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Journal of Natural History
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Discovery of the freshwater genus
Sicyopus (Teleostei: Gobioidei:
Sicydiinae) in Madagascar, with a
description of a new species and
comments on regional dispersal
Philippe Keith a , Gerard Marquet a & Laura Taillebois a
a Muséum national d'Histoire naturelle, DMPA, UMR 7208, CP026,
57, rue Cuvier, F-75231, Paris Cedex 05, France
Available online: 29 Sep 2011
To cite this article: Philippe Keith, Gerard Marquet & Laura Taillebois (2011): Discovery of the
freshwater genus Sicyopus (Teleostei: Gobioidei: Sicydiinae) in Madagascar, with a description of a
new species and comments on regional dispersal, Journal of Natural History, 45:43-44, 2725-2746
To link to this article: http://dx.doi.org/10.1080/00222933.2011.602479
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Journal of Natural History
Vol. 45, Nos. 43–44, November 2011, 2725–2746
Discovery of the freshwater genus Sicyopus (Teleostei: Gobioidei:
Sicydiinae) in Madagascar, with a description of a new species and
comments on regional dispersal
Philippe Keith*, Gerard Marquet and Laura Taillebois
Muséum national d’Histoire naturelle, DMPA, UMR 7208, CP026, 57, rue Cuvier, F-75231 Paris
Cedex 05, France
(Received 26 November 2010; final version received 24 June 2011; printed 27 September 2011)
In the Indo-Pacific area, insular rivers are mainly colonized by gobiids of the
Sicydiinae subfamily. These species spawn in freshwater, where the free embryos
drift downstream to the sea before returning to rivers to reproduce; they are
amphidromous. These gobies are the greatest contributors to the fish diversity and
have the highest levels of endemism. Among the nine known genera of Sicydiinae,
only two have been found in the West Indian Ocean, and only one (Sicyopterus)
is known from Madagascar. Recently, two surveys discovered a new species in the
genus Sicyopus.Sicyopus lord sp. nov. differs from other species by a combination
of characters that includes a particular colour pattern, similar in both sexes; more
scales than its congeners in lateral series (35–43), transverse back series (12–18) and
a shorter caudal fin length in male fish. This paper describes the new species and
discusses the presence of this genus in Madagascar.
Keywords: Sicyopus; new species; Gobiidae; Madagascar; dispersal
Introduction
In the Indo-Pacific area, insular river systems are colonized by freshwater gobies with
a life cycle adapted to the conditions in these distinctive habitats, which are young
oligotrophic rivers subject to extreme climatic and hydrological seasonal variation.
These species spawn in freshwaters, where the free embryos drift downstream to the
sea where they undergo a planktonic phase, before returning to rivers to grow and
reproduce (Keith 2003), hence they are described as amphidromous (McDowall 2007).
These gobies contribute most to the diversity of fish communities in the Indo-Pacific
insular systems, and have the highest levels of endemism (Keith et al. 2006; Lord and
Keith 2008). Amphidromous gobies belong mainly to the Sicydiinae sub-family, with
nine genera and nearly 110 species. Among them, seven genera are present in the
Indo-Pacific area (Keith et al. 2011). All of these genera have particular geographic dis-
tributions, but only two (Sicyopterus,Cotylopus) are known to occur in the West Indian
Ocean, and only one (Sicyopterus) with two species, is known to inhabit Malagasy
freshwaters.
Madagascar is the biggest island in the Indian Ocean. Many papers have been
written about the freshwater fish of this country since 1850 (Bleeker and Pollen 1875;
Sauvage 1891; Pellegrin 1933; Arnoult 1959; Kiener 1963, 1965), and it was often
*Corresponding author. Email: keith@mnhn.fr
ISSN 0022-2933 print/ISSN 1464-5262 online
© 2011 Taylor & Francis
http://dx.doi.org/10.1080/00222933.2011.602479
http://www.tandfonline.com
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2726 P. Keith et al.
considered that its fauna was well known. Nevertheless, recent studies have shown that
this is not the case and new species have been described from remote areas (Stiassny
and Raminosoa 1994; Loiselle 2005). In April 2004, the Reunion Association for
Aquaculture Development (ARDA) and the National Museum of Natural History
of Paris (MNHN), made specific inventories in the eastern rivers of Madagascar to
improve knowledge on freshwater biodiversity. During this survey, one specimen of
a new species of Sicyopus (Gobiidae, Sicydiinae), a genus never before caught so far
west in the Indian Ocean, was collected. Recently (May 2010), the MNHN conducted
freshwater inventories in the northeastern rivers of Madagascar, for the DIAMSOI
(Diadromy in South Indian Ocean) program. Six other specimens of this new species
were caught.
