On the paraphyly of Homaloptera (Teleostei: Balitoridae) and description of a new genus of hillstream loaches from the Western Ghats of India

Abstract

Homaloptera van Hasselt 1823 as treated historically exhibits substantial morphological diversity and is paraphyletic based on both morphological and molecular data. The morphological diversity and phylogenetic relationships of Homaloptera, Homalopteroides Fowler 1905, Homalopterula Fowler 1940, and Balitoropsis Smith 1945, are elucidated. Pseudohomaloptera Silas 1953 is removed from the synonymy of Homaloptera. Homalopteroidini is created for the monophyly of Homalopteroides and Homalopterula; it is the sister group to balitorini Swainson 1839. Ghatsa n. gen. is created for species previously assigned to Homaloptera from the Western Ghats of India, and a redescription of Ghatsa montana (Herre 1945) is provided.

Full text

Accepted by R. Pethiyagoda: 31 Dec. 2014; published: 4 Mar. 2015
ZOOTAXA
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Copyright © 2015 Magnolia Press
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http://dx.doi.org/10.11646/zootaxa.3926.1.2
http://zoobank.org/urn:lsid:zoobank.org:pub:20666BE9-1457-41A6-9727-AC0077203595
On the paraphyly of Homaloptera (Teleostei: Balitoridae) and description of
a new genus of hillstream loaches from the Western Ghats of India
ZACHARY S. RANDALL & LAWRENCE M. PAGE
Florida Museum of Natural History, University of Florida, Dickinson Hall, Gainesville, FL 32611, USA.
E-mails: zrandall@flmnh.ufl.edu, lpage1@ufl.edu
Abstract
Homaloptera van Hasselt 1823 as treated historically exhibits substantial morphological diversity and is paraphyletic
based on both morphological and molecular data. The morphological diversity and phylogenetic relationships of
Homaloptera, Homalopteroides Fowler 1905, Homalopterula Fowler 1940, and Balitoropsis Smith 1945, are elucidated.
Pseudohomaloptera Silas 1953 is removed from the synonymy of Homaloptera. Homalopteroidini is created for the
monophyly of Homalopteroides and Homalopterula; it is the sister group to balitorini Swainson 1839. Ghatsa n. gen. is
created for species previously assigned to Homaloptera from the Western Ghats of India, and a redescription of Ghatsa
montana (Herre 1945) is provided.
Key words: Ghatsa, Balitoropsis, Homalopterula, Pseudohomaloptera, Homalopteroides, Helgia, Chopraia, Loaches,
Southeast Asia
Introduction
The limited information on phylogenetic relationships of hillstream loaches has led to inconsistent and often
transient recognition of genera. Relationships among species historically assigned to Homaloptera van Hasselt
1823 have been particularly problematic, as noted by Fang (1930), Hora (1932), Kottelat (1998), Tan & Ng (2005),
Tan (2009), and Randall & Page (2012). Several names historically synonymized with Homaloptera (Helgia
Vinciguerra 1890, Homalopteroides Fowler 1905, Chopraia Prashad & Mukerji 1929, Homalopterula Fowler
1940, Balitoropsis Smith 1945, and Pseudohomaloptera Silas 1953) have been recognized as genera or junior
synonyms in recent years (Randall 2012; Kottelat 2012; Kottelat 2013). Some of these recent classifications are
without supporting data or diagnoses, which only adds to the confusion of balitorid classification. The objectives of
this study were to test the most recent classifications (Randall 2012; Kottelat 2012; Kottelat 2013) by analyzing
genetic and morphological data (including type species when available) to identify clades and to diagnose well-
supported clades as genera using morphological criteria.
Material and methods
Morphological. Measurements follow Hubbs & Lagler (2004) or Kottelat (1984) (see Randall & Page 2012 for
measurements from each source), and counts follow Randall & Page (2014). The definition of a rostral cap follows
Roberts (1982). A central furrow refers to an indentation on the ventral surface of the head at the branchiostegal
membrane just anterior to the isthmus. The terms origin and insertion refer, respectively, to the anterior and
posterior ends of fin bases for paired and unpaired fins. Total lateral-line scale count includes scales on the caudal
fin. Counts are given as ranges where taxa are distinguished by the mode of that range (M). Lengths were measured
to the nearest 0.1 mm using digital calipers and taken on the left side when possible. All measurements are given in
millimeters (mm). Head length and measurements of the body are given as proportions of standard length (SL).
Measurements of the head are presented as proportions of head length (HL).

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Institutional abbreviations follow Sabaj Pérez (2012). A ‘(?)’ represents a lack of data or an uncertain locality.
Photographs of preserved specimens were taken using a Visionary Digital system (Palmyra, Virginia) with Canon
40D and 5D cameras at the Florida Museum of Natural History and edited using Photoshop CS3. When
coordinates for localities were unavailable, they were estimated using maps and GEOlocate. Maps were
constructed using ArcMap Version 9.3.1 in ArcGIS 9th edition.
Three-hundred and three individuals were examined in this study representing all but four species of
Homaloptera (sensu lato) (as recognized by Kottelat 1998) from the Western Ghats of India. The taxonomic status
of these species from the Western Ghats of India was assessed using only original descriptions, due to the
inaccessibility of specimens.
Molecular. The supraspecific relationships of balitorids were reconstructed using the nuclear recombination
activating gene 1 (RAG1) based on recently collected tissues (Table 1) and sequences available on Genbank (Table
2). Total genomic DNA was extracted from fin clips or muscle tissue preserved in 95% ethanol using a DNeasy
Tissue Kit (Qiagen) or with a 5% Chelex solution and 3 µl of proteinase K with an overnight digestion. RAG1 was
amplified and sequenced using the primers RAG-1F: 5’–AGC TGT AGT CAG TAY CAC AAR ATG–3’
(Quenouille et al. 2004) and RAG-RV1: 5’–TCC TGR AAG ATY TTG TAG AA–3’ (Ŝlechtová et al. 2007).
Individual samples for the polymerase chain reaction (PCR) consisted of a total 25 µl reaction containing 16.5 µl of
sterile water, 2 µl of genomic DNA, 4 µl of MyTaq Reaction Buffer (BioLine USA, Boston, MA), 0.5 µl of MyTaq
DNA Polymerase (BioLine USA, Boston, MA), and 1 µl of 10 µm of each primer. The PCR cycling parameters for
RAG1 followed Ŝlechtová et al. (2007). PCR cleaning with ExoSAP-IT and sequencing took place at the
Interdisciplinary Center for Biotechnology Research (ICBR), University of Florida, Gainesville. Geneious version
7.1.4 (Kearse et al. 2012) was used to view chromatograms, assemble consensus sequences, and generate a final
alignment using the Geneious global alignment tool which was corrected manually. All sequences produced from
this study are available on Genbank (Accession numbers given in Table 1).
A Bayesian inference (BI) analysis using MrBayes 3.2.2 (Ronquist et al. 2012) and a Maximum Likelihood
(ML) analysis using RAxML 7.2 (Stamatakis 2014) were performed through the Cyberinfrastructure for
Phylogenetic Research (CIPRES) (Miller, Pfeiffer, & Schwartz 2010). In both analyses the cyprinid Cyprinus
carpio was used to root the tree.
For the BI analysis, SYM+I+G was the best fit substitution model of nucleotide evolution, using the Akaike
Information Criterion with JModeltest version 2.1.5 (Darriba et al. 2012). Two Markov chain Monte Carlo
(MCMC) analyses with four simultaneous chains were run for 5 million generations sampling every 1,000
generations resulting in a 10,002 sampled trees. The standard deviation for split frequencies was 0.002942 and the
average potential scale reduction factor for all parameters was 1.015, indicating clear convergence of the two runs.
A total of 25% of the first sampled trees were discarded as burn-in, sampling 7,502 trees. The best tree was viewed
in Figtree v 1.3.1 (http://tree.bio.ed.ac.uk/software/figtree/).
For the ML analysis, GTR+I+G was used as the substitution model of nucleotide evolution, being the best fit
substitution model using the Akaike Information Criterion with JModeltest version 2.1.5 (Darriba et al. 2012)
available for use in RAxML. The –f a option of RAxML was used to generate 1,000 bootstrap replicates
(Stamatakis, Hoover, & Rougemont 2008) followed by a search of the best scoring ML tree. The best tree was
viewed as in the BI analysis.
Bayes factor (BF), Shimodaira-Hasegawa (SH) and Approximately unbiased test (AU) were performed to
determine if various positive and negative topological constraints were significantly different from the optimal
topology (Table 3). It was not possible to implement negative constraints in the ML analysis. Homaloptera (sensu
lato) was constrained as monophyletic to include all species of Homaloptera, Homalopteroides, Balitoropsis,
Pseudohomaloptera, and Homalopterula. BF was used for the BI analysis while SH and AU tests for the ML
analysis using Paup 4.0 (Swofford 2003). The Bayes factor (2lnBfs) for competing models was evaluated using the
criteria of Kass & Raftery (1995) where a 2lnBfs of 0–2 is not worth more than a bare mention, a 2lnBfs of 2–6
provides positive support, a 2lnBfs of 6–10 provides strong support, and a 2lnBfs of > 10 provides very strong
support for the unconstrained best tree.

