Barbus sensitivus, a new species with extensive pitlines from the Sanaga River (Teleostei: Cyprinidae).
A number of small Old World (African and Asian) cyprinid fishes or barbs are known with extensively developed "sensory papillae" especially on the head. The African ones are documented in papers by Herre (1936), Barnard (1948), Dejen et al. (2002), Greenwood (1962), Loiselle & Welcomme (1971), Loiselle (1973), Stewart (1977) and Tweddle et al. (2006). The sensory papillae usually are restricted to, or most highly developed on the lower surfaces of the head. In some, as in the new species described here, they also occur on the body. They have not been observed on the fins.
In Africa these fishes generally are placed in the genus Barbus, a catchall taxon with about 300 species recognized as valid (Daget et al. 1984; Eschmeyer 2010). Several of these African Barbus species sometimes were referred either to Enteromius or Beirabarbus because of their superficial sensory papillae but this practice has waned and currently they are all assigned to Barbus (Eschmeyer 2010).
In Asia there are no cyprinids assigned at present to the genus Barbus, but there are at least three barb-like genera with extensively developed rows of cephalic sensory papillae large enough to be seen with the naked eye: Cyclocheilichthys, Eirmotus and Oreichthys (pers. obs.). As with the African barbs having them, these structures have not yet been examined with scanning electron microscopy.
Another point about the presence of possibly similar exposed ciliated mechanoreceptors in a variety of Asian as well African barbs is that they may have evolved independently, hence may represent an ancient recurrent adaptive feature of cyprinoids. Just how ancient is unclear. Some African and Neotropical dischodontid characoids may have similar organelles. These also have not been adequately studied.
Experimental observations indicate that exposed neuromasts facilitate feeding on zooplankton in crepuscular (poorly lit) conditions by larvae of the cyprinid Gnathopogon elongatus caerulescens (Mukai et al. 1994; Mukai 2006). Larvae of a variety of distantly related cyprinids including Cyprinus carpio have exposed pitline organs distributed on the head and body like those in B. sensitivus. I suggest that the presence of exposed pitline organs in various lineages of Old World barbines, Asian as well as African, may result from a widespread neotenic tendency to retain exposed pitline organs usually present only in larval stages. In other words, the presence of pit lines and of exposed pitline organs in adults of diverse Old World cyprinids may result from numerous instances of independent retention of a larval characteristic. For this reason the present new species is assigned to the genus Barbus. At such time as the African barbs are generically split, Enteromius and Beirabarbus are available names, and it may be necessary to recognize them both as valid. Certainly none of the tropical African barbs are truly congeneric with the highly distinctive European genus Barbus.
Barbus sensitivus n. sp.
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Holotype (Figs 1-2): CAS 95344, 40.6 mm, nontuberculate female, Sanaga River at Belabo, 60 km NW of Bertoua, Cameroon, 4[degrees]56'N', 13[degrees]18'E, February 1980, Tyson R. Roberts. The river was very swift, with bottom of coarse gravel and smooth water-worn stones.
Paratypes: CAS 95345, 2: 38.1-40.7 mm, tuberculate males, collected with the holotype (40.7-mm specimen used for SEM examination).
Diagnosis: Barbus sensitivus differs from all other small species of African Barbus by its very slender caudal peduncle. It is distinguished from the great majority of small African barbs by its exceptionally well developed sensory papillae or pitline organelles which cover most of the head in both sexes. Similar pit lines occur on the scales covering the body (not known in other African barbs, but perhaps under-reported). Males differ from those of most other species of African barbs of similar size in having the rostrum and dorsal surface of the head covered with numerous breeding tubercles.
Description: That B. sensitivus is a small barb (and does not grow to a much larger adult size than the three specimens reported here) is indicated by the marked sexual dimorphism of breeding tubercles on the heads of the two males (paratypes) versus their absence in the single female (holotype).
