Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 1, pp. 163–174, March 22, 2007
Two New Species of Trimma (Gobiidae) from the Western Pacific
Kiyoshi Hagiwara1 and Richard Winterbottom2
1
Yokosuka City Museum, 95 Fukada-dai, Yokosuka, Kanagawa, 238–0016, Japan
E-mail: kiyoshi-hagiwara@city.yokosuka.kanagawa.jp
2
Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, M5S 2C6, and
Department of Zoology, University of Toronto, Toronto, Ontario, M5S 3G5, Canada
E-mail: rickw@rom.on.ca
Abstract Two new species of Trimma from western Pacific coral reefs are described. Trimma
flavatrum is characterized by the presence of cheek and opercular scales, scales in the midline of
the predorsal region, no elongate dorsal fin spines, unbranched pectoral fin rays and a rather broad,
shallowly concave, interorbital region with no central fleshy ridge. When alive T. flavatrum has red
pectoral fins, yellow to reddish-orange dorsal and anal fins, and the caudal fin varies from hyaline
to yellow with hyaline to white margins. Preserved specimens are dusky with densely scattered
melanophores and chromatophores, the shade intensifying posteriorly to almost black in the peduncular region, and the fins are translucent, with a black basal stripe in the dorsal and anal fins.
Trimma flavatrum have been found throughout the western Pacific, ranging longitudinally from
Palawan to Samoa and latitudinally from the Nansei Islands to Fiji. Trimma hayashii is characterized by the absence of predorsal scales in the midline (except in some large specimens ⬎20 mm
SL, where 1 or 2 cycloid scales may be present), no opercular or cheek scales, no or only slightly
elongate dorsal spines, a few branched pectoral fin rays in the middle of the fin, a moderately developed interorbital trench and no postorbital trench. When alive T. hayashii has pupil-diameter
sized red spots on the head and nape, and 3 yellow or reddish-orange stripes on the body separated
by 2 grey stripes. Uniquely among species of Trimma known to date, this species has a ventrallydirected, ocellated, pupil-diameter sized spot on each of the branchiostegal membranes. Trimma
hayashii ranges longitudinally from the western tip of Java to Kosrae (Federated States of Micronesia), and latitudinally from the Nansei Islands to the Solomons.
Key words : Gobiidae, Trimma flavatrum, Trimma hayashii, New species, Pacific Ocean
Trimma contains some 85 species of small
(less than 30 mm SL), often colourful gobiids,
primarily associated with Indo-Pacific coral
reefs. Trimma may be recognized by the lack of
cephalic sensory canal pores, much reduced
cephalic sensory papillae pattern, wide gill opening extending to below the vertical limb of the
preopercle or anterior to this, lack of spicules on
the outer gill rakers of the first gill arch, less than
12 dorsal and anal rays, and a fifth pelvic fin ray
that is equal to or more than 40% the length of
the fourth pelvic fin ray. There are 48 valid
species (Table 1) of Trimma and approximately
40 additional species that have yet to be described. However, recent use of rebreather technology to collect fishes is uncovering a rich di-
versity of deep-dwelling (to 100 m) and undescribed species, and the total number of species
in the genus can be expected to rise considerably.
Methods
Methods follow Winterbottom (2002), except
pectoral and pelvic fin ray branching is described
from preserved material stained with a cyanineblue solution as outlined in Saruwatari et al.
(1997). Values for holotype in bold where appropriate. Lengths given are standard lengths (SL),
and are given in millimeters. Numbers in parentheses after the catalogue numbers for non-type
material listed represent the number of specimens in that lot. Institutional codes for reposito-
164
K. Hagiwara and R. Winterbotton
Table 1. List of valid species of Trimma.
No. species
Species
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
Trimma caesiura
Trimma grammistes
Trimma macrophthalma
Trimma striaa
Trimma okinawae
Trimma naudei
Trimma unisquamis
Trimma corallina
Trimma flammeum
Trimma necopinus
Trimma tevegae
Trimma avidori
Trimma mendelssohni
Trimma taylori
Trimma flavicaudatus
Trimma dalerocheila
Trimma haima
Trimma sheppardi
Trimma fraena
Trimma griffithsi
Trimma hoesei
Trimma winchi
Trimma fishelsoni
Trimma emeryi
Trimma winterbottomi
Trimma rubromaculatus
Trimma filamentosus
Trimma barralli
Trimma benjamini
Trimma anaima
Trimma bisella
Trimma omanensis
Trimma halonevum
Trimma stobbsi
Trimma woutsi
Trimma milta
Trimma volcana
Trimma annosum
Trimma lantana
Trimma cana
Trimma sostra
Trimma squamicana
Trimma agrena
Trimma fangi
Trimma marinae
Trimma nasa
Trimma anthrenum
Trimma preclarum
ries of material examined follow Leviton et al.
