Jean H. Huber*
* Laboratoire d'Ichtyologie générale, Museum national d'Histoire naturelle, 43 rue Cuvier, 75231 Paris Cedex 05, France.
Abstract: The identity and status of a since long forgotten taxon, Haplochylus lacazei Rochebrune, 1885, are discussed. Because the description does not permit to allocate lacazei to any known Cyprinodont species, the author proposes to fix the status of the available name lacazei as an invalid name, following the new article 23.9 of the Code of the International Commission of the Zoological Nomenclature, valid from January 1. 2000. Knowing Rochebrune's works and the Gambian Cyprinodont fauna, and as the most reasonable conservative measure, lacazei might be considered as a junior synonym of Epiplatys spilargyreius, and then it would belong to the subgenus Parepiplatys (sensu Huber, 1998).
Résumé: L'identité et le statut d'un taxon oublié depuis longtemps, Haplochylus lacazei Rochebrune, 1885, sont discutés. Parce que la description ne permet pas d'allouer lacazei à une espèce connue de Cyprinodonte, l'auteur propose de fixer le statut du nom utilisable lacazei comme un nom invalide au sens du nouvel article 23.9 du Code de la Commission Internationale de Nomenclature Zoologique, valide depuis le premier Janvier 2000. Connaissant les travaux de Rochebrune et la faune Cyprinodonte de Gambie, lacazei pourrait être un synonyme junior d'Epiplatys spilargyreius en tant que mesure conservatoire la plus raisonable et alors il appartiendrait au sous-genre Parepiplatys (sensu Huber, 1998).
Introduction. Eschmeyer (1998), in his monumental "Catalogue of Fishes" (1), has remarkably pinpointed the availability of disregarded or simply forgotten publications of old times, which comprise descriptions of taxa at the species or at the generic level. This is the case of Rochebrune's paper (1885) which deals with several new species said to have been collected in Gambia and elsewhere in western Africa. One of these may relate to Cyprinodonts, Haplochylus lacazei, and the purpose of this paper is to try to identify that taxon on the basis of all the information available and propose for it a systematic status.
Analysis of the genus name and considerations on the putative belonging of lacazei to the Cyprinodonts. The genus Haplochilus Agassiz, 1846 (Haplochylus, erroneously by Rochebrune) is an unjustified emendation of Aplocheilus Mc Clelland, 1839. This name was constantly used to designate non deep-bodied forms of tropical Cyprinodonts and describe new species from both the Old and New Worlds, up to 1934 (Haplochilus carlislei Horst): over 50 taxa, mainly in Epiplatys but also in Aplocheilichthyins, have been originally described in that genus (Huber, 2000a). Rochebrune was well aware of that name and of the morphology of the fishes it represents, for two reasons:
- already before 1885, 8 taxa had been described from the Old World: H. fasciolatus Günther, 1866; H. infrafasciatus Günther, 1866; H. playfairii Günther, 1866; H. rubropunctatus Steindachner, 1867; H. senegalensis Steindachner, 1870; H. bifasciatus Steindachner, 1881; H. chaperi Sauvage, 1882; H. antinorii Vinciguerra, 1883;
- in 1882, Rochebrune himself studied infrafasciatus, fasciolatus and spilargyreius; and he placed the latter in Haplochilus, the first and third species being surprisingly synonymized.
Therefore we may reasonably acknowledge H. lacazei as a Cyprinodont in relation to the experience of the above taxa, but nothing in the following description can demonstrate this with full certainty.