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TABLE 1. Samples produced from this study included in the phylogenetic analysis,
Genus Species Vial # Voucher Origin (region, province, river) Genbank accession #
Balitora meridionalis 332 ANSP 179834 Thailand, Kanchanaburi, Ulong KP322550
Balitoropsis ophiolepis 2006-0588 UF 166109 Sumatra, Lampung, Way Besai KP322540
zollingeri 2005-0948 UF 161715 Sumatra, Lampung, Way Rarem KP322535
2005-0962 UF 161715 Sumatra, Lampung, Way Rarem KP322537
Homaloptera bilineata DAN09-182.02 CAS 231723 Myanmar, Sagaing, Irrawaddy KP322549
confuzona 2007-1185 UF 169906 Thailand, Changwat, Khlong Sato KP322543
344 ANSP 178721 Thailand, Chanthaburi, shop KP322551
ocellata 2005-0955 UF 161719 Sumatra, Lampung, Way Rarem KP322536
2005-0980 UF 161605 Sumatra, Lampung, Way Abung KP322539
parclitella 336 ANSP 179982 Thailand, Surat Thani, Tapi KP322552
Homalopteroides wassinkii 2005-0975 UF 161619 Sumatra, Lampung, Way Seputhi KP322538
nebulosus 2012-0600 UF 235748 Malaysia, Kelantan, Kelantan KP322548
smithi 2012-0157 UF 182840 Thailand, Nakhon Si Thammarat, Pong KP322546
stephensoni E97 USNM 393671 Borneo, S Kalimantan, Aib KP322553
Homalopterula gymnogaster SN25 UF 185031 Sumatra, Sumatra Utara, Aek Tongguran KP322554
Pseudohomaloptera leonardi 2012-0597 UF 235746 Malaysia, Kelantan, Kelantan KP322547
2007-1071 UF 169909 Thailand, Chanthaburi, Khruv Wui KP322541
2007-1104 UF 169909 Thailand, Chanthaburi, Khruv Wui KP322542
sexmaculata 2010-0211 UF 177819 Thailand, Chiang Mai, Mae Tiaen KP322544
2011-0203 UF 181170 Thailand, Kanchanaburi, Kroeng Krawia KP322545

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TABLE 2. Genbank accession numbers and sources for samples included in the phylogenetic analysis.
TABLE 3. Bayes factor (2lnBF), Shimodaira-Hasegawa (SH) and Approximately Unbiased (AU) tests performed on
various positive (+) and negative (-) topological constraints. Bayes factor was evaluated using the criteria of Kass and
Raftery (1995). A bolded value represents a significantly better optimal topology.
Genus Species Accession # Source
Acantopsis choirorhynchos EU711139 Mayden et al. 2008
Balitora annamitica EF056359 Ŝlechtová et al. 2007
Balitoropsis zollingeri EF056388 Ŝlechtová et al. 2007
Cyprinus carpio EF458304 Mayden et al. 2007
Formosania fascicauda JN177173 Liu et al. 2012
Hemimyzon yaotanensis JN177053 Tang et al. 2012
Homaloptera parclitella EF056358 Ŝlechtová et al. 2007
Homalopteroides smithi EF056356 Ŝlechtová et al. 2007
Jinshaia abbreviata JN177051 Tang et al. 2012
sinensis JN177052 Tang et al. 2012
Lepturichthys dolichopterus JN177035 Tang et al. 2012
fimbriata JN177039 Tang et al. 2012
Pangio kuhlii EF056331 Ŝlechtová et al. 2007
Sewellia lineolata HM224068 Tang et al. 2010
Sinogastromyzon sichangensis JN177040 Tang et al. 2012
szechuanensis JN177055 Tang et al. 2012
hsiashiensis JN177054 Tang et al. 2012
Metahomaloptera omeiensis JN177041 Tang et al. 2012
Pseudogastromyzon changtingensis tungpeiensis JN177175 Liu et al. 2012
cheni JN177176 Liu et al. 2012
cheni EF056357 Ŝlechtová et al. 2007
fangi JN177177 Liu et al. 2012
Vanmanenia pingchowensis JN177174 Liu et al. 2012
Constraint BI ln score 2lnBF ML ln score SH AU
Optimal topology -5402.97 - -5577.940582 - -
Homaloptera (sensu lato) (+) -5412.18 18.42 -5591.793874
0.0449 0.0158
Homalopteroidini (-) -5417.85 29.66 ---
Balitorini (-) -5409.02 12.1 ---
Homalopteroides (-) -5411.32 16.7 ---
Balitoropsis (-) -5405.45 4.96 - - -
Homaloptera (sensu stricto) (-) -5431.55 57.16 ---
Pseudohomaloptera (-) -5404.87 3.8 - - -
(Balitoropsis (Homaloptera,
Pseudohomaloptera)) (-)
-5404.66 3.38 - - -
(Homaloptera, Pseudohomaloptera) (-) -5404.00 2.06 - - -

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Phylogenetic results
The optimal topology (i.e. the one with the greatest likelihood) was generated in the BI analysis (Table 3) and is
represented in Figure 1.
FIGURE 1. Balitorid phylogeny based on Bayesian analysis of RAG1 with support values indicated at the branch lengths (PP/
BS). Posterior probability values ≥ 0.95 and bootstrap support ≥ 90 are represented by an asterisk (*). Posterior probability and
bootstrap support less than 50 are represented by a hyphen (-).Type species are in bold. A) Balitoridae; B) Gastromyzontinae;
C) Balitorinae; D) Homalopteroidini; E) Balitorini.

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Homaloptera (sensu lato) is not resolved as monophyletic in either analysis. Homaloptera, Homalopteroides,
Balitoropsis, and Pseudohomaloptera formed monophyletic groups with the same interrelationships among them
and the single species of Homalopterula included in the analysis in both BI and ML analyses (Fig. 1). Balitoropsis,
Homaloptera and Pseudohomaloptera are more closely related to Balitora and its close relatives than they are to
Homalopteriodes and Homalopterula. Within Balitorinae two major clades are resolved: Homalopteroidini, newly
proposed tribe name (Fig. 1, D) (Homalopteroides, Homalopterula), and Balitorini Swainson 1839 (Fig. 1, E)
(Balitora, Lepturichthys, Sinogastromyzon, Hemimyzon, Metahomaloptera, Jinshaia, Balitoropsis,
Pseudohomaloptera, Homaloptera). Within Balitorini, two major clades are resolved: a strongly-supported clade
consisting of Balitora as the sister group to Lepturichthys, Sinogastromyzon, Hemimyzon, Metahomaloptera,
Jinshaia; and a weakly-supported clade consisting of Balitoropsis as the sister group to Homaloptera plus
Pseudohomaloptera. Species recognized in Balitoropsis by Kottelat (2012) (B. leonardi, B. sexmaculata, B.
stephensoni) are resolved in the genera Pseudohomaloptera (P. leonardi, P. sexmaculata) and Homalopteroides (H.
stephensoni).
Nearly all negative and positive constraints were significantly worse than the optimal topology (Table 3). The
constrained Homaloptera (sensu lato) Bayesian best tree had a Likelihood Harmonic mean of -5412.18, a 2lnBfs of
18.42 (very strong support for a paraphyletic Homaloptera [s.l.]). The SH and AU tests for the likelihood analysis
resulted in a constrained tree that was significantly worse (SH score: 0.0449; AU score: 0.0158). The negative
constrained Homalopteroides best tree resulted in Homalopterula gymnogaster being nested within
Homalopteroides. The Likelihood Harmonic mean was -5411.32, a 2lnBfs of 16.7 (very strong support for
Homalopteroides). The negative constrained clade containing Homaloptera and Pseudohomaloptera resulted in a
best tree with Pseudohomaloptera as the sister group to Balitoropsis. The Likelihood Harmonic mean was
-5404.00, a 2lnBfs of 2.06 (positive support for Homaloptera being the sister group to Pseudohomaloptera).
Based on consistent monophyly, clades recognized herein as genera are Homaloptera, Homalopteriodes,
Balitoropsis, Pseudohomaloptera, and Homalopterula. Homalopterula is represented in the phylogeny by only one
individual, but based on its long branch length and morphological diagnosibilty described below, it is recognized as
a genus. Although species of Balitoropsis and Pseudohomaloptera are most similar morphologically, they were not
found to form a monophyletic group in any of the unconstrained analyses of genetic data and are easily separated
morphologically as described below.
Homaloptera van Hasselt 1823
(Figures 2, 3A, 4D, 5A)
Homaloptera van Hasselt, 1823:133 (type species: Homaloptera ocellata van der Hoeven 1830, by subsequent monotypy in
van der Hoeven, 1830:211). Gender feminine.
Helgia Vinciguerra, 1890:328 (type species: Homaloptera bilineata Blyth, 1860:172, by subsequent designation by Jordan
1920:448). Gender feminine.
Remarks. Two species names were listed in Homaloptera by van Hasselt (1823), H. javanica and H. fasciata. Both
lacked a description or figure and are unavailable (nomina nuda) (Hora 1932; Kottelat 1987). The first species
described and figured under Homaloptera was H. ocellata van der Hoeven 1830 (see Roberts 1993:24 for 1830 as
publication date), making H. ocellata the type species of Homaloptera. Homaloptera bilineata Blyth 1860 was
placed in the genus Helgia Vinciguerra 1890 and subsequently designated as the type species of Helgia by Jordan
(1920). Helgia Vinciguerra 1890 is a synonym of Homaloptera van Hasselt 1823 (Hora 1932; Kottelat 1998).
Kottelat (1998) recognized Homaloptera ocellata van der Hoeven 1830, H. bilineata Blyth 1860, H. orthogoniata
Vaillant 1902, and H. ogilviei Alfred 1967 as possibly forming a clade (Homaloptera sensu stricto) based on the
following characters: “unique color patterns (having in common reddish tints and similar dark markings on the
head and the fins), a slightly compressed body, 56–65 lateral-line scales and the dorsal-fin origin situated in
advance of the pelvic-fin origin.” Based on this diagnosis and an elongated head and a lateral-line scale count of
61–77, Tan & Ng (2005) treated this same clade as a subgenus of Homaloptera. With the availability of more data
and analyses, the only character provided by Kottelat (1998) that seems to be apomorphic for Homaloptera (sensu
stricto) is color pattern.

