Review of the Species of Achias (Diptera : Platystomatidae)
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Inverteba Tauon, 1994, 8, 117-281 Review of the Species of Achias (Diptera :Platystomatidae) David K. McAlpine Entomology Section, Australian Museum, Box A285, Sydney South, N.S.W. 2000, Australia. Abstract The genus Achias Fabricius occurs in New Guinea, northern Australia, and some islands adjacent to these areas. Evidence is presented as to the disputed publication dates for Hendel's two significant reviews of the Platystomatidae as these affect nomenclature. Available information on Achias biology is summarised, but includes no larval records. A method of estimating the approximate number of undiscovered species is presented, and indicates that the register of Achias species is very incomplete. A description of the genus Achias is given. Generic synonyms and means of generic determination are discussed. Certain taxonomic characters are discussed and these are related to taxonomic placement of the genus, informal infrageneric groupings of species, and species delimitation. A key to all recognised species is given. In total, 67 species (listed in index) are described as new. No synonyms among nominal species of Achias are recognised. Four apparent species represented by inadequate available material are assigned provisional numbers. An etymology of Achias nomenclature is appended. Introduction The genus Achias is of interest as exemplifying the rich diversity of platystomatid species in New Guinea forests, and also for the grotesque modifications of the head in males of many species, often including stalked eyes (McAlpine 1979, 1982). On the present species count, it is the largest platystomatid genus in the New Guinea area, and one of the largest in the world, though undoubtedly exceeded in number of world species by Rivellia Robineau-Desvoidy and Euprosopia Macquart. Achias is geographically much more restricted than these genera, very few species being found outside New Guinea and the most closely adjacent islands. The limits of distribution in the west are Waigeu, Aru, and Melville Island, in the east New Britain and the Louisiade Archipelago, and in the south the Townsville district of Queensland (see Fig. 1). Methods The morphological terms used here are those previously employed by me for platystomatids (McAlpine 1973). In addition, the cell-4 index is defined as the ratio of the length of the antepenultimate section of vein 4 to the full length of the discal cell along vein 4. In listing material, the following collectors' names are abbreviated to the initials: J. H. Ardley, L. Bir6, W. W. Brandt, L. E. Cheesman, I. F. Common, R. W. Crosskey, F. P. Dodd, J. L. Gressitt, G. A. Holloway, J. W. Ismay, N. L. Krauss, R. B. Lachlan, H. A. Lorentz, D. K. McAlpine, A. S. Meek, B. J. Moulds, M. S. Moulds, H. Roberts, J. SedlaEek, M. SedlaEek, S. Shinonaga, R. Straatman, M. S. Upton, A. R. Wallace. The following abbreviations refer to collections housing specimens. AM Australian Museum, Sydney AMNH American Museum of Natural History, New York City AMST Zoological Museum, Amsterdam ANIC Division of Entomology, CSIRO, Canberra 118 D. K. McAlpine BM British Museum (Natural History) (The Natural History Museum), London BPB Bernice P. Bishop Museum, Honolulu CNC Canadian National Collection, Agriculture Canada, Ottawa DEI Institut fiir Pflanzenschutzforschung der Akademie der Landwirtschaftswissenschaften, Eberswalde FRIL Forest Research Institute, Lae (formerly at Bulolo) HELS Zoological Museum, Helsinki KONE Department of Agriculture and Livestock, Konedobu, Port Moresby MCG Museo Civico, Genoa MNB Museum of Natural Science at Humboldt University, Berlin MNM Hungarian Natural History Museum, Budapest NAT Natal Museum, Pietermaritzburg NSMT National Science Museum, Tokyo OX University Museum, Oxford PM National Museum of Natural History, Paris SAM South Australian Museum, Adelaide SPB Zoological Institute, Academy of Sciences, Saint Petersburg UQ University of Queensland Insect Collection, Brisbane USNM National Museum of Natural History, Washington WM Natural History Museum, Vienna Publication Dates of Hendel's (1914) Works Different views have been expressed on priority of publication between Hendel's two major works on Platystomatidae, that in Genera Insectorurn here referred to as Hendel (1914a), and that in Abhandlungen der K.K. 2001.-Botan. Gesellschaft in Wien here referred to as Hendel (1914b) (see Neave 1940: 72, under Rhytidortalis, and 0" 10"s zoos Fig. 1. Map of New Guinea and northern Australia showing numbers of Achias species recorded for each grid square (2' long. x 2" lat.), reference notation used in Distribution under each species description. Review of Achias Species 119 other places in that work; McAlpine 1973; Steyskal 1977; Evenhuis 1989). It is important to establish which of these works was published first because of the large number of new taxa established therein and the numerous consequent decisions to be made affecting nomenclature and interpretation of type material. Some of the problems have been discussed by McAlpine (1973: 49, 70, 141, 183), and a further relevant problem concerns the type material of Achias furcatus Hendel. Hendel (1914a): Diptera Fam. Muscaridae Platystominae. At the end of the text @. 