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Accepted by R. Pethiyagoda: 8 Oct. 2009; published: 30 Oct. 2009 53
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2009 · Magnolia Press
Zootaxa 2277: 53–60 (2009)
www.mapress.com/zootaxa/Article
Danionella priapus, a new species of miniature cyprinid fish from West Bengal,
India (Teleostei: Cypriniformes: Cyprinidae)
RALF BRITZ
Department of Zoology, The Natural History Museum, Cromwell Road, London, SW75BD, United Kingdom.
E-mail: r.britz@nhm.ac.uk
Abstract
Danionella priapus, a new species of sexually dimorphic miniature cyprinid from the Brahmaputra drainage in India, is
distinguished from the other three species in the genus by the presence in adult males of a conical projection of the
genital papilla situated between funnel-shaped pelvic fins, the number of anal- and pectoral-fin rays, and the position of
insertion of the last anal-fin pterygiophore. It differs further from D. translucida and D. mirifica in details of the colour
pattern, from D. dracula and D. mirifica in number of procurrent caudal-fin rays, from D. translucida in number of
vertebrae and from D. dracula in several skeletal characters. Like the other species in the genus, D. priapus shows a
developmentally truncated skeleton that is associated with several evolutionary morphological novelties. The present
distribution of the Danionella species may be the result of a vicariance event in the early Miocene, when the tectonic
uplift of eastern Tibet and the Indo-Burman ranges lead to the interruption of the palaeo-connection between the Tsangpo
(Brahmaputra) and the upper Irrawaddy Rivers.
Key words: miniature fishes, developmental truncation, evolutionary novelty, vicariance
Introduction
Species of the genus Danionella are completely transparent, miniaturized cyprinids. Three species have been
known so far, all distributed in Myanmar: D. translucida Roberts, 1986, from the Bago Yoma in southern
Myanmar, and D. mirifica Britz, 2003 and D. dracula Britz et al., 2009, from the Myitkina district in northern
Myanmar. All species of Danionella show a highly developmentally truncated skeleton, but exhibit at the
same time astonishing morphological novelties (Britz et al., 2009). The same is true for a Danionella species
from the Jorai River in West Bengal, material of which I received recently. A detailed investigation revealed
that this Indian species is new to science and the present paper serves to make a name available for it and
report its unusual characters.
Danionella priapus, sp. nov.
(Figs. 1–4)
Danionella sp.: Jorai River, Britz et al., 2009
Type material. Holotype: BMNH 2009.9.9.1, male, 14.4 mm SL; India, West Bengal, Jalpaiguri District,
Brahmaputra drainage, Jorai River, a tributary of the Sankosh at Laskarpara, outskirts of Barobisha town. 26°
28' 52.3''N, 89° 49' 29.8''E; M. Das, 2 Apr 2008.
Paratypes: BMNH 2009.9.9.2-37 (36), 11.8–16.0 mm SL; data as for holotype. BMNH 2009.9.9.38–43
(6), cleared and double stained, 13.6–15.9 mm SL; data as for holotype.
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FIGURE 1. Lateral view of Danionella priapus: a. BMNH 2009.9.9.1, holotype, male, 14.4 mm SL; b. BMNH
2009.9.9.2-37, paratype, female, 15.0 mm SL. Note presence of eggs and larval preanal fin fold in female, and difference
in size and shape of pelvic fins and in position of anus and genital papilla between male and female.
FIGURE 2. Difference in pelvic-fin structure in Danionella priapus: a holotype, BMNH 2009.9.9.1, male, 14.4 mm SL,
close-up of modified pelvic fins and conically projecting genital papilla; b, BMNH 2009.9.9.2-37, paratype, female, 15.0
mm SL, close-up of pelvic fins and preanal fin fold.
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DANIONELLA PRIAPUS, A NEW CYPRINID FROM INDIA
FIGURE 3. Dorsal and ventral view of colour pattern of head and anterior body in Danionella priapus, holotype,
BMNH 2009.9.9.1, male, 14.4 mm SL.