Watson (1999) defined three subgenera (Juxtastiphodon,Sicyopus and
Smilosicyopus) as belonging to Sicyopus Gill, 1863, based largely on dental charac-
teristics found in both jaws. Juxtastiphodon has closely set conical teeth in both jaws,
none recurved and without canines. Sicyopus has widely spaced conical teeth in both
jaws with most teeth sharply recurved and without canines. Smilosicyopus has slightly
recurved conical teeth anteriorly in both jaws, needle-like teeth laterally with none
recurved, and between anterior and lateral teeth, at least one (one to three) canine
tooth, well developed in males. Recently, Keith et al. (2011) elevated Smilosicyopus to
genus level.
The new species belongs to the genus Sicyopus, which currently includes seven
species: Sicyopus auxilimentus Watson and Kottelat, 1994; Sicyopus cebuensis Chen
and Shao, 1998; Sicyopus discordipinnis Watson, 1995; Sicyopus exallisquamulus
Watson and Kottelat, 2006; Sicyopus jonklaasi (Axelrod, 1972); Sicyopus multisqua-
matus de Beaufort, 1912 and Sicyopus zosterophorum (Bleeker, 1857).
The aim of this paper is to describe the new species of Sicyopus found in
Madagascar and to discuss the presence of this genus in this area in terms of dispersal
in the Indian Ocean.
Materials and methods
Methods follow Watson (1995) and Keith and Marquet (2005). Measurements were
taken with dial callipers to the nearest tenth of a millimetre. All counts were taken
from the right side. Preanal length, distance from origin of anal fin to tip of snout;
predorsal length, distance from origin of first dorsal fin to tip of snout; jaw length,
distance from posterior edge of upper jaw to anterior edge of upper jaw at symphysis;
caudal peduncle length, distance from base of posterior ray of second dorsal fin to
central hypural base; caudal peduncle depth, vertical distance across narrowest point
of caudal peduncle; body depth, vertical distance from origin of second dorsal fin to
belly (value only given in males because females may vary considerably from gravid to
non-gravid condition); second dorsal and anal fin lengths, distance from base of spine
to tip of last ray when depressed; caudal fin length, and distance from central hypural
base to tip of longest ray were recorded. The size is given in standard length (SL).
Teeth were counted to the right of the symphysis.
Abbreviations used in the descriptive account follow Watson (1995) and Keith and
Marquet (2005): A, anal fin; C, caudal fin (only branched rays are reported); D, dorsal
fins; D1, first dorsal fin; D2, second dorsal fin; LS, scales in lateral series counted from
upper pectora1 base, or anteriormost scale along lateral midline, to central hypural
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Journal of Natural History 2727
base; P, pectoral fin; PD, predorsal midline counted from scale directly anterior to first
dorsal fin insertion to the anteriormost scale; TRB, transverse series back, refers to
scales counted from the first scale anterior to second dorsal fin, in a diagonal manner,
posteriorly and ventrally to the anal fin base or ventralmost scale; TRF, transverse
series forward refers to scales counted from the first scale anterior to second dorsal
fin, in a diagonal manner, anteriorly and ventrally to the centre of belly or ventralmost
scale; ZZ, zigzag series, refers to scales on the narrowest region of the caudal peduncle
counted from the dorsalmost scale to the ventralmost scale in a zigzag (alternating)
manner.
Abbreviations for institutions and collections cited follow Leviton et al. (1985),
except LICPP, which is now BLIP (Biological Laboratory, Imperial Palace, Akasaka
Imperial Palace, Tokyo) and CMK (Collection of Maurice Kottelat, Cornol,
Switzerland). Abbreviations for the cephalic sensory pore system follow Akihito
(1986).
Meristics and morphometrics are summarized in Tables 1–4.
Comparative material
Sicyopus jonklaasi
SMF 20398, male (37.5 mm SL); SMF 20400, 2 males (35.5–38.7 mm SL); fast flowing
mountain stream at Atweltota, 5 km from Badurelia, Sabaragamuwa Province, Sri
Lanka; 1 September 1982, P. Beyer.