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FIGURE 2. Dorsal, lateral, and ventral views of preserved Homaloptera ocellata, UF 161718, 64.1 mm SL, Way Seputhi,
Lampung Province, Sumatra, Indonesia.
Diagnosis. Distinguishing characters are given in Table 4 and shown in Figures 2, 3A, 4D and 5A.
Homaloptera is distinguished by the following combination of characters: reddish tints on fins in life (Fig. 3A);
dorsal-fin origin anterior to pelvic-fin origin; 7–8½, 8½ (M) branched dorsal-fin rays; 7 branched pelvic-fin rays;
forked caudal fin; keeled scales (Fig. 4D); 20–27 predorsal scales; 59–73 total lateral-line scales; no adipose keel
on caudal peduncle; anus closer to anal-fin origin than to pelvic-fin insertion; large rostral cap; 2 thick rostral
barbels in close proximity to each other; thick triangular upper lip (Fig. 5A); fleshy pad between lateral portions of
lower lip (Fig. 5A); absence of central furrow at isthmus.
Species included. Homaloptera ocellata van der Hoeven 1830, H. bilineata Blyth 1860, H. orthogoniata
Vaillant 1902, H. ogilviei Alfred 1967, H. confuzona Kottelat 2000, and H. parclitella Tan & Ng 2005. The type
localities for species of Homaloptera are shown in Figure 6.
Comparison. Homaloptera is distinguished from Homalopteroides by presence vs. absence of reddish tints on
fins in life, having a dorsal-fin origin anterior vs. posterior to pelvic-fin origin; 7–8½, 8½ (M) vs. 6–8½, 7½ (M)
branched dorsal-fin rays; 59–73 vs. 33–52 total lateral-line scales; large vs. small rostral cap; medial- and lateral-
rostral barbels in close proximity to one another vs. widely separated from one another at base; triangular vs.
crescentic upper lip; thick vs. thin upper lip; presence vs. absence of pad between lateral portions of lower lip; and
absence vs. presence of a central furrow at isthmus.
Homaloptera is distinguished from Homalopterula by presence of reddish tints on fins in life vs. without red
color; dorsal-fin origin anterior vs. posterior to pelvic-fin origin; 7–8½, 8½ (M) vs. 5½ and 7½, 7½ (M) branched
dorsal-fin rays; keeled vs. smooth scales; 20–27 vs. 28–56 predorsal scales; forked vs. truncated or deeply
emarginated caudal fin; absence vs. presence of adipose keel on caudal peduncle; large vs. small rostral cap;
medial- and lateral-rostral barbels in close proximity vs. widely separated at base; triangular vs. crescentic upper
lip; presence of fleshy pad vs. lobes between lateral portions of lower lip; and absence vs. presence of a central
furrow at isthmus.

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TABLE 4. Characters distinguishing Homaloptera, Homalopteroides, Homalopterula, Balitoropsis, Pseudohomaloptera and Ghatsa. Values are given as ranges; numbers in
parentheses represent modal values. For Ghatsa, data from original descriptions are included; values in parentheses are for G. montana (CAS-SU 39871).
Character Homaloptera Homalopteroides Homalopterula Balitoropsis Pseudohomaloptera Ghatsa (G. montana)
No. specimens examined 51 148 24 42 37 1
Reddish tints on fins in life Yes No No No No ?
Dorsal-fin origin relative to pelvic-
fin origin
Anterior Posterior Posterior Anterior or
above
Anterior or above Posterior (Posterior)
Branched dorsal-fin rays 7–8½ (8½) 6-8½ (7½) 5½, 7½ (7½) 8½ 8½ 7–9½ (7½)
Branched pelvic-fin rays 7 5–8 (7–8) 7 7–9 (8) 8–9 (8) 6–9 (7–8)
Caudal-fin shape Forked Forked Truncate/deeply
emarginated
Forked Forked Slightly emarginated-
emarginated (truncate)
Scales keeled Yes Yes/No No Yes Yes ? (No)
Predorsal scales 20–27 14–25 28–56 13–15 13–19 ? (ca. 53)
Total lateral-line scales 59–73 33–52 57–75 42–55 50–61 59–95 (ca. 101)
Adipose keel No No Yes No No ? (Yes)
Anus closer to pelvic-fin base or
anal fin
Anal fin Anal fin Anal fin Pelvic-fin base Anal fin Anal fin
Rostral cap Large Small Small Large Large Small
Rostral barbels separated at base No Yes Yes No No Yes
Upper lip shape/width Triangular/Thick Crescent/Thin Crescent/Thick Crescent/Thick Triangular and
crescent/Thick
Crescent/Thin
Large central furrow at isthmus No Yes Yes Yes/No No ?
Structure between portions of lower
lip (Pad/Lobes/None)
Pad None Lobes Pad Pad None

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Homaloptera is distinguished from Balitoropsis and Pseudohomaloptera by presence vs. absence of reddish
tints on fins in life; 7 vs. 8–9 branched pelvic-fin rays; 20–27 vs. 13–15 and 13–19 predorsal scales, respectively. It
is further distinguished from Balitoropsis by having 59–73 vs. 42–55 total lateral-line scales; anus closer to anal-fin
origin vs. closer to pelvic-fin insertion; triangular vs. crescentic upper lip.
FIGURE 3. Lateral views of living (A) Homaloptera orthogoniata, not cataloged, Kalimantan, Borneo, Indonesia. Strong
reddish tints on body likely breeding colors; (B) Homalopteroides smithi, UF 235740, 45.0 mm SL, Khlong Tasae, Salui
Subdistrict, Chumphon, Thailand; (C) Homalopterula cf. ripleyi, not cataloged, Sumatra, Indonesia; (D) Balitoropsis
zollingeri, not cataloged, Kalimantan, Borneo, Indonesia; (E) Pseudohomaloptera leonardi, UF 235735, 34.2 mm SL, Ta Pi
River, Nakhon Si Thammarat, Thailand. Photos (A) & (D) by Renny Hadiaty. Photo (C) by Daniel Lumbantobing. Specimens
not to scale.