(1985) with an additional abbreviation: Biological Labotatory, Imperial Household (BLIH).
Author (s)
Date
Jordan and Seale
(Tomiyama)
(Tomiyama)
(Herre)
(Aoyagi)
Smith
(Gosline)
(Smith)
(Smith)
(Whitley)
Cohen and Davis
(Goren)
(Goren)
Lobel
(Goren)
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Goren
Winterbottom
Randall and Downing
Allen and Munday
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom
Winterbottom and Villa
Winterbottom
Winterbottom
Winterbottom
Winterbottom and Chen
Winterbottom and Chen
Winterbottom
Winterbottom
Winterbottm
Winterbottom
1906
1936
1936
1945
1949
1957
1959
1959
1959
1959
1969
1978
1978
1979
1982
1984
1984
1984
1984
1984
1984
1984
1985
1985
1994
1995
1995
1995
1996
2000
2000
2000
2000
2001
2002
2002
2003a
2003b
2003
2004
2004
2004
2004
2004
2005a
2005a
2006
2006
Trimma flavatrum sp. nov.
(New Japanese name: Hime-aogi-haze)
(New English name: Wasp pygmy goby)
(Figs. 1–6)
Trimma sp. 10: Senou et al., 2004: 114 (Okinawa Islands,
Miyako Island, Ishigaki Island, Iriomote Island).
Two New Trimma from Western Pacific
165
Fig. 1. Trimma flavatrum, fresh specimen, 18.1 mm SL male paratype, Amami-oshima Island, Nishikomi Cave,
YCM-P39240. Photo by K. Hagiwara
Fig. 2. Trimma flavatrum, stock specimen, 23.0 mm SL male holotype, Amami-oshima Island, Nishikomi Cave,
YCM-P42599. Photo by K. Hagiwara
Holotype. YCM-P42599, 23.0 mm SL male, Japan,
Nansei Islands, Amami-oshima Island, Nishikomi Cave,
28°13⬘54⬙N; 129°10⬘35⬙E, 8 m, K. Hagiwara, 14 Sep.
2005.
Paratypes. 21 specimens. Japan (Nansei Islands):
YCM-P34476 (20.6 SL), Amami-oshima Island, Nishikomi Cave, 8 m, Sagami-bay Marine Biological-research
Club (SMBC), 31 Aug. 1994. YCM-P36357, (17.3),
Kakeroma-jima Island, reef edge in front of Saneku village, 28°11⬘00⬙N, 129°11⬘36⬙E, 25 m, SMBC, 23 Aug.
1995. YCM-P39240, (18.1), Amami-oshima Island,
Nishikomi Cave, 8 m, K. Hagiwara and T. Itoh, 5 Nov.
1999. YCM-P41475, (19.0), Kakeroma Island, reef edge
in front of Adachi village, 28°08⬘37⬙N, 129°11⬘48⬙E, 10
m, SMBC, 2 Sep. 2000. NSMT-P61915, (16.3), Ishigakijima Island, Urazoko Bay, 24°27⬘31⬙N, 124°14⬘00⬙E, 15
m, K. Matsuura, 22 Aug. 1996. BLIH 20010029, (16.9),
Ishigaki-jima Island, Yonehara reef, 24°27⬘36⬙N,
124°11⬘16⬙E, 8 m, Y. Ikeda, K. Sugiyama and K. Hagiwara, 2 July 2001. BLIH 20030044, (21.4), Amami-oshima Island, Nishikomi, 28°13⬘54⬙N, 129°10⬘35⬙E, T.
Yonezawa, 9 Apr. 2003. Caroline Islands: ROM 74801, 3
(14.0–15.0), Philippine Sea, Short Drop-off SE of Koror
Island, SW corner of Augulpelu Reef, 07°16⬘28.7⬙N,
134°31⬘32.1⬙E, 13.7–25.9 m, R. Winterbottom, B. Hubley,
D. Winterbottom and A. Bauman, 21 May 2004.
ROM 74802, 4 (6.7-15.8), Philippine Sea, Short Drop-off
SE of Koror Island, SW corner of Augulpelu Reef,
07°16⬘54.3⬙N, 134°31⬘38.7⬙E, 15.2–26.5 m, R. Winterbottom, W. Holleman, B. Hubley and D. Winterbottom, 28
May 2004. ROM 74926, 3 (14.0–17.1), Philippine Sea,
off SE corner of Koror Island. along Short Drop-off, Augulpelu Reef, SW corner, 07°16⬘26.9⬙N, 134°31⬘29.3⬙E,
R. Winterbottom, B. Hubley, A. Bauman and S. Kiefer, 20
May 2004. Papua New Guinea: ROM 73442, (12.7),
Greater Kimbe Bay, New Britain, 05°53⬘S, 150°10⬘E, M
Beger, 12 Oct. 2002. ROM 60747, (16.4), Bismark Sea,
5°09⬘S, 145°50⬙E, G. Allen, 1 Jan. 1993 Indonesia: ROM
64646, (16.6), Sulawesi, Manado, 1°38⬘N, 124°46⬙E, 16
m, J. Randall and M. Severans, 29 Oct. 1991. BPBM
36559, 2 (18.8–20.1), Banda Sea, Penyu Islands, Maisel
Island, 14 m, J. L. Earle, 23 Oct. 1990.
