J.H. Huber *
* author's address : Laboratoire d'Ichtyologie, Museum national d'Histoire naturelle, 43 rue Cuvier, 75231 Paris Cedex 05, France.
Note: this article has been initially written in English and kindly translated into German by Wolfgang Eberl, then D.K.G.'s editor; for its first publication in August 1998 in:
Das Aquarium, 32, 350 (Aug.): 15-23.
The English version is kindly published herein in Killi News by the British Killifish Association (B.K.A.) to open its access to English speaking readers.
Abstract: A new species of the subgenus Chromaphyosemion Radda, 1971 is described, Aphyosemion alpha n. sp. from Cap Estérias in Northwestern Gabon, at the most Southern part of the subgenus distribution. It is diagnosed by a distinctive colour pattern character -males and females show, on the pre-and-post opercular region, a unique alpha-shaped marking drawn in red, like the Greek letter (hence the name!)- and by some minor morphomeristical differences, especially the dorsal fin deviation over the anal fin. Results of crossings and of caryotyping separate the species from the other members of the Chromaphyosemion subgenus. Another member of the subgenus from the Engong Kouamé area in the interior of the coastal plain is not named because it lacks significant differences with A.loennbergii s.l. (Boulenger, 1903), according to our present diagnostic tools.
Aphyosemion alpha, une nouvelle espèce du sous-genre Chromaphyosemion avec un patron de coloration distinctif et une distribution à l'extrême limite méridionale.
Résumé: une nouvelle espèce du sous-genre Chromaphyosemion Radda, 1971 est décrite, Aphyosemion alpha n. sp. du Cap Estérias au Nord-ouest du Gabon, à l'extrême limite méridionale de la distribution du sous-genre. Elle est caractérisée par un trait particulier de la coloration -les mâles et les femelles présentent, sur la région pré et post-operculaire, une tache en forme d'alpha comme la lettre grecque (d'où le nom!) colorée en rouge- et par certaines différences morphoméristiques mineures, en particulier la position relative de la dorsale par rapport à l'anale; les résultats de croisements et les caryotypes séparent l'espèce des autres membres du sous-genre Chromaphyosemion. Un autre membre du sous-genre du secteur de Engong Kouamé, à l'intérieur de la plaine côtière n'est pas nommé, en l'absence de différences significatives avec A.loennbergii s.l. (Boulenger, 1903), selon nos outils diagnostiques actuels.
Contrary to most Aphyosemion species, the history of this fish belongs to local aquarists from Libreville : actually, at the end of my discovery trip of 1976, together with Dr Alfred C. Radda throughout Gabon, I met M. Merssemann who showed in his aquarium, with other indigenous fishes, two Chromaphyosemion species which he had collected "more than three years before": a blue form (the new species) and a red form "from inland". The history of collections of the blue form by European Killi-hobbyists is long but concerns only a pocket handkerchief area between Cap Estérias and Cap Santa Clara: Herzog and Bochtler (Germany, 1975), Pürzl (Austria, 1985), Buon, Mallet and Ragot (France, 1986), Nümrich (Germany, 1988), Fourdrinier (France, 1988), Harz (Germany, 1992), Legros and Cerfontaine (Belgium, 1993), together with Eberl (Germany), who also caught the fish with Passaro (Germany, 1993, 1994). Although this history is long, the formal description of the fish has been delayed due to the difficulty of its breeding (sex-ratio strongly unbalanced in favour of males), which precluded a serious hybridisation programme until the strain was better established. However, I was personally attached to the distinctive position of this fish since I used in my earlier publications the derived naming "affinis": A. aff. splendopleure. During this interval, roughly from 1975 until to-day, I was supported by many aquarists and notably by the three renown experts on the Chromaphyosemion subgenus, Daniel Poliak (France), Olivier Legros (Belgium) and Wolfgang Eberl (Germany). These three wrote respectively in 1981-86, 1990-1995, 1996, remarkable detailed papers on the distributions, colour patterns, breeding and maintaining, and history of collections of the various valid components of the subgenus: A. bivittatum (Lönnberg, 1895), A. loennbergii (Boulenger, 1903), A. riggenbachi (Ahl, 1924), A. bitaeniatum (Ahl, 1924), A. splendopleure (Meinken, 1930), A. volcanum Radda & Wildekamp, 1977 (?) (1), A.poliaki Amiet, 1991, A.lugens Amiet, 1991, distributed from Togo to Ecuatorial Guinea. Without their support, their valuable observations in aquarium and their advice, this description would not have been possible and they are warmly thanked. Moreover, the species name "alpha" derives from a pertinent observation and suggestion by Olivier Legros and we are pleased to honour him that way. It is no use to repeat their findings here and the reader is referred to their publications quoted in the bibliography. I would also extend those thanks to the additional active breeders, Maurice Chauche (France) and Jean-Paul Cicéron (France) who took patience and time to prepare the crossing experiments. All these dynamic and enthusiastic aquarists have patiently waited the too long -but wise- time necessary to finalize this paper, as usual in institutional circles ! The manuscript has benefited from positive contributions by the 3 quoted experts, by Prof. Jean-Louis Amiet (now retired in the South of France), whose diagnostic key of the Cameroonian Chromaphyosemion species still stands firm (1987, 1991), by Dr. Glen E. Collier (Tulsa University, U.S.A.) who kindly caryotyped the species for us, by Prof Jacques Daget (M.N.H.N, Paris, France) and by Univ.-Prof. Dr Alfred C. Radda (Wien, Austria) and by anonymous reviewers. They all receive my gratitude.