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FIGURE 4. Scale sizes and keel patterns of predorsal region: Large, and keeled scales, (A) Balitoropsis zollingeri, UF 166094,
52.1 mm SL; (B) Pseudohomaloptera tatereganii, RMNH 7632 (holotype), 64.6 mm SL. Large, and wart-like/keeled scales (C)
Homalopteroides wassinkii, UF 183330, 43.6 mm SL. Medium, and keeled scales, (D) Homaloptera cf. ocellata, UF 161718,
64.1 mm SL. Small, and smooth scales, (E) Homalopterula ripleyi, ANSP 188908, 43.1 mm SL; (F) Ghatsa montana, CAS-SU
39871 (holotype), 46.6 mm SL. Scale bar represents 1 mm. Left side of photo is anterior.
Material examined. Homaloptera ocellata: Java: RMNH 2723 (holotype); ZMA 100.645 (2), 103.205 (1).
Sumatra: UF 161605 (1), 161719 (4), 161718 (2), 166096 (4), 166104 (2), 166106 (2), 166107 (4). H. bilineata:
Myanmar: RMNH 10263 (1); CAS 231723 (4). H. orthogoniata: Borneo (Kalimantan): RMNH 7790 (lectotype);
CAS 49326 (1). H. ogilviei: Peninsular Malaysia: ZRC 1555 (holotype); BMNH 1966.9.26.1 (paratype); RMNH
25443 (paratype); UF 235405 (1), 235408 (1). Sumatra: UF 161716 (2), 166085 (2), 166091 (2). Thailand: INHS

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93605 (1). H. confuzona: Thailand: UF 169906 (1); INHS 93493 (5). H. parclitella: Peninsular Malaysia: ZRC
49257 (holotype); CAS-SU 39390 (2).
FIGURE 5. Oral morphology of (A) Homaloptera cf. ocellata, UF 161718, 64.1 mm SL; (B) Homalopteroides wassinkii,
UMMZ 155660, 46.2 mm SL; (C) Homalopterula ripleyi, ANSP 188908, 43.1 mm SL; (D) Balitoropsis zollingeri, UF 166094,
52.1 mm SL; (E) Pseudohomaloptera tatereganii, RMNH 7632 (holotype), 64.6 mm SL; (F) Ghatsa montana, (holotype) CAS-
SU 39871, 46.4 mm SL. Abbreviations: CF, central furrow; FL, fleshy lobe; FP, fleshy pad; LJ, lower jaw; LL, lower lip; LRB,
lateral-rostral barbel; MB, maxillary barbel; MRB, medial-rostral barbel; RC, rostral cap; UJ, upper jaw; UL, upper lip.
Homalopteroides Fowler 1905
(Figures 3B, 4C, 5B, 7)
Homalopteroides Fowler, 1905:476. (Type species: Homaloptera wassinkii Bleeker 1853, by original designation; see Randall
& Page, 2012:335 fixed type species as H. wassinkii, under Code art. 70.3.1). Gender masculine.
Chopraia Prashad & Mukerji, 1929:188 (Type species:Chopraia rupicola Prashad & Mukerji, 1929, by original designation).
Gender feminine.
Remarks. Homalopteroides rupicola was originally designated as the type species of Chopraia (Prashad &
Mukerji 1929). Chopraia was distinguished from other balitorines by “shape of the head, the situation and better
development of the eyes, the branchial openings and the fins (Prashad & Mukerji 1929).” Since these characters are
shared with species of Homalopteroides, we treat Chopraia as a junior synonym of Homalopteroides (Randall &
Page 2012).

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FIGURE 6. Type localities for species of Homaloptera. Asterisk represents the type species of the genus.
Diagnosis. Distinguishing characters are given in Table 4 and shown in Figures 3B, 4C, 5B and 7.
Homalopteroides is distinguished by the following combination of characters: without reddish tints on fins (Fig.
3B) in life, dorsal-fin origin posterior to pelvic-fin origin; forked caudal fin; 6–8½, 7½ (M) branched dorsal-fin
rays; wart-like/keeled scales (Fig. 4C), 14–25 predorsal scales; 33–52 total lateral-line scales; anus closer to anal-
fin origin than to pelvic-fin insertion; no adipose keel on caudal peduncle; small rostral cap; 2 thin rostral barbels,
widely separated from one another; thin crescentic upper lip; no fleshy pad or lobes between lateral portions of
lower lip (Fig. 5B); and presence of a central furrow at isthmus.

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ON THE PARAPHYLY OF HOMALOPTERA
FIGURE 7. Dorsal, lateral, and ventral views of preserved Homalopteroides wassinkii, UMMZ 155660, 46.2 mm SL,
Tjisokan, Java, Indonesia.
Species included. Homalopteroides wassinkii (Bleeker 1853), H. modestus (Vinciguerra 1890), H. rupicola
(Prashad & Mukerji 1929), H. smithi (Hora 1932), H. stephensoni (Hora 1932), H. weberi (Hora 1932), H. tweediei
(Herre 1940), H. indochinensis (Silas 1953), H. nebulosus (Alfred 1969), H. yuwonoi (Kottelat 1998), and H. avii
Randall & Page 2014. The type localities for species of Homalopteroides are shown in Figure 8.
Comparison. Homalopteroides is distinguished from Homaloptera by absence vs. presence of reddish tints on
fins in life; dorsal-fin origin posterior vs. anterior to pelvic-fin origin; 6–8½, 7½ (M) vs. 7–8½, 8½ (M) branched
dorsal-fin rays; 33–52 vs. 59–73 total lateral-line scales; small vs. large rostral cap; medial- and lateral-rostral
barbels widely separated from one another at base vs. barbels in close proximity to one another; crescentic rather
than triangular upper lip; thin vs. thick upper lip; presence vs. absence of a central furrow at the isthmus; absence
vs. presence of fleshy pad between lateral portions of lower lip.
Homalopteroides is distinguished from Homalopterula by having forked vs. truncated or emarginated caudal
fin; wart-like/keeled vs. smooth scales; 14–25 vs. 28–56 predorsal scales; 33–52 vs. 57–75 total lateral-line scales;
absence vs. presence of adipose keel on caudal peduncle; thin vs. thick upper lip; absence vs. presence of fleshy
lobes between lateral portions of lower lip.
Homalopteroides is distinguished from Balitoropsis and Pseudohomaloptera by having dorsal-fin origin
posterior vs. anterior to or above the pelvic-fin origin; 6–8½, 7½ (M) vs. 7–9½, 8½ (M) branched dorsal-fin rays;
small vs. large rostral cap; medial- and lateral-rostral barbels widely separated from one another at base vs. barbels
in close proximity to one another; thin vs. thick upper lip; absence vs. presence of pad between lateral portions of
lower lip. It is further distinguished from Balitoropsis by having the anus closer to anal-fin origin than to pelvic-fin
insertion.

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FIGURE 8. Type localities for species of Homalopteroides. Asterisk represents the type species of the genus.
Material examined. Homalopteroides wassinkii: Java: RMNH 4987 (lectotype of Homaloptera wassinkii),
1934 (paralectotypes of Homaloptera wassinkii) (2), 4627 (2); BMNH 1866.5.2.52 (1); ZMA 103.206 (2); UMMZ
155660 (1); MNHN 3122 (syntype of Balitora ocellata). Sumatra: UF 161619 (7). H. modestus: Thailand: ANSP
179826 (5); NIFI 4517 (1), 3786 (1), 4514 (1); ROM 51147 (2); UF 172926 (1), 173067 (1), 176377 (10), 176408
(2), 176438 (8), 176454 (4), 176544 (1), 176557 (8), 181080 (5), 181160 (9), 181141 (1); ZRC 53385 (1), 53386
(1), 41272 (4). Myanmar: ZRC 22889 (1); BMNH 1893.2.16.50 (paralectotype of Helgia modesta); ZMA 100.982
(paralectotype of Helgia modesta). H. rupicola: Myanmar: CAS-SU 28726 (paratype of Chopraia rupicola); CAS