Body subcylindrical, laterally compressed, terete. Head moderately large and pointed. Snout narrow when viewed from the side, but short and blunt viewed from above. Dorsal profile of head evenly convex; no nuchal hump; ventral profile of head from tip of lower jaw to angle of preopercle very straight. Mouth termino-inferior, small, rictus of jaws lying anterior to a vertical through from margin of eye. Eyes large, oval, dorsally inclined; orbital length more than one-third head length. Nostrils moderately large, lying entirely on half of snout nearest eye. Rostral and mandibular barbels well developed, rostral barbels slightly longer than maxillary barbels, about equal to eye diameter. Rostral barbels originating almost at tip of snout, extending posteriorly to just beyond origin of maxillary barbels. Maxillary barbels extending posteriorly from below middle of eyes to slightly beyond them, their anterior halves when adpressed accommodated by deep postlabial groove.
Lateral line complete, nearly straight, slightly curved downwards anteriorly. Scales in lateral line series 33-35, each with single small lateral line canal pore near middle of its exposed portion. Scales of entire body nearly uniform in size and shape, with radiate striae; scales of lateral line series slightly enlarged. Posterior scale margin semicircular or very slightly arched. Predorsal scales 9. Scales absent on ventral body surface behind head, including isthmus, to slightly beyond bases of pectoral fins. Median abdominal scale rows, from just behind pectoral fin bases to pelvic fin bases, 9. Transverse scale rows 5/1/2-3. Circumpeduncular scales 9.
Dorsal fin very slightly falcate, with 3 simple and 8 branched rays. Simple rays very slender and flexible, last simple ray not serrated. Dorsal fin origin well in advance of vertical line drawn through pelvic fin origins. Anal fin noticeably more falcate than dorsal fin, with 3 simple and 5 branched rays. Pectoral fins with one simple and 12-15 branched rays. Pelvic fins with one simple and 7 branched rays. Body depth declining more or less regularly from its maximum near dorsal fin origin to end of anal fin base, then evening out; hence transition to slender caudal peduncle very abrupt. Caudal peduncle elongate and slender, its length 2.6-2.8 times its least depth. Caudal peduncle depth 13.4 times in standard length. Caudal fin deeply forked, with ovate lobes. Principal caudal fin rays 10/9.
Vertebral counts: The female holotype and intact male paratype were radiographed in the California Academy of Sciences Department of Ichthyology by Jon Fong. The female has 22+17=39 vertebrae (Fig. 2), the male 21+18=39.
Coloration in life: The three specimens (female holotype and two male paratypes), collected in two seine hauls in the late afternoon and observed alive only briefly an hour or two before dusk, were white or silvery laterally and ventrally with strong bluish reflections. No evident markings on head, body or fins (midlateral stripe, faint in preserved specimens, not noticed). Fins color-less. No sexual dichromatism observed.
Color pattern of preserved specimens: Head, body and fins without any distinctive or striking marks, melanophores on fins hardly visible to unaided eye. Head below middle of eye and body below lateral line scales row without any marks and almost devoid of melanophores. Numerous fin melanophores on dorsum of snout and near nostrils, on posterior part of cranial roof over expanded portion of brain, and on opercular insertion. Fine melanophores along occipital margin where anteriormost scales insert. Fine melanophores forming thin mid-dorsal stripe between occiput and dorsal fin origin. Fine melanophores in elliptic or crescentic patch covering posterior half of exposed part of each scale above lateral line scale row; very few much larger melanophores confined to small area near anteriormost exposed part of each scale (anteriorly) or to larger area of each scale (posteriorly). Faint midlateral longitudinal stripe closely associated with lateral line canal for its entire length composed of few large melanophores just above and below canal on anterior half of body, then same number or slightly more melanophores more evenly distributed over and near canal on its posterior half. Triangular base of caudal fin overlying hypural fan covered with fine melanophores.
Dorsal fin with small patch of larger melanophores just in front of and beside its origin. Dorsal fin rays with small melanophores (sometimes forming very thin continuous dark streak) along anterior and posterior margins of branched and unbranched portions, dorsal fin interradial margins otherwise without melanophores. Anal fin pigmentless except for few melanophores. Paired fins devoid or almost devoid of melanophores. Few tiny melanophores along inner margin of outermost (i.e. simple) pectoral fin ray, pectoral fins otherwise without melanophores. Pelvic fins pigmentless. Caudal fin with fine melanophores on edges of rays, sometimes also on lepidotrichial joints, rendering them more conspicuous.
