Records of the Museums and Art Galleries of the Northern Territory Volume 21 December 2005 The Beagle, Records of the Museums and Art Galleries of the Northern Territory (formerly ‘Records of the Northern Territory Museum ofArts and Sciences ) EDITORIAL COMMITTEE C. J. Glasby Editor R. C. Willan Associate Editor H. K. Larson D. Megirian R F. Murray The Beagle is a refereed journal published by the Museums and Art Galleries of the Northern Territory to disseminate the results of research in the following areas: Systematic and other studies of the terrestrial, marine and freshwater flora and fauna of the Northern Territory, tropical Australia, Southeast Asia and the Indo-Pacific; Australian Aboriginal, Southeast Asian and Oceanic art, material culture and archaeology; and Northern Territory and Oceanic history and archaeology. The Beagle is published once or twice a year, depending upon the material available and budgetary considerations. All contributions to The Beagle are reviewed by two referees and, where possible, at least one internationally-based referee is selected by the Editorial Committee. Whilst articles for The Beagle will normally be 10-50 pages in length, shorter communications, notes and review articles may also be acceptable. Longer articles, significant works or substantial revisions, which form integral studies, may be considered for separate publication as a Supplement. Prospective authors should contact the Editor, Academic Publications. The Beagle may be obtained by subscription or by exchange. The subscription rate for one year for individuals and institutions is $66.00 (includes postage and GST). Cheques or money orders should be make payable to ‘Museums and Art Gal leries of the Northern Territory’. All subscriptions, back numbers and exchange enquiries should be addressed to the Manager, Peter Spillett Library, Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin NT 0801, AUSTRALIA, or e-mail library.dam@nt.gov.au AUTHOR’S OFFPRINTS Twenty-five offprints are provided free for each published paper. Additional offprints may be ordered when returning proofs. SUBMISSION OF MANUSCRIPTS A Guide to Authors is provided on the inside back cover of each volume. Contributions should be posted or e-mailed to: The Editor, Academic Publications Museum and Art Galleiy of the Northern Territory GPO Box 4646, Darwin NT0801, AUSTRALIA E-mail: chris.glasby@nt.gov.au ©Museums and Art Galleries of the Northern Territory, 2005. ISSN 0811-3653 Printed by the Government Printing Office of the Northern Territory Northern Territory Government Department of Natural Resources, Environment and the Arts Front cover: Fragile and beautiful ascidian Perophora namei from the Solomon Islands (see Kott, pages 53-66). Photograph by Karen Gowlett-Holmes. The Beagle Records of the Museums and Art Galleries of the Northern Territory Volume 21, December 2005 CONTENTS SYMON, D. - Native tobaccos (Solanaceae: Nicotiana spp.) in Australia and their use by Aboriginal peoples .1 CRAVEN, L. A. - Seven new species of Heliotropium (Boraginaceae) from the monsoon and arid zones of Australia .11 GERSHWIN, L.-A. and ALDERSLADE, P. - A new genus and species of box jellyfish (Cubozoa: Carybdeidae) from tropical Australian waters .27 GIMIN, R., LUONG-VAN, T„ MOHAN, R. and GRIFFITHS, A. D. - Aspects of the reproductive biology of Polymesoda erosa (Solander, 1786) (Bivalvia: Corbiculidae) in northern Australia.37 KRONENBERG, G. C. and DHARMA, B. - New distributional records for four species of Stromboidea (Mollusca: Gastropoda) from Australasia . 47 KOTT, P. - Ascidians from the Solomon Islands .53 LARSON, H. K. and BUCKLE, D. A new species of the circumtropical goby genus Gnatholepis Bleeker (Teleostei: Gobiidae: Gobionellinae) from northern Australia.67 SCFIWARZHANS, W., M0LLER, P. R. and NIELSEN, J. G. - Review of the Dinematichthyini (Teleostei: Bythitidae) of the Indo-West Pacific. Part I. Diancistrus and two new genera with 26 new species.73 HORNER, P. - Gehyra koira sp. nov. (Reptilia: Gekkonidae), a new species of lizard with two allopatric subspecies from the Ord-Victoria region of north-western Australia and a key to the Gehyra australis species complex.165 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21 : 1-10 Native tobaccos (Solanaceae: Nicotiana spp.) in Australia and their use by Aboriginal peoples DAVID SYMON Honorary’ Research Associate State Herbarium of South Australia Hackney Road, Adelaide SA 5000, AUSTRALIA ABSTRACT The use of native tobaccos in Australia by the indigenous population is summarised. The choice of Nicotiana species is discussed in relation to their nicotine content. Addiction, ownership, trade and preparation for use, are all described. Keywords: Nicotiana, indigenous Australian tobacco, chewing tobacco, Nicotiana preparation, ownership, addiction, trade. INTRODUCTION The genus Nicotiana (Solanaceae) with about 71 species has its centre of diversity in South America and extends to North America, the Western Pacific and Australia, with a single species in Namibia. Like most species of the Solanaceae Nicotiana contains alkaloids that are physiologically active for many animals. Humans have used species of Atropa, Brugmansia, Datura, Duboisia, Mandragora, Nicotiana, Vestici and Withania for their alkaloid content since records began. In addition, species of Capsicum and Solatium (including Cyphomandra and Lycopersicon) are important crops worldwide, while species of Browallia, Brugmansia, Cestrum, Datura, Nicotiana, Petunia and Solandra decorate gardens. Weedy Datura, Lycium, Nicotiana, Physalis and Solarium may infest agricultural crops. The principal Nicotiana used by man, N. tabacum , arose and was cultivated by local peoples in South America and in pre-Columbian time had spread to North America. In the post-Columbian era, because of its addictive nature, it spread rapidly to the rest of the world and now dominates almost completely world trade in tobacco. The worldwide spread and use of tobacco is well covered by Gately (2001) and southern American use by Wilbert (1987). NICOTIANA IN AUSTRALIA In Australia there are about 22 native species plus three species in the Western Pacific all belonging to the section Suaveolentes, which is confined to this region. One species occurs in the Marquesas Islands, and two in New Caledonia and nearby islands. 1 have found no record of these being used and Riesenfeld (1952) says specifically that the species on the Isle of Pines (N. fragrans) was not cultivated or smoked. Smoking did not occur there until the arrival of the Europeans. In Africa a single species, N. africana, was named from Namibia as late as 1975, where it is confined to a few peaks. It is not known to be used. Cytological studies show it to be distantly related to section Suaveolentes rather than to American species (Merxmuller and Buttler 1975; Gerstel et at. 1979). The Australian species are widely spread over the continent excepting Tasmania [see maps, Horton (1981) or in Purdie et ah (1982)]. They are less common in the far north and have not been recorded from the Kimberleys, Arnhem Land or Cape York Peninsula. Principal speciation has been in the warmer arid and semi-arid mainland of Australia. No native tobacco occurs in New Guinea or New Zealand. The first Australian species named was N. suaveolens Lehm. (1818), the name based on plants cultivated at Malmaison (France) grown from seed from Port Jackson. Further species were named somewhat slowly. Even Bentham (1868) in Flora Auslraliensis, nearly 50 years later, included only N. suaveolens with four varieties. This name was applied loosely for many years. The first comprehensive account of the genus in Australia was by Wheeler (1935), a colleague of Goodspecd, whose monograph on the genus was published in 1954 (Goodspeed 1954). Wheeler’s account was followed by Burbidge (1960) and Horton (1981). The latter was incorporated, almost unchanged, into the Flora of Australia Vol. 29 (Purdie et ah 1982). Four more species have been described since then, giving a total of about 22 species. Most of the Australian species of Nicotiana are annuals, or may survive for a second year in exceptionally good seasons. A few of the southern species are short¬ lived perennials. It follows that supply of leaves will vary greatly from year to year. I have seen sand dunes virtually covered with N. velutina, which may be totally absent in 1 D. Symon a subsequent year. The supply of the two popular species for chewing - N. excelsior and N. gossei - growing in the Central Australian ranges may be a little more reliable but N. ingulba from the sand plains will be very dependant on irregular rains, especially after bushfires. A number of the early explorers comment on the abundance of plants in some years. Francis Gregory, in his journal of exploration to the Gascoyne River in Western Australia in 1858 (Gregory 1884), when in the vicinity of the Lyons River on 27 May 1858 stated: “We halted for the night amongst fine grass; melons and tobacco growing very luxuriantly" and on 31 May 1858, a little further on states: “Tobacco here grew to sufficient size for manufacture, occupying many hundred acres of the best land.” There is no mention of Aboriginal use. Later Francis Gregory, in his journal of exploration to the north-west coast in 1861 (Gregory 1884) stated: “Tobacco does not grow so luxuriantly here as on the Lyons River, but the natives collect it, and after preparation, chew it; but we did not on any occasion observe them to smoke.” This may have been A. benthamiana , which is scattered but widespread in the north-west and is a favoured species. The party was in the vicinity of the Lyons River on 25 June 1861, but no mention of tobacco was made at the time. The zoologist, chemist and explorer H.H. Finlayson records that dried tobacco as well as Duboisia hopwoodii were stored in caves at Uluru for poisoning smaller waterholes to stupefy emus (Finlayson 1943). lie also illustrated a vigorous stand of N. gossei in the George Gill Range in southern Northern Territory. Irregular supplies in one area or another are likely to have encouraged trading the leaves. Latz (1995), see also below, illustrated a large stand of N. excelsior in the Mann Ranges. Alkaloid content. The alkaloid content of the leaves and roots of sixty species of Nicotiana was published by Saitoh el al. (1985). Because of its relevance to the choice of Nicotiana species for chewing, the relevant section of Saitoh’s table is offered here (Table 1). fhe spread of tobacco, N. tabacum. The product tobacco travelled well in ships as compressed plug tobacco and as cigars (and seed). Its use was soon established in early Portuguese and Spanish outposts followed by cultivation. Haddon (1947) reports tobacco in the Philippines in about 1565, in Ternate (about 1599), Java (about 1601), Japan (about 1605) and Halmahera (about 1616). Cultivation soon spread to New Guinea which has no native species. It was in active use on the north-east coast before 1886 (Miklouho-Maclay 1886). The use of tobacco for smoking probably travelled from west to east in New Guinea and then across Torres Strait. Riesenfeld (1952) presented a detailed account of the spread of cultivation of tobacco in New Guinea and stated: “In Torres Strait tobacco is smoked as far south as Murolug Island (just off the tip of Cape York Peninsula) and its cultivation is recorded on many islands. A few tobacco pipes found on Cape York have been shown by Haddon (1947) to be of Torres Strait origin, as, no doubt, is also the habit of smoking on Cape York, the only Australian region where tobacco is smoked”. The latter statement is not strictly true, as tobacco seems to have been smoked further west along the north coast of Australia (see below). Riesenfeld’s map clearly indicates the passage of smoking across Torres Strait to Cape York. It is of note, too, that there is no mention of chewing tobacco in this detailed account. McCarthy (1939) states specifically that tobacco was amongst the goods traded by the Malays from Makassar (Celebes) and Kupang (Timor) from an early date. The chief centres in northern Australia where the Makassans fished were at Malay Bay, Elcho Island, Caddcll Strait, Milingimbi Island, Caledon Bay and Groote Eylandt. Capt. J.B. Jukes while surveying on I I.M.S. Fly in 1847 recorded its use in Cape York (Jukes 1847). Here it was smoked, and Jukes notes that the informant knew exactly what to do with a cigar. Tobacco in Australia. Ships trading to the Americas soon introduced tobacco to Europe and it had become well established by 1600 (Gately 2001). Tobacco was chewed, snuffed and smoked; the dominance of cigarettes was a later development. The historian Samantha Fabry slated that tobacco did not form part of the provisions of the First Fleet to Australia, but was included in the stores held by the ships’ captains. Shortly after the First Fleet’s arrival in 1788, Major Ross purchased a quantity of tobacco from the Master (Captain) of one of the returning ships for the marines who were, he said, “so much distress’d for tobacco” (Fabry 2004). Early urbanised Aborigines would soon have used tobacco. Such use is illustrated in Dutton (1974) in a picture by Augustus Earle of a native family dated about 1826 in which the man is smoking a pipe. This aspect of smoking by urbanised native peoples will not be considered further here. Early records of smoking by non-urbanised native peoples are all from the northern near-coastal regions of Australia. This is almost surely the result of contact with Macassan and other possible Asian visitors in addition to the progress from New Guinea across Torres Strait. Macknight (1972) considered that the Macassan voyages to the north coast of Australia ceased about 1906- 07. Records of smoking in northern Australia using bamboo pipes may be found in several accounts. J. McGillivray, doctor and naturalist, together with T.H. Huxley under Owen Stanley on 1 I.M.S. Rattlesnake , while surveying Torres Strait in 1850, noted: “The custom of smoking, so general throughout Torres Strait, has been introduced to Cape York. The most addicted to it were the Papuans above mentioned, but many of the Australians joined them, and were equally 2 Native tobacco use in Australia Table 1. Alkaloid contents of Nicotiana species. After Saitoh et al. (1985). tr = trace. Leaves Roots Nicotiana section Suaveolentes Total content mg/g dry weight % of total Total content mg/g dry weight % of total Nicotine Nornicotine Anabasine Anatabine Nicotine Nornicotine Anabasine Anatabine N. africana Merxmuller and Buttler* 6776 4.7 92.4 0.3 2.6 7698 45.0 45.1 1.0 8.9 N. amplexicaulis Burbidge 4959 98.5 0.3 tr 1.2 7648 71.8 3.1 11.9 13.2 N. benthamiana Domin 3602 80.8 0.6 16.1 2.5 3826 68.4 1.4 26.3 3.9 N. cavicola Burbidge 285 16.8 76.9 6.3 tr 3798 58.0 9.1 25.4 7.5 N. debneyi Domin 2457 31.1 15.8 46.0 7.1 3038 34.7 1.4 53.2 10.7 N. excelsior (J. M. Black) 4772 95.9 1.1 0.9 2.1 4772 66.5 1.4 20.0 12.1 N. exigua Wheeler* 7692 70.0 26.0 1.5 2.5 12040 60.1 1.8 21.2 16.9 N. fragans Hooker 14985 91.0 0.4 0.4 8.2 13344 81.4 0.8 1.1 16.7 N. goodspeedii Wheeler 730 5.5 68.4 21.1 5.0 4284 67.8 1.3 26.6 4.3 N. gossei Domin 12169 98.8 0.3 0.2 0.7 7222 73.8 0.9 17.9 7.5 N. hesperis Burbidge 4108 52.1 0.4 44.3 3.2 1930 22.1 1.2 74.9 1.8 N. ingulba J. M. Black 669 7.5 44.8 42.2 5.5 4804 46.3 4.5 42.3 6.9 N. maritima Wheeler 608 7.2 70.4 15.8 6.6 14030 20.8 30.0 44.5 4.6 N. megalosiphon lleurck and Mull. 5566 18.5 42.0 39.5 tr 5566 50.5 1.8 43.7 4.0 N. occidentalis Wheeler 519 9.4 75.1 15.4 tr 6490 41.5 14.6 39.5 4.4 N. rosulata (S. Moore) Domin 992 24.9 70.8 3.0 1.3 3376 85.6 1.7 10.5 2.2 N. rotundifolia Lindley 4949 96.6 0.8 2.1 0.5 4912 53.9 1.1 40.7 4.3 N. simulans Burbidge 258 14.0 67.0 19.0 tr 6624 45.1 6.4 44.3 4.2 N. sauveolens Lehmann 4954 85.0 13.6 0.9 0.5 6658 51.7 9.9 29.0 9.4 N. umbratica Burbidge 43 51.2 48.8 tr tr 9932 43.3 23.5 27.8 5.3 N. velutina Wheeler 5276 2.8 88.4 8.1 0.7 24817 52.9 13.4 32.3 1.4 * Species did not bloom clamorous for tobacco. ... The pipe is a piece of bamboo as thick as the arm and two or three feet long, is first filled with tobacco-smoke, and then handed round the company seated on the ground in a ring-each takes a long inhalation, and passes the pipe to his neighbour slowly allowing the smoke to exhale. On several occasions at Cape York, I have seen a native so affected by a single inhalation, as to be rendered nearly senseless with the perspirations bursting out at every pore, and require a draught of water to restore him.” (McGillivray 1852: 126-127). The explorer F. Jardine, when near the apex of Cape York, Queensland, in 1865 makes the comment: “The black guides were not forgotten, and received their reward of biscuit and tobacco. The manner in which they use this latter is curious and worthy of notice. Not satisfied with the ordinary ‘cutty’ [a short pipe] of the whites, they inhale it in volumes through a bamboo cane. The effect is profound stupefaction, which appears to be their acme of enjoyment.” (Jardine 1998). J. Bancroft, doctor and chemist in Brisbane, records a rather sad request for tobacco: “I was much struck with this tobacco-want when passing through Torres Straits lately. Steaming slowly among the islands of that calm sea the vessel encountered a native and his wife in a bark canoe. The only word they used was ‘Tabac, (abac’. A loaf of bread was thrown to them, but this did not satisfy; and in the wake of the steamer there could still be heard the cry, ‘Tabac, tabac!’