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284
Accepted by E. Hilton: 17 Jun. 2015 published: 10 Jul. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3985 (2): 284
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290
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Article
http://dx.doi.org/10.11646/zootaxa.3985.2.7
http://zoobank.org/urn:lsid:zoobank.org:pub:A275D0B9-F3C7-4BA4-B82E-E0B611F373C8
Garra robertsi, a new cyprinid (Cypriniformes: Cyprinidae)
fish species from Borneo
RYAN J. THONI
1
& RICHARD L. MAYDEN
Department of Biology, Saint Louis University, St. Louis, Missouri 63103 USA
1
Corresponding author. E-mail: rthoni@slu.edu
Abstract
Garra robertsi is described from specimens collected from the Sungai Bongan and Tempassuk rivers in Sabah, Borneo.
The species is differentiated from G. borneensis, its only congener on the island of Borneo, in having five (versus four)
transverse scale rows above lateral line, the first branched dorsal-fin ray extending beyond the posterior-most extent of
any other part of the dorsal fin when depressed (versus not extending posteriorly beyond last ray when depressed), breast
with deeply embedded scales (versus exposed scales), fewer tubercles on snout, thin (versus thick) anteromedial fold on
the lower lip, absence (versus presence) of a lateral stripe, absence (versus presence) of a stark, contrasting black stripe
on lower caudal-fin rays, and other pigmentation characteristics.
Key words: Garrinae, Malaysia, Sungai Bongan River, Tempassuk River
Introduction
Fishes of the genus Garra are diverse, widely distributed, and are of economic and cultural importance. There are
more than 100 species throughout its trans-Afro-Asian range (Eschmeyer 2015), with the species diversity being
the greatest in south and Southeast Asia (Kottelat 2013). Despite being a source of immense biodiversity, Borneo is
disproportionately depauperate with regard to species of Garra. Only one species has been recognized, G.
borneensis (Vaillant). There are no other species that have been synonymized with G. borneensis, suggesting that
variation in Garra throughout Borneo has remained largely unstudied and G. borneensis has been treated as a name
used for all Garra encountered on the island. While examining specimens of G. borneensis collected throughout
the island, a unique and distinct form was found from several lots collected at localities in the northern river
drainages in Sabah. One locality referred to as Marak Parak, likely refers to a road named Marak Parak which
parallels the Sungai Bongan River in Kota Manudu Sabah, has several lots collected by R.B. Stuebing in October,
1988. Additionally, P. K. Chin collected a single lot of this form in February 1950 from the Tempassuk River. This
species is described herein as Garra robertsi.
Methods
Museum abbreviations follow Sabaj Perez (2014). Measurements were taken using digital calipers to the nearest
0.1 mm. All measurements follow Kullander and Fang (2004) and Nebeshwar et al. (2009). Meristic data used in
this analysis include the following: lateral-line scale rows were taken from the first pored scale to the posterior-
most scale on the hypural plate; predorsal scale rows were counted from the first scale at occiput to the posterior-
most predorsal scale at dorsal-fin origin; transverse scale rows above lateral line were those horizontal scale rows
beginning at dorsal-fin origin and continuing ventrally to include the scale row above lateral line; transverse scale
rows below lateral line were those one horizontal scale row beneath the lateral line to the vent; circumpeduncular
scale rows above the lateral line included scale rows from the left lateral line scale row dorsally to, and including,
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GARRA ROBERTSI, A NEW CYPRINID FISH FROM BORNEO
right lateral-line scale; circumpeduncular scale rows below lateral line included the first scale row below the left
lateral line, ventrally around to first scale below right lateral; dorsal-fin rays and anal-fin rays excluded any
“spinelet” or incomplete primary rays; caudal-fin rays included both branched and principle rays; pectoral-fin rays
and pelvic-fin rays were broken down between branched and unbranched rays.
Garra robertsi sp. nov.
(Fig. 1)
Garra borneensis (Vaillant) 1902, originally Discognathus borneensis
Holotype. FMNH 99403, 70.4 mm SL; Malaysia: Sabah: Kota Marudu, Marak Parak, Sungai Bongan River.
Collected by R. B. Stuebing, October 1988.
Paratypes: FMNH 126067, 3 specimens; UF 237241, 1 specimen; same data as holotype.