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Diagnosis. Danionella priapus is distinguished from its congeners, D. dracula, D. translucida and D.
mirifica, by the presence in adult males of a conical projection of the genital papilla situated between the
pelvic fins, which form a funnel-like structure (vs. genital papilla not developed as a conical projection and
pelvic fins not funnel-shaped), by possessing 8 pectoral-fin rays (vs. 6–7) and 20–21 anal–fin rays (vs. 12–14
in D. dracula, 12–16 in D. translucida, and 17–19, rarely 20 in D. mirifica), and by having the last anal-fin
pterygiophore inserted in front of haemal spine of vertebra 27 or 28 (vs. 21 or 22 in D. translucida, 23–25 in
D. mirifica, and 22–24 in D. dracula). It differs further from D. translucida and D. mirifica in the presence of
a median and two paramedian rows of pigment cells on the dorsal side of the body (vs. pigment rows absent).
It is further distinguished from D. dracula and D. mirifica by 7–8 dorsal caudal procurrent rays (vs. 5–6) and
6–8 ventral procurrent rays (vs. 4–5). In addition it differs from D. dracula in having two upper jaw bones (vs.
one); edges of jaw bones entire, without processes (vs. dorsal face of dentary and ventral face of upper jaw in
males each with a single row of 6–13 odontoid processes, anterior most large and canine-like); presence of
maxillary-mandibular cartilage (vs. absence); absence of membrane bone flanges on basipterygium (vs.
presence); and 9+9 principal caudal fin rays (vs. 7–8 + 7–8). It differs also from D. translucida in having 15–
16 abdominal (vs. 13–14) and 22–23 caudal (vs. 19–20) vertebrae and a well developed lateral stripe
extending from ear capsule to caudal peduncle (vs. a row of few melanophores midlaterally, restricted to
posterior part of body).
Description. Morphometric information, based on 15 specimens, is presented in Table 1. Known
maximum size 16.0 mm SL. Head and eye large, mouth supraterminal (Fig. 1). Body laterally compressed,
elongate; dorsal fin short, situated opposite posterior half of long anal fin. Caudal fin furcate, with remnants of
larval fin fold in front of its dorsal and ventral margins. Remnant of pre-anal larval fin fold present in females
only (Fig. 1b). Anus and genital papilla of adult males located between pelvic fins, in females at normal
position in front of anal fin. Genital opening of adult males located at tip of short, conical projection at
proximal end of funnel-like structure formed by pelvic fins being fused along their midline (Figs. 1a, 2).
Pseudotypanum present in body musculature at lateral side of anterior gas-bladder chamber, rendering its
pigmented surface visible (Fig. 1). Body muscles greatly thinned out at lateral side of posterior gas bladder
chamber. Scales absent. Lateral-line canals and pores on head and body absent.
Skull, hyopalatine arch, gill arches, and endoskeletal shoulder girdle mostly cartilaginous with
perichondral ossifications giving skeleton overall larval appearance. Numerous bones and cartilages absent:
kinethmoid (and kinethmoid cartilage), preethmoid, vomer, dermethmoid, nasal, parietal, intercalar,
extrascapular, infraorbitals 2–5, ectopterygoid, metapterygoid, urohyal, hypobranchials 1–3, sclerotic bones,
posttemporal, postcleithrum, mesocoracoid, pectoral radials 3–4 (and pectoral radial cartilages 3–4), pelvic
radials 1–3 (and pelvic radial cartilages 2–3), supraneurals 2 and 5–9 (and supraneural cartilages 5–9), middle
and distal radials in dorsal and anal fins, epineurals, epipleurals, uroneural 2, and scales. Ceratobranchial 5
heavily ossified, bearing 4–5 tri- to quadricuspid teeth plus the tips of one or two replacement teeth. Outer arm
of os suspensorium sexually dimorphic, being enlarged and hyperossified in males and in confluence with
lateral process of second vertebra via anterior process and with inner arm of os suspensorium below tripus via
broad median flange. Males with a relatively large conical nodule of cartilage between fifth rib and outer arm
of suspensorium, its proximal surface approaching swimbladder wall.