.” (Bancroft 1878-82). The Gregory brothers, Augustus and Francis, conducted several expeditions in northern Australia between 1846 and 1861. The journals of these explorations were published in a single volume in 1884 under the authorship of Augustus and Francis Gregory. Augustus Gregory in 1855, when 3 D. Symon in the vicinity of Port Albany Island and the mainland of Cape York, states: “Some canoes with natives came to the vessels. They evidently have frequent communication with vessels passing through the Straits, and are well acquainted with the use and name of tobacco, which they smoke in large bamboo pipes.” (Gregory and Gregory 1884). Telegraphist and anthropologist F. J. Gillen recorded smoking in his ‘Camp jottings’ while on an expedition to northern Australia with Spencer in 1901-1902. Gillen states: “Another of their adaptations is the Chinese pipe which is modelled on the pipe used for smoking opium by the Chinese. We first saw these pipes at Powells Creek and since then there have been one or more in every camp visited.... The method of smoking this pipe is to take half a dozen rapid whiffs and then inhale as much as they can swallow retaining it for a minute or two and then puffing it off through the mouth and nostrils. ... The Chingilli Umbai and Gnanji powder up tobacco mix it with an equal quantity of white ashes and chew the mixture. Curiously none of these tribes appear to have had a subst itute for tobacco before the white man penetrated the interior. ... We have not seen a plant of Nicotiana suaveolens ... nor of Duboisia hopwoodii north of the Barron.” (Gillen 1901-1902). Howard (1933) described early English trading activities on the north coast of Australia. Among the list of items to be traded was tobacco and short-lived trading posts were established at Melville Island in 1824, Raffles Bay in 1827, and Port Essington in 1838. The anthropologist, Donald Thomson, devoted an article on the smoking pipes of north Queensland and the Northern Territory (Thomson 1939). He stated that: “There is evidence for the belief that the use of tobacco has been known in North-Eastern Australia for a considerable time; nevertheless there is nothing to suggest that any attempt to grow tobacco was ever made by any of these people, although it was grown by their neighbours in Torres Strait.” Thomson goes on to describe the bamboo pipes as well as the use of a large crab claw as a short stocky pipe. He also notes the power of addiction and adds: “Under the ‘drive’ for tobacco the natives will undertake long journeys and endure unbelievable hardships, to obtain a few ounces of trade tobacco.... a native will sell anything he possesses for a small quantity of tobacco.” While smoking with pipes of Asian origin became established in northern Australia, smoking with pipes of European origin was soon practised in southern Australia. Thomson (1939) illustrates examples of bamboo and crab- claw pipes. By these dates the tobacco likely to be traded is Nicotiana tabacum (American tobacco) as there is no evidence that local native peoples cultivated the plant, nor that native Australian tobacco was traded so far north. Carl Wilhelmi, a German horticulturist and botanist, visited South Australia and in late 1849 made a journey to the lower River Murray. When in the vicinity of Wellington he observed the Aboriginal canoes and noted, “A small fire was burning in the middle of the small craft, from which they could light their pipes”. Later in 1850-1851 Wilhelmi travelled to Moorunde and then down the River Murray. During part of this journey their goods were carried on a boat crewed by five Aborigines and “... they were willing to take our things with them when they heard that they were to receive tobacco and white money” (Wilhelmi 1857). Later, when crossing Lake Alexandrina to Goolwa. the party exchanged half a stick of tobacco for a large Murray cod. These few incidents show that pipe smoking was practised and tobacco readily traded in the mid 1800s by Aboriginal people still largely practising their original life style. All these examples show that smoking and the use of tobacco was established in the north coast of Australia before the use of indigenous tobacco was recognised by the European population. The use of native tobacco, Nicotiana spp. The first Central Australian records of the use of native Australian species of Nicotiana appear to be those of Helms (1891) on the Elder Expedition and Stirling (1896) on the Horn Expedition. Since then many writers have described the collection and preparation of Nicotiana. David Carnegie, engineer, gold miner and explorer, on his epic journey almost due north of Menzies (Western Australia) in 1896 records the making and chewing of a quid with ashes but does not name the plant used (Carnegie 1898). H. Basedow, anthropologist and explorer, using the term ‘pitjuri’ confused Nicotiana and Duboisia. but was clearly referring to Nicotiana when describing its use in the valley of the Finke River and the MacDonnell Ranges. He described the typical preparation of a quid and comments: “... they look upon pitjuri-chewing in company as a social comforter, which fosters mirthfulness and friendly fellow feeling” (Basedow 1925). In 1933 J. B. Cleland, pathologist, botanist and anthropologist, together with T. H. Johnston, zoologist and anthropologist, published a major paper on pituri. They attempted to sort out the confusion in the use of the word ‘pituri’. Their paper is a major source of early references and although the emphasis is on Duboisia it does include useful material on Nicotiana (Johnston and Cleland 1933-1934). C. Chewings, in a popular book on the Aboriginal peoples, again described the preparation of a quid and also included the use of plant roots (Chewings 1936: 31-32). Likewise A. Groom, author of another popular book, records pituri growing in stream bank sands and that: “We travelled through about a hundred acres of the pituri, growing about three leet high, very much like tobacco. Tamalji and Njunowa darted through it. selected and pulled leaves, and piled them in bundles on the camels 4 Native tobacco use in Australia until we resembled a travelling market garden; I was left walking beside Tiger, who was leading the team. ‘Pituri,’ Tiger explained, ‘we take plenty back to Areongonga people. We get three shillings a sugar bag, sometime more.” [This was N. gossei ] (Groom 1950). Later Donald Thomson spent time with the Bindibu (= Pintubi) Aboriginal peoples in the Western Desert. This tribal area is in the vicinity of Lake Mackay on the border between Western Australia and the Northern Territory. This is extremely harsh country and Thomson gives a detailed account of the enthusiastic collection of Nicotiana ingidba and its preparation. His notes show the avidity with which it was appreciated: “Soon after they had chewed the concoction the men, invariably tired after hunting, would rest reclining on their elbows or loll on the ground in other relaxed attitudes. Invariably, on their return to camp, they would devote themselves to the preparation of the chewing quid if they had collected the ingredients, rather than attend to the less laborious and seemingly more immediate task of preparing food.” (Thomson 1975: 81-91). C. Godard, linguist, and A. Kalotas, botanist, in an account of plant use by the Yankunytjatjara quote an account by Sam Pumani, again on the demand for and satisfaction of chewing tobacco (Godard and Kalotas 1985): “Suppose they were poorly off for tobacco, without any leaf. ‘We’re out of leaf.’ In this case the people who have passed away would turn to the stalks. And grind them and chew them dry, suck on them bite them. ‘Well, I'll just have to chew this. 1 can sort of chew the stalks! My mouth’s gone dry. Since I don’t have any leaf I’ll just have to carry stalks around with me.’ The people who have passed away would take stalks around with them, when they did not have leaf. They’d grind the stalks into a quid. After grinding them to a powder they’d carry the quid around with them. The remaining stalks would be kept wrapped in emu feathers. They put them in emu feathers. And after grinding and softening them, they used to chew them, stringing it out to make it last. And as the hills got close, fresh tobacco would get close. Then they would throw away the stalks. Yes, throw the stalks away. Break off some fresh tobacco, and after that, singe the leaves on a fire. As they were doing that, one might say: ‘Hold on. I’ll just suck on a fresh leaf!’ He'd suck it alright, the one without tobacco. Yes, then his eyes’d lift up, from chewing that tobacco. ‘Oh, yes! 1 am chewing tobacco at last! I’ve been without it forages!’ ‘Oh, quick, 1 can sleep with it in my mouth.' They would talk about sleeping with it in the mouth. ‘Oh, to sleep with the stuff!’ As it gets dark, they’d be saying: ‘Oh I'm going to sleep with it!’ And he’d sleep with it holding it in mouth. He doesn’t sleep without it. When day breaks they would tuck it behind the ear, and carry it around with them. And would travel around contented, with it in the mouth. They’d feel satisfied. They put some in the mouth and travel around, looking out for game.” P. Brokensha, in his book on Pitjantjatjara crafts, provided some pictures of tobacco preparation in the Musgrave Ranges. He stated that few of the community smoked but that many men and women chewed tobacco (Brokensha 1975). Finally, the authoritative book by Peter Latz, botanist, ethnobotanist and ecologist, devotes a chapter to the use of Nicotiana with illustrations of the principal species in Central Australia (Latz 1995). Preparation of tobacco. Collecting could involve the gathering of the whole plant, sometimes including the roots, or just stripping leaves off large plants. Usually the material was wilted or partially dried on a clean surface, warm sand or by a fire before use. Occasionally it was air dried and stored. It may be ground down or broken into sufficiently small pieces to chew and make into a quid. A Imost invariably clean white ash prepared from a number of shrubs. Acacia anenra, A. coriacea, A. kempeana, A. lignlata, A. pruinocarpa, Grevillea striata, and some Eucalyptus barks was added to the quid (Meggitt 1966). An analysis of the ash of A. salicina was published by Higgin (1903) and it was shown to be strongly alkaline. The ash is known to facilitate the release of alkaloids as is the practice of using lime with betel nut. Lime was also used by South American tobacco addicts (Wilbert 1987) who, like the betel nut chewers, used a small gourd to contain the lime. The use of Nicotiana root. In his description of the use of N. ingulba by the desert Bindibu people, Thomson (1961) stated specifically that the entire plant complete with taproot was pulled up and this can be seen in his figure 1. In his description of the preparation and use he makes no mention of stalks or roots but elsewhere the use of stalks as second grade tobacco has been described (Goddard and Kalotas 1985). There is every reason to suggest that the roots might well be included with the stalks. The roots of N. ingulba have a moderate yield of alkaloids, of which 46% is nicotine. An early record of the use of Nicotiana roots was given by Cleland and Johnston (1933). They stated that leaves, stems and roots of N. ingulba are all used, being dried and ground up on a stone. This is also repeated by Chewings (1936), but here the suggestion is that it might have been N. gossei, of which the leaves are very popular. This is a convenient place to note the record by George French Angas, artist, traveller and author, on the use of a root with narcotic properties. He states: “The Taltayarra Tribe: An intoxicating root is also frequently used by them: it grows in the scrub and when taken has much the same effect as opium.” (Angas 1847). No other information is given. The identity of this root has not been confirmed. Cleland (1966) suggested that it might be Cyphanthera myosotidea but there are no other records of its use. No Nicotiana is known from the Tatiara region but two species, N. goodspeedii and N. maritima, both reach the areas near the Murray River mouth and N. goodspeedii can be abundant on flats and mallee areas 5 D. Symon on the lower River Murray. Both species may be short-lived perennials and develop a tap root. Both species would be within easy trading distance of the Tatiara region. That the roots of N. goodspeedii may have been used is supported by the account in Cleland (1957). He records that: “Early in 1954 Mr W. B. MacDougall forwarded me material which Mrs E. Robertson identified as Nicotiana goodspeedii. He did not know how reliable his information was, but one native told him that only the roots were used as pituri for chewing, and another that the leaves were dried and then used as well as the roots. Both said that it was very potent. The Kokota name given was ‘towell- towelf or ‘dowell-dowel!’. Mr MacDougall found it growing in sheoak valleys from Lake Maurice to the east-west railway line and in ‘crab-holes’ on the edge of the Nu Harbor Plain as far south as Lake Tallacootra. The natives said that it grew as far east as Wynbring Rocks. It is abundant on Yalata.” Mr MacDougall was, I believe, a ranger employed at the Woomera Rocket Range. Nicotiana goodspeedii grows in the area indicated to the virtual exclusion of any other Nicotiana. The high nicotine content and yield indicate that the roots of N. goodspeedii may well have been appreciated (Table 1). This species is not particularly leafy and in dry seasons persists as an almost leafless crown surmounted by dead seed stalks. The tap root would still be available for use when all leaves had withered. As trade in American tobacco via Europe increased and pipe smoking spread to the pastoral stations, native tobacco was sometimes combined with traded tobacco. The botanist and explorer, Alan Cunningham, in his journal during Oxley’s land journey (Lee 1925), reported the interesting and early use of native tobacco by white men. He states: “June 21” 1817 in the vicinity of Peel’s Range NSW, Nicotiana undulata [=A. suaveolens] is very frequent on these flats, the lower leaves of which our people gathered, and when dried found them not a bad substitute for its congener N. tabacnm, although not so strong a narcotic.” The use of native tobacco by station hands in pipes was recorded by Johnston and Cleland (1933-1934). Station managers also appreciated the value of native tobacco. On herbarium sheets in the State Herbarium of South Australia, C. T. Madigan in 1944 recorded the collection and storage of pituri by the station owner at Huckitta Station. His label states: “A central Australian tobacco used by the Aboriginals. Note capsule fruits. Pituri from Huckitta Stn. NE of Alice Springs. This material gathered and stored by the station owner especially to distribute to the natives. An unusually stout specimen.” The specimen is Nicotiana ingnlba. R.A. Gould described the overlap in the use of native tobacco with traded European tobacco (Gould 1969: 181, 185-186, 207). At Warburton he exchanged plugs of tobacco and cigarettes for information and favours. He found that the older men preferred to tear open a cigarette and chew the tobacco and that younger men preferred to smoke a cigarette, as wearing a cowboy hat interfered with placing a quid behind the ear. Premasticated balls of tobacco were stored in circular cans and were passed around at meetings or ceremonies. With knowledge of cigarettes, native tobacco was sometimes rolled in newspaper to make a cigarette as described by Wallace and Wallace (1973). O’Connell et al. (1983) in their paper on traditional and modern plant use among the Alyawara of Central Australia, stated: “Since World War 11 the use of native tobacco has declined in favour of the pressed plugs of cultivated varieties distributed in government rations or sold in local cash stores. However, the more powerful local forms are still collected whenever encountered.” Addiction. An example of Aboriginal craving and probable addiction to tobacco is indicated in this letter, which is attached to a specimen of A. gossei in the Melbourne Herbarium. It is from E. Ryans and dated 24 January 1887: “I am forwarding by this mail a sample of ‘Engoulba’, native name for ‘Pitcheri’ or native tobacco that grows in this locality, it only grows in two places here and near Doctors Stones, the natives come many miles for it and some of them preserve it from one season to another. The Blacks partly roast it and chew and suck and swallow the juice which acts on them like opium on Chinese, it makes them dead drunk for about 24 hours. They save the pulp after sucking the juice for smoking, they take an extra dose when they are hungry or