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ON THE PARAPHYLY OF HOMALOPTERA
231681 (2), 231726 (3), 231835 (1), 61338 (1); USNM 378433 (3); ZRC 43569 (2). H. smithi: Thailand USNM
109821 (syntypes of Homaloptera smithi) (5); ANSP 76852 (1), 76851 (3); BMNH 1934.12.18.34 (1); UF 183330
(3), 183411 (2), 183915 (1). H. stephensoni: Borneo: RMNH 7633 (holotype of Homaloptera stephensoni);
USNM 393671 (3). H. weberi: East Malaysia: BMNH 1895.7.2.81 (syntypes of Homaloptera weberi) (7); ZMA
100990 (syntype of Homaloptera weberi). H. tweediei: Malaysia: BMNH 1938.12.1.132 (paratype of
Homaloptera tweediei); CAS-SU 33012 (holotype of Homaloptera tweediei), 33013 (paratype of Homaloptera
tweediei) (2). H. indochinensis: Vietnam?: BMNH 1933.8.19.50 (holotype). H. nebulosus: Malaysia: BMNH
1967.11.15.15 (paratype of Homaloptera nebulosa); CAS-SU 66428 (paratype of Homaloptera nebulosa); ZRC
1759 (paratype of Homaloptera nebulosa); ZRC 2020 (holotype of Homaloptera nebulosa); UF 235748 (6). H.
yuwonoi: East Malaysia: MZB 5938 (holotype of Homaloptera yuwonoi). H. avii: East Malaysia: USNM 323875
(holotype of Homalopteroides avii), 323878 (paratype of Homalopteroides avii), USNM 323879 (paratypes of
Homalopteroides avii) (2); UF 185293 (paratype of Homalopteroides avii).
Homalopterula Fowler 1940
(Figures 3C, 4E, 5C, 9)
Homalopterula Fowler, 1940:379 (type species: Homaloptera ripleyi Fowler 1940:379) by original designation. Gender
feminine.
Remarks. Homalopterula was created for the new species Homalopterula ripleyi Fowler 1940 and distinguished
from other balitorids by the “peculiar shape of its jaws, in combination with its truncated caudal and entirely naked
medial under surface of the abdomen” (Fowler 1940:379). Homalopterula was treated as a junior synonym of
Homaloptera based on the illustration provided in Fowler’s type description and the variation that “exists regarding
the nature of the caudal and presence or absence of scales on the ventral surface of the abdomen in species of
Homaloptera” by Silas (1953). This decision was followed by Roberts (1989).
Kottelat (1998) recognized Homaloptera gymnogaster Bleeker 1853, H. heterolepis Weber & de Beaufort
1916, H. ripleyi (Fowler 1940), and H. vanderbilti Fowler 1940 as possibly forming a clade (Homalopterula) based
on having “a more cylindrical body, a relatively wide mouth, short paired fins, and a truncated or slightly
emarginated caudal fin.” Based on this description, Tan & Ng (2005) and Ott (2009) treated Homalopterula as a
subgenus of Homaloptera. Randall & Page (2012) recognized Homalopterula as a subgenus based on “mouth
morphology, dorsal-fin origin over pelvic fin, ≥ 60 lateral-line scales, and ≥ 30 predorsal scales.” They did not
include H. modiglianii, then recognized as a junior synonym of H. gymnogaster (Kottelat 1993). Including it in this
study following Kottelat (2012, 2013) changes the lateral-line and predorsal scale count range found in
Homalopterula to 57–75 and 28–56, respectively.
Diagnosis. Distinguishing characters are given in Table 4 and shown in Figures 3C, 4E, 5C and 9.
Homalopterula is distinguished by the following combination of characters: without reddish tints on fins in life
(Fig. 3C); dorsal-fin origin posterior to pelvic-fin origin; 5½ and 7½, 7½ (M) branched dorsal-fin rays; 7 pelvic-fin
rays; truncated or emarginated caudal fin; smooth scales (Fig. 4E), 57–75 total lateral-line scales, 28–56 predorsal
scales; anus closer to anal-fin origin than to pelvic-fin base; adipose keel on caudal peduncle; small rostral cap; 2
thick and widely separated rostral barbels; thick crescentic upper lip; 2 fleshy lobes between lateral portions of
lower lip (Fig. 5C); and presence of a central furrow at the isthmus.
Species included. Homalopterula gymnogaster (Bleeker 1853), H. modiglianii (Perugia 1893), H.
amphisquamata (Weber & Beaufort 1916), H. heterolepis (Weber & de Beaufort 1916), H. ripleyi Fowler 1940,
and H. vanderbilti (Fowler 1940). The type localities for species of Homalopterula are shown in Figure 10.
Comparison. Homalopterula is distinguished from Homaloptera by absence vs. presence of reddish tints on
fins in life; dorsal-fin origin posterior vs. anterior to pelvic-fin origin; 5½ and 7½, 7½ (M) vs. 7–8½, 8½ (M)
branched dorsal-fin rays; 28–56 vs. 20–27 predorsal scales; smooth vs. keeled scales; truncated or emarginated vs.
forked caudal fin; presence vs. absence of adipose keel on caudal peduncle; small vs. large rostral cap; medial- and
lateral-rostral barbels widely separated from one another at base vs. barbels in close proximity to one another;
crescentic rather than triangular upper lip; presence vs. absence of a central furrow at the isthmus; fleshy lobes vs.
pad between lateral portions of lower lip.

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FIGURE 9. Dorsal, lateral, and ventral views of preserved Homalopterula ripleyi, ANSP 188908, 43.1 mm SL, Kampung
Bassam, Sumatera Utara Province, Sumatra, Indonesia.
Homalopterula is distinguished from Homalopteroides by having truncated or emarginated vs. forked caudal
fin; smooth vs. wart-like/keeled scales; 28–56 vs. 14–25 predorsal scales; 57–75 vs. 33–52 total lateral-line scales;
presence vs. absence of adipose keel on caudal peduncle; thick vs. thin lips; presence vs. absence of fleshy lobes
between lateral portions of lower lip.
Homalopterula is distinguished from Balitoropsis and Pseudohomaloptera by having dorsal-fin origin
posterior vs. anterior to or above pelvic-fin origin; 5½ and 7½, 7½ (M) vs. 8½ branched dorsal-fin rays; 7 vs. 7–9,
8 (M) branched pelvic-fin rays; truncated or emarginated vs. forked caudal fin; smooth vs. keeled scales; 28–56 vs.
13–15 and 13–19 predorsal scales, respectively; presence vs. absence of adipose keel on caudal peduncle; small vs.
large rostral cap; medial- and lateral-rostral barbels widely separated from one another at the base vs. barbels in
close proximity to one another; presence of fleshy lobes vs. pad between lateral portions of lower lip. It is further
distinguished from Balitoropsis by having the anus closer to anal-fin origin than to pelvic-fin insertion and 57–75
vs. 42–55 lateral-line scales.
Material examined. Homalopterula gymnogaster: Sumatra: BMNH 1866.5.2.49 (holotype of Homaloptera
gymnogaster); ZMA 100256 (syntypes of Homaloptera lepidogaster) (3); UF 185031 (1). H. modiglianii:
Sumatra: BMNH 1931.10.29.1-2 (syntypes of Homaloptera modiglianii) (2). H. amphisquamata: Sumatra: ZMA.
100.998 (syntype of Homaloptera amphisquamata), 100.994 (syntypes of Homaloptera amphisquamata) (2). H.
heterolepis: ZMA 100.999 (syntypes of Homaloptera heterolepis) (3). H. ripleyi: Sumatra: ANSP 68713
(holotype), 187003 (2), 187004 (1), 188907 (1), 188908 (4). H. vanderbilti: Sumatra: ANSP 68688 (holotype of
Homaloptera vanderbilti), 68700 (Holotype of Homaloptera ulmeri).

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FIGURE 10. Type localities for species of Homalopterula. Asterisk represents the type species of the genus.
Balitoropsis Smith 1945
(Figures 3D, 4A, 5D, 11, and 12)
Balitoropsis Smith, 1945:278. (type species: Balitoropsis bartschi Smith 1945:279, by original designation). Gender feminine.
Remarks. The genus Balitoropsis was created for the species B. bartschi Smith 1945 and distinguished from

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Homaloptera (sensu lato) by having a deep preoral groove extending around the corners of the mouth and
papillated lips. Kottelat & Chu (1988) noted that all members of Homaloptera (sensu lato) have a preoral groove to
a varying degree and recognized Balitoropsis bartschi as a junior synonym of H. zollingeri. The papillated lips of
Smith (1945) refer to unculi found on the lips of most balitorids, not the large diagnosable papillae of some
balitorid genera (e.g., Balitora, Hemimyzon, Metahomaloptera). The holotype of B. bartschi (USNM 107963) is
identified as a gravid female of Homaloptera zollingeri Bleeker 1853, as assumed by Kottelat & Chu (1988) (Fig.
12).
Kottelat (1998) recognized Balitoropsis as possibly warranting recognition as a genus based on having “an
elongate body, a slender caudal peduncle, carinated scales, short paired fins (pectorals usually not reaching pelvic
base, pelvics not reaching anal), a dark body with a series of saddles along the back.” Balitoropsis was recognized
as a genus by Kottelat (2012, 2013) and as a subgenus by Tan & Ng (2005) and Randall & Page (2012). It is
recognized herein as a genus containing two species (B. zollingeri and B. ophiolepis).
Diagnosis. Distinguishing characters are given in Table 4 and shown in Figures 3D, 4A, 5D, and 11.
Balitoropsis is distinguished by the following combination of characters: without reddish tints on fins in life (Fig.
3D); dorsal-fin origin anterior to or above pelvic-fin origin; 8½ branched dorsal-fin rays; 7–9, 8 (M) branched
pelvic-fin rays; forked caudal fin; keeled scales (Fig. 4A); 42–55 total lateral-line scales; 13–15 predorsal scales;
large rostral cap; 2 thick rostral barbels in close proximity to one another; thick crescentic upper lip; fleshy pad
between lateral portions of lower lip (Fig. 5D); anus closer to pelvic-fin insertion than to anal-fin origin.
Species included. Balitoropsis zollingeri (Bleeker 1853) and B. ophiolepis (Bleeker 1853). Type localities for
species of Balitoropsis are shown in Figure 13.
FIGURE 11. Dorsal, lateral, and ventral views of preserved Balitoropsis zollingeri, UF 166094, 52.1 mm SL, Air Ogan,
Sumatera Selatan Province, Sumatra, Indonesia.