Synonym: A. splendopleure (non
Meinken, 1930); Radda, 1975.
Code name: Aphyosemion sp. n°2; Legros, 1990; Eberl, 1996.
Holotype: MNHN 1994-1114, a male of 32.9mm S.L. and 42.8mm T.L. from P.K.17.1 of the road Libreville airport (Hotel Gamba) to Cap Estérias, locality code LEC 93/26, Northwestern Gabon. Legros, Eberl, Cerfontaine, Coll. January 14. 1993.
Paratypes: MNHN 1994-1115, 5 specimens from the same locality and collected with the holotype.
Paratypes: MNHN 1997-183, 14 specimens, MRAC 97-44-P-1-6, 6 specimens, UFRJ 3879, 6 specimens, ANSP 176180, 6 specimens, BMNH 1922.214.171.124-6, 6 specimens, ZMB 32774, 6 specimens; all from the same creek as the holotype, but collected later (code PEG 94/48), Northwestern Gabon. Passaro and Eberl, Coll. August, 1994.
Derivatio nominis: alpha, an invariable noun in apposition, refers to the typical red marking similar to the Greek letter.
Diagnosis: A rather large Chromaphyosemion species, larger than all other components except riggenbachi and showing the most advanced Dorsal insertion (D/A= -1 compared to between 0 and +4); a strictly blue phenotype, with a unique red marking situated on the sides, apart from the opercle, with the two dark longitudinal mood-driven bands being conspicuous, with a red dotted Dorsal, a flamed Caudal and an "unmarked" Anal, except a thin red submargin and white margin, with a more trapezoidic form of vertical unpaired fins and a Caudal with short streamers, in male; female, also with the alpha marking: with conspicuous dark bands and an immaculate Anal, but no dark reticulation around the scales on sides.
Colour in life: male, strongly blue on sides and fins, with rather numerous red dots along sides, in Dorsal and upper Caudal; in all specimens, the "alpha" mark is prolonged by two continuous, variable, short lines of red dots on sides; the Caudal fin is red flamed; a red submargin and a white or light blue margin, both thin, are present in Ventrals, Anal, and lower Caudal; the excited male shows a yellow orange zone from lower snout to mid Anal level of lower sides; streamers, if any, are extremely short and light yellow to white; the dark shield on lower lip and lower snout is extremely conspicuous, nearly as in Epiplatys. Female, overall brown with a few lines of darker spots and with the two black longitudinal bands often visible on sides, with an immaculate anal fin, a punctuated dorsal fin and an orange zone on lower caudal fin. The alpha diagnostic marking is also present, to the contrary to all known females of other Chromaphyosemion species (Legros, pers. comm.).
Color in alcohol: male, with the same markings than in life except the blue and yellow colours; in addition, all fins are darkly flamed along rays; a dark reticulation on upper sides from ahead of dorsal insertion up to caudal peduncle; a dark band on lower sides, near base, from behind the anal base ending up to the caudal peduncle; the lower lip, dark and conspicuous. Female, not known fully adult, but not distinguishable from the other species of the subgenus, apart from the Anal which is free of markings.