hard pressed for water. It would also save a white mans life if hard up for water by chewing a little bit of it raw. 1 am informed it has killed some bullocks on Mr Stokes’ run eating it while it was green. Gov. [government] Well Party, Old Depot, OT [Overland Telegraph] line via Charlotte Waters.” Donald Thomson (1961) also comments on addiction to tobacco by the Bindibu (= Pintubi) people of the Western Desert in Western Australia: “Observations on the behaviour of the tobacco-chewing groups among the Bindibu, with whom I lived in close proximity for some months, led me the conclusion that the chewing of the Nicotiana quid described in this paper was practised under a compulsion or craving which appeared to have a physiological rather than a social basis. The urge to chew would begin to manifest itself when the hunting party was far out on the dunes and while the men were still hunting ... The preparation of the chewing quid would take precedence over the preparation of food.” An example of enthusiasm, with some social comment, is provided by Hansen and Hansen (1974) from the Pintubi people of the Western Desert in a Pintubi post-primary reader: 6 Native tobacco use in Australia “Pluck mingkulpa plants. Let us cook and eat it. Pluck mingkulpa plants. Don’t bring back the weak leaves -bring back the strong ones. Let us try it first. Don't bring back the weak leaves without trying it. Let us bring back ash tree to mix with the mingkulpa. Let us eat it together with the ash, we who are starving for mingkulpa. Let us eat it, so it can burn our throats. When walking without water, chewing mingkulpa is good to keep one alert. Let us cook and eat mingkulpa. One should break his lump in half and give it to another. After preparing it. let us hide it in the shelter, so the women won't grab it from us. Let us carry it in our pockets. If you keep it where people can see it, they ask you for it, and finish it all up. Not only mingkulpa but tin of tobacco and cigarettes as well. We are all eating that mingkulpa which is from the white man, cigarettes and tobacco, it is very good mingkulpa. That which grows on the hills is called pinatilypa [possibly N. gossei]." Ownership. When writing of pitjuri, Basedow (1925) in his book on the Australian Aborigines, confused Duboisia and Nicotiana. However, as he stated: “The Arunndta and latterly the Aluridja as well regularly collect as much pitjuri as they want in the valley of the Finke and other gorges of the MacDonnell Ranges while the Wongapiteha have their sources in the Musgrave and Everard Ranges.” These are prime areas for Nicotiana. Basedow goes on to state: “The collecting grounds are as a rule owned by a circle of old men, each of whom clearly defines his boundaries by placing a number of stones upon the ground. A proprietor may give another person necessary permission to gather leaves on his plot according to certain terms agreed upon.” The concept of ownership is also recorded by the anthropologist, Tindale (1974). He states: “Trespassing to hunt was one of the main causes of fights between tribes, as well as persons of local groups within tribes all over Australia." He quotes an example: “The thought that native tobacco growing in the cave mouths and other sheltered places after rain, might have been the object of plundering was anathema to the old man . . . [he] was very much disturbed because these plants were considered particularly precious possessions by the horde.” Tindale also recorded the attention of the Aborigines to the natural growths of tobacco. “In the arid environment of the Western Desert ... the men know most of the places where the tobacco flourishes and watch over its growth. They are zealous in watching over maturing plants. The plants are associated with specific ‘kulpi’ or rock shelters linked with clan totems, hence can be very particular possessions of the clan. Thus trespass to take tobacco is considered a serious matter.” (Tindale 1974). Trade. Aborginal trade routes were extensively developed through arid Australia (e.g., pearl shell from the north-western coast was traded as far south as Ooldea in South Australia). Duboisia. pituri, from the Mulligan River area of south-western Queensland was traded as far south as the northern Flinders Ranges in South Australia, west to the Central Australian Ranges and north-east into central Queensland. Early records of trade in Duboisia may be found in Bancroft (1878) and Gregory and Gregory (1884). The anthropologist F. D. McCarthy (1939) presented a number of maps of trade routes in Australia extending over most of the continent. His map number 13 combines the routes for both Duboisia and Nicotiana. at least in the principal central region. In fact, they may have extended further west than shown there (McCarthy 1939). The Duboisia trade is further extended in Watson (1983). Trade in Duboisia will not be considered further here. The principal trade in Australian Nicotiana species originated further west than that of Duboisia in the Central Australian Ranges and included the trade route to Ooldea, south of the eastern borders of the Nullarbor Plain. This epic route required following 27 rockholes and could take approximately 35 days to reach Ooldea (Juldi). Tobacco was exchanged for wombat fur, and for red, white and yellow ochre (Berndt 1942). From Ooldea, the route went west, at least touching Fraser Range and eventually joined a trade route from the north-west coast, and east towards Lake Torrens. An interesting side effect of the trade in tobacco is a few records of N. excelsior (a popular species) being collected long distances from the main occurrence of the species with no evidence that they represent disjunct local populations (Fig. I). These sporadic occurrences are almost surely the result of the distribution of seed from the trade tobacco. The following specimens, all in the State Herbarium, Adelaide, represent such plants. 1. R. I Iclms, 23 December, 1891, Fraser Range (south¬ eastern Western Australia). [AD 97433242] 2. M. Koch 118, Augudt 1899, Mount Lyndhurst. [AD 97602403] 3. E. H. Ising, 20 September, 1951, Evelyn Downs. Called Pituri and chewed by the natives. Growing near S M homestead [?station manager] and said by native Hector and Jerry not native to this place but coming from a long way N W (?Everard Park). Evidently grown from material obtained for chewing. [AD 97413288] 4. Mrs Waskett, 14 December, 1955, Oodnadatta. [A D97413289] 5. P. Horton 209,28 August, 1980, FarNW Aboriginal family settlement, 68 km SSE of Mt Davies Camp and 73 km north-west of Mount Lindsay. Single erect herb in red sand near bore in low sand dune country. Chewed by local Pitjantjatjarra people, possibly brought here by them. [AD 98565306] Nicotiana excelsior is an easily recognised species as is N. gossei. If N. ingulba was also traded, disjunct occurrences may not be so easy to recognise as its distribution appears to overlap that of N. rosulata. 1 D. Symon However, no strays were recognised in the collections of these species at Adelaide. Species of Nicotiana used for chewing. Latz (1995) gives a detailed account of the species used in Central Australia. The plants are illustrated. Aboriginal names are given and the notes below are his, except where indicated by square brackets. N. benthamiana: This species is used as a chewing tobacco throughout its range. It is considered inferior if other ‘stronger’ species are available. [Also recorded for Western Australia by Reid and Betts (1977)]. N. cavicola: [Mainly western in its distribution and recorded for Western Australia by Reid and Betts (1977)]. N. excelsior: This is an important plant for the Pitjantjatjara people. They know where to find the best stands (usually in caves) and often travel considerable distances to obtain them. [Also recorded for Western Australia by Reid and Betts (1977). While plants may be found in cave mouths, large populations are in open sites]. N. g/anca: Found in the southern quarter it is reported to be chewed by the Pitjantjatjara people as a drug. [An introduced Argentinian species], N. goodspeedii: [Cleland (1957) records this being used, both tops and roots, by Aboriginal peoples at the eastern end of the Nullarbor Plain and to the south], N. gossei: Occurs in most of the major ranges of the area. This is probably the single most important plant to the Central Australian Aborigines. The literal translation of the Alyawarr name for this plant is ‘bone-marrow tobacco’ and bone-marrow is considered the ultimate delicacy. [The name N. suaveolens was wrongly applied to this species in early records]. N. megalosiphon: Treated with disinterest. N. occidentalis: Treated with disinterest. N. simulans: Treated with disinterest. [Peterson (1979) reports the use of this species amongst the southern Walpiri people but ranked the last of three], N. rosulata subsp. ingulba “Because it is available over a wide area and large amounts can be gathered in a good season, this is the plant most often used for pituri. Its potency varies somewhat from area to area, but is generally considered to be less potent than species found growing in the hills ... This plant is especially important for the Pintubi and features in their mythology”. [Horton’s (1981) treatment presents the names as given here. Other taxonomists prefer to maintain them separately. In which case the name N. ingulba applies to Central Australian populations and N. rosulata to more southern and western populations.]. N. rotundifolia: [Cleland and Tindale (1959) list this species as not used in the vicinity of Haasts Bluff, Central Australia. The species does not grow in this region and the name was wrongly applied to what may be N. simulans.]. N. species: [Reid and Betts (1977) record N. suaveolens being used in the Coolgardie area but on p. 145 quote D. A. Herbert stating that: “The natives in the country behind Port Hedland go out of their way to collect the leaves . . .” Two species of Nicotiana are recorded in the vicinity of Port Hedland. The first is a strongly glandular species, N. occidentalis, which Latz (1995) says is treated with disinterest. The second is N. benthamiana, which is cne of the favoured species (see above) and is highly likely to be the one referred to by Herbert. See also Gregory (1884) who probably refers to this species.]. N. suaveolens: [The species, recorded by A Cunningham in Lee (1925) was used by ‘our people’ and it is not clear whether this included Aboriginal peoples.]. N. velutina: Occurs throughout Central Australia. It grows in most habitats, but is especially abundant on limestone river banks or above salt lakes [and on sand dunes]. It is a weedy plant. It is rarely if ever used by Aboriginal people. Favoured species. Latz (1995) writes of the popular species in Central Australia roughly in the order shown in Table 2 to which N. tabacum has been added. It is of interest to add the chemical analysis from Saitoh et al. (1985) and see the close relationship of choice with nicotine content. Note that the spurned species, N. velutina, has the ‘wrong’ alkaloid nornicotine. Well may Norman Tindale (1974) say: “One of the few luxury crops harvested by the Aborigines and that chiefly in the central desert regions is native tobacco”. Fig. I. Distribution of Nicotiana excelsior. Native tobacco use in Australia Table 2. Most favoured species of Nicotiana. tr = trace. Nicotiana species Total Yield (mg/g) Nicotine Nornicotine Anabasine Anatabine N. gossei 12169 98.8 0.3 0.2 0.7 N. excelsior 18902 95.9 1.1 0.9 2.1 N. benthamiana 3602 80.8 0.6 16.1 2.5 N. rosulata 992 24.9 70.8 3.0 1.3 N. megalosiphon 319 18.5 42.0 39.5 tr N. simulans 258 14.0 67.0 19.0 tr N. occidentalis 519 9.4 75.1 15.4 tr N. ingulba 669 7.5 44.8 42.2 5.5 N. velutina 5276 2.8 88.4 8.1 0.7 N. tabacum 11462 94.8 3.0 0.3 1.9 And Latz (pers. comm. 2005) says “I consider ( Nicotiana ) was the most important plant substance to the arid- and semiarid-land Aborigines”. ACKNOWLEDGMENTS I am grateful to my wife Judith for assistance, to Robyn Barker for many helpful criticisms and for supplying internet information and to Evelyn Bourne for typing the manuscript. REFERENCES Angas, G.F. 1847. Savage life and scenes in Australia and New Zealand, Vol 1. Smith Elder and Co.: London. Bancroft, J. 1878-1882. 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Accepted 22 July 2005 10 The Beagle, Records of the Museums and Art Galleries of the Northern Territory. 2005 21 : 11-25 Seven new species of Heliotropium (Boraginaceae) from the monsoon and arid zones of Australia LYN A. CRAVEN Australian National Herbarium, CPBR, CSIRO Plant Industry, GPO Box 1600, Canberra ACT 2601, AUSTRALIA ABSTRACT Seven new Australian species of Heliotropium, H. albrechtii, H. arenitense, H. argyreum, H. lapidicola, II. microsalsoloides, H. mitchellii and //. viator, are described and their distributions mapped. Partial identification keys are provided to enable the new taxa to be integrated in a recent revision (Craven) of the Australian species. The generic circumscription of Heliotropium is discussed and a broad definition of the genus is retained. Keywords: Boraginaceae, Heliotropium, Euploca , new species, Australia, Northern Territory, Western Australia. INTRODUCTION The Australian species of Heliotropium L. were revised by Craven (1996) who recognised 81 species for the region of which 78 are indigenous (with 75 of these being endemic) with the remaining three species being introduced and naturalised as a result of the activities of European settlers and/or traders. The Australian species typically are annuals to perennials, and in habit may be cushion forming (Fig. 1) but more usually are subshrubs (Fig. 2). Since the revision was finalised, this being several years before its ultimate publication in 1996, material of several distinctive new Australian species became available and several more have been collected in more recent years. Reconsideration of the taxonomic status of some anomalous collections included in H. ovalifolium Forsskal in the 1996 revision has resulted in the conclusion Fig. I . Heliotropiumpiilvinum Craven, a perennial that forms small cushions. that two additional species should be recognised. With the impending publication of a volume in the Flora of Australia series that will include Boraginaceae, seven new Australian species of Heliotropium are described below in advance of the flora account. The codes of the cited herbaria arc as given in Index Herbariorum (Anonymous 2005). The species are numbered herein as to where they should be inserted in the revision by Craven (1996). For example, species 8a. H. argyreum should be inserted after species 8 in the revision, H. pleiopterum F. Muell., and so on. But note, however, that the positioning of a new species in the sequence does not imply that it is necessarily related to the preceding species; in the case of H. argyreum, its relationships are with the II. ovalifolium subgroup, not with sect. Heliotropium. The relationships of the new species are indicated under the subheading Remarks. Fig. 2. Heliotropium temtifolium R.Br., a short-lived perennial subshrub. L. A. Craven GENERIC CONCEPTS Hilger and Diane (2003) proposed that Boraginaceae subfam. Heliotropioideae (as Heliotropiaceae) be redefined to comprise five genera: Euploca Nutt., Heliotropium, Ixorhea Fenzl, Myriopus Small and Tournefortia L. In their scheme, the majority of the Australian species of Heliotropium including those being treated here, belong to Euploca, with the balance remaining in Heliotropium (which on the data of Hilger and Diane (2003) includes Tournefortia s.s., although they advocated the continued recognition of this taxon (apparently in part due to its long¬ standing acceptance by authors, Hilger, pers. comm.)). The clade names used by Hilger and Diane (2003) will be used where appropriate in the following discussion; these are: EUPLOCA (= Euploca), and HELIOTROPIUM I and HELIOTROPIUM II (both = Heliotropium). The defining morphological and anatomical features of EUPLOCA sensu Diane (2003), Diane etal. (2003), Diane et al. (in press) and Hilger and Diane (2003) apparently are the following: leaves lacking glandular hairs; strigose leaf hairs on a pedestal of enlarged epidermal cells; Kranz- chlorenchyma organisation in the leaves of most species; leaves lacking crystal druses but with crystal needles present; corolla lobes with the margin involute; anther apices cohering and closing the corolla tube; mericarp with surface pits; embryo curved. Contrary to Diane (2003) and Diane et al. (2003), glandular trichomes do occur in EUPLOCA, being present in H. aenigmatum Craven and IT. glanduliferum Craven (Craven 1996). Stout leaf hairs on a pedestal of enlarged epidermal cells occur in H. asperrimum R. Br. (HELIOTROPIUM II) and the feature may be of sporadic occurrence outside EUPLOCA. Of 14 species of EUPLOCA examined by Diane etal., all lack crystal druses and tubes with crystal needles occurring in only two species (Diane et al. 2003) whereas in HELIOTROPIUM I and II druses and tubes occur more commonly and needles have not yet been reported. Even so, of the 29 species of