SMF 20399, female (37.4 mm SL); SMF 20401, 3 females (31.6–36.4 mm SL); SMF
20402, male (30.2 mm SL); SMF 20403, 6 males (29.3–34.1 mm SL); SMF 20404,
2 females (30.0–32.0 mm SL), SMF 20411, 4 males (30.5–32.4 mm SL); mountain
stream in southwestern Sri Lanka; December 1985, Aquarium Dietzenback.
SMF 20427, 2 males (37.0–38.5 mm SL); Atweltota, Sabaragamuwa Province, Sri
Lanka; January 1981, R. Jonklaas and A. van den Nieuwenhuizen.
CMK 6401, 2 males (37.7–44.7 mm SL); above Bopath Ella Falls, Ratnapura District,
Sabaragamuwa Province, Sri Lanka; January 1989, R. Pethiyagoda.
CMK 6543, female (35.7 mm SL), Sitawaka River at Yogama, Sri Lanka; January
1989, R. Pethiyagoda.
Sicyopus zosterophorum
RMNH 4462, Holotype, male (37.2 mm SL), RMNH 4766, Syntypes of Sicydium bali-
nense, 2 females (37.9–39.7 mm SL); freshwaterStream, Boleling, Bali; ca. 1856,
P. L. van BloemenWaanders.
ANSP96648, male (45.0 mm SL); Tenaru river, Betilange village, Guadalcanal,
Solomon Islands; 8 April 1953, M. Laird.
MNHN (uncatalogued), 3 males; New Caledonia, Napoemien river; January 2010,
P. Keith, C. Lord, L. Taillebois, P. Feutry.
MNHN (uncatalogued), 3 females; Vanuatu, Lasunuwé river, Malekula, Vanuatu;
January 2010, P. Keith, C. Lord, L. Taillebois, P. Feutry.
CMK 9780, 20 males, 25 females (22.1–32.9 mm SL); Lagu Lagu creek, southern mar-
gin of Visayan State College of Agriculture, about 7 km northof Baybay, Leyte
Island, Philippines; 25 June 1993, M. Kottelat and J. Margraf.
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2728 P. Keith et al.
Sicyopus discordipinnis
WAM P.27834-004 (Holotype), male (25.4 mm SL); LetakCreek, 25 km southeast of
Wewak, New Guinea, Papua New Guinea, 17 October 1982, G. R. Allen and
D. Coates.
WAM P.32372-006, 3 males, 4 females, Papua New Guinea, Apatabuia River, above
village; 30 January 2003, G. Allen and T. Stevenson.
WAM P.27834-005 (Paratypes), 2 females (26.0–27.6 mm SL); same data as Holotype.
WAM P.28167-001, 2 males, 4 females (29.3–34.3 mm SL); about 18 km southwest
of Arawa on Panguna Road, Bougainville, Papua New Guinea; 4 October 1983,
G. R. Allenand R. Steene.
NMBA 5075, male (24.4 mm SL); Namamosa, New Hanover, Bismarck Archipelago,
Papua New Guinea; November 1931, A. Bühler.
Sicyopus exallisquamulus
MZB 5918 (Holotype), male, 31.4 mm SL; Indonesia: Maluku: Halmahera, Sungei
Okitai, about 10 km upstream from coast, just below waterfalls; Robb, August
1994.
CMK 11294 (Paratypes), 2 males, 2 females, 36.0–51.3 mm SL; Indonesia: Maluku:
Halmahera, Sungei Tolawi about 10 km upstream from coast; D. Robb, August
1994.
CMK 11349, 2 males, 33.8–39.5 mm SL; same data as holotype.
CMK 11378, 1 male, 1 female, 30.0–31.1 mm SL, MZB 5926, female, 29.4 mm SL;
Indonesia: Maluku: Halmahera, Sungei Ifis about 8 km upstream from coastand
about 400 m below waterfalls; D. Robb, August 1994.
Sicyopus auxilimentus
ZRC 38286/CMK 10047 (Holotype), male (29.2 mm SL);Philippines: Leyte Island:
Lagu Lagu creek, about2 km from sea, southern margin of Visayan StateCollege
of Agriculture, about 7 km north of Baybay; 23 March 1991, J. Margraf.
ZRC 38287/CMK 10015 (Paratype), female (25.9 mm SL);Philippines: Leyte Island:
Lagu Lagu creek about 2 km from sea, southern margin of Visayan State, College
of Agriculture, about 7 km north of Baybay; 11 July 1993, M. Kottelat and
J. Margra f.
Sicyopus cebuensis
ASIZP-057825 (Holotype), male (37.5 mm SL); Naga river, Cebu Island, Philippines.