Vertebrae totalling 37 (2) or 38 (4), abdominal vertebrae 15 (3) or 16 (3); caudal vertebrae 22 (5) or 23 (1).
Ribs present on vertebrae 5 to 14. Dorsal-fin rays ii,6,i (5) or ii,7,i (1). First dorsal-fin ray pterygiophore
inserted behind neural spine of vertebra 21 (3) or 22 (3), and last in front of neural spine of vertebra 25 (2) or
26 (4). Anal-fin rays ii,17,i (2) or ii,18,i (4). Number of anal-fin pterygiophores in front of first hemal spine: 0
(2), 1 (2) or 2 (2). Last anal-fin pterygiphore inserted in front of neural spine of vertebra 27 (3) or 28 (3).
Principal caudal-fin rays 9+9 (6) plus 7 (5) or 8 (1) dorsal and 6 (2), 7 (1) or 8 (3) ventral procurrent rays.
Pectoral-fin rays 8 (6) and pelvic-fin rays 5 (6).
Colouration. Pigmentation in alcohol specimens restricted to several rows of melanophores (Figs. 1, 3):
mid-lateral row along horizontal septum from shoulder girdle to hypural plate; ventral row from in front of
and slightly above anal-fin base along ventral larval fin fold to end of hypural plate; row along anal-fin base;
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DANIONELLA PRIAPUS, A NEW CYPRINID FROM INDIA
mid-dorsal row starting behind head with several enlarged melanophores forming small blotch and extending
usually halfway between head and anterior base of dorsal fin, rarely to anterior dorsal-fin base; paired dorsal
paramedian rows from head to base of caudal fin; dorsal-fin base row; abdominal mid-ventral row from
ventral tip of cleithrum to anus; several melanophores on otic capsule; melanophores capping dorsal and
dorso-lateral face of gasbladder chambers and their connecting duct and lining peritoneum covering gonads;
streaks of melanophores lining anterior edge of lower cleithrum, ventral edge of opercle and preopercle,
branchiostegal rays and fin rays in dorsal, anal and caudal fins and anterior two or three rays of pectoral fin;
and several larger melanophores near symphysis of lower jaw. Head with supraoccipital blotch formed by
several large contiguous melanophores, with numerous large scattered melanophores between supraoccipital
blotch and epiphyseal bar, with a preepiphyseal blotch, and a few prenasal melanophores.
In life, body colourless and largely translucent (Fig. 4), except for melanophore patterns described above,
a thin yellowish-greenish line running along body at level of neural tube; golden chromatophores forming ring
around pupil, capping swimbladder chambers and connecting duct.
Distribution. The species is known only from the type locality, Jorai River in West Bengal, India (Fig. 5).
Etymology. The species name priapus is derived from the greek , the god of fertility of Greek
mythology. It was inspired by the conical projection of the genital papilla in males, which—at a superficial
level—is reminiscent of the penis of mammals. A noun in apposition.
TABLE 1. Selected morphometric information on 15 specimens (7 males, 8 females) of Danionella priapus, including
the holotype.