Morphomeristic data of the first quoted six types, all males, the holotype first in bold (after confirmation by radiophotographs): D= 13, 12, 12, 13, 12, 12 (mean: 12.33). A= 14, 14, 13, 14, 15, 14 (mean: 14.00). D/A= -1, -1, -1, -1, -1, -1 (mean: -1). LL= 26+1; 26+2, 25+2; 25+1; 26+2; 26+2 (mean: 25.67 + 1.67). Predorsal scales= 12, 13, 12, 13, 12, 13 (mean: 12.5). Transversal scales (TRAV.)= 9, 9, 9, 9, 8, 8 (mean: 8.67). Circumpeduncular scales (CIR)= 14, 13, 14, 14, 14, 14 (mean: 13.83). S.L. (in mm)= 32.9; 26.2; 25.9; 26.1; 30.0; 29.8. T.L. (in % of S.L.)= 130%, 129%, 125%, 127%, 122%, 127% (mean: 126.7%). P.D. (predorsal length)= 57%; 60%, 60%, 62%, 57%, 61% (mean: 59.4%). P.A. (preanal length)= 64%, 62%, 63%, 62%, 60%, 62% (mean: 62.1%). P.V. (preventral length)= 48%, 49%, 50%, 48%, 46%, 48% (mean: 48.1%). Height at Anal level= 19%, 19%, 17%, 17%, 16%, 18% (mean: 17.7%). Height at peduncle level= 12%, 11%, 12%, 12%, 12%, 13% (mean: 12.1%). Head length= 30%, 33%, 30%, 31%, 28%, 30% (mean: 30.3%). Interorbitar= 14%, 15%, 16%, 14%, 15%, 15% (mean: 14.8%). Eye diameter= 8%, 8%, 8%, 9%, 9%, 9% (mean: 8.7%). Snout= 7%, 8%, 8%, 9%, 7%, 8% (mean: 7.9%). Vertebrae (abdominal+caudal)= 11+14, 11+14, 11+13, 11+13, 12+13, 11+14 (mean: 24.7). The dorsal fin is inserted nearer to the snout than any other Chromaphyosemion species (D/A negative, -1 and even -2 in some specimens; shorter predorsal length); this may be linked to the shape of the dorsal fin, more like a trapezoid than a triangle. Teeth are conical and standard for the subgenus. The neuromast system is fully open and standard; the frontal scalation is of the normal G type; no ctenoidy on scale or fin has been disclosed; scales on the belly are much smaller than on sides.
The biotope is typical of the subgenus Chromaphyosemion: a shady forest brook ("marigot") with crystal clear water on sand or fine gravel; the water is of low conductivity, slightly acid, very similar to rain water. The Cyprinodonts living in the area are, according to Eberl (1996), Aphyosemion australe, A. microphtalmum, A. striatum, Epiplatys sexfasciatus, Ep. singa and Procatopus (Plataplochilus) ngaensis; all species are found in slightly different subniches: e.g. A. australe and A. striatum prefers warmer and less clear part of the biotope (dead leaves), A. alpha, the clear parts on sand, the juveniles and subadults of the Epiplatys species dwell more open parts, the Procatopus species is schooling in deeper and flowing waters. At the type locality, on January 14, 1993, the biochemical data were (noon time): air temperature: 25°C; water temperature: 23°C; pH: 5.7; water colour: strongly amber; hardness (dH): 0; conductivity: 25µS. The fish have been collected with Barbs, softwater shrimps, A. australe, Ep.sexfasciatus: 59 specimens with only one subadult pair and 57 fry of a few millimetres size, which turn out, all, in males in aquarium. In aquarium, the new species is, according to experts, difficult to maintain due to the relative aggressiveness of males, compared to other Chromaphyosemion . According to Legros (1995), the maintenance was difficult and the productivity low with the 1988 strain collected by Fourdrinier; it is somewhat better with the LEC 93/26 strain, provided that only one male is placed with females at the same time in the aquarium: else, fin bites between males are frequent. Productivity is low compared to the easier populations of A. bitaeniatum from the Lagos area . Eggs are collected from floating and sinking mops; they are relatively larger than those of bitaeniatum and incubation time last the standard 2-3 weeks at 21°C; better crops are obtained with a pH value of between 4.5 and 5: below 4, eggs fungus and fish get sick with Oodinium frequently; fortunately, these constraints with the wild stock have been less stringent with the F1 generation; the fry eats first Artemia nauplies, like for all Chromaphyosemion; growth is very slow. The greatest difficulty is the odd sex ratio of offspring: nearly 100% of males with O. Legros, B. Drake (a U.K. aquarist) and several other killi-hobbyists in France, Germany and U.K. However, A. Cerfontaine, also from Belgium, obtain nearly 100% of females, for unknown reasons, and exchanges permit the maintenance of the strain over time.