ASIZP-057826 (Paratype), female (39.1 mm SL); same data as Holotype.
Sicyopus multisquamatus
ZMA 110.982 (Holotype), female (46.2 mm SL); Ceram, Indonesia.
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Journal of Natural History 2729
Sicyopus lord sp. nov.
(Figures 1, 2; Tables 1–4)
Material examined
Seven specimens from Madagascar, totalling three males, four females; size range
35.2–47.4 mm SL (40.8–56.7 mm, total length), largest male 39.2 mm SL, largest
female 47.4 mm SL.
Holotype MNHN 2010-0925, male (37.2 mm SL), Ambodiforaha River, altitude: 50 m,
14 May 2010, G. Marquet, T. Robinet, Roger and H. Grondin.
Paratypes MNHN 2010-0926, 1 male, 1 female, size range 35.2–35.9 mm SL; same
data as Holotype. MNHN 2010-0927, female, 47.4 mm SL; Manoumpana River, April
2004, ARDA. MNHN 2010-0953, 1 male, 2 females, size range 39.3–44.9 mm SL;
Andraka River, 16–18 May 2010, G. Marquet, T. Robinet, Roger and H. Grondin.
Diagnosis
ASicyopus species which is distinguished from all the other species with the following
set of characters: (i) a particular colour pattern, being similar in both sexes with three
blackish bands present on the flanks: the first band extends from the tip of snout and
upper lip, under the eye to the posterior edge of operculum and pectoral base, and
from pectoral base to hypural; the second extends from behind the eye to the caudal
fin; and the third runs along the upper part of flank from above the pectoral fin to the
second dorsal fin base; (ii) a second dorsal fin with nine soft rays; (iii) a shorter caudal
fin length in male; (iv) more scales in lateral series (35–43) and transverse back scale
series (12–18).
Description
Scale counts in S. lord sp. nov. and related species are given in Table 2, number of
upper jaw teeth in Table 1, morphometrics in Table 3 and fin length in Table 4. Below,
the holotype counts are given first, followed, in brackets if different, by the paratypes’
counts.
Dorsal fins D VI–I, 9, spines slightly filamentous in males and less so in females,
spines 3 and 4 longer. First dorsal fin not contacting second dorsal fin basally (when
depressed). Anal fin I, 10. Pectoral fin rays 15. Caudal fin with 13 branched rays
with posterior margin slightly rounded. Pelvic disc with one spine and five strongly
branched rays. Scales in lateral series 35 (35–43) (Table 2); scales may extend midlat-
erally over the origin of first dorsal fin in male and second dorsal fin in female, and
posteriorly to hypural base. Scales usually ctenoid from hypural base to origin of the
second dorsal fin and cycloid elsewhere. Scales along dorsum usually extending ante-
riorly along medial base of second dorsal fin (may extend to base of first dorsal fin).
Scales in zigzag series 7 (6–10), transverse back series 13 (12–18), transverse forward
series 13 (11–16) (Table 2). Predorsal midline naked. Head, breast and pectoral base
usually naked. Belly usually naked or with few cycloid scales. Upper jaw teeth mostly
conical in the female with more and shorter teeth (10–15) than male (7–9) (mostly
caniniform). Lower jaw teeth conical in female (range 7–9) and males (4–6) (Table 1).
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2730 P. Keith et al.
Table 1. Number of upper jaw teeth in Sicyopus lord and related species.
Upper jaw teeth
6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
S. lord M111
S. lord F 111––1
S. discordipinnis M 1–1112–1
S. discordipinnis F 111231–1
S. jonklaasi M 3272– 2 1 1
S. jonklaasi F 1122–1
S. zosterophorum M 238221111–––1
S. zosterophorum F 11–13553352
S. auxilimentus M1
S. auxilimentus F 1
S. multisquamatus M
S. multisquamatus F1
S. exallisquamulus M 31––1 1
S. exallisquamulus F 21––1
S. cebuensis M1
S. cebuensis F 1
Notes: M, male; F, female.
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Journal of Natural History 2731
Table 2. Scale counts in Sicyopus lord and related species.