HELIOTROPIUM I and II studied by Diane etal (2003), 12 altogether lack crystals of any form. Kranz-chlorenchyma tissues were observed by Diane etal. (2003) in all studied EUPLOCA species with the exception of two species belonging to H. subsect. Ebracteata I.M. Johnst. (77. ovalifolium and H. procumbens Mill.); Kranz-typc anatomy was not observed in HELIOTROPIUM I and II. Diane et al. (in press) give cochlear aestivation as being the most common type of corolla aestivation in HELIOTROPIUM I and II but they note that quincuncial, apert-introfiex, apert-duplicative and vicinal-cochlear aestivation may also be found. They state that EU PLOCA is characterised by “usually apert aestivation (rarely indistinctly cochlear) and pre-anthetically involute corolla lobes”. The coherent anthers that tend to close the corolla tube in many species of EUPLOCA were believed by Craven (1996) to be associated with inbreeding as the anthers form a ‘cap’ over the stigma, effectively inhibiting the removal and/or deposition of pollen by insects. The presence of two surface pits together with the presence of oil bodies on the adaxial mericarp surface in many species of EUPLOCA may represent a specialised evolutionary trend towards myrmecochory from a more generalised form. Heliotropium europaeum L. (HELIOTROPIUM 11) lacks surface pits but does possess on the adaxial surface of each mericarp a central, distinctly oily tissue which, together with the subfleshy exocarp, may be an adaptation to endozoochory. Craven (1996) discussed oil bodies (as ‘food bodies’) with respect to their occurrence in EUPLOCA but did not record the presence ofthe analogous oily tissue in HELIOTROPIUM II (i.e., in 77. europaeum). My observations ofthe embryo are that it is straight in H. europaeum (HELIOTROPIUM II), slightly curved in H. amplexicaule Vahl (HELIOTROPIUM II), moderately curved in H. asperrimum (HELIOTROPIUM II), and distinctly curved in II. inexplicitum Craven and H. ovalifolium (both EUPLOCA). None of the differences between EUPLOCA and HELIOTROPIUM I and II noted above are particularly strong and, even when taken in combination, they do not make a convincing argument for the recognition of Euploca as a genus distinct from Heliotropium. The presence/ absence of crystals is quite variable and the different structural forms ofthe crystals are perhaps more applicable to infrageneric than to generic classification. Kranz-type anatomy is an evolutionary innovation that may have some significance for classification but this is lessened by its absence in subsect. Ebracteata of EUPLOCA. Corolla aestivation in HELIOTROPIUM I and II apparently is quite variable and it seems that overlapping variation may occur in EUPLOCA. While the bifacial mericarps, commonly with oil bodies, are a very characteristic feature of EUPLOCA, the comparable features in HELIOTROPIUM II are less developed and are indicative that in these aspects EUPLOCA species are more highly specialised. The curvature in the embryo in EUPLOCA is very pronounced relative to that in II. europaeum but this distinction is lessened by the curvature present in other HELIOTROPIUM II species such as 7/. amplexicaule and H. asperrimum. Based on the available evidence from literature, and on my own observations of the flower and fruit morphology of the Australian indigenous and naturalised species of Heliotropium as the genus is traditionally circumscribed, it is my belief that all of them should be classified in the same genus. Consequently, the following new species are described in Heliotropium. 12 New species of Heliotropium (Boraginaceae) SYSTEMATICS Boraginaceae Heliotropium L. 8a. Heliotropium argyreum Craven sp. nov. (Fig. 3) A H. oval folio Forsskal ramulis trichomatibus classis majoris pernumerosis et eis minoris persparsis differt. TYPUS: Australia, Western Australia, Hillside Road, 3.2 km from Woodstock Homestead, 23 April 1958, Burbidge 5825 (holotype: CANI3; isotype: PERTH). Description. Perennial subshrub, hermaphroditic, ascending, up to c. 20 cm tall. Branchlets with hairs predominantly of one class (sparse small curved hairs also occur), the hairs dense, appressed to ascending. Leaves 7-12 mm long, 1.5-2 mm wide, short-petiolate. Leaf blade revolute, narrowly oblong-elliptic, narrowly elliptic to narrowly obovate; abaxial surface with the hairs uniformly distributed, the hairs of one class and dense and appressed. Cymes simple or 2-branched, without bracts or leaves. Calyx lobes with the hairs on the abaxial surface as on the leaf blade. Corolla white, 3.5 mm long. Anthers very narrowly ovate, 1.2 mm long, the apex long acuminate, coherent at the apex. Gynoecium 1 mm long; ovary 0.3 mm long; style 0.1 mm long; stigmatic apparatus 0.5 mm long, the disc relatively short (0.1 mm) and the cone long (0.4 mm). Mericarps with macrohairs, the hairs appressed to spreading-ascending and up to c. 0.5 mm long; elliptic-ovate, 1.2 mm long, 1.3 times as long as wide, the apex acute; commissures not pitted, with a very weakly developed food body. Distribution and ecology. Heliotropium argyreum has been recorded from a single locality in the northern Pilbara region of Western Australia (Fig. 10) where it was noted to grow on loose rubble on a limestone ridge. Remarks. Craven (1996) included material of this species in the concept of H. ovalifolium adopted in the revision, albeit with reservations due to the indumentum of dense, appressed to ascending large hairs and paucity of small hairs. The opportunity has now been taken to reconsider the taxonomic status of both Pilbara forms of H. ovalifolium mentioned by Craven (1996: 560) and it is concluded that separate recognition is warranted for both. The second form, from the Meekatharra region, is described below as H. mitchellii Craven. Etymology. The epithet is derived from the Greek word, argyros, silver, in reference to the silvery foliage that this species possesses. 12a. Heliotropium mitchellii Craven sp. nov. (Fig. 4) A H. ovalifolio Forsskal ramulis trichomatibus classis majoris plerumque sparsis et eis minoris pernumerosis, et mericarpiis majoris (2 mm longis) differt. TYPUS: Australia, Western Australia, 42 km SE of Meekatharra on the Sandstone road, lower slopes of Mt. Yagahong, 9 September 1982, Craven 7531 (holotype: CANB; isotypes: A, E, L, MEL, PERTH, W). Other specimens examined. Australia. Western Australia: c. 18 km W of Meekatharra, 3 September 1957, Speck 592 (CANB); 15 km W of Hillvicw Homestead on slopes of the Yagahong basalt, 25 March 1984, Mitchell 1217 (CANB); Mt. Meekatharra, 1 August 1966, Fairall 2183 (CANB); Gabanintha Hill, SE of Meekatharra, August 1963, Gardner 14474 (CANB, PERTH). Description. Perennial subshrub, dioecious, ascending to spreading, up to c. 40 cm tall. Branchlets with hairs of two classes, the hairs of the larger class usually sparse, appressed to ascending, and of the smaller class very dense, curled and appressed to recurved. Leaves 5-22 mm long, 3-6 mm wide, short-petiolate. Leaf blade flat, elliptic to obovate, sometimes narrowly so; abaxial surface with the hairs uniformly distributed, the hairs of the larger class appressed and of the smaller class curled to straight and appressed to recurved. Cymes during anthesis curled, simple or 2-branched, without bracts or leaves. Calyx lobes with the hairs on the abaxial surface as on the leaf blade. Corolla white to creamy white with a yellow throat, 5.3-6 mm long. Anthers (in male functional flowers) very narrowly ovate to sublinear, 1 mm long, the apex long acuminate, coherent at the apex. Gynoecium (in female functional flowers) 1.7 mm long; ovary 0.7 mm long; style 0.3 mm long; stigmatic apparatus 0.7 mm long, the disc relatively short (0.2 mm) and the cone long (0.5 mm). Mericarps with macrohairs, the hairs appressed to ascending and up to c. 0.5 mm long; oblong-elliptic, 2 mm long, 1.7 times as long as wide, the apex acute to obtuse; commissures only slightly pitted, with a weakly developed food body. Distribution and ecology. Heliotropium mitchelli has been recorded in the Meekatharra region of Western Australia (Fig. 10) and has been noted to grow on sandstone lalus/scree with open mulga shrubland, on very stony ground, under Acacia linophylla , and on boulder-strewn basaltic soil with Eremophila-Acacia scrub. Remarks. Craven (1996) included this plant in his concept of H. ovalifolium although, as noted above under H. argyreum , it was considered atypical for that species. The Meekatharra plant has indumentum with the hairs of the larger class mainly distributed sparsely and of the smaller class distributed very densely. In addition, the meriearps are larger than in H. ovalifolium s.s. These differences are now considered to be taxonomically significant and accordingly a new species is described. Etymology. The species is named in honour of Andrew Arthur Mitchell, formerly of Meekatharra, Western Australia, and presently of Darwin, Northern Territory. Andrew developed considerable familiarity with the flora of the eastern Pilbara region while conducting rangeland assessment studies there and has made many important 13 L. A. Craven SfUM •UEN$E 537*9 ^NSERUA w.& HERBARIUM AUSTRALIENSE *•. ^825 Jw2j5.iv.l95S Nam* H^lotropiu^iOja^^lluo Hob. loose rubble on linestone ridge ^.Hillside Hd., 2 ailes irom Woodstocic H.S.t ff.A. deicrraiaivit LA. Craven Australian National Herbarium (CANB) 200£ C.SI.R.O. Division <,r Puir I.vocsntv •J+-r >C Oeirrminaut HrC/it-Mr v Vc* C.U. N.T.Burbidge „ erect perennial herb.Flowers white 25mm ^LCclw* * Fig. 3. Heliotropium argyreum Craven, holotype. 14 New species of Heliotropium (Boraginaceae) AUSTRALIAN NATIONAL HERBARIUM {CANB) nonwealth Scientific A Industrial Researcl Organisation; Australia Heliotropium /v^-A<£L»_S2^>A — AUSTRALIA; Mestcrn Australia 42 ka SE of Meekatharra on the Sandstone road, lower slopes of Xt Yagahong. 26* 55 S 1 IB* 40 E Sandstone talus/scree with open nulga shrubland. Perennial, corolla creasy white with yellow throat. L.A.Craven 7531 » Sep 1982 Fig. 4. Heliotropium mitchelli Craven, part of liolotype. 15 L. A. Craven contributions to our knowledge of the heliotrope flora of the Australian western arid zone, including making the second and third collections of the unusual species H. muticum Dornin that was first collected in the latter part of the 19 lh century. 23a. Heliotropium arenitense Craven sp. nov. (Fig. 5) A H.frohlichii Craven ramulis foliisque trichomatibus patentiascendentibus usque patentibus, cymis 1-2-floris, et mericarpiis glabris differt. TYPUS: Australia, Northern Territory, Kakadu National Park, 6.5 km NE of El Sharana, on a tributary of Koolpin Creek, 22 April 1990, S/ee & Craven 2755 (holotype: CANB; isotypes: A, B, BRI, DNA, E, G, L, M, MEL, NSW, P, US, W). Other specimens examined. Australia. Northern Territory: Kakadu National Park, [near type locality,] 22 April 1990, Dunlop c£ Munns 8610 (BRI, CANB, DNA). Description. Perennial subshrub, hermaphroditic, sprawling, up to c. 30 cm tall. Branchlets with hairs of one class, the hairs dense, spreading. Leaves 6-15 mm long, 0.7-1.2 mm wide, short-petiolate. Leaf blade revolute, linear; hairs on the abaxial surface uniformly distributed, spreading-ascending to spreading. Cymes 1-2-flowered, bracteate. Inflorescence bracts on the abaxial surface with the hairs fine, and as on the leaf blade. Calyx lobes with the hairs on the abaxial surface as on the inflorescence bracts. Corolla white with a yellow throat, 4.5-5 mm long. Anthers ovate, 0.7 mm long, the apex acuminate, coherent at the apex. Gynoecium 1.4 mm long; ovary 0.4 mm long; style 0.5 mm long; stigmatic apparatus 0.5 mm long, the disc (0.2 mm) fairly equal in length to the cone (0.3 mm). Mericarps glabrous; oblong-ovate, 1 mm long, 1.4 times as long as wide, the apex obtuse; commissures strongly pitted, the pit suborbicular, with a very weakly developed food body. Distribution and ecology. Heliotropium arenitense has been recorded only on the sandstone plateau in the El Sharana region of the Northern Territory (Fig. 10) where it grows among rocks on scree in very open Eucalyptus miniata woodland. Remarks. The affinities of this species are difficult to assess. The combination of short, spreading hairs, one to two-flowered cymes, hermaphroditic flowers and glabrous mericarps set 11 . arenitense apart from its other Australian congeners. Possibly it is related to the species of the H. tenuifolium subgroup but it could well prove to be a hermaphroditic member of the H. epacrideum subgroup as it has some similarities with H.frohlichii Craven and H. lapidicola Craven in the foliage; it is tentatively assigned to the latter subgroup. In the Northern Territory, it is the only species of the genus that appears to be restricted to unmetamorphosed sandstones; other species that may occur on these substrates in the Territory are not confined to them. However, it is acknowledged that the species is very poorly known, the two collections known to me having been made on the same day and from localities very close together, and it may be found to be more widely distributed than is presently known. Etymology. The specific epithet is derived from the geological name for sandstone, arenite, and refers to the colloquial name used for the sandstone plateau country of northern Australia, i.e. ‘the sandstone’. 28a. Heliotropium lapidicola Craven sp. nov. (Fig. 6) A H. frohlichii Craven ramulis foliisque trichomatibus patentiascendentibus usque patentibus, et mericarpiis ovatis et commissuris cum penu differt. TYPUS: Australia. Queensland, Maronan Station, Fullarton River garnet locality (148.6 km ESE ofMount Isa by air), 15 May 2001, Fraser357 (holotype: BRI; isotypes: A, CANB, E, L, MEL). Other specimens examined. Australia. Queensland: Maronan Station, Fullarton River garnet locality (148.6 km ESE ofMount Isa by air), 29 March 2002, Fraser 372 (AD, BRI, CANB, DNA, G, K, L, MEL, P, W), 31 March 2002, Fraser 374 (A, BRI, CANB, L, M, P, US). Description. Perennial subshrub, dioecious, sprawling to bushy, up to 40 cm tall. Branchlets with the hairs of one class, the hairs dense, spreading-ascending to spreading. Leaves 8-45 mm long, 0.7-3 mm wide, sessile. Leaf blade revolute, linear to narrowly oblong; hairs on the abaxial surface coarse and uniformly distributed, ascending on the midrib, oppressed on the lamina, and ascending to spreading-ascending on the margin. Cymes simple, straight at anthesis, bracteate. lnfloresence bracts with the hairs on the abaxial surface uniformly distributed and oriented as on the leaves. Calyx lobes with the hairs on the abaxial surface coarse in female plants and much finer in male plants, uniformly distributed, the hairs oppressed to ascending on the lamina and midrib and ascending to spreading-ascending on the margin, with those on the margin generally shorter in length than those on the lamina. Corolla white, 2.8-3 mm long. Anthers (in male functional flowers) ovate, 1 mm long, the apex acuminate, coherent at the apex. Gynoecium (in female functional flowers) 2.2 mm long; ovary 0.6 mm long; style 1.1 mm long; stigmatic apparatus 0.5 mm long, the disc (0.2 mm) fairly equal in length to the cone (0.3 mm). Mericarps with microhairs, the hairs appressed and up to c. 0.2 mm long; ovate, 1.5-1.6 mm long, 1.5-1.6 times as long as wide, the apex obtuse; commissures moderately pitted, the pit elliptic, with a food body. Distribution and ecology. Heliotropium lapidicola has been recorded from the Fullarton River region in Queensland (Fig. 10), growing on a rocky schistose ridge with acacias, low shrubs and herbs, and a few stunted eucalypts. Remarks. This species is clearly closely related to H. frohlichii, having leaves that are similar in shape, 16 New species of Heliotropium (Boraginaceae) Flora of AUSTRALIA AUSTRALIAN NATIONAL HERBARIUM (CANS) BORAGINACEAE Heliotropium AUSTRALIA: Northern Territory Kakadu NatoasJ Park. 6 5 km NE of El Sharana. (on tributary of Kodp«n Creek) 3r*J ret JIM JIM sheet 55471-337109 (1:100000) 13* 2TS 132*35 E Edge c* sandstone pavement /u« up out of shakrw creek, very open Eucalyptus numata woodland Denser, 1 branched sub-shrub to 30 cm tan. rooted amongst rocks: flowers white with yeaow throat A.V. Slee. LA. Craven 2755 22 Apr 1990 !!lillllllll ! |]||||l11ii:illlili!IIIEIlil CANO 397789.1 25mm Fig. 5. Heliotropium arenitense Craven, part of holotype. 17 L. A. Craven Scattered '«>>• top A.Fraser 357 15 May 2001 CANB 641866 25mm Fig. 6. Heliotropium lapidicola Craven, holotype. New species of Heliotropium (Boraginaceae) similarly structured cymes, generally similar gynoecium, etc. The two species differ especially in the possession by H. lapidicola of spreading/spreading-ascending hairs on most of the organs, the comparable indumentum in H.frohlichii being appressed. The mericarps of H. lapidicola have food bodies, unlike H.frohlichii. Etymology. The specific epithet is derived arbitrarily from the Latin lapis, precious stone, gem, jewel, and - cola , dweller, inhabitant, in reference to the occurrence of this plant at the Fullarton River garnet locality. 