Non-type material. 116 specimens. Australia (Great
Barrier Reef): AMS I.19472-089 (1); AMS I.20757-067
166
K. Hagiwara and R. Winterbotton
Fig. 4. Trimma flavatrum, composite dorsal view
of head of cyanine-blue stained 15.8 mm SL
male paratype, Augulpelu Reef, Palau, ROM
74802. Photo and imagine enhancement by R.
Winterbottom
Fig. 3. Trimma flavatrum, live, Amami-oshima
Island, Nishikomi Cave, 8 m. Photo by K.
Hagiwara
(1); AMS I.20775-056 (1); AMS I.20779-142 (1); AMS
I.22580-025 (1); Rowley Shoals: WAM P.28024-33 (1).
Caroline Islands: USNM 223193 (3); USNM 223195
(3); USNM 298765 (3); BPBM 31421 (4). Indonesia:
NSMT-P71254 (3); USNM 210240 (3); USNM 244186
(8). Fiji: ROM 46001 (2); ROM 46002 (2); USNM
236756 (8). Japan (Nansei Islands): YCM-P34470 (1);
YCM-P36317 (3); YCM-P41476 (1); YCM-P41475 (5);
NSMT-P61946 (7); NSMT-P64899 (2); NSMT-P65034
(2); BLIH 20010413 (1); BLIH 20010019 (1). Marshall
Islands: BPBM 12181 (1). Papua New Guinea: USNM
244098 (1). Philippines: ROM 52976 (2); ROM 52977
(1); ROM 52978 (1); USNM 243918 (28); USNM
243919 (8); USNM 264524 (1); USNM 295300 (1);
YCM-P40675 (1); YCM-P40691 (1). Samoa: AMS
I.21998-002 (2).
Diagnosis. Trimma flavatrum is characterized by the presence of scales in the predorsal
midline and on the cheek and opercle; no elongate dorsal spines; a broad and somewhat concave interorbital region with a bony interorbital
equal to about half or more of the width of the
pupil with no raised, longitudinal, fleshy ridge in
the midline; no postorbital trench; fifth pelvic fin
ray usually unbranched (may be branched once
sequentially); and, in preserved specimens, a
dusky body with densely scattered melanophores
and chromatophores, the shade intensifying posteriorly to almost black in the peduncular region.
When alive or freshly collected, the body is
yellow to dirty yellow-orange with scattered
melanophores intensifying to a blackish or dark
brown caudal peduncle, the median fins are reddish-orange to yellow with a dark basal stripe,
except for the caudal fin, which varies from almost translucent to yellow with whitish margins
(Figs. 1, 3).
Description. The description is based on the
holotype and a variable number of additional
specimens. Dorsal fins VI-I, 7–8 (mean⫽7.9,
n⫽11), second spine longest and reaching the interspace between the dorsal fins when adpressed,
all dorsal rays branched; anal fin I, 7–8
(mean⫽7.9, n⫽11), all rays branched; pectoral
fin 13–15 (mean⫽14.1, n⫽11), rays unbranched,
reaching posteriorly to a vertical line with the anterior few elements of the anal fin; pelvic fin I, 5,
no frenum or basal membrane, first 4 rays with 1
sequential branch, fifth ray unbranched or with 1
sequential branch and 40–60% the length of the
fourth, fourth ray reaching posteriorly to a vertical line with the first to third element of the anal
fin; caudal fin with 11 branched rays, 8 dorsal
and 7–8 ventral segmented rays, with 5–7 dorsal
(6 in holotype) and 4–6 ventral unsegmented
rays visible externally. Lateral scales 22–26
(mean⫽23.5, n⫽15), anterior transverse scales
7–9 (mean⫽8.2, n⫽15), posterior transverse
scales 6–8, (7 in holotype, mean⫽7.4, n⫽15);
predorsal scales 7–8 (mean⫽7.8, n⫽6), scales on
Two New Trimma from Western Pacific
breast cycloid, 2 rows of cycloid scales on pectoral base, with posterior row of 2–3 scales, the
dorsalmost scale very large and occupying about
80% of the height of the fin base, scales on the
head extending into the interorbital region to
about level with the posterior third of the pupil,
2–4 cycloid (3 in holotype) scales on cheek, 2–3
rows of cycloid scales (up to 7 scales) on opercle
(Fig. 5).