Another dimension of doubt is raised from Rochebrune himself and from his works themselves. Rochebrune was, even for his years of activity, an unusual generalist, dealing with very many families of zoology: for example in his 1885 paper, he describes many new species among which are mammals, birds, reptiles, amphibians and not less than 11 new fishes of various families from Casamance, the Mellacorée river, the Congo River, Loanda, Bathurst and/or all Gambia. Rochebrune, from the region of Angoulême, southwest of France, published first in botany (1860-1862), then he participated to some archeological field research (from 1865), lived in Saint-Louis (Senegal) as a colonial medicine doctor between 1875 and 1877 and finally became assistant naturalist in Malacology at the Paris M.N.H.N. in 1879. Up to 1885, he dealt with fishes among other animals and was criticized about the weakness of his diagnoses (see Rochebrune, 1885; and comments on papers from the Zoological Record). In addition, his new species from various families and origins came from traders in Paris (e.g., Louis Petit, from various regions of western Africa, including "Landana") who were regularly showing him their novelties, usually before selling them: this explains the poor quality of information regarding collection origins and also the lack of types (the Paris M.N.H.N. holds only two types of fish by Rochebrune from his 1880 paper and none from the 1885 paper). The only regions in which he specialized and sampled were the Senegambia. Between 1885 and 1904, Rochebrune only published in Malacology and notably the coming detailed follow up that he announced in his 1885 paper with Senegambia distribution details and the full diagnosis of his new taxa never occurred. It is not known or understood why he withdrawn from that project, but he was accustomed to that behaviour.
Original description and its analysis. The next step in the identification of that taxon is to analyse the description which appears to be in Latin (translation into french by Prof. Daget; author's comments between brackets).
"24. Haplochylus lacazei. D= 10; A= 16; LL= 26".
"Body compressed posteriorily from the Pectoral fins insertion; body depth four times in (standard) length (i.e., depth= 25% in S.L.); head strongly flattened, three and a half times in (standard) length (i.e., head length= 29% in S.L.); eye diameter, three and one fifth times in head length (i.e., eye= 9% in S.L.); short subprotractile snout, hardly equal to eye diameter (snout= 7-8% in S.L.). Dorsal fin insertion at the level of the eighth Anal fin ray (i.e., D/A= +8); last rays of Dorsal, Anal and Ventral fins are filamentous.
Upper part (of body), purple red with yellow markings. Lower part (of body), yellow orange, with vermilion red markings and six, intensively blue, bars. Dorsal fin, yellow with scattered blue dots. Caudal fin, yellow with a blue bar near to its base. Anal fin, yellow, with a blue border. Iris, blue.
Maximal size (total length): 7.5 cm (near to 3 inches)".
"Habitat: marshy biotopes, near a location named Landana".
"Registered in Bouvier's personal collection".
"We are pleased to dedicate this very beautiful species to the knowledgeable Professor H. Lacaze-Duthiers".
"It is very distinctive from its congeners".
The first comment is that Bouvier's collection is probably lost: Eugène-Louis Bouvier (1856-1944) was a Professor of Entomology at the Paris Museum (MNHN) from 1895 till 1931. In 1885, he was only 29 year old in the early steps of his career. No Bouvier's collection is registered in the Paris MNHN. Therefore, types (if any) may be hypothesized as lost.
The second comment concerns the type locality of Landana. Investigations by Prof. Daget (pers. comm., November 1999) disclosed that Louis Petit (cf. supra) sent birds to Bouvier that were said to come from Landana, too. A priori, it may be hypothesized that Landana is in Senegambia, as many other fishes of Rochebrune's paper of 1885 e.g., Isistius marmoratus from "Landana and the entire Gambia". Moreover, Rochebrune's main focus was Senegambia and nowhere else in Africa. Landana should not be far from the sea, because, in the same paper, other fishes are quoted with the same location and a habitat in mangroves. However, Landana does not exist in the present maps of Gambia, nor in the place-name changes (Room, 1993), nor in the Gazetteers of Gambia, Sénégal, Guinea and Guinea-Bissau. The embassy executive of the