Lateral scales
18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47
S. lord 211–1–1–1
S. discordipinnis 1–11242211––1
S. jonklaasi 121452322111
S. zosterophorum 379101482
S. auxilimentus 1––1
S. multisquamatus 1
S. exallisquamulus 2–43
S. cebuensis 11
Predorsal scales
0123456789101112
S. lord 7
S. discordipinnis 121211
S. jonklaasi 25
S. zosterophorum 171323421142211
S. auxilimentus 2
S. multisquamatus 1
S. exallisquamulus 55
S. cebuensis 2
(Continued)
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2732 P. Keith et al.
Table 2. (Continued).
Transverse back series
4 5 6 7 8 9 101112131415161718
S. lord 122–1–1
S. discordipinnis 12743
S. jonklaasi 1–3966
S. zosterophorum 92813
S. auxilimentus 1––1
S. multisquamatus 1
S. exallisquamulus 1121––211–1
S. cebuensis 11
Transverse forward series
2 3 4 5 6 7 8 9 101112131415161718
S. lord 121111
S. discordipinnis 123343
S. jonklaasi 134563–1––1
S. zosterophorum 1–5551155751
S. auxilimentus 11
S. multisquamatus 1
S. exallisquamulus 13–211––11
S. cebuensis 11
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Journal of Natural History 2733
Zigzag scales
678910111213
S. lord 13111
S. discordipinnis 15 2
S. jonklaasi 11284
S. zosterophorum 20 30 3
S. auxilimentus 11
S. multisquamatus 1
S. exallisquamulus 451
S. cebuensis 11
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2734 P. Keith et al.
Table 3. Morphometrics in Sicyopus lord and related species expressed to the nearest whole percent of standard length.
Predorsal length
33 34 35 36 37 38 39 40 41 42
S. lord 1–13–11
S. discordipinnis 43541
S. jonklaasi 233872
S. zosterophorum 11211141771
S. auxilimentus 11
S. multisquamatus 1
S. exallisquamulus 21233
S. cebuensis 2
Preanal length
53 54 55 56 57 58 59 60 61 62 63 64 65
S. lord 12–1–11–1
S. discordipinnis 1–255111
S. jonklaasi 2159231–1
S. zosterophorum 11510111664
S. auxilimentus 1––1
S. multisquamatus 1
S. exallisquamulus 23–111––1
S. cebuensis 1–––1
Head length
22 23 24 25 26 27 28 29
S. lord 142
S. discordipinnis 564211
S. jonklaasi 54673
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Journal of Natural History 2735
S. zosterophorum 1 – 1 8 18 19 7
S. auxilimentus 2
S. multisquamatus 1
S. exallisquamulus 254
S. cebuensis 11
Jaw length
8 9 10 11 12 13 14 15
S. lord M1–2
S. lord F 3–1
S. discordipinnis M25
S. discordipinnis F37
S. jonklaasi M2493
S. jonklaasi F232
S. zosterophorum M 49623
S. zosterophorum F9156
S. auxilimentus M1
S. auxilimentus F1
S. multisquamatus M
S. multisquamatus F1
S. exallisquamulus M231
S. exallisquamulus F4
(Continued)
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2736 P. Keith et al.
Table 3. (Continued).
Caudal peduncle length
17 18 19 20 21 22 23 24 25 26
S. lord 31–12
S. discordipinnis 2662
S. jonklaasi 41533
S. zosterophorum 26101810521
S. auxilimentus 2
S. multisquamatus 1
S. exallisquamulus 14221
S. cebuensis 2
Caudal peduncle depth
8 9 10 11 12 13
S. lord 16
S. discordipinnis 881
S. jonklaasi 7162
S. zosterophorum 1 2 28 21 1
S. auxilimentus 11
S. multisquamatus 11
S. exallisquamulus 172
S. cebuensis 11
Body depth in males at origin of second dorsal fin
10 11 12 13 14 15 16 17
S. lord 12
S. discordipinnis 52
S. jonklaasi 4841
S. zosterophorum 46662
S. exallisquamulus 12111
S. cebuensis 1
Notes: M, male; F, female.
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Journal of Natural History 2737
Table 4. Fin lengths in Sicyopus lord and related species expressed to the nearest whole percent of standard length.