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ON THE PARAPHYLY OF HOMALOPTERA
FIGURE 12. Dorsal, lateral, and ventral views of preserved Balitoropsis bartschi, USNM 107963 (holotype), 76.0 mm SL,
Waterfall stream on Kao Chong, Trang Province, Thailand. Photos by Sandra Raredon, USNM, Ichthyology Division.
Comparison. Balitoropsis is distinguished from Homaloptera by absence vs. presence of reddish tints on fins
in life; 7–9, 8 (M) vs. 7 branched pelvic fin-rays; 13–15 vs. 20–27 predorsal scales; 42–55 vs. 59–73 total lateral-
line scales; crescentic vs. triangular upper lip; anus closer to pelvic-fin insertion than to anal-fin origin.
Balitoropsis is distinguished from Homalopteroides by having dorsal-fin origin anterior to or above the pelvic-
fin origin vs. posterior to pelvic-fin origin; 8½ vs. 6–8½, 7½ (M) branched dorsal-fin rays; anus closer to pelvic-fin
insertion vs. anal-fin origin; large vs. small rostral cap; medial- and lateral-rostral barbels in close proximity to one
another vs. barbels widely separated at base; thick vs. thin upper lip; presence vs. absence of fleshy pad between
lateral portions of lower lip.
Balitoropsis is distinguished from Homalopterula by having dorsal-fin origin anterior to or above the pelvic-
fin origin vs. posterior to pelvic-fin origin; 8½ vs. 5½ and 7½, 7½ (M) branched dorsal-fin rays; 7–9, 8 (M) vs. 7
branched pelvic fin-rays; forked vs. truncated or emarginated caudal fin; keeled vs. smooth scales; 13–15 vs. 28–56
predorsal scales; 42–55 vs. 57–75 total lateral-line scales; anus closer to pelvic-fin insertion than to anal-fin origin;
large vs. small rostral cap; medial- and lateral-rostral barbels in close proximity to one another vs. barbels widely
separated at base; presence of fleshy pad vs. lobes between lateral portions of lower lip.
Balitoropsis is distinguished from Pseudohomaloptera by having anus closer to pelvic-fin insertion than to
anal-fin origin.
Material examined. Balitoropsis zollingeri: Sumatra: BMNH 1866.5.2.53 (1); UF 161715 (3), 166094 (2),
166095 (1), 166102 (1), 166105 (1). Borneo: CAS 49331 (1); USNM 230253 (2). Thailand: USNM 107963
(Holotype of B. bartschi, examined photo); ANSP 68004 (Holotype of Homaloptera maxinae); UF 183727 (1),
235545 (1). Malaysia: CAS-SU 66420 (2), 66424 (Paratypes of Homaloptera nigra) (2); USNM 288456 (1); UF

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235547 (9), 235421 (2), 235420 (1); ZRC 2009 (Holotype of Homaloptera nigra). B. ophiolepis: Java: RMNH
4986 (lectotype of Homaloptera ophiolepis); BMNH 1866.5.2.49 (1). Sumatra: UF 166109 (3), 166103 (1),
166101 (1). Borneo: RMNH 28866 (1); USNM 230251 (1).
FIGURE 13. Type localities for species of Balitoropsis. Asterisk represents the type species of the genus. The type locality for
B. ophiolepis shares one of the type localities for B. zollingeri (Bandung, Java).

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Pseudohomaloptera Silas 1953
(Figures 3E, 4B, 5E, 14)
Pseudohomaloptera Silas, 1953:204. (type species: Homaloptera tatereganii Popta 1905:180, by original designation). Gender
feminine.
Remarks. Homaloptera tatereganii Popta 1905 was designated as the type species for the genus
Pseudohomaloptera by Silas (1953). Pseudohomaloptera was distinguished from Homaloptera by the “presence of
a rostral groove and other structures associated with the mouth” (Silas 1953:205). Tan (2009) recognized
Pseudohomaloptera as a junior synonym of Homaloptera, since all species of Homaloptera (sensu lato) have a
rostral and postoral groove to varying degrees. Kottelat (2012) recognized H. tatereganii as a species of
Balitoropsis and treated Pseudohomaloptera as a junior synonym of Balitoropsis.
Pseudohomaloptera is morphologically very similar to Balitoropsis, and the mouth characters given by Silas
(1953) cannot differentiate the two genera. Tan (2009) gave a simple pelvic-fin ray count of 3 for H. tatereganii to
distinguish it from species of Homaloptera (s.l.), which have 2 simple pelvic-fin rays. However, the holotype, the
only known specimen of H. tatereganii (RMNH 7632), has only 2 simple pelvic-fin rays (on both sides). Other
counts that differ from those given by Tan (2009) are the following: iii, 8½ vs. ii, 8 dorsal-fin rays; ii, 5½ vs. ii, 5
anal-fin rays; vii, 12 vs. viii, 12 pectoral-fin rays; 18 vs. 14 circumpeduncular scale count; and 6/7 vs. 5/6
transverse scale count. The following measurements differ from Tan (2009) (owing likely to different methods):
predorsal length 44.1% vs. 45.3% SL; body depth 12.5% vs. 10.4% SL; dorsal-fin base 16.7% vs. 18.8% SL;
pectoral-fin length 29.4% vs. 28.5% SL; head depth 48.2% vs. 42.8% HL; head width 81.3% vs. 78.3% HL; snout
length 61.9% vs. 57.2% HL; 15.1% vs. 14.5% HL; 40.3% vs. 37.7% HL.
FIGURE 14. Dorsal, lateral, and ventral views of preserved Pseudohomaloptera tatereganii, RMNH 7632 (holotype), 64.6
mm SL, Bo River, Upper Mahakam River basin, East Kalimantan, Borneo, Indonesia.

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FIGURE 15. Type localities for species of Pseudohomaloptera. Asterisk represents the type species of the genus.
Diagnosis. Distinguishing characters are given in Table 4 and shown in Figures 3E, 4B, 5E, and 14.
Pseudohomaloptera is distinguished by the following combination of characters: without reddish tints on fins in
life (Fig. 3E); dorsal-fin origin anterior to or above pelvic-fin origin; 8½ branched dorsal-fin rays; 8–9 branched
pelvic-fin rays; forked caudal fin; keeled scales (Fig. 4B); 50–61 total lateral-line scales; 13–19 predorsal scales;
anus closer to anal-fin origin than to pelvic-fin insertion; no adipose keel on caudal peduncle; large rostral cap; 2
thick rostral barbels in close proximity to one another; thick and triangular/crescentic upper lip; fleshy pad between
lateral portions of lower lip (Fig. 5E).

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Species included. Pseudohomaloptera tatereganii (Popta 1905), P. sexmaculata (Fowler 1934), P. leonardi
(Hora 1941), P. yunnanensis (Chen 1978), P. vulgaris (Kottelat & Chu 1988), and P. batek (Tan 2009). Type
localities for species of Pseudohomaloptera are shown in Figure 15.
Comparison. Pseudohomaloptera is distinguished from Homaloptera by absence vs. presence of reddish tints
on fins in life; 8–9, 8 (M) vs. 7 branched pelvic fin-rays; 13–19 vs. 20–27 predorsal scales.
Pseudohomaloptera is distinguished from Homalopteroides by having dorsal-fin origin anterior to or at pelvic-
fin origin vs. posterior to pelvic-fin origin; 8½ vs. 6–8½, 7½ (M) branched dorsal-fin rays; large vs. small rostral
cap; medial- and lateral-rostral barbels in close proximity to one another vs. barbels widely separated at base; thick
vs. thin upper lip; presence vs. absence of fleshy pad between lateral portions of lower lip.
Pseudohomaloptera is distinguished from Homalopterula by having dorsal-fin origin anterior to or at pelvic-
fin origin vs. posterior to pelvic-fin origin; 8½ vs. 5½ and 7½, 7½ (M) branched dorsal-fin rays; 8–9 vs. 7 branched
pelvic fin-rays; keeled vs. smooth scales; 13–19 vs. 28–56 predorsal scales; forked vs. truncated or emarginated
caudal fin; absence vs. presence of adipose keel on caudal peduncle; large vs. small rostral cap; medial- and lateral-
rostral barbels in close proximity to one another vs. widely separated at base; presence of fleshy pad vs. lobes
between lateral portions of lower lip.
Pseudohomaloptera is distinguished from Balitoropsis by having anus closer to anal-fin origin than to pelvic-
fin insertion.
Material examined. Pseudohomaloptera tatereganii: Borneo: RMNH 7632 (holotype of Homaloptera
tatereganii). P. sexmaculata: Thailand: ANSP 56374 (holotype of Homaloptera sexmaculata), 56375 (paratypes of
Homaloptera sexmaculata) (2), 56402 (holotype of Homaloptera septemmaculata), 56403 (paratype of
Homaloptera septemmaculata); UF 183358 (2), 177819 (3), 181170 (3). P. leonardi: Peninsular Malaysia: ZRC
1753 (paratype of Homaloptera leonardi); RMNH 23264 (6), 25921 (4); UF 169909 (3), 235746 (3). P.
yunnanensis: China: IHASW 60-VII-012 (holotype of Balitoropsis yunnanensis). P. vulgaris: China: 788229 (KIZ
1978001047) (holotype of Homaloptera vulgaris), 788225-788227 (KIZ 1978001048-50) (paratypes of
Homaloptera vulgaris). P. batek: Borneo: MZB 10990 (holotype of Homaloptera batek); ZRC 51743 (paratype of
Homaloptera batek).
Ghatsa, new genus
(Figures 4F, 5F, 16)
Remarks. The following five species of balitorines from the Western Ghats of India have been recognized as
belonging to the genus Homaloptera (Randall & Page 2012; Kottelat 2012): H. montana Herre 1945, H. pillaii
Indra & Rema Devi 1981, H. menoni Shaji & Easa 1995, H. santhamparaiensis Arunachalam et al. 2002, and H.
silasi Madhusoodana Kurup & Radhakrishnan 2011. Due to the inaccessibility of specimens from institutions in
India, this group has had an unresolved taxonomic status. It has been proposed by Pethiyagoda & Kottelat (1994)
[for the first two species listed above], and by Kottelat (1998) [for the first three species listed above] that these
species require a new genus or subgenus (Tan & Ng 2005). The only species examined in this study from this
Indian assemblage was H. montana (holotype, CAS-SU 39871) (Fig. 16). Homaloptera montana can be
distinguished from all species of Homaloptera (sensu lato) by the combined characters of placement of dorsal-fin
origin, tiny and smooth scales, truncated caudal fin, and features of the mouth. Based on these characters, H.
montana does not belong to any established balitorid genus and the newly created genus, Ghatsa is created for it.
The four other species formerly recognized in Homaloptera from the Western Ghats that were unavailable for
examination (H. pillaii Indra & Rema Devi 1981, H. menoni Shaji & Easa 1995, H. santhamparaiensis
Arunachalam et al. 2002, and H. silasi Madhusoodana Kurup & Radhakrishnan 2011) are tentatively placed in
Ghatsa, since they are morphologically (based on type descriptions) more similar to H. montana than to any other
balitorine. Further examination of these species is needed.
Type species. Homaloptera montana Herre, 1945:400.
Diagnosis. Distinguishing characters are given in Table 4 and shown in Figures 4F, 5F, and 16. Ghatsa is
distinguished by the combination of the following characters: dorsal-fin origin posterior to pelvic-fin origin;
emarginated-truncated caudal fin; small, smooth scales (Fig. 4F) (data only available for G. montana); 59–ca.101
lateral-line scales, ca. 53 predorsal scales (data only available for G. montana); anus closer to anal-fin origin than to