Gill opening to below mid-pupil. Upper jaw
with outermost and innermost rows of widely
spaced, enlarged, curved canines with a few irregular rows of small conical teeth between
them. Lower jaw with an outer row of enlarged,
widely spaced, curved canines across the front of
the jaw to the bend of the dentary, a middle row
of small closely spaced conical teeth, and an
inner row of widely spaced, somewhat enlarged
canines. Tongue rounded. Gill rakers on first arch
3–4⫹11–13⫽15–17 (holotype not counted, mean
⫽15.5, n⫽8). Nasal apparatus a slightly raised
sac, confined to the anterior half of the snout, anterior nasal opening a tube, posterior nasal opening a pore with a raised rim (Fig. 5). Positions of
sensory papillae are showed by Figure 6.
Head length 27.0–33.0% (28.6% in holotype,
mean 30.2%, n⫽11) SL, body depth 23.6–24.6%
(mean 24.5%, n⫽11), body width 13.3–16.3%
(mean 15.3%, n⫽10). Snout length 15.7–23.8%
(mean 18.8%, n⫽11) of head length, orbit diameter 36.8–51.1% (38.8% in holotype, mean
42.3%, n⫽11), upper jaw length 30.7–42.9%
(35.7% in holotype, mean 36.8%, n⫽11). Bony
interorbital 1/4–1/3 pupil width, with a interorbital trench and no postorbital trench (Fig. 5).
Vertebrae 10⫹16⫽26, first and second caudal
vertebra with the hemal arch expanded for a
length equal to 2/3 length of hemal spine, the
swim bladder extends into this space.
Color pattern (from slides of freshly collected
specimens from the Nansei Islands, Philippines,
and Palau, Fig. 1). A dark yellow to yellowishbrown body, heavily sprinkled with melanophores, without any conspicuous color patterns.
The body color darkens gradually posteriorly and
the caudal peduncle is dark brown to black. The
167
Fig. 5. Trimma flavatrum, composite left lateral
view snout of cyanine-blue stained 14.0 mm
SL female paratype, Augulpelu Reef, Palau,
ROM 74926. Photo and imagine enhancement
by R. Winterbottom
Fig. 6. Trimma flavatrum, diagrammatic figure of
sensory papillae. Drawn by K. Hagiwara
median fins, except the caudal fin, are yellow to
orange with a half-pupil width, dark or black
basal stripe formed by melanophores with scattered iridocytes; the tips of these fins are hyaline.
The caudal fin may be translucent with some
melanophores and chromatophores on the central
base, or with a white margin to the upper and
lower unbranched fin rays, sometimes confluent
with a white bar across the bases of the branched
fin rays, and, in some cases, the middle portion of
the fin yellow. The pectoral fins have a reddish
wash, the pelvic spine is red with the rest of the
fin yellow to orange. The iris is yellow to gold,
and usually heavily invested with melanophores.
Color in alcohol. The body color is as for
fresh specimens, except that all traces of yellow,
orange and red have disappeared, with a straw-
168
K. Hagiwara and R. Winterbotton
yellow background and a heavy sprinkling of
melanophores, which gradually become more
concentrated posteriorly on the caudal peduncle. Scales margins are not outlined with
melanophores. The dorsal and anal fins are
translucent with a basal stripe of melanophores.
NSMT-P71254 which collected in Ambon Island
(Indonesia) has melanophores on near the tips of
the dorsal and anal fins. Melanophores are absent
in the pectoral and pelvic fins. The caudal fin is
translucent with melanophores on the base. All
observed specimens have a black iris (Figs. 2, 5).
Affinities. Trimma flavatrum sp. nov. is very
similar to juveniles of Trimma tevegae Cohen
and Davis, 1969 when preserved in alcohol. The
2 share many characteristics including the number of dorsal, anal and pectoral fin rays (D 8–9,
A 8–9, P113–15 vs. D 7–8, A 7–8, P1 13–15 for
T. flavatrum), a few cycloid scales on the cheek
and opercle, and an almost black caudal peduncle. Trimma tevegae may have an elongated second dorsal spine (vs. never elongated), and has 9
to 12 predorsal scales (vs. 7–8), a longitudinal,
median, rounded, fleshy ridge in the center of the
interorbital region, and a bony interorbital width
about as wide as the pupil diameter [see Winter-
bottom (2005b); vs. no ridge and bony interorbital half to two-thirds pupil width]; extent of
hemal arch of first caudal vertebra equal to the
length of the hemal spine (vs. two-thirds neural
spine), a broad, diffuse stripe of chromatophores
on the dorsum, with the scales outlined with
melanophores creating a diamond-like pattern
(vs. stripe and scale outlines absent) and a relatively distinctly demarkated black caudal spot
(vs. a gradual darkening over the posterior half of
the body) (comparison with YCM materials).