Holotype Paratypes
Male Males Females
Range Mean St. Dev Range Mean St. Dev
Standard length (SL) 14.4 12.5–14.1 13.7–15.0
In % of SL
Head length 17.4 17.4–20.0 18.3 0.9 19.0–20.7 19.8 0.7
Predorsal-fin length 70.1 68.7–70.4 69.4 0.7 68.7–71.0 70.1 0.8
Prepelvic-fin length 30.6 29.8–32.8 30.8 1.1 34.0–35.6 34.8 0.5
Preanal-fin length 52.1 51.1–52.1 51.5 0.4 53.3–54.8 54.2 0.6
Snout to anus 29.9 29.1–32.0 30.1 1.0 51.4–52.7 51.9 0.4
Body depth at dorsal-fin origin 13.2 13.1–15.2 14.1 0.7 14.4–16.0 15.4 0.5
Caudal-peduncle depth 7.6 7.5–8.5 8.0 0.3 7.5–8.9 8.0 0.5
Caudal-peduncle length 18.8 17.9–21.6 19.6 1.4 18.1–21.3 19.6 1.1
Length of dorsal-fin base 9.7 9.7–11.3 10.7 0.5 10.3–11.7 10.9 0.5
Length of anal-fin base 30.6 28.4–30.7 29.5 0.9 26.0–28.3 27.4 0.7
Length of pelvic fin 8.7 7.2–8.7 7.4 0.3 6.3–8.3 7.0 0.6
In % of HL
Eye diameter 44.0 40.0–44.0 40.8 1.5 35.0–39.3 37.3 1.3
Snout length 16.0 16.0–16.7 16.1 0.3 17.9–21.4 20.0 1.2
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FIGURE 4. Live colouration of male (above) and female (below) Danionella priapus. Specimens not preserved. Note
eggs in body cavity of female.
FIGURE 5. Distribution of the four species of Danionella in India and Myanmar.
Discussion
Species of the genus Danionella are among the smallest fishes with reported maximum sizes of 12 mm in D.
translucida (Roberts, 1986), 14.1 mm in D. mirifica (Britz, 2003), and 16.7 mm in D. dracula (Britz et al.,
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DANIONELLA PRIAPUS, A NEW CYPRINID FROM INDIA
2009). With the largest specimen in the type series measuring 16 mm, D. priapus is one of the larger species in
the genus.
Danionella priapus is easily distinguished from its congeners by the conically projecting genital papilla
and the funnel-shaped pelvic fins of males, and by its 8 pectoral-fin rays, but also by its longer anal fin, which
consists of 20–21 rays, a number that is never reached in D. translucida and D. dracula, and only rarely in D.
mirifica (only in 1 out of 20 specimens, see Britz, 2003). The three rows of dorsal-pigment pattern of the body
of D. priapus are also diagnostic, as neither D. translucida nor D. mirifica shows any dorsal body
pigmentation. Danionella dracula usually has only a single short row of melanophores, but occasionally
individuals may have a denser covering with melanophores that resembles the condition in D. priapus.
Britz et al. (2009) argued that D. dracula with its massive tooth-like jaw projections, but otherwise larval
skeleton, exemplifies in a compelling way the link between evolution of morphological novelties and
progenetic paedomorphosis via heterochronic change in developmental timing. Danionella priapus can
certainly be mentioned in this context too, as it combines a developmentally truncated skeleton with highly
modified anatomical structures. While males of all Danionella species have their anus and genital opening
shifted anteriorly between the pelvic fins, the conical projection of the genital papilla of male D. priapus is not
only unique among the Danionella species, but also does not seem to be present in any other cypriniform. The
pelvic fins are modified in some Danionella species, like D. dracula, but the funnel-like formation of these
fins in D. priapus is highly unusual, if not unique, among cypriniforms. It seems extremely surprising to find
such a variety of highly specialized and modified anatomical structures among only four species of closely
related and highly miniaturized fishes. This is even more astonishing when at the same time living Danionella
translucida, D. mirifica, D. dracula, and D. priapus are superficially very similar and easily confused.
As already stressed by Britz et al. (2009: 2183) for D. dracula, it seems, however, that precisely this
evolutionary process of developmental truncation may have also facilitated the evolution of the conically
projecting genital papilla and funnel-shaped pelvic fins of D. priapus “by freeing large parts of the anatomical
structures from developmental constraints, dissociating developmentally linked pathways and creating a
greater potential for more dramatic changes”.