28b. Heliotropium microsalsoloides Craven sp. nov. (Fig. 7) A H. epacrideo F. Muell. ex Benth. foliis angustioribus (0.4-1.3 mm latis), et mericarpiis oblongo-obovatis usque oblongo-ovatis 1.5—1.8 mm longis et commissuris sine penu differt. TYPUS: Australia, Western Australia, King Sound, Sunday Island, at site of abandoned mission station, 2 June 1993, Craven Stewart 9271 (holotype: CANB; isotypes: A, AD, B, DNA, E, G, K, L, M, MEL, P, PERTH, UC, US, W). Other specimens examined. Australia. Western Australia: King Sound, Sunday Island, 24 September 1988, Carter 328 (PERTH), 1 April 1992, Mitchell 2234 (CANB, PERTH), 25 August 1993. Carter 673 (PERTH), Mitchell 32S9 (CANB, PERTH), 4 April 2000, Mitchell 6098 (CANB); Cone Bay Hermits Camp, 4 July 1996, Kenneally 11710 (PERTH); Xerxes camp about 1 km NW of Alcatraz Island in Cone Bay, 31 May 1995, Mitchell 3980 (CANB); 30 km ESE of Sale River mouth at junction of unnamed creek, 13 May 1986, Kenneally 9615 (PERTH). Description. Perennial subshrub, dioecious, erect to spreading, up to c. 35 cm tall. Branchlets with the hairs fine relative to those on the leaves and calyx, the hairs of two classes (sometimes appearing to be one class when the hairs of the larger class are sparse to absent), the hairs dense to moderately dense, those of the larger class spreading-ascending and of the smaller class ascending, spreading or recurved. Leaves 1.5-6 mm long,0.4-1.3 mm wide, sessile. Leaf blade very slightly revolute, narrowly elliptic to narrowly ovate; hairs on the abaxial surface coarse and usually distributed more towards the midrib and central part of the lamina than towards the margin, appressed on the lamina and midrib and spreading- ascending on the margin. Cymes 1-flowered. Calyx lobes with the hairs on the abaxial surface coarse and uniformly distributed or sometimes absent on the lamina towards the margin, the hairs appressed on the lamina and midrib and spreading-ascending on the margin, with those on the margin generally equal in length to those on the lamina. Corolla white, 2.5-3 mm long. Anthers (in male functional flowers) ovate, 0.8 mm long, the apex acuminate, not coherent at the apex. Gynoecium (in female functional flowers) 0.8 mm long; ovary 0.3 mm long; style 0.25 mm long; stigmatic apparatus 0.25 mm long, the cone obsolete. Mericarps with microhairs, the hairs appressed to ascending and up to c. 0.15 mm long; oblong-obovate to oblong-ovate, 1.5-1.8 mm long, 1.4-1.5 times as long as wide, the apex obtuse; commissures strongly pitted, the pit elliptic, without a food body. Distribution and ecology. Heliotropium microsalsoloides has been recorded from the Buccaneer Archipelago-Sale River region in Western Australia (Fig. 10), growing among rocks and in fissures on sandstone, and in hummock grassland on sandstone cliffs. Remarks. The gewfa// of this remarkable little plant at first suggests it might be a species ofChenopodiaceae. The leaves are closely clustered on very short, secondary shoots and, together with the bristly leaf hairs, give the plant a very chenopodiaceous aspect. Its relationships are with H. epacrideum F. Muell. ex Benth. and H. muticum. Etymology. The specific epithet is derived from the Greek mikros, small, little and - eides , like, resembling and Salsola , a genus of Chenopodiaceae, as the plant superficially resembles a dwarf version of the widespread S. tragus L. 36a. Heliotropium albrechtii Ci aven sp. nov. (Fig. 8) A H. geochare Domin mericarpiis minoris (1.3 mm longis), et trichomatibus patentiascendentibus usque patentibus et minoribus (usque 0.2 mm longis) differt. TYPUS: Australia, Northern Territory, 14 km NW of Hamilton Downs homestead, near CSIRO exclosure, 10 March 1997, Albrecht 8010 (holotype: NT; isotype: CANB). Description. Annual herb, hermaphroditic, erect, up to c. 25 cm tall. Branchlets with hairs of one class, the hairs moderately dense, appressed. Leaves 12-40 mm long, 1.2-3.5 mm wide, short-petiolate. Leaf blade revolute, linear-elliptic; hairs on the abaxial surface uniformly distributed, appressed. Cymes simple, bracteate. Inflorescence bracts on the abaxial surface with the hairs coarse, uniformly distributed, and appressed on the lamina and spreading-ascending on the margin. Calyx lobes with the hairs on the abaxial surface as on the inflorescence bracts, and with the hairs on the margin generally longer than those on the lamina. Corolla white, 4.5 mm long. Anthers ovate, 0.7 mm long, the apex acuminate, coherent at the apex. Gynoecium 1.1 mm long; ovary 0.4 mm long; style 0.2 mm long; stigmatic apparatus 0.5 mm long, the disc (0.2 mm) fairly equal in length to the cone (0.3 mm). Mericarps with microhairs, the hairs spreading-ascending to spreading and up to c. 0.2 mm long; suborbicular, 1.3 mm long, 1.1 times as long as wide, the apex obtusely acute; commissures strongly pitted, the pit circular, with a very weakly developed food body (no fatty material observed). 19 L. A. Craven -U STKAI UVN VTIONU. Ill kllVKIl M (Civil Commonwealth Soentirtc i Industrial Research Or jarusancai. Australia BORACBNACEAE Heliotropium AUSTRALIA: Western Australia Sunday I viand. Kin* Sound 16* 23 * S 123’ 10 ' E Anion* rums of abandoned mission station on undstcnc-: burnt the previous year. dctcrmirvjui LA. Craven Australian Malawi*] Herbarium (CANS) 20U^ 25mm Exnccatac from collaborative plant exploration by the bertnna CANS. DMA. ISC. and Mac Stewart of Arkansas. with the major sponsor being the U S Department of Agriculture (ARS - GermpUsm Unit) Dioecious shrubs to 30 a tC cm; corolla white. LA. Craven and J.McD. Stewart 9271 2JuaI993 Fig. 7. Heliotropium microsalsoloides Craven, part of holotype. New species of Heliotropium (Boraginaceae) 25mm D€Q SOto dfttrminiMl L-A- Craven Australian National Herbarium (CANB) 200 Aibrecfet 0 Z IG1C &2RAGINACEAE V7 can A0094882 10 nxn Ml Fig. 8. Heliotropium albrechtii Craven, holotype. 21 L. A. Craven lictermiruvit LA. Craven Australian National Herbarium (CANB) 25mm WESTERN AUSTRALIAN HERBARIUM. PERTH Flora of Western Australia Snail, sparse herb to 40 co. Foliage silvery. Flowers off-white. Frequent on open scree slopes. Loc. E facing slopes of the Napier Range just N of Wombarella Gap. Lat. I 70 1 7 ‘ S Long. 124° 46* E Coll. E.M. Goble-Gar ratt 569 12 May 19 88 OUPUCATES DISTRIBUTED H&1 ' CRnB < Fig. 9. Heliotropium viator Craven, holotype. 22 New species of Heliotropium (Boraginaceae) Distribution and ecology. Heliotropium albrechtii has been recorded only in the Hamilton Downs Station region of the Northern Territory (Fig. 10) where it grows on a heavy soil plain with Astreblapectinata. Glycine falcata and Sauropus trachyspermus. Remarks. The species is a member of the II. tenuifolium subgroup within which its closest relative may be H. geocharis Domin, a species that occurs in similar habitats on the northern black soil plains of Western Australia, the Northern Territory and Queensland. Heliotropium geocharis , however, has bigger mericarps with appressed hairs up to 0.4 mm long. Etymology. The specific epithet honours the collector of the type material, David Edward Albrecht of Alice Springs, Northern Territory. David has made many interesting collections during the course of his work on the Australian arid-zone flora. 62a. Heliotropium viator Craven sp. nov. (Fig. 9) A H. tenuifolio R. Br. lobis calycis trichomatibus grossis, et mericarpiis trichomatibus apice hamato differt. TYPUS: Australia, Western Australia, E facing slopes of the Napier Range just N of Wombarella Gap, 12 May 1988, Goble-Garratt 569 (holotype: PERTH; isotypes: CANB, K h.v.). Other specimens examined. Australia. Western Australia: Napier Range, 2 km NNW of Napier Downs Homestead, 13 April 1988, Cranfieid6332 (MEL, PERTH); 1 km NNW of Barker River Gorge, 20 April 1988, Cranfieid 6470 (CANB, PERTH). Description. Annual (or short-lived perennial?) herb, hermaphroditic, ascending, up to c. 40 cm tall. Branchlets with hairs of one class, the hairs dense, appressed. Leaves 5-15 mm long, 0.7-1.2 mm wide, short-petiolate. Leafblade revolute, linear; hairs on the abaxial surface uniformly distributed, appressed. Cymes during anthesis straight, simple to several-branched, bracteate. Inflorescence bracts on the abaxial surface with the hairs coarse, uniformly distributed, appressed on the lamina and spreading- ascending on the margin. Calyx lobes with the hairs on the abaxial surface of the lamina generally as long as those on the margin, coarse, uniformly distributed, appressed on the lamina and spreading-ascending on the margin. Corolla white to off-white, 5 mm long. Anthers ovate, 0.8 mm long, the apex acuminate, coherent at the apex. Gynoecium 2.2 mm long; ovary 0.3 mm long; style 1.4 mm long; stigmatic apparatus 0.5 mm long, the disc (0.2 mm) fairly equal in length to the cone (0.3 mm). Mericarps with microhairs, the hairs ascending to spreading-ascending and up to c. 0.2 mm long and generally hooked at the apex; ovate, 0.9-1.1 mm long, 1.3-1.4 times as long as 120 MO Fig. 10. Distributions of Heliotropium species. H. argyreum O' H. mitchellii V; H. arenitense I; H. lapidicola ^; H. microsalsoloides H. albrechtii *tk; H. viator 23 L. A. Craven wide, the apex acute; commissures strongly pitted, the pit suborbicular, without a food body. Distribution and ecology. Heliotropium viator has been recorded in the Napier Range-Barker River region of Western Australia (Fig. 10) and has been noted to grow on open scree slopes, in shallow brown clay over limestone on a rocky hill-top and slope, and in open tussock grassland with Adansonia and Sorghum on red clay with exposed limestone outcrops. Remarks. This taxon is a member of the H. tenuifolium subgroup as circumscribed by Craven (1996). It is characterised by its limestone-related ecology and the hooked mericarp hairs, the latter feature being unusual in the Australian species of the genus and shared with H. paniculatum R.Br. Etymology. The epithet is a noun in apposition, Latin, viator, traveller, wayfarer, and refers to the hooked mericarp hairs whereby this species is believed to adhere to animals and thus be dispersed. Partial keys to species of Heliotropium The species described above have been inserted in the keys given in Craven (1996) as follows: I. Key to H. argyreum and H. mitchellii Replace the first lead of couplet 3 in Key 1 (Craven 1996: 534) with the following: 3. Mericarp hairs appressed to spreading-ascending A. Branchlet indumentum seemingly of a single class of hair, these appressed to ascending, sparse small hairs also occur .8a. H. argyreum A. Branchlet indumentum consisting of hairs of two classes B. Branchlets with hairs of the larger class frequent; mericarps 1-1.4 mm long. ...13. H. ovalifolium B. Branchlets with hairs of the larger class usually sparse; mericarps 2 mm long. .12a . H. mitchellii 2. Keys to //. arenitense Replace couplet 5 in Key 2 (Craven 1996: 535) with the following: 5. Mericarps 1 mm long.23a. H. arenitense 5. Mericarps more than 1.2 mm long A. Mericarps 1.6-1.7 mm long, 1.4-1.5 times as long as wide.34. H. transforme A. Mericarps 1.3-1.6 mm long, 1.4-2 times as long as wide.35. H. vestitum Replace the first lead of couplet 7 in Key 4 (Craven 1996: 544) with the following: 7. Mericarps with the commissure pit orbicular to suborbicular A. Mericarps with microhairs, ovate to broadly ovate, 1.2-1.4 mm long.27. H. filaginoides A. Mericarps glabrous, oblone-ovate, 1 mm long. 23a. H. arenitense 3. Keys to H. lapidicola Insert after the second lead of couplet 23 in Key 2 (Craven 1996: 536) the following: A. Plant a perennial, dioecious subshrub. .28a. H. lapidicola A. Plant an hermaphroditic annual. Insert after the first lead of couplet 51 in Key 2 (Craven 1996: 537) the following: A. Mericarp pits elliptic.28a. H. lapidicola A. Mericarp pits orbicular to suborbicular. 4. Key to //. microsalsoloides Replace the first lead of couplet 1 and couplet 2 in Key 4 (Craven 1996: 544) with the following: 1. Leaf blade on the abaxial surface with the hairs not distributed fairly uniformly 2. Leaf blade on the abaxial surface with the hairs all appressed.24. H. cupressinum 2. Leaf blade on the abaxial surface with the hairs either ascending to spreading or else appressed on the lamina and midrib and spreading-ascending on the margin A. Leaf blade on the abaxial surface with the hairs ascending to spreading. .26. //. epacrideum A. Leaf blade on the abaxial surface with the hairs appressed on the lamina and midrib and spreading-ascending on the margin... 28b. H. microsalsoloides 5. Keys to H. ulhrechtii Replace the second lead of couplet 22 in Key 2 (Craven 1996: 536) with the following: 22 Mericarps broadly ovate or suborbicular, the commissures moderately to strongly pitied A. Mericarps 0.8-1.1 mm long; calyx lobes with the hairs on the margin generally equal in length to shorter than those on the lamina. .15 . H. paniculatum A. Mericarps 1.3 mm long; calyx lobes with the hairs on the margin generally longer than those on the lamina.36a. H. albrechtii Replace the second lead of couplet 61 in Key 2 (Craven 1996: 538) with the following: 61. Mericarps 1-1.4 mm long A. Mericarps with weakly developed food bodies; leaf blade on the abaxial surface with the hairs appressed; gynoecium 1.1 mm long. ...36a. H. alhrechtii A. Mericarps without food bodies; leaf blade on the abaxial surface with the hairs appressed on the lamina and appressed to spreading-ascending on the margin; gynoecium 2 mm long. .53. H. melanopedii 24 New species of Heliotropium (Boraginaceae) 6. Key to H. viator Insert after the first lead of couplet 17 in Key 2 (Craven 1996: 535) the following: 17A. Mericarps with at least some of the hairs hooked at the apex B. Gynoecium 0.8-1.2 mm long; mericarps broadly ovate, 1-1.1 times as long as wide. .. 15. H. pcmiculatum B. Gynoecium 2.2 mm long; mericarps ovate, 1.3-1.4 times as long as wide. 62a. H. viator A. Mericarps without hooked hairs. Insert after the second lead of couplet 17 in Key 2 (Craven 1996: 537) the following: A. Mericarps with at least some of the hairs hooked at the apex.62a. H. viator A. Mericarps without hooked hairs. ACKNOWLEDGMENTS The Directors and/or Curators of the following herbaria are thanked for the opportunity to study collections in their care: CANB,NT, PERTH. I especially wish to acknowledge the many people who have been collecting and/or sending specimens, doubtless in the desire to further extend my frustration with this taxonomically complex group of plants: without your interest our knowledge of this rich component of the Australian flora would be significantly poorer. Prof. Hartmut Hilger is thanked for his generous and speedy provision of publications, and for presenting a seminar in Canberra on the phylogenetic studies of Boraginaceae s.l. being conducted in his laboratory. The images were taken and prepared by Carl Davies. REFERENCES Anonymous. 