Second dorsal fin length
25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46
S. lord M21
S. lord F 2–1–1
S. discordipinnis M 1–231
S. discordipinnis F1531
S. jonklaasi M36621
S. jonklaasi F 11131
S. zosterophorum M 1––11122346111
S. zosterophorum F 12366623
S. auxilimentus M 1
S. auxilimentus F1
S. multisquamatus M
S. multisquamatus F1
S. exallisquamulus M 121––11
S. exallisquamulus F11–2
S. cebuensis M1
S. cebuensis F1
Anal fin length
23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41
S. lord M21
S. lord F111––1
S. discordipinnis M2221
S. discordipinnis F 11341
S. jonklaasi M 1–1–1251325
S. jonklaasi F 11–311
S. zosterophorum M149412
S. zosterophorum F 24466342
(Continued)
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2738 P. Keith et al.
Table 4. (Continued).
Anal fin length
23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41
S. auxilimentus M 1
S. auxilimentus F1
S. multisquamatus M
S. multisquamatus F1
S. exallisquamulus M 21111
S. exallisquamulus F1–3
S. cebuensis M1
S. cebuensis F1
Caudal fin length
16 17 18 19 20 21 22 23 24 25 26 27 28 29
S. lord M111
S. lord F111–1
S. discordipinnis M 11221
S. discordipinnis F 11233
S. jonklaasi M5643
S. jonklaasi F124
S. zosterophorum M 246642
S. zosterophorum F31781
S. auxilimentus M1
S. auxilimentus F1
S. multisquamatus M
S. multisquamatus F1
S. exallisquamulus M 121–11
S. exallisquamulus F211
S. cebuensis M1
S. cebuensis F1
Notes: M, male; F, female.
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Journal of Natural History 2739
Cephalic sensory pore system A, B, C, D, F, H, K, L, N and O, D singular, with
all others paired, oculoscapular canal separated into anterior and posterior canals
between pores H and K (Figure 1). Urogenital papilla in male long and thin with
fairly pointed to rounded tip. Urogenital papilla in female with two rounded lobes.
Colour in preservation
The sexual dichromatism is not well developed. Background of head and body grey-
ish. Body greyish dorsal to midline. Head greyish dorsally with small blackish dots.
A blackish band extending from the tip of snout and upper lip, under the eye to the
posterior edge of operculum and pectoral base, and from pectoral base to hypural.
Head whitish under this band and ventrally. Blackish horizontal band above the pec-
toral fin to the second dorsal fin base. Snout blackish. Nape greyish. Faint and thin
blackish band extending from pectoral base to caudal fin along body midline. First and
second dorsal fin hyaline, with several blackish dots. Dorsal fin spines highlighted by
black pigments. Caudal fin dusky to greyish, with three to four vertical blackish bands.
Anal fin dusky to hyaline with black pigments. Pelvic disc entirely white. Pectoral
fin dusky becoming whitish dorsally and ventrally. Pectoral base blackish medially,
slightly dusky ventrally and dorsally.
Colour in life
The sexual dichromatism is not well developed.
Body dusky to slightly yellowish. Three horizontal blackish bands on the flanks.
The first one from the tip of snout and upper lip, under the eye, to the posterior edge
of operculum and pectoral base, and from pectoral base to hypural. Colour above this
band iridescent gold in the last third of the body. Second black band from the back
of the eye to the caudal fin. Third one along the upper part of flank from above the
pectoral fin to the second dorsal fin base. Dorsal and anal fins dusky to hyaline with
black pigments on the rays. Caudal fin greyish with three to four vertical blackish
bands. Pectoral fins hyaline.
Belly of male fish whitish to greyish (Figure 2A); belly bright orange in gravid
female fish (Figure 2B).
Ecology
Like other Sicydiinae (see Keith 2003), S. lord is found in clear, high-gradient streams
with rocky bottoms. It lives on the bottom of the river, on top of rocks, but it is also
often seen swimming in open water in the current between rocks or in large pools. This
new species is supposed, like the other Sicyopus, to be amphidromous (Keith et al.
2008; Lord et al. 2010).
Distribution
Sicyopus lord is known from streams of northeastern Madagascar above an altitude of
20–80 m. Its status is unknown but the species is probably rare, as few specimens have
been seen in the area surveyed.
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2740 P. Keith et al.
a
b
N
O
ABC
D
FHKL
FHKL
N
Figure 1. Diagrammatic illustration of the head in Sicyopus lord (MNHN 2010-0953) showing
head pores and sensory papillae. (A) Dorsal view; (B) lateral view. Scale bars, 5 mm.