Gambian State in Paris, M. Jallow, was kind enough to ask for an in-depth historical and geographical research of that locality name to the Gambian civil servants, but the search was not conclusive: Landana may be a mislabelling or an erroneous transcription for Latre Kunda, a village close to mangroves in the surroundings of Banjul, the capital city and single harbor of Gambia. An alternative option is to enlarge the search outside Senegambia, on the coast and this is conclusive: Lândana (geographical coordinates in hundredth of degrees: 5.22S; 12.13E) is today a village, near mangroves in Cabinda, northwestern Angola, about 50km southeast of Pointe Noire in Congo. However, senior ichthyologists in M.N.H.N. discard this hypothesis, because Rochebrune never deposited material from that region and that his paper of 1885 was the last he ever published on fishes. None the less, let's explore the Portuguese Landana as the correct type locality: lacazei's description partly corresponds to the single Epiplatys species of Cabinda, which reaches 7 cms and exhibits bars on sides. It is yet an unnamed component of the multifasciatus superspecies or identical to ansorgii (sensu Wildekamp, 1996) or to boulengerior to huberi, reported from not too far away, in southwestern Gabon/Congo. The meristics of the three taxa (D= 9; A= 15; D/A= +10; LL= 30) somewhat fit with lacazei, except the scale count in lateral series; however, all components of the multifasciatus superspecies exhibits more than 7 bars on sides (usually 12 to 14) and these bars are black, never blue; besides, the color on the upper sides is not red but yellow or green. Equally, an undisclosed "blue-barred" component of the genus Aphyosemion in Cabinda, would show associated red bars on sides that are not mentioned for lacazei. Then, the most reasonable hypothesis for the type locality of lacazei is that it lies not far from Gambian mangroves of those days: near the sea or upstream the Gambia river influenced by tide (remembering that mangroves were then more extended than today).
The third comment concerns the description itself which is clear and detailed but not comprehensive at all and, within the scope of Senegambian Cyprinodonts, may refer to a male of a member of the genus Epiplatys or of the subgenus Scriptaphyosemion in Callopanchax or of the annual fishes of Fundulosoma or Nothobranchius; this discards the other Cyprinodont lineage of that region, the Aplocheilichthyins, because of their colors (no red or green colors and no dark bar in Aplocheilichthys spilauchen, or Poropanchax normani); the five Cyprinodont taxa of the above genera which are presently known, according to Killi-Data (Huber, 2000a) from the Gambian lowlands (and also Senegal) are (their size as T.L. and their average major meristic counts, D, A, D/A, LL, are given between brackets for comparison with lacazei): Cal. geryi (T.L.= 5cm; D= 13; A= 17; D/A= +5; LL= 33); Ep. bifasciatus (T.L.= 6cm; D= 8; A= 16; D/A= +10; LL= 27); Ep. spilargyreius (T.L.= 5cm; D= 8; A= 17; D/A= +11; LL= 27); Fundulosoma thierryi (T.L.= 3.5cm; D= 11; A= 15; D/A= +1; LL= 26); Nothobranchius kiyawensis (T.L.= 4.5cm; D=14 ; A= 14; D/A= -1; LL= 26). From the above diagnosis, it appears that the last two taxa can be excluded because of the lower D/A ratio and the smaller size and the first taxon, too, because of the higher Dorsal and LL counts.
The fourth comment, and a problem with the description of lacazei, is that it fits with some characteristics of the two Epiplatys from Senegambia, but not all: it presents more fin ray counts at Dorsal and Anal, which are consequently inserted less apart (a minor issue, though). Haplochylus lacazei shows the characteristics of filamentous fins which are to be seen in old dominant males of bifasciatus, however no fasciated pattern is reported for bifasciatus throughout its entire huge range; besides, it shows the key characteristics of fasciated pattern (blue oblique bars) of spilargyreius, however neither such red colorful pattern of male nor filamentous extensions are reported in spilargyreius throughout its entire huge range. The matter of filamentous rays is though not critical and it can be an artefact: when a fish is not properly fixed, its fin rays are stuck together and can simulate such a condition (e.g., Pellegrin has described in 1913 Haplochilus senegalensis var. acuticaudata on such a basis, but it turned up to be a mere synonym of spilargyreius).