2005. Index herbariorum: a global directory of public herbaria and associated staff. URL: http://sciweb.nybg. org/sciencc2/IndexIlerbariorum.asp (Accessed 31 January 2005) Craven, L.A. 1996. A taxonomic revision of Heliotropium (Boraginaceae) in Australia. Australian Systematic Botany 9: 521-657. Diane, N. 2003. Systematic analysis of the lleliotropiaceae based on molecular and morphological-anatomical data. PhD thesis, FU Berlin. URL: http://www.diss.fu-berlin.de/2003/197/ index.html Diane, N.Jacob, C. and Hilger, 11.11.2003. Leaf anatomy and foliar trichomes in Heliotropiaceae and their systematic relevance. Flora 198: 468-485. Diane, N„ Forthcr, if, Hilger, H.H. and Weigend, M. In press. Heliotropiaceae Schrad. In: Kubitzki, K. (ed.) Families and genera of the flowering plants. Springer: Berlin. Hilger, H.H. and Diane, N. 2003. A systematic analysis of Heliotropiaceae (Boraginales) based on trnL and ITS1 sequence data. Botanische Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzengeographie 125: 19-51. Accepted 24 November 2004 25 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21: 27-36 A new genus and species of box jellyfish (Cubozoa: Carybdeidae) from tropical Australian waters LISA-ANN GERSHWIN 1 AND PHILIP ALDERSLADE 2 'School of Marine Biology and Aquaculture, James Cook University Townsville OLD 4811, AUSTRALIA and Australian Institute of Marine Science Townsville QLD 4810, AUSTRALIA lisa.gershwin@jcu.edu.au 2 Museum and Art Galleiy of the Northern Territory GPO Box 4646, Darwin NT0801, AUSTRALIA pltil. alderslade@nt.gov. an ABSTRACT A new genus and species of cubozoan is reported from the Northern Territory and Queensland. It possesses the following autapomorphic characters with respect to other known forms in the 7amoya/Irukandji group ofcarybdeids: a robust, matchbox-sized body; broad and overhanging pedalial inner wings; laminar-branching velarial canals; and qualitatively and quantatively unique nematocysts. Furthermore, while it shares some characters with other taxa in the group, it is unique in its combination of major morphological characters: like Tamoya, Carukia, Malo and ‘Morbakka’, it has frown-shaped rhopalial niche ostia; like Carukia , Malo and ‘Morbakka’, it has rhopalial horns, perradial lappets, and lacks gastric cirri; like Tamoya and ‘Morbakka’, it has a prominent thorn-shaped upward- pointing diverticulum near the base of the pedalial canal and greatly flared tentacle bases; like Malo and ‘Morbakka’, it has two rows of nematocyst warts on the perradial lappets; like Malo, the rhopaliar horns are short and broad. Its cnidome comprises Type 4 microbasic p-mastigophores on the tentacles, and spherical isorhizas on the bell. Recent molecular phylogenetic studies place it, along with ‘Morbakka’, intermediate between the Carukia clade and the Malo clade. It is considered a mild health risk to humans. Keywords: Gerongia rifkinae, new genus, new species, Cubozoa, Cnidaria, Coelenlerata, Northern Territory, Queensland, Australia, taxonomy, Irukandji syndrome. INTRODUCTION Jellyfishes of the Southern Hemisphere are poorly known. In particular, the Cubozoa, which comprises some of the world’s most dangerous animals, remains little studied. The cubozoans of Australia were investigated taxonomically by Haacke (1886, 1887), Southcott (1956, 1967), Barnes (1965), Moore (1988), Hartwick (1991), and Gershwin (2005b). Presently, more cubozoan species are documented from Australia (6) than from any other country; however, recent studies have indicated that the Australian cubozoan fauna remains poorly documented and that at least 17 new species remain undescribed (Gershwin 2005a). Of particular interest in Australia are the dangerous jellyfishes, i.e., the so-called ‘box jellies and Irukandjis’ (note that the term ‘box jellies’ or ‘box jellyfish’ is used outside Australia to refer to cubozoans in general, whereas in north Queensland it is often used to refer to chirodropids only - a circumstance that we do not support). Up until recently, only a single species of Irukandji was named and classified, namely Carukia barnesi Southcott. Other species have recently been identified as causing systemic symptoms, in some cases even life-threatening Irukandji syndrome (Gershwin 2005a, b, and references therein), but most have yet to be described. In some species ambiguous data exist e.g., Barnes tested a form that he called ‘Pseudo- Irukandji’ (Kinsey 1988), which looked to him like an Irukandji but gave only fleeting neuralgias (it should be noted, however, that the animals were nearly dead, after being kept in a hot plastic pool for 12-14 hours, by which time the poor condition of the animals was thought to be due to the strong plastic smell being given off by the pool (Nick Barnes, pers. comm., January 2000). This same species has been linked through known-specimen events to a sting with mild Irukandji symptoms and a sting with no systemic symptoms whatsoever (Gershwin, unpublished notes), and has been linked to a fatal sting 27 L.-A. Gershwin and P. Alderslade by its nematocysts (Gershwin 2005a). However, it does not appear that all Irukandji syndrome-producing species pose a significant health risk, i.e., there may be species- specific variation in syndrome intensity. For example, the undescribed group ‘Morbakka’ has been known for two decades to give mild Irukandji symptoms (Fenner et al. 1985), as has the ‘Darwin carybdeid’ more recently (Currie 2000a; O’Reilly et al. 2001). The ‘Darwin carybdeid’ has caused only localized pain or mild Irukandji syndrome in the few recorded instances where known stinging has occurred (see Williamson et al. 1996;Currie 2000a; O’Reilly etal. 2001). Possibly because it is regarded as only a mild health threat, this species has been mentioned only a handful of times in the medical literature (Currie and Wood 1995; Williamson et al. 1996; Currie 2000a; Currie 2000b; O’Reilly et al. 2001). However, because it is morphologically and genetically in the Irukandji group (Gershwin 2005a), it may prove comparatively informative toward a better understanding of Irukandji venom and the Irukandji syndrome. The purpose of this paper is to formally describe the ‘Darwin carybdeid’ as a new genus and species of cubozoan from the Northern Territory and Queensland. MATERIALS AND METHODS Most specimens were obtained by dip-netting at high tide in the vicinity of mangroves; similar specific data do not exist for the remainder. Morphology of preserved specimens was studied under a dissecting microscope, following the criteria of Gershwin (2005a). Sex was determined by the visible presence of eggs on female gonads or patterns resembling human finger-prints on the male gonads. Measurements were made on preserved material with Max-Cal digital calipers, to the nearest 0.01 mm. Nematocysts were examined and measured with a Leica DM LB compound microscope and Leica IM-50 Image Manager v. 1.20 for Windows; all observations and photographs were made through a 40x objective. Nematocysts were identified following the keys of Calder (1974), Mariscal (1971), and Williamson et al. (1996). Institutional abbreviations used: specimen numbers prefixed ‘NTM’ indicate Museum and Art Gallery of the Northern Territory; specimen numbers prefixed ‘AM’ indicate Australian Museum; specimen numbers prefixed ‘SAM’ indicate South Australian Museum; those prefixed with ‘RVS’ are from the collection of the late Ronald V. Southcott, now housed along with his notes at the South Australian Museum. The prefix ‘JHB’ indicates reference to the collection and notes of the late Jack Barnes; the collection is housed at the Museum ofTropical Queensland, and the notes are archived at the James Cook University Library in Townsville, Queensland. SYSTEMATICS Phylum Cnidaria Verrill, 1865 (Hatschek, 1888) Subphylum Medusozoa Petersen, 1979 Class Cubozoa Werner, 1973 Order Carybdeida Gegenbaur, 1857 (Werner, 1984) Family Tarnoyidae Haeckel, 1880 Gerongia gen. nov. Type species. Gerongia rifkinae sp. nov., here designated. Genus feminine. Diagnosis. Tarnoyidae with gastric phacellae absent; rhopalial horns short, broad, curved inward; adaxial pedalial keels broadly rounded, overhanging; upward- pointing, blind-ending thorn-shaped pocket at bend ol pedalial canal; base of tentacles greatly flared; large, ballooned stomach attached to subumbrellar walls by moderately well-developed perradial mesenteries; two parallel rows of low, rounded, ncmatocyst warts on the perradial lappets of the velarium. Remarks. Gerongia gen. nov. would not be easily mistaken for other genera in the Tarnoyidae (Table 1). It is distinct from all others in possessing the following autapomorphic characters: a robustly developed matchbox¬ sized body, broad and overhanging pedalial canal inner keels, laminar-branching velarial canals, and elongate club- shaped nematocysts with a progressively coiled tubule. Like all but Tamoya Muller, it lacks gastric phacellae and possesses perradial lappets and rhopaliar horns. Like Tamoya and ‘Morbakka’ only, it possesses a conspicuous pedalial canal ‘thorn’ and flared tentacle bases, but unlike Tamoya and ‘Morbakka’, the tentacles are round in cross section, the velarial canals lack lateral diverticula, and the inner pedalial keels are greatly flared and overhanging. Like Carukia Southcott, it has a frown-shaped rhopalial niche ostia and lacks gastric phacellae, but the tentacles are entirely different, having rounded, evenly spaced bands compared with tailed and widely spaced bands in Carukia. Additionally, Carukia spp. are diminutive in height and bulk, with a rounded pyramidal body, whereas Gerongia gen nov. is about 4-5 times taller, with a heavy, robust, more boxy body, and in Carukia the exumbrellar warts are bright red, whereas those of Gerongia gen. nov. are pale and uncoloured. Like Malo Gershwin, 2005b, it has broad, short, curved rhopaliar horns; rows of nematocyst warts on the perradial lappets; and the tentacles are round in cross section; however, the body is an entirely different shape, being robustly boxy in Gerongia but tall and relatively slim in Malo. The tentacles are heavy and flared at the base in Gerongia but fine and straight in Malo ; the perradial lappet warts are twice as numerous in Gerongia as in Maio\ the mesenteries in Gerongia lack a cord-like extension to the rhopalium but possess it in Malo ; and the velarial canals are entirely different, being seven laminar and crowded in Gerongia but one, palmate and uncrowded, in Malo. 28 New box jellyfish from tropical Australia Table 1 . Comparison of characters useful for distinguishing the genera of the Tamoyidae and the undescribed ‘Morbakka’ group. Parenthetical numbers after genera indicate number of species known at this time for each group (Gershwin 2005a). Carukia (3) Gerongia (1) Malo (4) ‘Morbakka' (4) Tamoya (2) Maximum bell height 1-2 cm 6 cm 2-5 cm 9-15 cm 12-22 cm Bell shape Small and pyramidal, with rounded apex Cuboid and robust, with rounded apex Taller than wide, with flat apex Taller than wide, with flat apex Taller than wide, with flat apex Exumbrellar warts Red warts Pale freckles Purple freckles Bright pink warts White warts Rhopaliar horns Narrow, long, straight; thread- shaped Broad, short, curved; devil- horn-shaped Broad, short, curved; devil- horn-shaped Broad, long, straight, pointy; rabbit-ear-shaped Absent Number of eyes per rhopalium 6 (2 median, plus 4 lateral) Unknown, possibly 6 2 median lensed eyes only, lacking laterals 2 median lensed eyes only, lacking laterals 6 (2 median, plus 4 lateral) Pcdalial shape Narrow Broadly rounded, overhanging Narrow Scalpel Scalpel Pedalial canal bend Simple Thorn Knee-shaped Thorn Thorn Tentacles Round in cross section, with tailed bands Round in cross section, heavy, with flared base Round in cross section, fine Flat in cross section, heavy, with flared base Flat in cross section, heavy, with flared base Gastric phacellae Absent Absent Absent Absent Bands arranged vertically along stomach wall Mesenteries Flap-like half way; cord-like to rhopalium Robust, flap¬ like halfway to rhopalium, without cord-like extension Flap-like one-third way to rhopalium; cord-like to rhopalium Robust, flap¬ like halfway to rhopalium, with fine cord-like extension to rhopalium Robust, flap¬ like halfway to rhopalium, cords extending to rhopalium Velarial canals (per octant) 2, simple or somewhat branched, lacking lateral diverticula 7, with laminar branching, lacking lateral diverticula 1 root, with 3-4 unbranched fingers, lacking lateral diverticula Very complexly branched; too many to easily count, with lateral diverticula Numerous, dendritic and diverticulated Perradial lappet warts Lacking or single on one side 2 rows of 3-6 (typically 5) 2 rows of 1-4 (typically 2) 2 rows of large warts plus scattered warts Lacking lappets Furthermore, the nematocysts of Gerongia are qualitatively and quantitatively distinctive: they are of a wholly different form to those found in Carukia (lemon¬ shaped tumiteles), and of a similar form (club-shaped Type 4 microbasic p-mastigophores) but different size or shape to those of Mala, Tamoya, and ‘Morbakka" (Gershwin 2005a,b) (Table 2). These characters and more (a total of 85 morphological characters) were extensively reviewed and compared among cubozoan species by Gershwin (2005a). Etymology. The generic name Gerongia is taken from the unpublished notes of Dr Ronald V. Southcott, who identified the present form as unique but never formally described it; the actual derivation of the name is not known, but may be derived from the word ‘geronggong,’ a Malaysian name for a dangerous jellyfish, as recorded by Cleland and Southcott (1965). Gerongia rifkinae sp. nov. (Figs 1-3, Table 3) Darwin carybdeid. - Currie and Wood 1995: 479; Williamson et al. 1996: 239, 242, 245-246, fig. 5.2, pi. 5.4, Table 9.2; Currie 2000a: 7; O’Reilly et al. 2001: 652-655, fig. lc, d; Collins 2002: Table 1, Figs 1-3; Gershwin 2005a: numerous pages throughout, PI. 4.6A; Gershwin 2005b: Table 1 and text; Gershwin in press: Table 1. Unnamed. Currie 2000b: 76. Material examined. HOLOTYPE - NTM Cl 1183, Shoal Bay, about 1 km east of the mouth of Buffalo Creek, Port Darwin, Northern Territory, at surface, coll. P. Alderslade, 23 April 1992. PARATYPES - NTM Cl 1176—Cl 1182, Cl 1184, Cl 1191, same data as holotype; NTM C4688, C4689, same locality as holotype, 28 March 1989; NTM Cl 1197-C11199, same locality as holotype, 25 May 1992; NTM 02547, same locality as holotype, 29 L.-A. Gershwin and P. Alderslade Fig. 1. Gerongia rifkinae gen. el sp. nov., holotype: A, whole medusa; B. interradial furrow (i) (the top of the pedalium can be seen in the lower right of the photo, for orientation); C, close-up of tentacle from live specimen now in collection at JCU, Cairns; I), pedalium. Note the prominent overhang of the inner wing (o), the flaring of the pedalial canal where it meets the tentacle (c). and the pronounced upward-pointing ‘thorn’ (t) at the bend of the pedalial canal after it leaves the body. The canal is strongly quadrate in cross section throughout its length. Note also the nematocyst freckles (n) scattered on the outer half of the pedalium, The tentacle in this photograph has been trimmed off. 15 March 1995; NTM 02566, same locality as holotype, coll. P. Alderslade, S. Horner, G. Dally, 15 September 1995; NTM 01385, Fannie Bay, Port Darwin, Northern Territory, at surface, coll. B. Currie and E. McCombie, 25 April 1992; SAM 11922 (= RVS A79I), Darwin Harbour, Northern Territory, at surface at night, 7 April 1964; SAM H920 (= RVS A2390 = JHB J629), on beach, Karumba Lodge, Queensland, coll. J. H. Barnes, between 5-20 December 1960; SAM 11921 (= RVS A734), Dudley Point, Port Darwin, Northern Territory, swimming against tide, 8:30 -8:45 am, 22 February 1964; AM G16036, Norman River, Karumba, Gulf of Carpentaria, Queensland, 17°14'S. 140°I9’E, CSIRO Prawn Survey, 1963-1965, 3 specimens. Measurements and sex determination of each specimen are given in Table 3. 30 New box jellyfish from tropical Australia Fig. 2. Gerongia rifkinae gen. el sp. nov„ holotype: A, rhopaliar niche. Note frown shape of the ostium comprised of one upper and one lower covering scale, both undivided. In this specimen, the ‘upper lip’ of the ‘frown’ is slightly distorted so that the ostium takes on a slight hour-glass shape as well. Note too, the blind-ending ‘rhopaliar horns’ extending upward from the niche (h). The rhopalium is visible as the white structure dangling inside the cavity; B, velarial canals. Note how their growth has completely filled the velarium so that their number cannot be easily counted. For reference, the outer portion of the perradial lappets can be seen approximately 2/3 of the way across the photo (1). Note also laminar nature of canal branching, and lack of lateral diverticula; C, subumbrellar view of mouth and stomach. Note the wide lips of the mouth (m), the well-defined perradial mesenteries (p), and the muscular ‘area corrugata’ (a) of the stomach wall, deflated in this specimen. Gastric cirri are absent; D. perradial lappet, showing conspicuous pattern of two vertical rows of nematocyst warts (w). The perradial frenulum (f) is visible as a shadow behind the middle of the lappet, reaching approximately halfway to the margin. Other material. Unregistered specimen, in collection of James Cook University, Cairns, captured at Fannie Bay Beach, Northern Territory, on 4 January 1999, and examined live. NTM C12064, NE ofCape Hotham, Beagle Gulf, Northern Territory. 11°59.7’S, 131°26.0’E, trawled, 15 m depth, coll. R. Williams, 9 October 1993. Diagnosis. As for the genus. Description. Bell to at least 60 mm tall, and 24 mm wide, with shallow coronal furrow just below apex (Fig. 1A); thickened interradially into vertical pillars, each corner with moderate to deep median, vertical furrow (Fig. IB). The immediate rhopaliar region of the bell is somewhat demarcated from the rest of the body by a pair of adradial furrows resembling human laugh-lines, bounding a flask-shaped raised area. Another pair of shallow adradial furrows demarcates each corner pillar from the flat sides of the body. Four thick, hollow tentacles, round in cross section through most of length but greatly flared at base, issuing singly from four simple interradial pedalia, and having nematocyst bands that are either of even width or alternating slightly thicker with slightly thinner (Fig. 1C). Pedalia longer than broad, with the outer keel modestly flared, and the inner keel broadly rounded, extending beyond the tentacle insertion (Fig. ID). Pedalial canals 31 L.