Identity of the Galaxioid Fishes of the Genus Aplochiton Jenyns
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Japanese Journal of Ichthyology 魚 類 学 雑 誌 Vol.34, No.3 1987 34巻3号1987年 Identity of the Galaxioid Fishes of the consistent with a report of a single elongate, Genus Aplochiton Jenyns from transparent juvenile of an undetermined species of Aplochiton from waters around the shores of Southern Chile Lago Villarica in central/southern Chile (Mc- Robert M. McDowall and Kazuhiro Nakaya Dowall, 1969a). Small juveniles, again of an undetermined species of Aplochiton were collected (Received November 6, 1986) from the sea in southern Chile (Puerto Aysen) by The genus Aplochiton Jenyns is one of two Mr. Akira Zama and reported on briefly (Mc- genera in the family Aplochitonidae. There are Dowall, 1984). two species of Aplochiton recognised from southern So little is known of these fishes that the capture South America and the Falkland Islands and, in of a small series of notably large Aplochiton in the addition, one species of Lovettia from Tasmania vicinity of Puerto Aysen is worth reporting and (McDowall, 1971a). A close phylogenetic rela- assists somewhat in clarifying the taxonomic tionship between Aplochiton and Lovettia should status of these fishes. not be assumed, although both genera clearly belong to the assemblage of galaxiid-aplochitonid Material examined fishes that is characteristic of cool southern tem- perate fresh waters (McDowall, 1969b; Fink, Aplochiton taeniatus: 2 individuals, HUMZ (Lab- 1984). The relationships between Aplochiton and oratory of Marine Zoology, Hokkaido University) Lovettia and their relationships with galaxiids are 095682, 095704, 21 January 1982, Fiordo Steele, Chile, not well enough understood to justify restructuring 48•K13'S, 73•K23'W, 361, 326.5mm SL. 1, SNP (PA) of these generic and familial arrangements al- (Servicio Nacional de Pesca (Puerto Aysen)) 76, 1 January 1982, Estero Ventisquero, Chile, 48•K15'S, though some authors present a contrived sim- 73•K35'W, 271mm SL. 1, HUMZ 095704, 1 January plification by placing all genera in Galaxiidae 1982, Fiordo Ventisquero, Chile, 48°13'S, 327 mm SL. (Nelson, 1972; Rosen, 1974). This does little to 1, Instituto de la Patagonica 46, 21 September 1977, clarify relationships. Estero Veto, Chile, 49•K32'S, 74•K30'W, 189mm SL. There are several nominal species of Aplochiton, A. zebra: 4, HUMZ 095696-9, 1 December 1982, the principle binomina being: Lago Quetro, Chile, 48•K7'S, 73•K7'W, 236-284mm SL. Aplochiton zebra Jenyns, 1842: collected from In addition data from the study of McDowall the Falkland Islands. (1971a) were reconsidered. Aplochiton taeniatus Jenyns, 1842: collected from Tierra del Fuego. Study methods Aplochiton marinus Eigenmann, 1928: collected from Estero Cutipai, near Valdivia, Chile. Study methods adopted were those consistently These fishes are poorly known. Their tax- in use for study of the southern freshwater salm- onomy was examined by McDowall (1971a) who, oniform fishes (Galaxiidae, Aplochitonidae etc.) somewhat tentatively, treated A. marinus as a most recently reviewed in McDowall and junior synonym of A. taeniatus, although this Frankenberg (1981). These methods, in general, action has not been accepted by Campos (1973), follow Hubbs and Lagler (1958); distinctive Bahamonde and Pequeno (1975), or Zama and features include: measurements taken point to Cardenas (1984). Neither Campos nor Baha- point with needle point dividers to the nearest monde and Pequeno gave reasons for their posi- 0.5 mm; fin ray counts include segmented rays tion but Zama and Cardenas discussed differences (excluding non-segmented procurrent rays); ver- in colouration between A. marinus and A. taeniatus. tebral counts exclude hypural centra. Biologically these fishes are also little known. Campos (1969) described the location of spawning, Meristic and morphometric variation and the development of the eggs of A. taeniatus in Lago Llanquihue, a southern Chilean lake. Systematics of fishes depends heavily on the He wrote of accumulations of juveniles in the use of meristic and morphometric variation to littoral zone of Lago Llanquihue-possibly an clarify variation within and differences between indication of shoaling juvenile stages. This is species. The salmoniform fishes, the southern •\ 377•\— 魚類学雑誌 Japan. J. Ichthyol. 34 (3), 1987 families like Aplochitonidae and Galaxiidae that were possibly diadromous. In one sample amongst them, are notoriously difficult in this (Lago Rinihue) vertebral number encompassed regard for several distinct reasons. 1. Meristic both ranges (59-65) suggesting that both dia- variation within populations tends to be high. dromous and non-diadromous stocks were present 2. Meristic variation between populations with in that lake. Campos (1974) demonstrated a the same life history strategy is also high. 3. similar broad range in vertebral number for Often there is clinal variation in features such as Galaxias maculatus (Jenyns) lower in that river vertebrae and fin ray numbers. 