Trimma milta Winterbottom, 2002 and Trimma
annosum Winterbottom, 2003 are similar to T.
flavatrum in having an interorbital trench, a poorly developed postorbital trench, no elongate dorsal spines and in the number of predorsal, lateral
and transverse scales. T. milta and T. annosum
have a pelvic fin basal membrane (vs. absent),
more than 17 pectoral fin rays (vs. less than 15
rays). Neither of these species possesses any dark
pigmentation on the caudal peduncle.
Habitat. Trimma flavatrum forms small
schools which are sometimes mixed with T. tevegae. The fishes hover in caves, recesses or under
the overhangs of coral reefs in the western Pacific. When hovering, the ventral side faces the
Fig. 7. Distribution map for Trimma flavatrum (inverted triangles) and T. hayashii (diamonds). Symbols underlined together indicate co-occurrence at that locality.
Two New Trimma from Western Pacific
nearest substrate surface (e.g. the wall or roof of
a cave) as occurs in T. tevegae. However, T. flavatrum tends to be found more towards the back of
the caves and recesses. When feeding, T. tevegae
tends to adopt a head-up, often vertical, posture
in the water column (Cohen and Davis, 1969;
Hagiwara, personal observation).
Distribution. Trimma flavatrum has been
collected at Japan, the Philippines, Palau, Micronesia, Indonesia, Papua New Guinea, the
Great Barrier Reef and Rowley Shoals in Australia, Fiji and Samoa (Fig. 7).
Etymology. From the Latin ‘flavus’, meaning yellow, and ‘atrum’, black, in allusion to the
unusual yellow and black coloration of the new
species.
169
Trimma hayashii sp. nov.
(New Japanese name: Eri-hoshi-beni-haze)
(New English name: Four-eye pygmy goby)
(Figs. 7–13)
Trimma sp. No.076: Hayashi and Shiratori, 2003: 45,
middle left figure (Amami-oshima Island).
Trimma sp. 4: Senou et al., 2004: 114 (Miyako Island, Iriomote Island).
Holotype. ROM79838 (ex-YCM-P26315), 22.2 mm
SL male, Kakeroma Island, Chino-ura, 28°09⬘42⬙N,
129°17⬘39⬙E , 10–20 m, SMBC, 28 Aug., 1991.
Paratypes. 135 specimens. Japan (Nansei Islands):
YCM-P26316 (19.8 SL) as for holotype. YCM-P 28274,
5 (20.3–23.4), Amami-oshima Island, Atetsu Bay,
28°13⬘00⬙N, 129°17⬘40⬙E , 5–17 m, SMBC, 28 Aug.
1992. YCM-P 29500, (25.8), Amami-oshima Island,
Fuka-ura, 28°12⬘29⬙N, 129°17⬘14⬙E, 5–15 m, SMBC, 3
Sep. 1993. YCM-P 42313 (18.3), Amami-oshima Island,
Atetsu Bay, 8 m, K. Hagiwara and T. Itoh, 6 Nov. 1999.
YCM-P 41238 (19.6), Amami-oshima Island, Atetsu Bay,
5–17 m, SMBC, 30 Aug. 2000. YCM-P 42551 (23.8,
Fig. 8. Trimma hayashii, fresh specimen, 22.2 mm SL female holotype, Kakeroma Island, Chino-ura,
ROM79838. Photo by M. Hayashi
Fig. 9.
Trimma hayashii, stock specimen, holotype. Photo by K. Hagiwara
170
K. Hagiwara and R. Winterbotton
female), Amami-oshima Island, Atetsu Bay, 5–17 m,
K. Hagiwara, 19 Oct. 2004. YCM-P42570, 4 (22.3–
23.3), Kakeroma Island, Nomino-ura, 28°06⬘49⬙N,
129°17⬘46⬙E, 20 m, K. Hagiwara, 11 Oct. 2004. NSMTP34813 (20.3), Amami-oshima Island, mouth of Atetsugawa River, 28°11⬘01⬙N, 129°17⬘05⬙E, 12 m, 12 June
1991. BLIH 19920341 (22.6), Ishigaki-jima Island, Kabira Bay, 24°26⬘46⬙N, 124°08⬘33⬙E, A. Iwata and S.
Hosoya, 27 Nov. 1992. BLIH 20030039 (24.1), Amamiosima Island, Kiyama-jima, 28°01⬘17⬙N, 129°16⬘27⬙E, T.
Yonezawa, 26 Mar. 2003. Caroline Islands: ROM 74937,
5 (15.9–21.2), west coast of Babeldaob Island. off Aimeliik, reef slope with corals, fine sand and silt,
07°29⬘20.3⬙N, 134°26⬘03⬙E, 9–15 m, R. Winterbottom, B.
Hubley, A. Bauman and D. Winterbottom, 19 May 2004.