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Journal of Natural History 2741
ab
Figure 2. Sicyopus lord,(A)malein vivo (holotype, MNHN 2010-0925) (photo: T. Robinet-
DIAMSOI); (B) mature female in vivo (paratype, MNHN 2010-0953) (photo: T. Robinet-
DIAMSOI).
Comparisons
Sicyopus lord differs from all other Sicyopus in having a particular colour pattern, sim-
ilar in both sexes, with three blackish bands on the flanks. The first band extends from
the tip of snout and upper lip, under the eye to the posterior edge of operculum and
pectoral base, and from pectoral base to hypural; the second extends from behind the
eye to the caudal fin; the third one runs along the upper part of flank from above
the pectoral fin to the second dorsal fin base. In other Sicyopus species, there is well-
developed sexual dichromatism with bright colours in males. Sicyopus lord also differs
from all other Sicyopus in having a shorter caudal fin in male (it represents 16–18%
of the SL versus 19–29% SL for the other Sicyopus). Moreover, it differs from S. aux-
ilimentus,S. exallisquamulus and S. cebuensis in having more scales in lateral series
(35–43) compared with,respectively, 20–23, 18–21 and 26–27 and from S. exallisqua-
mulus,S. cebuensis and S. zosterophorum in having more scales in transverse back series
(12–18) compared with, respectively, 5–8, 8–9 and 10–12. It differs from S. multisqua-
matus in having a second dorsal fin with nine soft rays rather than 10 and fewer scales
in zigzag series (6–10 compared with 13), and from S. jonklaasi and S. discordipinnis
in having a first dorsal fin with six spines in females versus five spines.
Key to Sicyopus species
1. Second dorsal fin with 10 soft rays (I10); more than 45 scales in lateral series
................................................S. multisquamatus
Second dorsal fin with nine soft rays (I9); fewer than 45 scales in lateral
series ................................................................... 2
2. More than 29 scales in lateral series .................................3
Fewer than 28 scales in lateral series ................................4
3. Caudal fin length of male equivalent to 16–18% of the standard length (SL);
no sexual dichromatism ......................................S. lord
Caudal fin length of male equivalent to 19–29% of the SL; sexual dichromatism
with bright colours in males ......................................5
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2742 P. Keith et al.
4. More than 25 scales in lateral series; second dorsal fin length equivalent to
25–30% of the SL .......................................S. cebuensis
Fewer than 25 scales in lateral series; second dorsal fin length equivalent to
30–45% of the SL ..............................................6
5. First dorsal fin of females with six spines; caudal fin length equivalent to
24–29% of the SL ..................................S. zosterophorum
First dorsal fin of females with five spines; caudal fin length equivalent to
17–23% of the SL ..............................................7
6. Nine to ten scales in zigzag series .......................S. auxillimentus
Six to eight scales in zigzag series .....................S. exallisquamulus
7. Male brownish to yellowish with bright red lips; jaw length equivalent to
8–10% of the SL .......................................S. jonklaasi
Male reddish from second dorsal fin to hypural; jaw length equivalent to
10–11% of the SL ....................................S. discordipinnis
Etymology
The new species is named after our friend Clara Lord, for her extensive and enthusias-
tic work on Sicydiinae, and is defined as a noun in apposition.
Discussion
All sicydiine genera have a specific distribution. Some have a very restricted range
and some are more widely distributed. Sicyopterus is distributed in the Indo-Pacific
area from the western Indian Ocean to the eastern Pacific Ocean (Watson et al. 2000;
Keith et al. 2004; Keith, Galewski et al. 2005; Berrebi et al. 2006). Stiphodon and
Lentipes are distributed from the eastern Indian Ocean to the eastern Pacific Ocean
(Watson et al. 2002; Keith and Marquet 2007). Cotylopus is restricted to the western
Indian Ocean (Keith, et al. 2005a). Smilosicyopus is distributed from Indonesia to the
Marquesas Islands (East Pacific) and from southern Japan to New Caledonia, and
Akihito (Watson et al. 2007; Keith et al. 2007) seems to be restricted to the west of
the Pacific Ocean. Finally, Sicyopus was known, before this paper, from the eastern
part of the Indian Ocean to Micronesia (East Pacific) and from southern Japan to
New Caledonia (Watson et al. 2001; Keith and Marquet, 2005) in swift clear, high-
gradient streams; but it was not known from the western Indian Ocean (Keith et al.
