioi, E. sparsa and E.
susanae. All of these species except E. susanae have a dorsal/anal-fin formula of 9/8, (8/8 in E. piperata). Eviota
susanae has a fimbriate genital papilla in both the male and female (non-flimbrate in E. piperata); the 5th pelvic-fin
ray is 53–90% of the length of the 4th ray in E. sparsa (vs. 7.2–16.1% in E. piperata); E. ocellifer has the
anteroventral region of the first dorsal fin black, bordered above with pale yellow, whereas E. piperata lacks that
dorsal fin coloration; the anterior tubular nares are black and very elongate, reaching well anterior to the upper lip
in E. pinocchioi (nares only reach to the upper lip in E. piperata).
Remarks. Eviota piperata is unusual in exhibiting variation in the presence of the preopercular pores (POP).
Even though the cephalic-sensory pore system groupings do not appear to reflect genetic relationships (Tornabene
et al. 2013), they are useful in differentiating different species and species groups and usually do not vary within a
species. We are aware of an undescribed species of Eviota in Indonesia that is similar in coloration to E. piperata
but differs in pore patterns. The relationship between these two species awaits the collection of more material,
especially DNA tissue. We note in passing that 1 of 234 specimens that we assigned to E. piperata possessed the IT
pore – we regard this specimen as an anomaly.
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GREENFIELD & WINTERBOTTOM
300 · Zootaxa 3755 (3) © 2014 Magnolia Press
Acknowledgments
The staff of the California Academy of Sciences as usual has provided continual support: D. Catania, J. Fong, M.
Hoang, and L. Rocha. The curatorial expertise of Don Stacey and Margaret Zur (both ROM) is gratefully
acknowledged. RW would like to express his sincerest thanks to the seven other members of the “fish” team (see
Material Examined for names), and Pat and Lori Colin of the Coral Reef Research Foundation (CRRF). Permission
to collect and export the specimens was graciously supplied by the members of the Palau Bureau of Marine
Resources, the Helen Reef Resource Management Project, the peoples of the South West Islands, and Youlsau
Bells and Joel Miles (Office of Environmental Response and Coordination). RW’s fieldwork was financially
supported by the ROM Foundation, the ROM’s Department of Natural History, NSERC Discovery Grant 7619, an
NSERC Ship Time grant, and a grant from The Nature Conservancy – deep gratitude to all these agencies and their
officers.
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