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pelvic-fin insertion; adipose keel on caudal peduncle (data only available for G. montana); small rostral cap; 2 thin,
widely separated rostral barbels; thin, smooth crescentic upper lip; absence of fleshy pad or lobes between lateral
portions of lower lip (Fig. 5F).
FIGURE 16. Ghatsa montana, CAS-SU 39871 (holotype), 46.4 mm SL. (A) Dorsal, lateral, and ventral views, Puthutotam
Estate, brook in Anamallai Hills, India; (B) radiograph. Photos by CAS, Ichthyology Section.
Species. Ghatsa montana (Herre 1945). The following species are tentatively recognized in Ghatsa: G. pillaii
(Indra & Rema Devi 1981), G. menoni (Shaji & Easa 1995), G. santhamparaiensis (Arunachalam et al. 2002), and
G. silasi (Madhusoodana Kurup & Radhakrishnan 2011). Type localities for species in Ghatsa are shown in Figure
17.
Comparison. Ghatsa is distinguished from Homaloptera by having the dorsal-fin origin posterior vs. anterior
to the pelvic-fin origin; scales small and smooth (data only available for G. montana) vs. medium and keeled; ca. 53
(data only available for G. montana) vs. 20–27 predorsal scales; truncated or slightly emarginated vs. forked caudal
fin; adipose keel present (data only available for G. montana) vs. absent; small vs. large rostral cap; medial- and
lateral-rostral barbels widely separated from one another at base vs. barbels in close proximity to one another;
crescentic rather than triangular upper lip; thin vs. thick upper lip; absence vs. presence of fleshy pad between
lateral portions of lower lip.

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Ghatsa is distinguished from Homalopteroides by having small and smooth scales (data only available for G.
montana) vs. large and wart-like/keeled scales; ca. 53 (data only available for G. montana) vs. 14–25 predorsal
scales; 59–ca. 101 vs. 33–52 lateral-line scales; truncated or slightly emarginated vs. forked caudal fin; and adipose
keel present (data only available for G. montana) vs. absent.
Ghatsa is distinguished from Homalopterula by having thin vs. thick barbels and upper lip; absence vs.
presence of fleshy lobes between lateral portions of lower lip; and by 59–ca. 101 vs. 57–75 lateral-line scales.
Ghatsa is distinguished from Balitoropsis and Pseudohomaloptera by having the dorsal-fin origin posterior vs.
anterior to pelvic-fin origin; smooth (data only available for G. montana) vs. keeled scales; ca. 53 (data only
available for G. montana) vs. 13–15 and 13–19 predorsal scales, respectively; slightly emarginated to truncated vs.
forked caudal fin; adipose keel present (data only available for G. montana) vs. absent; small vs. large rostral cap;
medial- and lateral-rostral barbels widely separated from one another at base vs. barbels in close proximity to one
another; absence vs. presence of a fleshy pad between lateral portions of the lower lip. It is further distinguished
from Balitoropsis by having an anus closer to anal-fin origin than to pelvic-fin insertion; 59–ca. 101 vs. 42–55
lateral-line scales.
Etymology. Named for the Western Ghats of India where species of this genus appear to be endemic. Gender
feminine.
Redescription of Ghatsa montana (Herre 1945)
(Figures 4F, 5F, 16)
Homaloptera montana Herre 1945:400 (no figure in original description); Journal of the Washington Academy of Sciences
35(12).
Type locality: brook on the Puthutotam Estate in Anamallai Hills at 3,600 feet altitude, Valparai Post Office, Madras
Presidency, South India. Holotype. CAS-SU 39871, Böhlke 1953:40.
Description. Dorsal, lateral, and ventral views are shown in Figure 16. Measurements and meristic counts are
given for the only specimen available to us, the holotype (CAS-SU 39871). The holotype is 46.4 mm SL, 54.8 mm
TL. Body (depth: 10.1% SL, width: 13.5% SL) arched predorsally, tapers posterior of anal-fin origin to caudal
base, flattened ventrally. Head (length: 21.7% SL, width: 76.2% HL, depth: 44.5% HL) triangular when viewed
dorsally; tubercles absent. Snout length 47.5% HL. Orbit (length: 22.8% HL) small, ovoid, positioned
dorsolaterally, shorter than interorbital width (31.7% HL).
Mouth (Fig. 5F) (width: 25.7% HL) subterminal, large portions of upper and lower jaws visible. Lips thin,
smooth, crescentic, continuous around corners of mouth; widest at corners of mouth. Lower lip medially
interrupted; the chin extending to most anterior portion of lip. Rostral and postlabial grooves present. Two pairs of
rostral barbels, 1 pair of maxillary barbels. Medial-rostral barbels widely separated from one another, distance
equal to that of medial interruption of lower lip. Medial and lateral-rostral barbels unequal in size, separated by
distance about equal to length of lateral-rostral barbel. Lateral-rostral barbel not reaching base of the barbel,
maxillary barbel reaching horizontally to vertical at anterior nostril. Gill opening extends to ventral surface of
body. Dorsal fin (base length: 9.7% SL, length: 18.5% SL) originates posterior to pelvic-fin base and closer to
caudal-fin base than to snout (predorsal length: 54.3% SL). Pectoral fin (length: 26.8% SL) longer than head, not
reaching pelvic-fin origin. Pelvic fin (length: 20.8% SL) lacks axillary pelvic lobe, not reaching anus; anus closer
to anal-fin origin than to pelvic-fin insertion. Anal fin length 13.5% SL. Adipose keel on caudal peduncle. Caudal
peduncle length 15.9% SL, depth 9.0% SL. Caudal fin truncate.
Body scaled except for ventral surface anterior to pelvic-fin insertion. Scales small, smooth, not obvious to
naked eye (Fig. 4F). Total lateral-line pores ca. 101, predorsal scales ca. 53, circumpeduncular scales ca. 60. Scales
above and below lateral line ca. 18 and ca. 15, respectively. Scales below lateral line to pelvic-fin origin ca. 12.
Dorsal-fin rays iii, 7½; anal-fin rays ii, 5; pectoral-fin rays vi, 9 for left side, and a total of 15 rays for the right side
(right fin damaged; simple and branched rays could not be differentiated); pelvic-fin rays ii, 8 for left side and iii, 7
for right side; total caudal-fin ray count 16. Pores of cephalic lateralis system: 7 supraorbital, 5+11 infraorbital, 7
preoperculomandibular, and 3 supratemporal. Total vertebrae count 38 (Fig. 16B).
Coloration. In 70% ethanol: The specimen is faded, and the general color of the body is brown with a black
stripe along the lateral-line. Herre (1945:400) gave the following description: “The color in alcohol is brown, the

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underside yellowish; 10 short dark brown bars across the back, but not extending down to the lateral-line; a poorly
defined dark longitudinal stripe below the lateral line from the eye to the caudal base; top of head very dark brown;
a blackish-brown spot on the ventral base; caudal with a blackish blotch on its base and another near its tip; other
fins all clear.”
Distribution. The type locality is the Puthutotam Estate, brook in Anamallai Hills, southern India, elevation
about 3,600 feet (Fig. 17). This is in the state of Tami Nadu close to the border with Kerala, Coimbatore District;
estimated coordinates are 10.36° N, 76.93° E. This drainage was reported to be adjacent and northeast to the
Chalakudy basin by Pethiyagoda & Kottelat (1994:110).
FIGURE 17. Type localities for species of Ghatsa. Asterisk represents the type species of the genus.