ROM 76084, 7 (20.3–23.1), W coast of Ngeruktabel Island., rock and coral wall to sand/coral base,
07°17⬘11.7⬙N, 134°25⬘34.5⬙E, 6–14 m, R. Winterbottom,
W. Holleman, B. Hubley, A. Bauman and D. Winterbottom, 26 May 2004. ROM 76085, 11 (9.0–20.5), west
coast of Babeldaob Island to north of main pass, reef
slope with mixed corals, sand and silt, 07°33⬘07.7⬙N,
134°29⬘17.9⬙E, 0–4.5 m, R. Winterbottom, W. Holleman,
B. Hubley and D. Winterbottom, 1 June 2004. ROM
76086, 23 (15.4–22.5), E. side of Nikko Bay just W. of
Kaibakku Island, coral slope to coral wall with silt/sand
slope below, 07°19⬘19.9⬙N, 134°29⬘58.4⬙E, 15–26 m, R.
Winterbottom, W. Holleman, B. Hubley, A. Bauman and
D. Winterbottom, 4 June 2004. ROM 76404, 62
(9.0–23.1) E. side of Nikko Bay about 50 m E of ROM
76086, very steep reef slope with huge variety of hard
corals, 07°19⬘22.8⬙N, 134°29⬘59.4⬙E, 9–16.7 m, R. Winterbottom, W. Holleman, B. Hubley, A. Bauman and D.
Winterbottom, 4 June 2004. ROM 1773CS (ex-USNM
223158, specimens cleared and stained), 10 (13.2–17.8),
Federated States of Micronesia, Pohnpei, reef just S of
Nanmatol Island. (6°59⬘30⬙N, 158°15⬘45⬙E), 0–13.7 m;
V.G. Springer et al., 6 Sep. 1980.
Non-type material 532 specimens. Caroline Islands: BPBM 28235 (3), BPBM 31378 (23), CAS 56595
(3), CAS 59890 (4), CAS 59884 (1), CAS 59886 (1),
CAS 59891 (3), CAS 59894 (3), CAS 59895 (2), CAS
59896 (5), CAS 59897 (1), CAS 59903 (3), CAS 59904
(1), CAS 59905 (4), CAS 59906 (8), CAS 59907 (1),
CAS 59909 (10), CAS 59925 (5), CAS 59926 (4), CAS
59927 (2), CAS 59928 (10), CAS 59930 (1), CAS 59932
(7), CAS 59933 (3), CAS 59936 (1), CAS 60047 (21),
CAS 60097 (10), CAS 60244 (1), CAS 60285 (1), USNM
223158 (131), USNM 233212 (78), USNM 233298 (1).
Indonesia: BPBM 30157 (2), USNM 209982 (3), USNM
263355 (13), USNM 264580 (3), USNM 294015 (2),
USNM 298750 (4), USNM 298790 (1). Japan (Nansei
Islands): YCM-P26273 (17, as for holotype), YCMP26442 (14), YCM-P 28012 (4), YCM-P 28438 (3),
Fig. 10. Trimma hayashii, live, Amami-oshima
Island, Aquarium. Photo by T. Shiratori
YCM-P 28559 (11), YCM-P 29337 (16), YCM-P 29493
(2), YCM-P 29551 (3), YCM-P 29552 (2), YCM-P36054
(1),YCM-P 38267 (1), YCM-P 41189 (5), YCM-P 41496
(1), YCM-P 42485 (2) and YCM-P 42548 (5, data as for
holotype). Marshall Islands: BPBM 29257 (4). Papua
New Guinea: BPBM 36888 (1), CAS 65406 (8), USNM
258774 (2), USNM 263465 (4), USNM 264529 (1),
USNM 264707 (3), USNM 297263 (2), WAM P.27825065 (1), WAM P.27827-043 (4). Philippines: AMS
I.24405-001 (4), NSMT-P72571 (3), USNM 244157 (1),
USNM 263463 (11), USNM 263470 (2), USNM 263557
(5), USNM 264678 (2), USNM 264709 (1), USNM
263557 (4). Solomon Islands: NTM S.12711-027 (1),
USNM 296031 (1).
Diagnosis. Trimma hayashii is characterized
by a little elongated or no elongated dorsal spine;
the presence of a basal membrane between the
pelvic fins; the absence of predorsal scales; an interorbital but no postorbital trench; absence of
cheek and opercular scales; fifth pelvic fin ray
branched sequentially once or twice and, uniquely among known species of Trimma, a bilateral,
pupil-sized black spot on the branchiostegal
membrane. When alive, T. hayashii has red spots
which are a little smaller than the pupil diameter
on the head and nape, the posterior half of body
has 3 yellow stripes separated by 2 grey stripes,
and the black spots on the branchiostegal membranes are ocellated with white or blue (Figs. 8,
10).