Genital papilla: The holotype and only known female specimen, apparently adult, has a pronounced genital pad (Fig. 1). This might indicate that only one or a few eggs are laid at a time and that the eggs are relatively large.
Distribution: Barbus sensitivus is known only from the three specimens collected in the Sanaga River in 1980. No additional specimens were collected during later ichthyological survey work in Cameroon (Randall E. Brummett, Cameroon Fish Project, World Fish Center, pers. comm., 4 Jan 2007).
Etymology: The name "sensitivus" is from the Latin sintio, sensus, to perceive by the senses or feel, in reference to the numerous pitline sensory organelles characteristic of this species.
SEM observations of epidermal structures
Methods: The three specimens of the new species were collected alive and immediately preserved in 10% formalin. They were not removed from the formalin until five years later, when one male specimen was selected and processed for SEM observation. Dehydration was done by gradually bringing the specimen up from deionized water to absolute alcohol. The specimen was then cut in half at the middle of the dorsal fin, the halves freeze-dried and gold-coated. The anterior half was mounted on a stub by its cut surface with its head projecting straight up, and the posterior half with the tail projecting straight up. Micrographs were taken on the left side of the specimen except for overall views of the ventral surface of the head and dorsal view of snout.
Breeding tubercles: The two male paratypes have exceptionally large and numerous breeding tubercles for such a small species, another character which may distinguish B. senstitivus from most other small African barbs (Figs 3-5). The tubercles on the head occur mainly on the rostrum and on the dorsal surface of the snout. Those on the rostrum have their apices directed laterally. In the one male studied with SEM, the dorsal rim of each nostril has five fairly large tubercles in two groups, an anterior group of three somewhat larger tubercles and a posterior group of two much smaller tubercles. A very few additional small tubercles are found on the snout between the nostrils. Particularly striking is the large number of shallow tubercle craters on the snout and posteriorly on the head between the eyes where almost no developed tubercles are present. This particular specimen may have recently shed most of the tubercles on the dorsal surface of its head posterior to the rostrum and nostrils. This suggests that it may have recently been in spawning condition, spawned, and then shed most of the tubercles previously present.
The largest tubercles, on the snout, about 120 micra high and 300 micra in diameter, are composed of several hundred cells, while the small tubercles on the pectoral fins are composed of about two dozen cells and are only 20-30 micra high. They are arranged in 4-5 rows on each interradial membrane. Epidermis on the dorsal surface of the fin rays is non-tuberculate, consisting of flattened hexagonal cells with well developed microvilli or microvillar ridges with an average of about one pore (presumably a mucus pore) for every two cells.
Tubercles of basically similar morphology, consisting of relatively small numbers of epidermal cells, occur on the dorsal surface of the pectoral fins of the males. The female has no tubercles on the head or on the pectoral fins.
Pit lines. The pit lines with their sensory papillae or ciliated pitline organs are even more evident than the tubercles. They are apparently restricted to the head and body (absent from barbels and fins). On the head they occur mainly in grooves or furrow on the ventrolateral surface. There also are numerous smaller pitline rows with much smaller pitline organelles on the dorsal surface of the head. These were not adequately recorded during the SEM work and are not included in the following quantitative observations.
Counting pitline organelles on the ventrolateral and ventral surfaces of the head of the 40.7 mm male yielded 1389 on the left side of the head. Doubling this number provides an estimated total number of 2,800 pitline organelles on the ventral and ventrolateral surfaces of the entire head.
Pit lines are present on all of the scales in the lateral line scale row and the scale row immediately above it, on nearly all of the scales on the scale row directly below it and on a few of the scales in the second and third rows above the lateral line scale row (Fig. 6). A direct count of pitline organs on the scales on the left side of the body of the 40.7 mm male yielded 903. Allowing for about a dozen missing scales in the middle of the body, including a few lost when the body was cut in two or covered with mounting media, the estimated total for the left side is 1,000-1,050. Doubling these numbers gives an estimated total of 2,000-2,100 pitline organs on the body.