The fifth and last comment is that the description even fits better with a population of Epiplatys fasciolatus. Indeed, fasciolatus shows the dark blue bars on sides and peduncle (although more than six plus one), the characteristic yellow markings on upper sides and red markings of lower sides, the extended last rays on fins and notably ventrals, the less slender form of body than the known Epiplatys species of the region, the larger size and the higher meristics (T.L.= 9cm; D= 12; A= 17; D/A= +9; LL= 28). Unfortunately, the present distribution of Ep. fasciolatus ranges from Liberia to Guinea (Wildekamp, 1996) and its most northern colllecting place is (Huber, 2000a) near Bagoro (10.52N; 13.55W) in western Guinea, about 400 km, south of Banjul, the main Gambian city and harbor, on the Gambia river mouth and putative approximate type-area of lacazei(13.47N; 16.65W).
If it is accepted, in spite of the lack of types or of a figure in the original description, that Haplochylus lacazei is a Cyprinodont, then our present knowledge of Rochebrune's works strongly favors its belonging to the genus Epiplatys.
From the incomplete description, from the ambiguous type locality and from our presently insufficient knowledge of the Senegambian or Cabindan Cyprinodonts, seven hypothesis may be forwarded:
- its identity with Epiplatys spilargyreius, as a junior synonym, and in that case some parts of the description are inacurate; knowing Rochebrune's works, in general, these inaccuracies are minor, notably the deviations in size and depth;
- its identity also with Epiplatys bifasciatus, as a junior synonym, and in that case several parts of the description are inacurate; if Rochebrune's knowledge of spilargyreius is tentatively thought comprehensive, then this second hypothesis is likely because he did not know that taxon described from the Nile basin and bifasciatus is very abundant in Gambia; however, the deviation in the color pattern appears to be too high;
- its identity, also as a junior synonym, with Epiplatys fasciolatus which fits the best with the description. In that case, collecting localities have to be discovered about 400km northwesterly to the most northern collections of fasciolatus but this possibility appears very unlikely to experts of the region;
- its validity as a distinct northern component of the fasciolatus superspecies that has to be collected for the second time after its description; the collection in western Africa of relict species in pocket regions linked to hypothesized refugia (Huber, 1998), such as Epiplatys barmoiensis or Ep. njalaensis or, if valid, Ep. lokoensis, does not make this hypothesis too theoretical; it seems, though, very unlikely to Etzel (1999 and comm. pers.), the expert of Gambia, a region which has been sampled a lot for Cyprinodonts and appears to be poor because it is situated at the northern most distribution of the Epiplatys genus (if it has not become extinct in the mean time). Detailed field surveys in the putative distribution of lacazei, near the coast of Senegambia, say between Dakar and Casamance, may only clarify that issue but it is an hazardeous venture;
- its possible identity with a component of the Epiplatys multifasciatus superspecies if the type locality, Landana in Cabinda, would be accepted as accurate and if the "7 blue" bars were changed into "12 to 14 black" ones: all its components have been described after lacazei's creation and the proper one would then be a junior synonym of lacazei;
- its status as a nomen dubium (or nomen vanum), as a consequence of its insufficient description data which prevent from clearly assigning the taxon to a known species.
A further eighth hypothesis would have been worth studying: its status as a nomen oblitum (or forgotten name), as a consequence of its non usage during more than one hundred years; however, this concept is now rejected by zoologists due to deviations in its usage (Huber, 2000b, in print) and it is unfortunate in that case.
Because it lacks traceable types, or a drawing of them, a proper description (by modern standards) and a clear and non ambiguous type locality, lacazei may be best considered as a nomen dubium and a probable junior synonym of Epiplatys spilargyreius, within the subgenus Parepiplatys (sensu Huber, 1998). Although this proposal does not tackle all possible issues, it was the one chosen for the first finalisation of the manuscript (late Summer 1999), when the author was informed of the coming publication of a new Code of Nomenclature which would tackle some cases of forgotten names.