The systematics of the genus Aphyosemion and especially of the subgenus Chromaphyosemion is made difficult, due to much stronger variability in genotypes than in phenotypes (Scheel, 1974; 1990) . For example, the number of haploid chromosomes and of arms (between parenthesis) of the splendopleure component vary from 13 (22) to 17 (25); in light of this, the ichthyologist can only name pragmatically those components with a minima a distinctive stable character, usually a chromatic one in the pattern of males, as discussed in details in Killi-Data (Huber, 1996) (2). The caryotype of A.alpha has been kindly studied for us by Glen E. Collier (pers. comm. in E-mail: July 16, 1996): it is distinctive from all other known components of the subgenus as published by Scheel (1990) and, in particular, from bitaeniatum and splendopleure, within the same systematic subgroup (details will be published by him later). The following crossing experiments have been undertaken; all are negative and confirm the deviation in genotype:
· male A. alpha - Cap Estérias X female A. bitaeniatum - Zagnanado: no development of the eggs, despite the proven fertility of parents (Legros, pers. comm. June 1995);
· male A.alpha - Cap Estérias (wild) X female A.loennbergii -Apou (C89/30): a few developed eggs which fungus rapidly in spite of several exchanges in breeding pairs; a lot of non developing fungused eggs (Legros, pers. comm. August 1994);
· male A. alpha - Cap Estérias X female A. splendopleure - Tiko (near the type locality): about 40 eggs which developed and hatched very (too?) quickly, within 10 days; the fry died after 48 hours or less, not being capable to swim or feed; eyes not visible; vital organs probably handicapped (Ciceron, pers. comm. December,1995);
· male A.alpha -Cap Estérias (wild) X female A. aff. loennbergii -Engong Kouamé, Gabon (LEC 93/24): many eggs with embryo, a few fungused; longer time of incubation (3-4 weeks); many embryos, with a huge vitelline bag, die within a few days and the available fry is crippled (crescent shaped) and dies after 2-3 days; several unsuccessful crossings, despite the repeated exchange of partners (Legros, pers. comm. August, 1994);
· male A. alpha - Cap Estérias X A.loennbergii - Kribi (near the type locality): no development of the numerous eggs, despite the proven fertility of parents (Chauche, pers. comm. April, 1996).
Because all components of the Chromaphyosemion subgenus are isomorphic and allopatric, i.e. cryptic species like for most other Rivulin superspecies, it is impossible to set their relationships based on morphology or meristics. Even caryotyping techniques have fallen short of this goal of phylogenetic relationships within a monophyletic group, like a superspecies, because of the large number of rearrangements; DNA techniques are very promising here but their results are too preliminary and fragmentary. To-day, it is only possible to compare and group similar phenotypes, with similar colour pattern characteristics within a superspecies . My preferred method for comparison is not based on photography (risky and inducing errors), but on direct facing of males, then females, in a small aquarium, two by two, then altogether, to exemplify instantaneous differences in temporary or (excited) permanent patterns . This has been undertaken in Olivier Legros's premices for all Chromaphyosemion species, but lugens, which was not available . Based on these results and on Legros's observations, he prepared in a comparative table for us, the Chromaphyosemion subgenus may be split into 4 artificial subgroups:
1- bitaeniatum - splendopleure - volcanum (a simple variation of splendopleure?) alpha: in male, no or very few markings in Anal, more conspicuous dark longitudinal bands on sides, lines on snout shield and post-opercular wound mark; less numerous small dots on sides; female, with similar key features; A.alpha is the largest component of the subgroup; this is the single subgroup living all along the distribution.