Diagnosis. Garra robertsi is distinguished from its only other Bornean congener, G. borneensis (Fig. 2), by the
following characters: five (versus four) transverse scale rows above lateral line, first dorsal-fin ray extending
beyond the posterior-most extent of any other part of the dorsal fin when depressed (versus not extending
posteriorly beyond last ray when depressed), deeply embedded scales (versus exposed scales) on the breast,
absence (versus presence) of a stark, contrasting black stripe on lower caudal-fin rays, two nearly continuous
patches (versus 3 discrete patches) of tubercles on snout, absence (versus presence) of a lateral stripe, presence
(versus absence) of broad suborbital bar, a thickened anteromedial fold on the lower lip, and fewer papillae
extending onto the callous pad from the posterior flap of the lower lip.
FIGURE 1. Dorsal, lateral and ventral views of Garra robertsi holotype. FMNH 99403.
THONI & MAYDEN
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FIGURE 2. Lateral comparison of Garra robertsi and G. borneensis. a) G. r o b e r t s i holotype, 70.4 mm SL; b) G. borneensis
FMNH 99300, 73.3 mm SL; c) G. borneensis holotype, RMNH.PISC.7698, 71.7 mm SL; d) drawing of G. borneensis holotype
taken from Vaillant (1902).
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GARRA ROBERTSI, A NEW CYPRINID FISH FROM BORNEO
FIGURE 3. Structural comparisons of mouth and disc of Garra borneensis and G robertsi. a) G. borneensis FMNH 94199,
81.9 mm SL; b) G. ro b ert s i holotype, 70.4 mm SL.
Description. Morphometric and meristic data are provided in Tables 1 and 2, respectively. The general
physical appearance and morphology of the species is presented in Figure 1. Body elongate; subcylindrical to
triangular in cross-section anteriorly, becoming laterally compressed posteriorly, with the caudal peduncle width
less than one half its depth. Snout sloping downward from nares to anterior-most point; small to large tubercles,
depending on breeding condition and sex, on rostral and lateral lobes of snout. Shallow transverse groove present
and proboscis absent; slight hump may be present immediately anterior to nares, hump becoming more prominent
in larger individuals. Dorsal profile rises steadily to dorsal-fin origin; slightly convex. Dorsal profile from dorsal-
fin origin to caudal base sloping downward, without curvature, to a level equal with dorsal edge of orbit. Entire
ventral surface flat except for a slight convex rise at caudal peduncle.
Head acute when viewed laterally; rounded from dorsal and ventral view. Head length (HL) about one fourth
of standard length (SL). Head width roughly 2/3 HL. Orbit roughly 1/5 of HL. Mouth with mental disc; disc
slightly less than 1/3 HL; disc roughly 2/3 as long as wide. Rostral cap with 22-27 fimbriae; posterior edge of cap
folds at connection with lower lip, creating a lobe with papillae at the widest point of mental disc; lobe may not be
homologous with anterolateral lobe of other Garra species. Rostral cap covers upper jaw, often leaving lower jaw
visible. Two pairs of barbels; rostral barbels two to three times the length of maxillary barbels.
Tubercles erect; appearing to be present on both males and females, but are considerably more conspicuous in
breeding males, occurring in two main patches with little delimitation between them. Rostral lobe with two
parallel, horizontal rows of small tubercles; no more than 8 tubercles per row. Tubercles on lateral lobe in circular
pattern and occurring immediately posterior and lateral to rostral lobe with only a small depression dividing the
tuberculate areas. In the most tuberculate males, some individuals may have one or two small tubercles
immediately posterior to nares.
Dorsal-fin origin slightly nearer to tip of snout than to caudal-fin base; margin of fin concave; first branched
ray the longest, slightly longer than HL, and extending beyond any other part of the depressed; rays ii8. Anal-fin
origin beginning at vertical to posterior extent of depressed dorsal fin; fin length about 1/4 to 1/5 SL; rays i5. Paired
fins inserted horizontally. Pectoral fins slightly shorter than head length; rays i13 (2 specimens), i14 (12), i15 (1).
Pelvic fins nearly equal in length to pectoral fins. Pelvic-fin rays ii7. Caudal fin forked; lobes roughly equal in
length; total rays 19, branched rays 17.
Preserved coloration. Dorsum of head and body uniformly brown; body darker than head. Head light brown,
transitioning to dark brown anterior to nares. Dark brown extending anteriorly to nares and grooves surrounding
tubercle patches where dark brown to black lines encircle nares, and is present in grooves and snout. Side of head
light brown. Broad, dark brown and anteroventrally angled suborbital bar present. Rostral lobe and dorsolateral
surface of rostral flap brown. Maxillary and rostral barbels brown to near tips.