Distribution of the pit lines on the head apparently follows a highly complex pattern not only characteristic of adults of B. sensitivus but also of the larvae of some other cyprinids. The main characteristics of these are: 1) a major system with a single continuous main line parallel to the lower mandibular-posterior circumorbital margin, from which numerous lesser curved lines radiate towards the eye; 2) a smaller continuous main line parallel to the lower margin of the anterior circumorbital margin, with similar curved lines radiating on the side of the mouth in front of the eye; 3) three or four large nearly vertical lines directly upwards from the ventral margin of the preopercle; 4) a number of variable lines directed upwards from the ventral margin of the opercle; and 5) an elongate oval patch of numerous close-set small pitline organelles on the ventral gular area between the mandibular rami.
The distribution and morphology of the pit lines and pit line organelles of adult B. sensitivus are similar to those of the pit lines and their organelles in larvae of Cyprinus carpio (Applebaum & Riehl 1997), Gnathopogon elongatus caerulescens (Mukai & Kobayashi 1995) and Danio rerio (Webb & Shirey 2003).
Status of nominal African barbin genera Beirabarbus and Enteromius: Beirabarbus Herre, 1936 (type species Beirabarbus palustris Herre, 1936, by original designation and monotypy) was distinguished by its author primarily on the basis of having cephalic pit lines. It was recognized as a valid subgenus of Barbus by Greenwood (1962: 193), who noted the presence of pit lines on the scales, and identified the pitline organs as neuromasts based on unpublished histological observations by P. J. Whitehead. If Beirabarbus is recognized, B. sensitivus might be referable to it.
Enteromius Cope, 1869 (type species Enteromius potamogalis Cope, 1869, by original designation and monotypy) was identified as a senior synonym of Beirabarbus by Karaman (1970: 189). Karaman based this synonymy on the identification of E. potamogalis as a junior synonym of Barbus (=Beirabarbus) ablabes Bleeker, 1863. This identification originated with Boulenger (1911: 156) and is mistaken. My examination of 8 of the 17 original syntypes of E. potamogalis deposited in the Academy of Natural Sciences of Philadelphia (ANSP 7607) indicates that it is distinct from B. ablabes, although it might be a Beirabarbus. Pit lines on the head are relatively few, and there seem to be none on the scales.
Beirabarbus also was recognized as a senior synonym of Enteromius by P. H. Greenwood (Greenwood 1970: 10-12), because the type species of both nominal genera or subgenera had a "peculiar" cephalic pitline system. This system still has not been adequately studied in any of the species referred to these nominal genera. The possibility that it evolved independently or that it re-appeared repeatedly in adults of small African barbins, and that therefore those species possessing it may not form a phyletic lineage, apparently has not been considered before now.
Type locality and trivial name of Enteromius potamogalis: A long-standing confusion confusion about the type locality of E. potamogalis (Daget et al. 1984; Bohlke 1984; Eschmeyer 2010) should be clarified. A remark also is in order concerning the origin of the specific name "potamogalis". In the original description Cope stated "This species was procured by the African traveler, P. B. Duchaillu [sic], in streams and rivulets fifty to sixty miles north of the equator, and the same distance from the ocean." This location has been identified as "Gaboon? (Daget et al. 1984: 219) and as "French Congo, streams and rivulets 50 to 60 mi N of the equator, and the same distance from the ocean" (Bohlke 1984: 87). Bill Eschmeyer simply cites the type locality as provided by Cope: "Streams and rivers 50-60 miles north of the equator and same distance from the ocean, West Africa" (Eschmeyer 2010).