2- loennbergii (and pappenheimi, if valid, according to Scheel, 1990) - riggenbachi: in male, a more or less densely red spotted or flamed Anal (and Caudal, plus Dorsal); generally weak longitudinal dark bands on sides; females show a dotted Anal in all populations; in riggenbachi, the largest component, the fry and juveniles do not show any dark longitudinal bands; this subgroup is distributed parallely to the previous one, but more in the interior of the coastal plain; not living far Northwest of the Sanaga River in Cameroon; the second Gabonese Chromaphyosemion species belongs here but cannot be separated from the Cameroonian phenotype of loennbergii.
3- bivittatum - lugens: a more melanophoric subgroup; male, with black or dark red blotches on (mainly upper) sides and unpaired fins near base, instead of dots or small spots; the longitudinal dark bands are conspicuous in both sexes; in female, Anal is virgin but not in male; the distribution is restricted to hilly regions of Biafra and the foothills of the Cameroonian plateau.
4- poliaki: strong red reticulation around scales of sides and all fins with many red dots, in both sexes; male, with gold yellow inside the reticulation on sides and no spot, but a strong post-opercular blotch; the distribution is restricted to the foothills of the Mont Cameroun volcano (may be fused with subgroup 3, in the future).
A. alpha is known only from a few localities, some kilometres distant, between Libreville and Cap Estérias . This does not imply that the species is relict to that very small area: actually, the more we know on Rivulin distribution (including the South American forms) the more it appears unclassical, i.e. made of discontinuous and isolated groups of populations, but the reason or rational is obscure. Therefore, the exploration of the regions North of Cap Estérias and South of the Komo estuary may provide new populations. The Cyprinodont fishes reported above from the same area are all present near the coast in Northern Gabon and Southern Ecuatorial Guinea: all but two (A. microphtalmum and Ep.singa) are known only from North of Lambaréné, where a change in fauna occurs . Northerly, in Central Ecuatorial Guinea, another change of fauna occurs for unknown reasons: A. alpha is replaced by a poorly known form of A.splendopleure, A.australe by A. ahli, Ep.singa by Ep. grahami and Ep. sexfasciatus by Ep.infrafasciatus and Aplocheilichthyins (or Procatopodins) of the subgenus Plataplochilus by those of the subgenus Procatopus. All these data suggest but do not prove that the distribution of A. alpha is similar, i.e. restricted between these two limits to the South and to the North. Easterly, it is replaced by the Chromaphyosemion sp., named here loennbergii, but their limits of distribution are obviously unknown since both species have only been collected in a few spots, geographically well separated.
Despite the difficulty of disclosing diagnostic characters in the Rivulin cryptic biospecies, i.e. for an isomorphic allopatric component of a given superspecies, as in Chromaphyosemion, A. alpha has been described on key characters which ascertain the concept of the biological species in practical terms as discussed in Killi-Data (1996):
· it is distinctive by at least one stable character in comparison with the other members of the superspecies: here, the red marking on the opercle and by the pure blue "phase" (or colour morph of other authors), especially on fins;
· it has a separate genotype demonstrated by the study of the caryotype and, in the future, of DNA contents: here, the number of haploid chromosomes is not shared by the other components;
· it is sterile in crossing experiments with the other components, especially those of its own subgroup; to be complete, a crossing experiment with A. riggenbachi, also with more trapezoidic vertical unpaired fins would be welcome;
· its distribution is clearly defined at its borders or by the sympatric fauna: here, it is replaced in Central Ecuatorial Guinea, at least, by a distinctive phenotype, not separable from A. splendopleure; besides, like in Cameroon, it is replaced in the interior part of the coastal plain by a component of A. loennbergii subgroup.
It is hoped that the delicate beauties of this species will be a spur for killi-hobbyists to go collecting and enlarge its distribution Northerly and, why not, Southerly, in the direction of Port Gentil.
(1) Radda (pers. comm. June 1997) when reviewing this paper has informed us that he has submitted an article which challenges Amiet's (1987), Legros's (1991e) and Eberl's (1996) opinions that volcanum is a junior synonym of splendopleure; he brings herein an argumentation in favour of volcanum being a valid species and poliaki being a subspecies of volcanum.