179) there appear the words 'Wien 15 April 1914'. This has sometimes been interpreted as a publication date, sometimes as a date of receipt of manuscript because Wien is the address of the author, not of the publisher/printer (Steyskal 1977). These city and date combinations appear on the last page of text of all fascicles of Genera Insectorum issued about this time. They do not indicate date of receipt of manuscript, as they are often too close to actual date of receipt of the finished work by libraries for complex works that must have often taken many months at the printers. On the other hand they cannot be taken as reliable dates of issue (i.e. publication). In the case of the relevant Fascicle 157, there is an author's proof copy in the Diptera library at the Natural History Museum, Vienna, with the date 15 April 1914 in the handwriting of F. Hendel (R. Contreras-Lichtenberg, in litt.). Thus, this date can be taken as that of return of the proof from Vienna, and the association of this date with 'Wien' as printed on the last page becomes meaningful. Dates of accession of this part in certain European libraries are given in Table 1. Hendel (19146): Die Arten der Platystominen. It is stated on the title page that the work was submitted for publication on 1 July 1912 and issued on 15 June 1914. Available dates of accession in European libraries are given in Table 1. Table 1. Subscribers' accession dates for Hendel 1914 publications Subscriber Date for Date for 1914a 19146 National Museum of Natural 10 June - History, Paris British Museum (Natural History) 20 June 21 July Zoological Museum, Berlin 23 June 9 July Zoological Society of London 24 June 30 June Limean Society of London - 30 June Royal Institute of Natural Sciences, 6 July - Brussels I regard the work on platystomatid genera (Hendel 1914a) as having been published before that on platystomatid species (Hendel 19146) because: (1) the earliest recorded librarian's accession date for Hendel (1914a) (10 June 1914) is earlier than both the printer's claimed date of issue (15 June 1914) and the first available library accession date (30 June 1914) for Hendel (1914b); (2) four libraries had accessioned their copies of Hendel (1914a) before any are known to have accessioned copies of Hendel (1914b); (3) each of the three recorded subscribers to both works accessioned Hendel (1914a) six or more days earlier than Hendel (19146). Habits and Biology Flies of the genus Achias live almost entirely in primary forests though they sometimes survive in small strips of forest. They have been found from near sea level to at least 3000 m, and some species occur through a wide range of altitude. At least 40% of the species are known only from the mid mountain to high mountain zones. In the author's experience, Achias have been found on trunks and foliage of rainforest trees, in the latter case generally on the lower surface of leaves. They have been observed to avoid direct sunlight. Achias kurandanus are sometimes gregarious on trunks of Ficus sp. 120 D. K. McAlpine Adults of some species have been observed to feed at or to be attracted to mammalian faeces (Osten Sacken 1881; McAlpine 1979). Traps baited with human faeces have yielded numerous Achias species in some areas. Parsons (1984) records Achias spp. feeding on sap at tree trunk wounds (including Trema orientalis, family Ulmaceae) and at cut banana plants. Some information on agonistic and other behaviour was given by McAlpine (1979), and a function for the eye-stalks was suggested. Larvae of Achias, like those of many other platystomatids, are unknown. Possibly, like those of some other platystomatids, they are associated with tree roots. Estimation of Number of Species Recent discussion of the numbers of the earth's species, particularly of invertebrates, has centred on the inhabitants of the tropical forests, a category that probably includes all Achias species. It is possible that estimates involving extrapolation from counts of samples of 'morphospecies' determined by non specialists have little reality (Monteith 1990). On the other hand, specialists may have made guesses as to numbers of yet unrecorded species on the basis of little more than a vague impression. The little information on platystomatid species in tropical forest canopies (Steyskal 1966) suggests that further collecting in this habitat will not greatly increase species counts obtained by other methods (e.g. baited traps near ground level), even though Malaysia, the area investigated, is not such a significant centre of platystomatid diversity as New Guinea or eastern Australia. The method of Steyskal (1965), who used trend curves to ascertain whether large numbers of undiscovered species still exist in certain animal groups, can only be used if there is some continuity in collecting and taxonomic activity with respect to time.