We propose here to apply the new article (23.9) of the Code, valid from January 1. 2000, and to regard lacazei as an invalid but available name, especially but not solely for the unlikely case when lacazei could be a senior synonym of Ep. multifasciatus or any other subsequent taxa of that superspecies: multifasciatus (and the other) are names which have been used as valid by all authors in many publications during at least the past 50 years and cannot be displaced by an earlier synonym which has not been used as valid since 1899 (in fact quoted only once without status, by Eschmeyer, 1998); "when an author has discovered that such conditions exist, and has published a statement of this fact which cites the relevant article (23.9) and gives appropriate evidence", then the later names (namely, multifasciatus, ansorgii, boulengeri or huberi) in prevailing use permanently takes precedence (without a ruling by the Commission) over the earlier but disused synonym ( lacazei).
Acknowledgements. We wish to thank the following individuals for advice and information, or for reviewing an earlier draft of this paper: J. Daget (MNHN, Paris), M.L. Bauchot (MNHN, Paris), and the anonymous reviewers of the manuscript. By courtoisy, the manuscript has been sent for information to the experts of the fishes of that region, V. Etzel (Cuxhaven), A.C. Radda (Vienna) and R.H. Wildekamp (Gemert) and their comments have been valuable.
Dedication. This paper is very friendly dedicated to Prof. Jacques Daget: it would have been fair that he co-authored it, owing to his decisive contribution. However, the subject interested him so much that he decided to start writing a full article to cover Rochebrune's life and his "new" taxa, in addition to lacazei (in prep.). At eighty year old, Prof. Daget is the memory of the Museum and is still very active in research and publications (his last book, last year, covers a major topic of Malacology). I shall never stress enough my debts to him as a researcher and as a man, since he kindly accepted to be assessor to my thesis, back in 1978.
Eschmeyer, W.N. 1998. Catalogue of Fishes. Calif. Acad. Sciences, 2805 pp.
Etzel, V. 1999. Gambia -abseits der Touristenpfade. Aquaristik aktuell, 9-10: 56-62, 13 figs.
Huber, J.H. 1998. A Comparison of Old World and New World Tropical Cyprinodonts. A parallel Outlook of similar and distinctive Characteristics regarding Distribution, Evolution, Ecology, Behavior, Morphomeristics. Soc. fr. Ichtyologie Ed., Paris (Oct. 10): 109 pp., 17 figs.
Huber, J.H. 2000a. Killi-Data 2000. Updated checklist of taxonomic names, collecting localities and bibliographic references of oviparous Cyprinodont fishes (Atherinomorpha); in french, english, german and spanish. Cybium, Soc. fr. Ichtyologie, Ed. , Paris: 538 pp.
Huber, J.H. 2000b. On nomina oblita among Cyprinodont species. J. Amer. Killifish. Ass., in print.
Room, A. 1993. Place-Name Changes (1900-1991). The Scarcecrow Press, Inc. Metuchen, N.J. and Canada, 293 pp.
Rochebrune, A.T. de. 1885. Vertebratorum novorum vel minuscognitorum orae Africae occidentalis incolarum. Diagnoses. Bull. Soc. Philomatique Paris, 7 (9): 86-89.
Wildekamp, R.H. 1996. A World of Killies. Atlas of oviparous cyprinodontiform fishes of the world, vol. III, American Killifish Association Publ., Mishawaka, Indiana, 330 pp.
Paris, January 1999-February 2000.
(1) I wish to thank warmly Prof. Eschmeyer to have listed my name as a contributor to the "Catalogue of Fishes", a monumental scope (55000 taxa of fish species, reported in total); for a further edition on CD-Rom, the cooperation has been extended and a special list of Killi-Data 2000 prepared for his team so that the oviparous Cyprinodonts be covered to the latest update.
Illustrations: fig. 1. Idealised sketches of the male of the 5 discussed Cyprinodont species of Gambia (Callopanchax geryi, Epiplatys spilargyreius, Ep. bifasciatus, Fundulosoma thierryi, Nohobranchius kiyawensis), of Ep. fasciolatus from Guinée-Liberia, and of the species of Mbosso River, near Cabinda, Ep. ansorgii, by R.H. Wildekamp.
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