It is still unclear if these greatly modified anatomical structures in D. priapus are part of an unusual
reproductive strategy and further studies will have to clarify this.
Despite their external similarity the evolutionary gap between D. priapus from the Brahmaputra drainage
in India and its sister group, D. mirifica plus D. translucida from the Irrawaddy/Sittang drainages in
Myanmar, is quite significant, if expressed as time since divergence. In the time tree published by Britz et al.
(2009), the earliest split in the genus is that of D. dracula from the other Danionella species at 29.5 million
years ago (95% confidence interval 27.4–31.9). The next split is that of D. priapus from its sister group, the
remaining Myanmar species, at around 22.6 million years ago (with a 95% confidence interval of 20.7–24.6
mya) at the Oligocene-Miocene boundary. This is a similar time frame as recovered in Rüber et al.’s (2004)
molecular phylogenetic analysis of the family Badidae for pairs of sister groups in the Bramaputra and
Irrawaddy/Sittang drainages: Dario dario vs. D. hysginon; Badis pyema+B. corycaeus vs. B.kanabos+B.
badis+B. sp. ‘‘Assam’’+B. assamensis+B. blosyrus; Colisa labiosa vs. other Colisa spp.; and Ctenops nobilis
vs. Parasphaerichthys spp. The present occurrence of D. priapus is thus consistent with a vicariant event
caused by the uplifting of eastern Tibet and the Indo-Burman ranges, which interrupted the hypothesized
palaeo-connection of the Tsangpo River with the Irrawaddy River in the early Miocene (see Rüber et al., 2004
for a comprehensive discussion).
Comparative material
Danionella translucida: NRM 32233 (5), paratypes, cleared and double stained; Myanmar, Pegu Division.
Sittang River basin, Dayame Chaung 1 mile from Daik-U. NRM 32234 (3), paratypes, cleared and double
stained; same data as NRM 32233. NRM 32235 (12), paratypes; same data as NRM 32233.
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Danionella mirifica: USNM 372847, holotype; Myanmar, Kachin State: hill stream, 8 miles from
Kamaing on road to Tanai. USNM 372848 (36), paratypes; same data as holotype. USNM 372849 (20),
paratypes, cleared and double stained; same data as holotype.
Danionella dracula: BMNH 2008.1.1.1, holotype; Myanmar: Kachin state: stream at Sha Du Zup
between Mogaung and Tanai. BMNH 2008.1.1.2-99 (98), paratypes; same data as holotype. BMNH
2008.1.1.100-119 (20), cleared and double stained; same data as holotype.
Acknowledgements
I am grateful to Andrew Rao, Malabar Tropicals, Calcutta, and Keith Lambert, Wildwoods, Crews Hill, who
made the specimens of D. priapus available. I also thank Sven Kullander (NRM), Jeff Williams (USNM) and
Oliver Crimmen (BMNH) for access to material under their care. The manuscript benefited from comments
by Sven Kullander and an anonymous reviewer.
References
Britz, R. (2003) Danionella mirifica, a new species of miniature fish from Upper Myanmar (Ostariophysi: Cyprinidae).
Ichthyological Exploration of Freshwaters, 14, 217–222.
Britz, R., Conway, K.W. & Rüber, L. (2009) Spectacular morphological novelty in a miniature cyprinid fish, Danionella
dracula n. sp. Proceedings of the Royal Society B, 276, 2179–2186.
Roberts, T. (1986) Danionella translucida, a new genus and species of cyprinid fish from Burma, one of the smallest
living vertebrates. Environmental Biology of Fishes, 16, 231–241.
Rüber, L., Britz, R., Kullander, S.O. & Zardoya, R. (2004) Evolutionary and biogeographic patterns of the Badidae
(Teleostei: Perciformes) inferred from mitochondrial and nuclear DNA sequence data. Molecular Phylogenetics and
Evolution, 32, 1010–1023.