Description. The description is based on the
holotype and up to 100 addition specimens. Dorsal fins VI-I, 8–9 (mean⫽8.5, n⫽98), second or
third spine longest and vriably elongate that extending posteriorly to between the base of second
to fourth dorsal fin rays when adpressed, with the
Two New Trimma from Western Pacific
third spine reaching as far as the fourth element
of the second dorsal fin, all fin rays branched;
anal fin I, 7–8 (mean⫽7.9, n⫽97), first ray
branched or unbranched, the others branched;
pectoral fin 14–17 (mean⫽15.8, n⫽96) with 2–9
(6 in holotype) branched rays in the approximate
center of the fin, the longest ray reaching posteriorly to a vertical line with the first few elements
of the anal fin; pelvic fin I, 5, no frenum, basal
membrane 50–100% the length of the fifth ray
(variation may be due to damage to this delicate
structure), first 4 rays with 1 sequential branch,
fifth ray unbranched or more often with 1 sequential branch and 60–90% the length of the
fourth, fourth ray reaching posteriorly to a verti-
171
Fig. 11. Trimma hayashii, composite dorsal view
of head of cyanine-blue stained specimen, 20.1
mm SL female paratype, Palau, ROM 76806.
Photo and image enhancement by R. Winterbottom
cal line with the first to second element of the
anal fin. Lateral scales 22–26 (24 in holotype,
mean⫽23.2, n⫽97), anterior transverse scales
7–10 (mean⫽9.2, n⫽97), posterior transverse
scales 7–8, (mean⫽7.7, n⫽97); predorsal without scales in the midline except in some large
specimens (⬎20 mm SL, Palau), in which there
may be 1 or 2 cycloid scales in the mid-region of
the nape; scales on breast cycloid, 2–3 cycloid
scales on pectoral base, scales on the side of the
head extend anteriorly to above the vertical limb
of the preopercle, cheek and opercle scaleless.
Gill opening to below mid pupil. Upper jaw with
an outer row of widely spaced, enlarged, curved
canines followed by a few irregular rows of small
conical teeth, the innermost row enlarged and directed medially. Lower jaw with an outer row of
widely spaced, enlarged, curved canines extending to the bend of the dentary, 1–2 middle rows
of small, closely spaced, conical teeth, and an innermost row of widely spaced, enlarged canines
about half as high as the outer row and directed
medially. Tongue round to roundly truncate. Gill
rakers on first arch 2–5⫹14–18⫽17–22, (holotype not counted, mean⫽19.0, n⫽30). Nasal apparatus a slightly elevated sac confined to the anterior half of the snout, anterior nasal opening a
tube, posterior nasal opening a pore with a raised
rim. Positions of sensory papillae are showed by
Figure 13.
Head length 25.9–33.3% (30.9% in holotype,
mean 30.3%, n⫽72) of standard length, body
Fig. 12. Trimma hayashii, composite ventral view
of head of cyanine-blue stained specimen, 20.1
mm SL female paratype, Palau, ROM 76806.
Photo and image enhancement by R. Winterbottom
Fig. 13. Trimma hayashii, diagrammatic figure of
sensory papillae. Drawn by K. Hagiwara
172
K. Hagiwara and R. Winterbotton
depth 20.8–27.5% (25.3% in holotype, mean
24.1%, n⫽70), body width 13.5–18.9% (15.6%
in holotype, mean 15.9%, n⫽72). Snout length
18.6–30.7% (mean 24.5%, n⫽68) of head length,
orbit diameter 29.5–43.3% (38.5% in holotype,
mean 35.1%, n⫽72), upper jaw length 38.043.3% (mean 35.1%, n⫽72). Bony interorbital
1/4–1/5 pupil width, with a interorbital trench
and no postorbital trench (Fig. 11). Vertebrae
10⫹16⫽26, hemal arches not expanded.
Color pattern (from slides of live and freshly
collected specimens from Nansei Islands, Indonesia, Palau, Marshall Islands, and Taiwan,
Fig. 8). Freshly collected specimens have a red to
reddish orange background, with pupil-sized or
slightly smaller, darker red spots on the head and
nape, and 3 yellow stripes separated by grey
stripes on the body. The uppermost yellow stripe
begins at the origin of the second scale row
below the first dorsal fin. The middle stripe follows the mid lateral septum and begins roughly
below the last elements of the first dorsal fin. The
ventralmost stripe begins just anterior to the anus
and passes posteriorly along the ventral portion
of the body. All 3 stripes extend onto the caudal
fin. An intense black spot, roughly pupil diameter
in size, with a white (or blue in life) ocellus is
present bilaterally of the branchiostegal membranes. Melanophores are sprinkled over body,
most intensely on the head. Scale pockets are diffusely outlined with melanophores, more so dorsally than ventrally. The dorsal fins have 2 orange
stripes, the upper stripe at the center of fin elements is diffuse, poorly developed, and about
one-third the length of fin rays in width, the
lower stripe is well defined, about one third
pupil-diameter in width, and situated distal to the
bases of the fin elements by about the same
width. A similar but more orange basal stripe is
present in the anal fin which may be expanded
distally over half the height of the fin, and is
sometimes separated into 2 stripes. Pectoral and
pelvic fins are light red or orange without pattern. The caudal fin rays may be margined with
yellow and orange, and the iris is red with scatterings of melanophores and/or dark chro-
matophores.