-A. Gershwin and P. Alderslade Table 2. Comparison of cnidomes in described forms in the Tamoyidae ( sensu Gershwin 2005a). Descriptions and measurements are from Gershwin (2005a). Nematocyst type Capsule shape Capsule size Tubule winding pattern Shaft spination Gerongia rifkinae gen. et sp. nov. Type 4 Microbasic p-mastigophores Elongate club- shaped (with the discharge end barely broader) 43.32-59.39pm long x 14.62-17.25gm wide, n=27 Coiled about three times in the long axis, progressively looser from the discharge end Long spines along the entire length of the shaft Carukia barnesi Southcott Tumiteles Lemon-shaped 22.88-26.93pm long x 14.59-16.46pm wide, n=8 Loosely coiled about 3 times with the long axis Long spines arising from a central shaft swelling Carukia shinju Gershwin Tumiteles Egg-shaped to Lemon-shaped 18.73-27.78pm long x 13.21-18.44pm wide, n=17 Loosely coiled about 6 times with the short axis Long spines arising from a central shaft swelling Malo maxima Gershwin Type 4 Microbasic p-mastigophores Quite elongate egg-shaped (i.e., noticeably broader at the discharge end) 34.55-49.32pm long x 14.59-19.65pm wide, n=58 About 4 coils in the long axis are confined to the centre region Long spines along the entire length of the shaft Tamoya haplonema Muller 1) Club-shaped mastigophores 2) Sub-spherical isorhizas Elongate oval (without noticeable broadening), tapered at non¬ discharge end 1) 56.42-59.47 pm long x 15.67-17.38 pm wide, n=6 2) 29.85-36.19 pm long x 19.73-28.44 pm wide, n=2 About 6 uneven coils in long axis, primarily in the centre region A few long spines along distal half of shaft ‘Morbakka’ sensu Fenner (1985) and Williamson et al. (1996) Type 4 Microbasic p-mastigophores, plus two types of oval isorhizas Tapered at both ends and rounded in the middle 60.99-69.97pm long x 13.72-18.62pm wide, n=28 About 3 uneven coils in the long axis, primarily in the centre region Long spines at the distal end only Table 3. Gerongia rifkinae gen. et sp. nov., table of measurements of known specimens. BH= bell height (from apex to velarial turnover), DBW= diagonal bell width (distance between diagonally opposing pedalia), IRW = inter-rhopalial width (between adjacent rhopalia), imm= immature, ?= not determined. Specimen no.sex_BH (mm) DBW (mm) IRW (mm) M 61.20 55.90 24.63 ? 40.00 37.37 18.97 M 43.24 32.43 20.92 SAM H920 =RVS A2390=JHB629 SAM H92I =RVS A734 SAM H922 =RVS A79I NTM C4689 NTM C4688 NTM Cl 1176 NTM Cl 1177 NTM Cl 1178 NTM Cl 1179 NTMC11180 NTM Cl 1181 NTM Cl 1182 NTM Cl 1183 NTM Cl 1184 NTM Cl 1191 NTM Cl 1197 NTM Cl 1198 NTM Cl 1199 NTM C12147 NTM C12566 NTM Cl 1385 9 damaged ? damaged ? damaged M 36.67 F 38.74 F 42.54 ? damaged M 45.12 M 37.73 M 46.26 F 39.23 M 23.44 M 41.26 F 44.64 M 42.57 F 36.38 M 42.44 imm. 31.03 39.47 18.84 43.70 20.43 44.95 21.12 47.39 22.74 36.99 21.65 48.17 24.35 38.27 18.70 33.48 15.22 42.67 20.73 51.35 26.37 38.74 20.78 30.35 14.04 39.24 22.87 33.17 20.37 32 New box jellyfish from tropical Australia Fig. 3. Gerongia rijkinae gen. et sp. nov., holotype: A, tentacular microbasic p-mastigophore nematocysts, undischarged (a), discharged (b); B, exumbrellar nematocysts from a bell freckle. flared at tentacle insertion, with upward-pointing ‘thorn’ at the bend, and strongly 4-sided in cross-section throughout entire length (Fig. ID). Two corners of the cross-section correspond one to each keel, and the other two corners correspond to the two broad sides. Interradial septa simple or with slight perforations. Rhopaliar niche with a frown-shaped ostium composed of two covering scales (I upper, 1 lower), both broadly convex in an opposing manner; the upper scale with down-turned ends, forming the shape of a frowning mouth (Fig. 2A). Two internal hollow, blind-ending, finger-like rhopaliar horns, short, broad, commencing from the base of the rhopalial stem and extending upward at about a 45° angle, to approximately even with the outer edge of the ostium, curved inward, having the appearance of‘devil-horns’ (Fig. 2A). Rhopalium with a conspicuous distal statocyst and 2 median lensed eyes; lateral pigment spots presently unknown; statolith shape unknown. Subumbrellar rhopalial windows concave. Velarial canals seven per octant, non-anastomosing, lacking lateral diverticula; branching laminar, with each canal divided into multiple finer fingers, not spreading laterally; in large specimens, canals have grown to cover the velarium, and are separated only by narrow lines of demarcation (Fig. 2B). Mouth almost perfectly quadrate, with four wide, rounded to square-cornered lips, slightly folded diagonally, on short manubrium (Fig. 2C). Gastric cirri absent. Stomach large and bag-like when relaxed, or with numerous parallel folds in each of four corners when contracted (Fig. 2C), connected to subumbrella by moderate mesenteries (Fig. 2C) which only reach about halfway to rhopalia. Gonads sheet-like, extending along entire length of interradial septa, smooth to rippled or pleated. Perradial lappets conspicuously developed, moderately triangular, with two rows of approximately 5 (3-6) broadly rounded, low nematocyst warts (Fig. 2D). Velarium connected to subumbrella with four well- developed perradial frenulae, which reach only about halfway to velarial margin (Fig. 2D). Nematocyst patches small, unraised, round, occuring over entire exumbrella and especially noticeable on outer keel of pedalia (Fig. ID); not in distinct rows. In life the medusa is transparent and colorless (P. Alderslade, unpublished observations), or with lavender nematocyst patches and tentacles (RVS unpublished notes for specimen A791). Cnidome. Tentacular nematocysts are of a single type, namely elongate, sub-ovate Type 4 microbasic p- mastigophores, with long spines along the entire length of the shaft, facing away from the capsule, 43.32-59.39 pm long by 14.62-17.25 pm wide (n = 27; Fig. 3A). The tubule is loosely coiled about three times, with one coil more closely near the proximal half of the capsule, another coil loosely extending the whole capsule length, and the other coil about midway between the two. Exumbrellar nematocysts are spherical isorhizas, 21.15-24.77 pm diameter (n = 21; Fig. 3B). Distribution. Most records are from the Darwin area, although the Southcott collection contained one (SAM H920) from Karumba, Queensland, and the CSIRO found three specimens at the Norman River near Karumba (AM G16036). Southcott’s unpublished notes record the species on two other occasions in Queensland ( A92 from Cleveland Moreton Bay 14 January 1957; A447 from Townsville Harbour 11 April 1960); however, the location of the two specimens is unknown and their identities cannot be confirmed. It is the senior author’s opinion, based on study of hundreds of Queensland carybdeid specimens, that probably the Moreton Bay specimen, and quite possibly the Townsville specimen, were erroneously identified. Phylogeny. In Collins’s (2002) nuclear rDNA 18S sequence analysis, Gerongia rifkinae (as the ‘Darwin carybdeid’) clustered most closely with Carukia barnesi, together forming a clade nested within the rest of the Carybdeida. While this pattern had very high support in Collins’ analysis of nine cubozoan species, those were the only two species he sampled in the Irukandji group. In more comprehensive molecular (six Irukandji species) and morphological analyses (10 Irukandji species), G. rifkinae 33 L.-A. Gershwin and P. Alderslade consistently and robustly clustered with ‘Morbakka’, external to both the Carukia and Malo clades (Gershwin 2005a). The tissues sequenced by Collins (2002) are from an unregistered specimen now in the collection at JCU, Cairns. Remarks. The number of eyes in G. rifkinae is unclear at this time, as whether they are faded or absent cannot be accurately determined from the preserved material. Most cubozoans have two median lensed eyes plus four lateral pigment-spot eyes on each rhopalium. However, species in the Malo maxima Gershwin, 2005 and ‘Pseudo- Irukandji’ clade have only two main lensed eyes, while other close relatives, such as species in the Carukia clade and ‘Morbakka’, have six. A single individual (NTM C12064) brought up in a trawl, is probably a juvenile of G. rifkinae, but this identification cannot presently be confirmed. The medusa possesses the characteristic thorn on the pedal ial canal and lacks any trace of gastric cirri. In addition, the exumbrella is covered with unraised nematocyst patches and the rhopaliar niche ostium is of the characteristic kidney-bean or frown shape. However, the velarial canals are of a form unlike mature G. rifkinae. In each octant, there is one large canal ‘root,’ which spans most of the space. This root has three ‘legs’: the one nearest the pedalium biforked and the other two simple. The forked canal is straight on the side away from the pedalium, and upside-down L-shaped nearest the pedalium, such that the two forks are parallel. Furthermore, the perradial lappets lack the nematocyst warts, which typically occur in two rows. The velarial canals are of a form that we would expect to observe in immature specimens; however, it is also possible that it belongs to a yet unknown species. Lacking a range of size classes for comparison we have not included the specimen as part of the type series, but hope that future studies will answer the question of whether it is G. rifkinae. The non-type specimen observed in life had the stomach greatly ballooned, extending approximately 1/3 the height of the bell cavity. The manubrium extended another 1/3 of the bell cavity height, reaching the level of the rhopalia. The mouth constantly twitched throughout observation. The medusa was colorless overall but slightly cloudy, with faintly pink tentacles. When first sighted, the medusa was swimming at the surface in turbid water of about 1 m depth, over sand, with the tentacles stretching several bell lengths behind. At this time, very little is known about the stinging power of Gerongia rifkinae. However, it does not appear to be a severe health threat to humans. Two stingings are confirmed from the retained specimens; both involved only localized pain, without any systemic symptoms; one is discussed in detail by Williamson et al. (1996). Additional stings are correlated based on nematocysts taken from the skin of the victims (O’Reilly et al. 2001), some with Irukandji syndrome overlap, but specimens were unavailable for positive identification. A single known experimental sting produced mild Irukandji syndrome symptoms (B. Currie pers. comm., March 2004). While it seems convincing that G. rifkinae can produce only mild Irukandji syndrome, it should nonetheless be handled with care, with the caution that specimen maturity, venom load, or personal sensitivity may conceivably produce a more severe reaction. Etymology. The species is named for Dr Jacquie Rifkin, who worked on Australian cubozoans prior to pursuing other interests. DISCUSSION In studying Gerongia rifkinae we came across many problems in the cubozoan taxonomic literature. Most workers seemed confused about several of the most obvious characters, leading to unclear and sometimes inaccurate classifications (Mayer 1910; Stiasny 1919; Uchida 1929; Stiasny 1937; Kramp 1961; Uchida 1970). Wc take this opportunity here to discuss some of these characters, in hope of clarifying cubozoan taxonomy for future studies. It has been common throughout cubozoological history to use the term ‘rhopalial niches’ when referring to the opening to the niche rather than the actual cavity. This is confusing, since the shape and depth ol the cavity are usable characters, as arc the number and shape of covering scales that form the opening to the niche. For this reason, we refer herein and elsewhere (Gershwin 2005a) specifically to the rhopalial niche ostia. The descriptive term ‘frown-shaped’ was coined by someone unknown to us, but we have adopted it because of its usefulness in describing the shape of the ostium, as if two lips were pursed incompletely, with slightly down-turned ends. In some specimens and taxa, the lips are even more ‘frown¬ shaped’ than in our holotype. Stiasny (1919, 1937) believed that all specimens with dumb-bell shaped rhopalial ostia belonged to a single very variable species. While his 1937 Arabian specimens may or may not have been conspecific, the shared dumb¬ bell-shaped ostium would not have been the reason. This character is shared by Tamoya haplonema Mtiller from the western tropical Atlantic, Carukia barnesi Southcott from north-eastern Australia, Malo maxima from north-western Australia, Tripedalia binata Moore from northern Australia, and now Gerongia rifkinae. However, each of these forms is easily separated from the others based on numerous other characters. The other rhopalial ostium forms have proven equally problematical, however, examination of worldwide material has helped us to elucidate the following: in Carybdea alata Reynaud [and its related forms (recently revised by Gershwin (In Press)], there is a single large covering scale above and two large covering scales below, one on each side, forming 34 New box jellyfish from tropical Australia a T-shaped opening; in Carybdea rastonii (Haacke) and Carybdea marsupialis (Linnaeus) and their related forms (C. morn Kishinouye and an undescribed species; and C. murrayana Haeckel; C. xciymacana Conant; and at least three undescribed species, respectively), the two lower covering scales are greatly reduced, forming a rather heart-shaped opening. In Carybdea sivickisi Stiasny, the covering scales are coalesced into a continuous ring, such that the opening is of a vertical oval appearance. Another character that has been the subject of much debate is the form of the gastric cirri in the four-tentacled carybdeids. Uchida (1929: 172) stated, “The direction of the gastral filaments often regarded as of generic importance can not be of such value.” This is incorrect. In brief, Carybdea marsupialis and its related species possess single-rooted or multiple-rooted, short phacellae; these appear to the unaided eye as little tassels in the four corners of the stomach. Carybdea alala and its related species have a crescentic arrangement of long cirri with one or more roots; the parallel arrangement of the individual cirri can be easily seen with the unaided eye. In C. sivickisi the crescentic phaccllus is inverted and the cirri are shorter and singly rooted; the species is unmistakable, however, on the basis of its aboral adhesive pads. Carybdea rastonii and its related species have a diagonal but straight row of short cirri in many small bunches; these appear to the unaided eye as a single fuzzy white line obliquely crossing each interradius. And in Tamoya haplonema the row is similar to that of C. rastonii , but is instead vertical along the stomach wall. Based on comparative morphology of numerous characters, the genus Carybdea appears to be made up of several groups identifiable, in part, by phacellus morphology (Gershwin, unpublished). This pattern was also observed by Collins’s (2002) and Gershwin’s (2005a) molecular studies. If the form ofthe phacellae is a reliable character, so then might be their absence. However, this has been questioned for Carukia barnesi, which was described as new based on the fact that it lacked cirri (Southcott 1967). Uchida (1970) thought that C. barnesi may have gastric filaments in the young stages, then they are lost as the individual ages, and he concluded the same for his own specimens. While this may or may not be true for Uchida’s specimens, it is not true for C. barnesi and Malo maxima , nor apparently for G. rifkinae. Gershwin has examined over 300 specimens of C. barnesi , ranging from 0.7 mm to 14 mm BH, about 200 specimens of Malo maxima, ranging from about 10 mm to 50 mm BH, and approximately 50 other specimens of at least seven other species in the Irukandji group; no trace of gastric cirri was found in any oflhese specimens. While the available material of G. rifkinae is more limited, none of the available specimens shows any sign of gastric cirri. Thus, it seems safe to state that, at least in these ten species, lack of cirri is not an artifact of age. Another confusing character is the nematocyst warts. Logically, the term ‘warts’ should be used to describe small, raised regions that can be felt with the fingertips (though this is not advisable without gloves). Indeed, many cubomedusae have exumbrellar warts (e.g., Tamoya haplonema , Carukia barnesi , and Chiropsalmus quadrumanus Miiller), and G. rifkinae and C. barnesi have such warts on the perradial lappets and velarial canals, respectively. However, in some other cubomedusae, including G. rifkinae, the exumbrella is sprinkled with nematocysts in patches, or freckles, rather than raised gelatinous warts. Finally, the number and form of the velarial canals has caused much confusion. According to Stiasny (1937), the velarial canals are subject to great changes during development and arc therefore of poor taxonomic use. While it is true that the canals develop as the medusa grows, and that in some species no two canals have precisely the same branching pattern, some generalizations can still be made on adult medusae. In most species, the number of canal roots is constant across a wide range of specimen sizes. The degree of branching too is diagnostic in many species. For example, in Carukia barnesi the canals are always simple, whereas in some members of the Carybdea alata species complex, the canals bifurcate in only the largest specimens, being simple in most (Gershwin, unpublished). In C. rastonii and C. xaymacana, the canals are always two per octant and bifurcated or more complexly branched, but they are 3-4 per octant in C. marsupialis (Gershwin, unpublished). In Gerongia, Tamoya, and the chirodropids, the canals are often highly diverticulated and always complexly branched, such that it is often difficult to get an exact count. At the generic level, character states which are taxonomically important in the Tamoyidae are set out in Tables 1 and 2. Cubozoan taxonomy has been virtually ignored in most regions, undoubtedly due in part to the unclear systematics hitherto. The cubomedusae of the world are at present poorly understood, and it seems likely that the number of recognized species will increase with modern techniques such as refined morphological studies and molecular studies. In the past, inappropriate emphasis on certain characters at the expense of others has led to progressive synonymy of species. If we consider the opinions of some past workers as outlined above, we find that we are unable to gain meaning from current descriptions, and are thus unable to resolve the taxa except at the coarsest level. However, sorting out the world’s cubozoan biodiversity is a necessary step in understanding them and managing their effect on human health. As our scientific understanding of evolutionary and biogcographical processes becomes more refined, so must our classification systems ofthe creatures that represent this understanding. 