Dorsal scales dark brown with no distinct pattern. Lateral-line scales and first rows above and below lateral
line brown and with distinct dark vertical bar in the posterior field. Bars on scales are very distinct to vertical near
anal fin; scales posterior to vertical of anal-fin origin have distinct dark margins creating crosshatched pattern.
Scales below horizontal to pectoral-fin insertion lightly pigmented to immaculate; venter of body around anal fin
and caudal peduncle immaculate.
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TABLE 1. Morphometric characters for Garra robertsi (n=15) expressed as rations of SL, HL, or vent-anal length.
Dorsal-fin membranes with light pigmentation basally and distally. Distal third of fin with distinct dark and
irregularly shaped streaks. Anal-fin rays with dark pigmentation along margins. First three anal fin membranes
with melanophores along posterior margin of rays. Remaining rays immaculate except for medial dark
pigmentation that together form medial band. Leading ray and first membranes of pelvic fin darkly pigmented from
min max mean sd
Standard Length (mm) 58.9 103.7 80.7 13.0
% standard length
body depth 18.2 26.1 20.8 1.7
head length 22.1 25.1 23.6 0.9
body width at dorsal 14.1 17.1 15.7 0.9
body width at anal 7.6 9.2 8.5 0.5
caudal peduncle width 3.2 4.5 3.9 0.5
caudal peduncle length 15.4 20.7 18.1 1.5
caudal peduncle depth 10.7 12.4 11.7 0.5
1st dorsal ray length 25.3 29.9 27.6 1.4
dorsal fin length 24.6 29.9 26.9 1.4
dorsal fin base length 13.6 16.6 14.9 0.9
pectoral fin length 20.4 22.5 21.6 0.7
pelvic fin length 20.1 24.9 22.0 1.2
anal fin length 18.2 25.3 20.5 1.8
anal fin base length 5.1 8.1 7.0 0.8
upper caudal fin lobe length 32.3 36.4 34.0 1.7
lower caudal fin lobe length 31.2 35.3 33.1 1.4
preanal-fin length 73.9 80.9 76.3 1.6
prevent length 60.9 67.4 63.4 1.7
prepelvic length 46.3 50.7 48.8 1.3
predorsal length 42.7 47.3 45.5 1.2
occiput-dorsal length 23.3 27.2 24.4 1.0
prepectoral length 21.1 23.2 21.9 0.7
pelvic-anal length 26.6 31.4 28.6 1.3
% pelvic-anal length
vent-anal length 40.1 49.4 46.5 2.9
% head length
head depth at occiput 59.3 67.5 62.2 2.4
head width 67.4 75.9 71.6 2.6
snout length 50.8 58.1 53.1 2.1
orbit diameter 19.1 25.8 21.6 2.3
interorbital width 41.4 45.2 42.8 1.1
disc length 25.4 33.8 29.6 2.1
disc width 46.8 66.3 53.9 5.9
callous pad length 18.6 24.7 21.5 1.7
callous pad width 28.6 40.4 33.6 3.4
rostral barbel length 16.5 23.2 19.4 2.2
maxillary barbel length 5.1 9.4 7.4 1.2
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GARRA ROBERTSI, A NEW CYPRINID FISH FROM BORNEO
base to margin. Remaining rays lightly pigmented; membranes posterior to ray 5 with melanophores concentrated
on basal 1/3. First five rays and membranes of pectoral fin dark with dense concentrations of melanophores;
remaining rays and membranes as in posterior-most rays of pelvic fin. Caudal-fin rays and membranes lightly
pigmented with no discernable bars or stripes; basal 1/3 of principal caudal ray and three lower-most branched rays
may be more dusky than other rays and membranes.
TABLE 2 . Frequency tables of meristic variables for Garra robertsi and G. borneensis. Values include the holotype for
both species. Holotype condition for G. borneensis signified with
*
; Holotype condition for G. robertsi signified with
+
.
Etymology. This species epithet “robertsi” is in honor of Dr. Tyson Roberts, a prominent ichthyologist with a
long-running focus on Bornean and Southeast Asian fishes.
Distribution. Garra robertsi is currently known to occur only in the Sungai Bongan and Tempassuk rivers in
northern Sabah, Malaysia. Due to its occurrence in separate drainages, it is likely present in other rivers in the
northern Borneo.