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Menon (1987) reported H. montana from Silent Valley and New Amarambalam of the Western Ghats, but this
is likely H. pillaii (see Remarks). Arunachalam et al. (2002) examined an individual from Kerala at Malakkapara,
headwaters of Chalakudy River. This specimen (KFRI F. 107) was not examined in this study, and this locality for
H. montana cannot be confirmed.
Remarks. Menon (1987) recognized H. pillaii (spelled pillai) as a junior synonym of H. montana. Menon
(1987) examined only the type series of H. pillaii but gave different counts than those in the original description
(also see Pethiyagoda & Kottelat 1994:109). Many authors (Indra & Rema Devi 1981; Madhusoodana Kurup &
Radhakrishnan 2010; Arunachalam et al. 2002; Silas 1953) have given a pectoral-fin ray count of iv, 8 for G.
montana. The pectoral-fin ray count here and in the original description (Herre 1945) is vi, 8.
Material examined. G. m o n t a n a : India: CAS-SU 39871 (holotype of Homaloptera montana).
Discussion
As recognized herein, Homaloptera consists of six species (Homaloptera ocellata, H. bilineata, H. orthogoniata,
H. ogilviei, H. confuzona, and H. parclitella) found in Myanmar, Thailand, Cambodia, Peninsular Malaysia,
Sumatra, Java, and Borneo. Of these six, H. parclitella and H. ogilviei are known to occur sympatrically, in the
Pahang (Alfred 1969) and Endau drainages (Ng & Tan 1999), Malaysia.
Homalopteroides, with 11 species (Homalopteroides wassinkii, H. modestus, H. rupicola, H. smithi, H.
stephensoni, H. weberi, H. tweediei , H. indochinensis, H. nebulosus, H. yuwonoi, and H. avii), is the most diverse
and widely distributed genus of species formerly in Homaloptera. It is known from northeast India, Myanmar,
Thailand, Laos, Cambodia, Vietnam (? see Kottelat 2012:51), Peninsular Malaysia, Sumatra, Java, and Borneo.
Homalopterula is only known to occur in Sumatra and consists of six species, five of which – H. heterolepis, H.
ripleyi, H. modiglianii, H. amphisquamata, and H. vanderbilti – were described from Aceh Province, while H.
gymnogaster was described from Sumatera Barat province. Homaloptera lepidogaster Weber & de Beaufort 1916
and H. ulmeri Fowler 1940 were recognized as junior synonyms of H. gymnogaster and H. vanderbilti,
respectively, by Kottelat et al. (1993), but without supporting data. Variation and relationships within
Homalopterula are unknown, and a taxonomic revision of the genus is needed.
Based on molecular and morphological data, Balitoropsis contains only two species. This differs from the
classification of Kottelat (2012) and Kottelat (2013), which recognizes ten and nine species in Balitoropsis,
respectively. Our results are consistent with the phylogenetic analysis of Randall & Page (2012, fig. 3), where B.
zollingeri is the sister species to a clade consisting of Pseudohomaloptera leonardi (previously designated in
Balitoropsis) and Homaloptera parclitella. Balitoropsis zollingeri has a wide distribution, having been reported
from Java, Sumatra, Borneo, Malay Peninsula, Thailand, Laos, and Cambodia
. Balitoropsis ophiolepis has been
reported from Java, Sumatra, and Borneo.
Pseudohomaloptera, which has not been recognized as a valid genus since its original description by Silas
(1953), contains six species (Pseudohomaloptera tatereganii, P. sexmaculata, P. leonardi, P. yunnanensis, P.
vulgaris, and P. batek) and is known to occur in southern China, Thailand, Laos, Cambodia, Peninsular Malaysia,
and Borneo. The relationships within Pseudohomaloptera are unknown. Based on limited access to specimens,
four species are tentatively assigned to Ghatsa (G. pillaii, G. menoni, G. santhamparaiensis, and G. silasi).
Intraspecific and interspecific variation and relationships among species of Ghatsa are unknown, and a revision of
this genus is needed. Based on a literature review, G. menoni is the most unusual, with a lateral-line scale count
much lower than that of other species (59–62 vs. 83–ca. 101).
Hora (1949) proposed the Satpura Hypothesis to explain the presence in peninsular India of taxa that seemed
mostly closely related to taxa in the Malay Peninsula. The disjunct populations of Homaloptera as understood at
the time in the Western Ghats and in the Malay Peninsula were used to support the hypothesis. However, based on
this study, Homaloptera as recognized by Hora is not monophyletic. In order to further test this hypothesis (see also
Karanth 2003; Dahanukar et al. 2013) and better understand the evolutionary history of Ghatsa and other endemic
hill stream loaches from the Western Ghats, phylogenetic analyses including these genera and those from
Indochina are required.

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Key to genera diagnosed in this study
1a. Origin of dorsal fin anterior to or above origin of pelvic fin; lateral- and medial-rostral barbels in close proximity to one
another; large rostral cap; fleshy pad between lateral portions of lower lip; 8½ (M) branched dorsal-fin rays . . . . . . . . . . . . . . 2
1b. Origin of dorsal fin posterior to origin of pelvic fin; rostral barbels widely separated; small rostral cap; fleshy pad between lat-
eral portions of lower lip absent; 7½ (M) branched dorsal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2a. Reddish tints on fins in life; predorsal scales ≥ 20; medium-sized keeled scales (Fig. 4D); 7 (M) branched pelvic-fin rays . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homaloptera
2b. Absence of reddish tints on fins in life; predorsal scales < 20; large keeled scales (Fig. 4A & B); 8 (M) branched pelvic-fin rays
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Anus closer to pelvic-fin base than to anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balitoropsis
3b. Anus closer to anal fin than to pelvic-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudohomaloptera
4a. Scales large (Fig. 4C); predorsal scales ≤ 25; total lateral-line scales ≤ 52; caudal fin forked; adipose keel absent . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homalopteroides
4b. Scales small (Fig. 4E & F); predorsal scales > 26; total lateral-line scales > 53; caudal fin truncated or slightly emarginated;
adipose keel present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Thick barbels and lips; fleshy lobes between lateral portions of lower lip; endemic to Sumatra . . . . . . . . . . . . . .Homalopterula
5b. Thin barbels and lips; no fleshy lobes between lateral portions of lower lip; endemic to Western Ghats of India . . . . . . Ghatsa
Comparative material. Balitora brucei: India: RMNH 11924 (neotype). Balitora sp: Thailand: NIFI 02927
(3). Bhavania australis: India: MNHN 50-79 (1); CAS 62052 (2). Cryptotora thamicola: Thailand: NIFI 3046 (1).
Hemimyzon yaotanensis: China: KU 21445 (1). Neohomaloptera johorensis: Peninsular Malaysia: CAS-SU
39840 (holotype), 39841 (paratype). Sewellia elongate: Laos: UF 185476 (3), 185488 (3). Travancoria jonesi:
India: MNHN 1950-0080 (1).
Acknowledgments
We would like to thank Renny Hadiaty (MZB) for images used in Figures 3A and 3D, Daniel Lumbantobing (UF)
for the image used in Figure 3C, Sandra Raredon (USNM) for images used in Figure 12, and CAS for images used
in Figure 16. We thank Mo Wang and Xiao-Yong Chen for providing data on Pseudohomaloptera vulgaris. For
specimen loans and access to institutional specimens, we thank Mark Sabaj Pérez (ANSP), James Maclain
(BMNH), David Catania (CAS-SU), E Zhang (IHASW), Zora Gabsi (MNHN), Sirwan Suksri (NIFI), Ronald de
Ruiter (RMNH), Robert Robins (UF), Jeffrey Williams (USNM), and Kelvin Lim (ZRC). We thank John Pfeiffer
(UF) and anonymous reviewers for their suggestions in improving this manuscript. The U.S. National Science
Foundation award (DEB 0845392) to D. Reed provided the Visionary Digital System used for Figures 2, 4, 5, 7,
and 11. Funding for this study was provided by the All Cypriniformes Species Inventory Project funded by the U.S.
National Science Foundation (DEB 1022720).
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