(2) Amiet (pers. comm. June 1997) when reviewing this paper makes the following remark of high interest: "in Ichthyology, the morphomeristic criteria are always put forward, but one must not forget that this is due, first, to the passing of the colours of the fishes when placed in alcohol ! Actually for organisms with a vision like Aphyosemion, colour criteria are THE good ones since they serve to interspecific recognition in normal conditions (I do not speak here of the prison-like conditions of the aquarium). When one is able to use the same chromatic criteria than the animals themselves for recognition, luck is there and isomorphy is nothing to regret. Besides one may argue that precisely the efficacy of the chromatic recognition may have laid in Aphyosemion evolution to "save" the morphological differentiation sensu stricto".
Amiet, J.L. 1987. Faune du Cameroun. 2. Le Genre Aphyosemion Myers. Sciences Nat. Compiègne: 262pp., 76 pls.
Amiet, J.L. 1991. Diagnoses de deux Espèces nouvelles d'Aphyosemion du Cameroun (Teleostei, Aplocheilidae). Ichthyol. Explor. Freshwaters, 2 (1): 83-95, figs.
Eberl, W. 1996. Die Untergattung Chromaphyosemion. D.K.G. Journal Supplementheft. NT 4: 88pp., figs.
Huber, J.H. 1981. A Review of the Cyprinodont Fauna of the Coastal Plain in Rio Muni, Gabon, Congo, Cabinda and Zaïre, with taxonomic Shifts in Aphyosemion, Epiplatys and West African Procatopodins. B.K.A. Publ.: 46 pp., 27figs., 16 photos.
Huber, J.H. 1996. Killi-Data 1996. Updated checklist of taxonomic Names, collecting Localities and bibliographic References of oviparous Cyprinodont Fishes (Atherinomorpha). Cybium, Soc. fr. Ichtyologie Ed., Paris: 400pp., 20 maps.
Legros, O. 1990a,b,c. Le sous-genre Chromaphyosemion. Killi Contact A.K.F.B. (4): 93-111; (5): 123-130; (6): 157-172.
Legros, O. 1991a,b,c,d. Le sous-genre Chromaphyosemion. Killi Contact A.K.F.B. (2): 7-18; (3): 9-19; (4) 12-25; (5): 7-16.
Legros, O. 1991e. Diskussion über die Gültigkeit der Art Aphyosemion volcanum. D.K.G.- Journal, 23 (4): 56-61, fig.
Legros, O. 1992a,b. Le sous-genre Chromaphyosemion. Killi Contact A.K.F.B. (1): 2-13; (2): 7-20.
Legros, O. 1995. Nouvelles populations de Chromaphyosemion. Killi Contact A.K.F.B. (6): 7-24.
Poliak, D. 1981. A. bivittatum. Killi Contact A.K.F.B. (4): 75-80.
Poliak, D. 1982a. A. multicolor. Killi Contact A.K.F.B. (1): 13-24.
Poliak, D. 1982b. A. volcanum. Killi Contact A.K.F.B. (3-4): 65-78.
Poliak, D. 1985a. A. riggenbachi. Killi Revue K.C.F. (1): 13-22.
Poliak, D. 1985b. A. loennbergii. Killi Revue K.C.F. (2): 13-23.
Poliak, D. 1986a. Le droit à la différence. Killi Revue K.C.F. (1): 10-19.
Poliak, D. 1986b. A. splendopleure. Killi Revue K.C.F. (1), fiche technique N° 96.
Radda, A.C. 1975. Contribution to the Knowledge of the Cyprinodonts of Gabon with the Description of four new species and one new subspecies of the Genus Aphyosemion Myers. B.K.A. Separatum, 20pp., 12 pls, tab.
Scheel, J.J. 1974. Rivuline Studies. Taxonomic studies of Rivuline Cyprinodonts from tropical Atlantic Africa. Ann. Mus. Roy. Afr. Centr. Tervueren, ser.8, Zool., No 211: 148 pp., 18 figs.
Scheel, J.J. 1990. Atlas of the Killifishes of the Old World. T.F.H. Publ., Neptune City, 448 pp., figs.
Illustrations from Eberl (1996), courtoisy of D.K.G.: male, PK 16.3, Fourdrinier Coll. Photo M. Chauche (p66) male, PK 17.1, Passaro and Eberl Coll. Photo W. Eigelshofen (p67, lower) distribution of A. alpha, A. aff. loennbergii (p69) head detail drawing with the alpha marking (p70) biotope of LEC93/26 with Olivier Legros netting (p12, lower)
Paris, March 1994 - October 1996.
Additional reading of this author's publications ? Select any ONLINE article.