4. Furthermore, system which drains Lago Rinihue (Rio Valdivia- many salmoniforms have alternative life history Rio Calle Calle), although his bimodality did not strategies within species. Many species have both extend that far upstream. diadromous (sea migratory) and entirely freshwater Similar variability was also demonstrated for stocks and sometimes these stocks are sympatric A. taeniatus by McDowall (1971a) who described (Campos, 1974; McDowall, 1972, 1979; Northcote the following: Eye size decreased and snout length and Ward, 1985). The meristic features of di- increased with absolute size. Vertebral number adromous and non-diadromous stocks of a species was found to be bimodal with some populations may be distinctly different, eg. diadromous stocks at 62-66 and others at 69-73. The lower counts of Galaxias maculatus (Jenyns) may have 57 to were regarded as being from non-diadromous 61 vertebrae at the northern extremities of that stocks. Bimodality was also evident in gill raker species' range in New South Wales, Australia counts, but this feature subdivided the popula- (McDowall and Frankenberg, 1981) or up to tions into different subsets than those generated 60-66 in populations in far southern Chile by vertebral count. (McDowall, 1971b). Lacustrine populations have On the basis of vertebral count, the Rio Calle many fewer vertebrae, with 50-53 in northern Calle stocks (69-73) were different from all others Australian populations, but up to 54-60 in (62-66). Using gill rakers, however, the Rio southern Chile. Similar sorts of variation are Calle Calle fish grouped with most other popula- described for other galaxiids and also for Re- tions (17-23 gill rakers) fish from Lago Todos los tropinna (McDowall, 1979). Morphometric fea- Santos had higher counts (26-31) while those tures also tend to vary in relation to the above from Lago Llanquihue had fish belonging to both factors, differences between diadromous and non- subsets (21-23 and 30-34 gill rakers) . Thus diadromous stocks being particularly well re- various characters are varying in different ways cognised for characters like head length and eye between and within populations (in Rio Calle size. 5. In addition, allometric growth during Calle vertebral number is high gill raker number somatic growth of salmoniform fishes is widely low; in Lago Todos los Santos both are low; recognised in addition to which there is often ex- in Lago Llanquihue vertebral number is low but aggeration of some morphological features in gill raker numbers are both high and low). relation to the attainment of sexual maturity, In view of these sorts of variations, resolution especially in males. of the taxonomy and the identification of the All these problems relating to between in- new series of specimens is not simple. Two dividual, within and between population, and plausible scenarios can be envisaged. One is within and between species variability do or can that there is a complex of species (or perhaps be expected to apply to Aplochiton. In view of subspecies) of Aplochiton in the rivers and lakes the meagre knowledge of the life history strategies of Patagonian South America and the Falkland of these fishes, resolution of their taxonomy has Islands, for which existing material is inadequate its difficulties. to allow resolution. Alternatively there are two McDowall (1971a) has already examined some (A. zebra, A. taeniatus) or perhaps three (A. of these questions for a limited array of Aplochiton marinus) species of Aplochiton, which have al- samples. In A. zebra, it was shown that eye size ternative within-species life history strategies decreases with growth. Vertebral numbers in (diadromous and non-diadromous) and that there the samples appeared to be bimodal with number are morphological differences within species as- being low (about 57-62) in populations that ap- sociated with each of the alternate life history peared to be landlocked and high (63-66) in those strategies. Material available for study and •\ 378•\— McDowall and Nakaya: Galaxioids from Chile knowledge of the life histories of Aplochiton Identification species do not permit absolute clarification of these hypotheses. Material in the present study is easily divided As knowledge of within-species morphological into two taxa (see Table 1). variation related to similarly alternative life history Differences are quite clear in body depth, shape strategies in several species of Galaxias (McDowall, of the caudal peduncle, the sizes of the dorsal 1972) and Retropinna (McDowall, 1979; Northcote and anal fins, head length and depth, eye diameter, and Ward, 1985) is entirely consistent with the mouth size and other features. The four more types of variation discussed above for Aplochiton slender large-mouthed fish (Fig. 1) are typically we see no reason to modify the taxonomic ar- heavily covered on the upper sides and back with rangement of Aplochiton proposed by McDowall small round spots, and in some there is a general (1971a) in which two species A. zebra and A. peppering of small widely spaced melanophores. taeniatus were recognised and redescribed. They have 72-73 vertebrae. When alive the fish Table 1. Morphometric and meristic variation in Aplochiton species (figures given as percentages of denominators in ratios).