Discussion
A comparison of the lateral external physical differences between G. robertsi and G. borneensis can be seen in
Figure 2. Garra robertsi was examined from several lots previously identified as G. borneensis from Marak Parak,
Kota Manudu, Sabah, and the Tempassuk River in Sabah. All specimens are consistently diagnosed from Vaillant’s
(1902) type specimen (Fig. 2c), original description, and illustration of G. borneensis (Fig. 2d). Vaillant’s
illustration and type specimen both have dark bands on the upper and lower rays of the caudal fin, four (versus
five) transverse scale rows above the lateral line, and scales present on the belly and breast. Additionally, on
Vaillant’s type specimen and all other comparative materials of G. borneensis examined, the first dorsal-fin ray
never extended beyond the last dorsal ray when depressed.
Head morphology between the two species differs as well. Differences exist in the tuberculation of the head
and the size and shape of the rostral lobe. The tubercles of G. borneensis are found in three conspicuous patches on
the rostral lobe, the lateral lobe and the inner narial region. Tubercles on the rostral lobe of G. borneensis are
aligned in 3 rows: the dorsal-most row havening 4-6 large tubercles; center row with 4-6 large tubercles and one
laterally-projecting tubercle at the lateral terminus of each side of rostral lobe; and ventral-most row consisting of
several smaller tubercles (6-10). Tubercles on G. robertsi are consistently arranged in two patches; highly
tuberculate individuals may have a faint third patch. The most common and conspicuous patch is on the lateral
lobes of the snout, where there is a small circular patch. The tubercles are small, erect, and sparse. The second
Transverse scale rows above lateral line 5 6
Garra robertsi 15
+
Garra borneensis 10
*
Lateral-line scale rows 27 28 29 30
Garra robertsi 2 5 8
+
Garra borneensis 1
*
63
Predorsal-scale rows 9 10 11
Garra robertsi 6
+
9
Garra borneensis 4 6
*
Branched pectoral-fin Rays 12 13 14 15
Garra robertsi 2 12
+
1
Garra borneensis 2 5
*
3
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tuberculate area is demarked by a thin, sparse row of small tubercles along the rostral lobe. In a few of the most
tuberculate males there is a very small patch consisting of only 1-3 tubercles immediately anteromedial to the
nares. Upon examination of photographs of the specimens after our analysis, we noticed that there is a notable
difference in the thickness of the anteromedial fold on the mental disc; the fold being much thicker and square-
shaped in G. borneensis than in G. robertsi (Fig. 3)
Garra is one of the most diverse genera of freshwater fishes in Asia. It is rather surprising the genus has been
represented by only one species on the hyperdiverse island of Borneo. This is a doubtful assumption, and the
finding of G. robertsi, a clearly distinct species previously identified as G. borneensis and undetected for more than
25 years, is a testament to such doubt. It is likely that there are more species of Garra present in the hillstreams of
Borneo.
Comparative materials
Garra borneensis: RMNH.PISC.7698: holotype: Bloeoe River Borneo, collected by A.W. Nieuwunhuis, August
1896. FMNH94199: Sungai Pinoh, Sungau Tebelian, Borneo, Kalimantan, Indonesia. Collected July 1976, by
Tyson Roberts. FMNH 99300: 1 specimen, Palum Tambun, Danum Conservation area, Borneo, Indonesia.
Collected October 21, 1986 by Emerson and Inger. FMNH 68505: 1 specimen, Deramakot Camp station,
Kinabatangan District, Borneo, Kalimantan, Indonesia. Collected May 2, 1956 by P.K. Chin. Garra robertsi:
FMNH 99378: 1 specimen, Marak Parak, Kota Marudu, Sabah, Malaysia. Collected October 1988, by R.B.
Stuebing. FMNH 99395: 2 specimens, Marak Parak, Kota Marudu, Sabah, Malaysia. Collected October 1988, by
R.B. Stuebing. FMNH 9940: 5 specimens, Marak Parak, Kota Marudu, Sabah, Malaysia. Collected October 1988,
by R.B. Stuebing. FMNH 99384: 1 specimen, Marak Parak, Kota Marudu, Sabah, Malaysia. Collected October
1988, by R.B. Stuebing. FMNH 99409: 1 Specimen, Marak Parak, Kota Marudu, Sabah, Malaysia. Collected
October 1988, by R.B. Stuebing. FMNH 44788: 10 specimens, Tempassuk River, Kota Belud, Sabah, Malaysia.
Acknowledgments
The authors express gratitude to Caleb MacMahen and others at the FMNH Fishes Division for accommodations in
the laboratory while examining these specimens, and to Ronald DeRuiter at RMNH, Leyden, for accommodating a
last minute visit to the museum to examine the holotype specimen, and for providing pictures of the G. borneensis
holotype. This research was funded by USA NSF DEB-1021840.
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