Color in alcohol. Body straw-yellow with
sprinkled chromatophores, with heavier concentrations on anterior sections of body. Scale pattern on dorsal half of body is diffusely outlined
with chromatophores and melanophores. The red
spots on the head and nape are pale, the ocellus
of the black spot on the branchiostegals is not
visible, and the spot itself is somewhat diffuse
around the edges (Fig. 12). The stripes on the
body are indistinct. The anal fin has a basal stripe
and is darker at the distal tip, with sparse
melanophores between them. The caudal fin is
translucent with melanophores along some fin
ray, and melanophores are sparsely present in
pectoral and pelvic fins. All observed specimens
have a simple black iris.
Affinities. Trimma fangi Winterbottom and
Chen, 2004 is very similar to T. hayashii sp.nov.,
but lacks the unique (among members of the
genus) dark branchiostegal spot. Trimma bisella
Winterbottom, 2000 and Trimma woutsi Winterbottom, 2002 are similar to T. hayashii in the
presence of an interorbital trench present, no postorbital trench, dorsal spines often elongated, and
in the number of predorsal, lateral and transverse
scales. Trimma bisella has no pelvic fin basal
membrane (vs. present). Trimma woutsi has 9
anal fin rays (vs. 7–8 rays), 1–3⫹11–12⫽13–15
gill rakers of first gill arch (vs. 2–5⫹14–18⫽
17–22). Trimma preclarum Winterbottom, 2006
also has 3 yellow stripes separated by grey
stripes on the posterior half of the body, similar
to those of T. hayashii. But Trimma preclarum is
distinguished from T. hayashii by the presence of
5–9 scales in the predorsal midline, 4 yellow
spots on the iris, and lack of an ocellated black
spot on the branchiostegal membrane and of red
spots on anterior half of body.
Habitat. Trimma hayashii lives under corals
in embayments, which are sometimes covered by
siltation. It often positions itself upside-down on
the roof of the recess.
Distribution. Trimma hayashii has been collected at Japan, the Philippines, Palau, Micronesia, Indonesia, Papua New Guinea and the
Two New Trimma from Western Pacific
Solomon Islands.
Etymology. Trimma hayashii is named for
Mr. Masayoshi Hayashi. It has been referred to
by Hagiwara and Hayashi (1997) as an undescribed species, and Mr. M. Hayashi provided a
photograph of holotype for this paper.
Comparative materials. Trimma tevegae (74 specimens). Japan (Nansei Islands): YCM-P4473, 1 (22.1
mm SL). YCM-P4768, 1 (19.2). YCM-P6034, 3 (18.3–
19.0). YCM-7534, 3 (14.7–17.1). YCM-P7564, 1 (18.2).
YCM-P10227, 1 (15.4). YCM-P24653, 1 (17.9). YCMP26391, 1 (28.3). YCM-P26436, 3 (20.0–21.1). YCMP28135, 9 (14.9–27.2). YCM-P28318, 9 (9.9–27.4).
YCM-P28603, 1 (26.5). YCM-P29044, 12 (12.5–28.3).
YCM-P29153, 1 (25.1). YCM-P29308, 3 (26.0–29.1).
YCM-P29434, 4 (25.2–32.0). YCM-P34048, 7 (12.0–
27.8). YCM-P34059, 1 (30.3). YCM-P34369, 6 (15.5–
27.5). YCM-P38336, 1 (29.3). YCM-P38463, 1 (22.0).
YCM-P38502, 2 (19.5–20.7). YCM-P39049, 1 (21.1).
Malaysia: YCM-P36862, 1 (23.1). See also material in
Winterbottom (2005b).
Acknowledgments
We thank Keiichi Matsuura, Koichi Sibukawa
(NSMT), Yuji Ikeda (BLIH) and members of
SMBC for provided materials, Masayoshi
Hayashi and Taketomo Shiratori for provided the
photographs. And we would like to thank Marg
Zur for data collation, and Doug Hoese for freely
providing many details of the distributions of the
new species. Support for fieldwork from the
Members Volunteer Committee through the
ROM Foundation is gratefully acknowledged.
Support for the research was provided by the
people of Canada through NSERC Discovery
Grant A 7619 and the ROM.
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Manuscript received 8 February 2006; revised 23 October
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Associate editor: S. Kimura.