The American Paul Belloni Du Chaillu (1831-1903) is known chiefly for his early explorations in Gabon and Rio Muni, during which he discovered numerous undescribed birds and mammals and obtained the first complete skeletons and skins of Gorilla gorilla. Narrative accounts of his African travels occur in two books, Explorations and Adventures in Equatorial Africa (1861) and A Journey to Ashango-Land: and Further Penetration into Equatorial Africa (1867). The first described a journey in July-August 1856 up the river Rio Muni and on up the Muni tributaries and on up its tributaries the Ntambonay (=Temboni or Utamboni) and Noonday (Du Chaillu, 1861, Chapter IV). Of his recorded travels, this corresponds best to the description "50 to 60 miles N of the equator and an equal distance from the ocean." Part of the Ntambounay travelled by Du Chaillu is in fact only about 60 miles N of the equator. At this point he would have been only some 20 or 30 miles inland, but anyone familiar with cross-country travel in this densely forested and very rocky country will know how difficult it is to calculate precise distances travelled, as Du Chaillu explains in the preface to the second book on his travels in equatorial Africa (Du Chaillu 1867: vii).
That he recorded distances travelled carefully is evident from Du Chaillu's description of his travel on the river Ntambounay. Shortly after 27 July 1856 he entered the Ntambounay, ascending 20 miles in an ESE direction, then 29 miles up the it and into the Noonday River, which he then ascended for an additional 12 miles (Du Chaillu, 1867). Modern maps show that the Ntambounay (or Tamboni or Utamboni) courses for some 20 miles ESE before it bends abruptly to ENE. At this point it is about 60 miles N of the equator and 40 miles inland. Fish or fishing activities are mentioned at least twice in this part of the narrative, including an incident shooting fish with a rifle which caused a stampede of elephants. Du Chaillu did not actually record preserving any fish specimens.
The type specimens of E. potamogalis apparently are the only fish specimens he collected during this trip; at least no other specimens were deposited in the fish collection of the Academy of Natural Sciences of Philadelphia (W. B. Saul, pers. comm., 10 July 1997). The river Rio Muni is in what is now officially known as the Republic of Equatorial Guinea (formerly the country of Rio Muni). Nearly the entire country is occupied by the basin of the river Rio Muni. Although Karaman (1970) suggested it is a junior synonym of Barbus camtacanthus, Enteromius potamogalis probably is a valid species, perhaps endemic to the Rio Muni basin, or to the Rio Muni and Ogooue and smaller river systems adjacent to them. It may be referred to as Barbus potamogalis.
In naming his new species Enteromius potamogalis, Cope (1869) did not provide any explanation for the generic or species terms. "Enteromius" presumably refers to the intestine, which Cope considered to be comparatively short. He did not mention the cephalic pitline system. The species or trivial name "potamogalis" evidently refers to a highly distinctive fish-eating mammal, the West African water shrew Potamogale velox Du Chaillu (1868). Cope presumably knew of the fish-eating habits of this animal directly from the explorer also from the fine print of a Potamogale on the bank a river with a large barb in its forepaws on the title page of Du Chaillu's second book on his African travels.
The Ministry for Scientific and Technological Research of the Federal Republic of Cameroon granted permission to the author to do fieldwork and collect fish specimens for scientific study in 1980. Fieldwork was funded by a grant from the Fonds Leopold III pour la Conservation et la Exploration de la Nature; the late Jean-Pierre Gosse kindly helped me obtain this grant. The resulting collections are deposited in the California Academy of Sciences. The paper was prepared in the Department of Ichthyology of the California Academy of Sciences and in the Center for Conservation Biology, Institute for Molecular Biosciences, Mahidol University, Nakhon Pathom, Thailand. The holotype was photographed and the holotype and one paratype radiographed by Jon Fong in the Department of Ichthyology, California Academy of Sciences. Scanning electron microscopy was done by Darryl Ubick and the author using the Academy's SEM facility. Funding for SEM work was provided by the Research Fund of the California Academy of Sciences; the author is grateful to William N. Eschmeyer and Patrick J. Kociolek for arranging this funding. Jacqueline F. Webb, Biology Department, Villanova University, commented on an early version of the manuscript, particularly on identification of the pitline organelles.
Received: 29 June 2010 - Accepted: 10 August 2010
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Tyson R. Roberts
Research Associate, Smithsonian Tropical Research Institute. Email: email@example.com
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|Author:||Roberts, Tyson R.|
|Publication:||aqua: International Journal of Ichthyology|
|Date:||Oct 15, 2010|
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