2011). The discovery of the genus Sicyopus in the western Indian Ocean is of great
importance to the understanding of the evolution of the subfamily and the disper-
sal route used between the Indian and Pacific oceans. Indeed, as with many other
taxonomic groups (see Planes and Galzin 1997) the Sicydiinae subfamily probably
originated in the western Pacific as discussed by Parenti (1991) and Keith et al. (2011)
as 90% of the species inhabit this area, whereas the other 10% are from the Indian
Ocean, western and eastern Middle America and West Africa. Among the seven sicy-
diine genera occurring in the Indo-Pacific area, only three occur in the western Indian
Ocean: Sicyopterus (Indo-Pacific), Cotylopus (Indian Ocean) and Sicyopus, this paper
(Indo-Pacific).
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Journal of Natural History 2743
Keith et al. (2011), in their phylogeny of the Sicydiinae, estimate the divergence
for the Akihito and Sicyopus clade at 1.57 to 3.73 million years ago. But they could
not explain the dispersal of Sicyopus as the relative position of the genus in the phy-
logeny was poorly supported. It is probable that a complex system including marine
currents, duration of the larval phase, period of emergence of islands, vicariousness
and variation of the sea level, in addition to the peculiarities of Madagascar (altitude,
substratum, age, velocity of river flow), explain the dispersal of Sicyopus from the
Pacific to the western Indian Ocean and the presence of Sicyopus in this island. Among
the factors cited above, the pelagic larval duration has been most studied for the
understanding of dispersal. Arai et al. (2001) and McDowall (2007) suggested that the
oceanic life stages and larval duration of Sicydiinae could be regarded as key elements
in explaining their dispersal abilities and distribution. However, few studies have inves-
tigated the relationship between the migration strategies of Sicydiinae with prolonged
oceanic life stages and their distribution (Iida et al. 2010). Lord et al. (2010), working
on Sicyopterus species, have shown that the mean marine larval duration is signifi-
cantly longer for the widespread species, Sicyopterus lagocephalus, around 130 days,
than for species with a much more restricted range (around 80 days). All Sicydiinae
species that have been studied have a significantly longer pelagic larval duration than
the 30–50 days usually reported for reef fish (Wellington and Victor 1989; Wilson and
McCormick 1999), and the 20–50 days typically reported for marine gobies (except for
Gobionellinae) (Shafer 1998). The longer marine larval duration for amphidromous
species may be a developmental adaptation to their special and complex life cycle
that requires them to complete the two migrations to and from marine environments.
Larval phase duration could therefore partly explain the evolution of Sicydiinae and
the distribution range observed for each genus. A shorter marine phase would suggest
limited dispersal abilities, hence a restricted distribution range (Murphy and Cowan
2007). The species having a longer marine phase would be able to delay recruitment
(Victor 1986; Keith et al. 2008) and could therefore colonize distant islands, spreading
their geographic distribution to all islands, colonizing them gradually. These abilities
could have allowed some Sicydiinae species to reach other oceans over evolutionary
times, and particularly Sicyopus to reach Madagascar. But no pelagic larval duration
studies are available for Sicyopus species, and these are needed to compare with other
species. For eels, also diadromous fish, the larval phase is known to be shorter in trop-
ical ancestral species than in the most recent temperate species (Kuroki et al. 2006).
This plasticity in the duration of the larval phase might correspond to the expres-
sion of selected strategies, which are defined as genetically determined life histories or
behaviours (Robinet et al. 2007).
The variability of the duration of the marine larval phase could be one factor
explaining the species’ distribution by favouring or limiting their dispersal, but, as
suggested by Lord et al. (2010), length of larval duration is not sufficient to explain
species distribution. There are undoubtedly other factors of biogeographical (variation
of sea level and terrestrial barriers), physical (currents used), and ecological (behaviour
and swimming depth of larvae) origin that impact on species distribution range, and
particularly, on Sicyopus range. These must be studied.
Acknowledgements
We are grateful to Joe Aride from Madagascar National Parcs (MNP-ANGAP) and to the
manager of Masoala National Parc at Maroantsetra. We thanks the DIAMSOI team and
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2744 P. Keith et al.
particularly ARDA (P. Bosc and P. Valade), T. Robinet, Roger, and H. Grondin. Lastly, we want
to thank all the Responsible Chiefs of the areas concerned for their kind permission, and the
villages and communities who have always heartily received us and helped us in our prospecting
of rivers. This research was financed by the IFB (Institut Français de la Biodiversité) for the
DIAMSOI programme (2007–2010).
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