35 L.-A. Gershwin and P. Alderslade ACKNOWLEDGMENTS We are grateful to Jenny Roberts for access to the Barnes Collection and Notes, Bart Currie for sting information, and Wolfgang Zeidler for patiently improving the manuscript. LG also wishes to thank the following institutions for their support (in alphabetical order): the Australian-Anierican Fulbright Commission, CRC Reef Research Centre, CSIRO C.R.I.M.R, Environment Australia (ABRS Grant No. 20045 to L.G. and W. Zeidler), James Cook University, Museum and Art Galleries of the Northern Territory, South Australian Museum, Surf Life Saving Australia, Surf Life Saving Queensland, and University of California Berkeley. REFERENCES Barnes, J.H. 1965. Chironexjleckeri & Chiropsahmis quadrigatus - morphological distinctions. North Queensland Naturalist 32(137): 13-22. Calder, D.R. 1974. Ncmatocysts of the coronate scyphomedusa, Linuche unguiculata , with a brief reexamination of scyphozoan nematocyst classification. Chesapeake Science 15: 170-173. 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Revision of some Carybdeidae (Scyphozoa: Cubomedusae), including a description of the jellyfish responsible for the ‘Irukandji syndrome'. Australian Journal of Zoology 15: 651-671. Stiasny.G. 1919. Die Scyphomedusen-SammlungdesNaturhistorischen Reichsmuseums in Leiden. 1, Die Carybdcidcn (Cubomedusen). Zoologische Mededelingen (Leiden) 5: 33-58, text-figs. [In German], Stiasny, G. 1937. Scyphomedusae. Scientific Report on the John Murray Expeditions 1933-1934 4(Pt. 7): 203-242, 14 text- figs., PI. I. Uchida, T. 1929. Studies on the Stauromedusae and Cubomedusae, with special reference to their metamorphosis. Japanese Journal of Zoology 2: 103-193. Uchida, T. 1970. Revision of Japanese Cubomedusae. Publications of the Seto Marine Biological Laboratory 17(5): 289-297. Verrill, A.E. 1865. Classification of polyps (extract condensed from a synopsis of the Polypi of the North Pacific Exploring Expedition, under Captains Ringgold and Rodgers, U.S.N.). Annals and Magazine of Natural History 16(Third Scries): 191-197. Werner, B. 1973. New investigations on systematics and evolution of the class Scyphozoa and the phylum Cnidaria. Publications of the Seto Marine Biological Laboratory 20 (Proceedings of the Second International Symposium on Cnidaria): 35-61. Werner, B. 1984. Klasse Cubozoa. Pp 106 133. In: Gruner, H.E. (ed.), Lehrbuch der Speziellen Zoologie. Gustav Fischer Verlag: Stuttgart. Williamson, J., Fenner, P„ Burnett, J. and Rifkin, .1. 1996. Venomous and poisonous marine animals: a medical and biological handbook. NSW University Press: Sydney, Australia. Accepted 9 October 2005 36 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21 : 37-46 Aspects of the reproductive biology of Polymesoda erosa (Solander, 1786) (Bivalvia: Corbiculidae) in northern Australia RICKY GIMIN 1 *, LUONG-VAN THINH'\ RAM MOHAN 2 , ANTHONY D. GRIFFITHS 3 'School of Science and Primary Industries, Charles Darwin University, Darwin NT 0909, AUSTRALIA 2 School of Life Sciences, University of Queensland, Brisbane QLD 4072, AUSTRALIA 3 Key Centre for Tropical Wildlife Management, Charles Darwin University, Darwin NT 0909, AUSTRALIA *Corresponding author: jim.luong-van@cdu.edu.au "Present address: University Nusa Cendana, Kupang, NTT, INDONESIA 85117 ABSTRACT The reproductive biology of Polymesoda erosa (Solander, 1786) (Bivalvia: Corbiculidae), a staple food for Aborigines in the Maningrida region of northern Australia, was studied between June 2001 and September 2002 as part of a program to manage its subsistence fishery. Polymesoda erosa is dioecious (gonochoristic) and matures at a shell length of approximately 45 mm. Within the range of shell lengths of 55 to 90 mm. the sex ratio in the population was 1 male:l female. However, at shell lengths less than 55 mm, males were significantly more common. Females were more common at shell lengths greater than 80 mm, but not statistically so. At shell length greater than 90 mm, the production of new gametes is reduced. Based on the gonadosotnatic index (GS1) and histological study, P. erosa spawned during the mid- to late-wet season (February and May). Males and females showed spawning synchrony. Females within the shell range of 65 to 85 mm produced 1.20 x I0 6 eggs per individual (SD = 0.43 x 10\ n = 10). The high frequency of spawning was probably related to long inundation during the wet season. The present practice of not gathering P. erosa during the wet season may be important in sustaining the current subsistence harvest. Keywords: clams, gonadosomatic index, sex ratio, harvest. Aboriginal, mangroves. INTRODUCTION The mangrove clam, Polymesoda erosa Solander, 1786 (Bivalvia: Corbiculidae) (Fig. 1) is a large bivalve, reaching more than 120 mm in length (Morton 1985). This bivalve is naturally distributed throughout the tropical Indo- West Pacific from Sri Lanka in the west to the Solomon Islands in the east, from Japan in the north and to northern Australia and New Caledonia in the south (Morton 1984). Polymesoda erosa inhabits firm mud substrata surrounding streams draining through the mangroves and is capable of withstanding long periods of desiccation and wide ranging salinities. During emersion, it is able to exchange water and suspended material with subterranean water through the pedal gape, and then resume suspension feeding when inundated (Morton 1976). Polymesoda erosa is harvested for food in various Southeast Asian countries including the Indonesian archipelago (Morton 1976, 1984). In northern Australia, P. erosa is a staple food of Aborigines living in coastal Fig. I. Polymesoda erosa from Maningrida spawning: A, male clam ejecting sperm; B, female clam releasing eggs. 37 R. Gimin, T. Luong-Van, R. Mohan and A. D. Griffiths regions (Meehan 1982; Isaac 2002). It is the most prominent non-fish group numerically in the indigenous catch (Coleman et al. 2003; Willan and Dredge 2004). Despite this, P. erosa is not commercially harvested, although there is some potential for indigenous communities to develop commercial enterprises based on the wild harvest of species such as P. erosci (Ray Hall, pers. comm.). In order to do this successfully, it is important to understand, among other things, the timing of reproductive events because it is an essential prerequisite for various aspects of commercial farming and harvesting including collection of wild spat, timing of collecting season and hatchery production. Information on the reproductive biology of Polymesoda erosa is scarce. To date, the study of Morton (1985) on the ecology and reproductive strategy of the species in Hong Kong is the only published work available. In Hong Kong, P. erosa has a single but extended spawning season during summer. Because the reproductive cycle of a species may vary considerably over its geographical range (Sastry 1979; Fournier 1992; Sato 1999; Ward and Davis 2002), a species that has a wide distribution covering several latitudes, such as P. erosa, is likely to exhibit a variety of reproductive strategies (Cardenas and Aranda 2000). The present study investigated the reproductive biology of P. erosa from the Maningrida region of northern Australia. The results provide the basic information necessary for the farming and harvesting of this species and allows for a comparison of the reproductive strategy of the species in Hong Kong (22° 20’ N, 114° 11’ E) (Morton 1985) in the northern hemisphere and the Northern Territory coast (i.e. Maningrida) in the southern. MATERIALS AND METHODS Collections. Live Polymesoda erosa were collected from the mangrove forest bordering the Tomkinson River in the Maningrida region. Northern Territory, Australia (12° 3’ S, 134° 13’ E; Fig. 2). Temperatures and rainfalls during the study period were provided by the Bureau of Meteorology for the Maningrida region (Fig.3B, C). Salinity data (Fig. 3A) were obtained indirectly by measuring the salinity, using a calibrated hand-refractometer Vista A366ATC, of the water entrapped inside the mantle cavity when the collected individuals reached Charles Darwin Fig. 2. Map of Maningrida region, Northern Territory, Australia, showing the collection site (x) for Polymesoda erosa adjacent to the Tomkinson River. 38 Reproduction of Polymesoda erosa in northern Australia A 50 Month B ,$■ o' & $■ S' & & i? A’ ^ cf* ^ yP ^ v 8 ' ^ s? s* Month c Fig. 3. A, Salinity of the water entrapped in the mantle cavity of Polymesoda erosa sampled at Maningrida between June 2001 and September 2002. Broken line indicates no sampling was done; B, Minimum and maximum air temperature recordings from June 2001 to September 2002; C, Total monthly rainfall (mm) recorded at Maningrida between June 2001 and September 2002. University (see below). This was necessary because during collection the habitat of/ 5 , erosa was dry. From 40 to 100 individuals (31 to 112 mm in size) were taken at each sampling time spanning from June 2001 to September 2002, by digging the mud under mangrove vegetation in the hinterland margin during low tide. The animals were airfreighted dry in styrofoam boxes to an experimental hatchery at Charles Darwin University (12° 28’ S, 130° 51’ E). On each sampling occasion, shell length was recorded using vernier callipers. Sex determination. The shells were opened, tissues removed and sexes recorded based on the colour of the gonad: black in females and creamy white in males. When in doubt, a gonadal smear was examined under the microscope for sperm or eggs. Gonadosomatic index. Tissues were dissected and classified into the gonad and non-gonadal. The gonad, inclusive of stomach and intestine, was separated from other organs. Non-gonadal parts included the mantle, anterior and posterior adductor muscles, ctenidia and labial palps, pericardial organs, digestive diverticula, crystalline style and foot. Both gonadal and non-gonadal tissue were blotted dry on paper towels and weighed separately to the nearest 0.001 g with a Sarlorius B310S electronic balance. Total tissue weight (TTW) was obtained by combining gonad weight (GW) and non-gonadal tissue weight (NGW). The gonadosomatic index (GS1) was calculated using the equation used by Gonzalez et al. (2002): GSJ = GO/x 100 Histological analysis. After measurements for gonad index, the dissected gonads were fixed in aqueous Bouin’s solution for 48-72 h. A central portion about 5 mm thick was dissected from each fixed gonad, placed in a tissue cassette and stored in 70% (v/v) ethanol prior to dehydration. The gonad portion was later dehydrated in FAA (40% formaldehyde, 95% ethanol, and glacial acetic acid in the ratio of 1:5:0.2) and in ascending an ethanol series ranging from 70 to 100% in a tissue processor (Shandon Duplex) overnight. The tissue was cleared with histoclear and impregnated with paraffin wax. The resulting block was sectioned at 5 mm, and sections mounted onto slides, stained with Mayer’s Acid Hemalum and counterstained with eosin Y (Bancroft and Stevens 1982). The gonadal developmental stages were assigned to criteria modified from Morton (1985) for Polymesoda erosa in Flong Kong (Table 1). Gonad index. Each developmental stage was assigned a numerical score as follows: primordial = 1, developing = 2, maturing and partial spawned = 3, ripe = 4. and spent = 0. This scoring was modified from Sause et al. (1987) and Fladfield and Anderson (1988). For each monthly sample, a mean gonad index was determined following calculations described by Gosling (2003). The number of individuals in each stage was multiplied by the numerical ranking of the stage and the sum of these products divided by the total number of individuals in each sample. Fecundity. Estimates of fecundity (the number of mature oocytes contained in female gonads) were made following the technique described by Dredge (1981). Mature ovaries were selected by macroscopic appearance from individuals of greater than 65 mm shell length and were fixed in Bouin’s solution until hard. After washing with 70% ethanol, the membrane encasing the eggs mass was peeled off and carefully separated from the stomach and intestine. The mass of eggs was blotted dry and later weighed to the nearest 0.0001 g. A small portion from this mass was weighed and placed individually into a 250 ml beaker containing 100 ml of distilled water. A mixer was used to detach the eggs from the mass and disperse them through the water. The contents of the beaker were then transferred to a graduated cylinder and water added to a required volume. The number of eggs in the cylinder was estimated using a Sedgwick-rafter cell. The results represented the number of eggs per mass of gonad. This was multiplied by the weight of the gonad to estimate the total number of contained oocytes. Statistical analyses. Observed sex ratios were tested against a 1:1 ratio using a chi-square goodness-of-fit (Fowler et ai 2000). Data forGSI from females and males were treated separately. ANCOVA with shell volume as a 39 R. Gimin, T. Luong-Van, R. Mohan and A. D. Griffiths Table 1. Microscopic characteristics of the reproductive stages of male and female Polymesoda erosa from Maningrida. Stage Characteristics Males Primordial Developing Maturing Ripe Partially spawned Spent Females Primordial Developing Maturing Ripe Partially spawned Spent Small follicles scatter gonadal area. Interl'ollicular space large. Phagocytes present in some follicles. Germinal cells start lining up inside of the follicle. Dense spermatogonia in germinal layer of the follicle. Spermatozoa appear at the centre of the follicular lumen. Spermatozoa occupy greater area at the central lumen. The spermatozoa are in orderly stripe-configuration with tails pointing toward the centre of the lumen. Spermatocytes and spermatids are dense at the periphery of the follicle. Stripes of spermatozoa occupy almost the whole area of the follicle leaving a very narrow strip along the periphery for the spermatogonia and spermatids. The number of spermatozoa filling the lumen decreases in some follicles. Spermatids occupy the area vacated by the spermatozoa. Most of follicles are empty. Empty follicles with irregular and elongated shape present in the gonadal matrix. Phagocytes are active. Follicles have large empty lumen. Some lytic oocytes and phagocytes present in the lumen. Few oogonia start lining follicular wall. Follicular lumen is large. Follicles increase in size and have well defined walls. Oogonia and early-developmental- stage oocytes densely pack periphery of the follicle. Young oocytes attached to the wall by thick stalk. Follicles are large. Round vitellogenic-oocytes increase in size, hut many still attach to follicular wall by thin Stalked. Some mature oocytes with prominent nucleus and nucleolus arc free in the lumen. Proportion of free oocytes in the lumen increase. Small number oocytes are still attached to follicular wall by thin stalk. Most oocytes are polygonal. Some follicles contain mature ova. Others are empty or contain small number of ova. Follicles reduce in size particularly in those devoid of ova. Follicles are empty and follicular wall collapses in places. Some residual mature oocytes still present in empty lumen. covariate was used to check for any differences in GSI over the whole year. To check whether the gonads develop in the same way in both males and females, a two-way ANCOVA with month and sex as fixed factors and shell volume as a covariate was performed (Grant and Tyler 1983). Percentage data were arcsine transformed. The frequency of individuals in the six developmental stages was tested using a contingency chi-square test (Fowler et at. 2000). To check to what extent the GSI can be used to follow the reproductive cycle in Polymesoda erosa , the GSI values were analysed using ANCOVA with the gametogenic stage as a factor and shell volume as a covariate. RESULTS Seven-hundred and ten Polymesoda erosa were collected; 361 males, 323 females, and 26 undifferentiated. No hermaphrodites were detected. The sex ratios were not significantly different from the expected 1:1 ratio for any of the 12 samples, although there were more males than females in most of them (Fig. 4). Individuals of45-55 mm had a sex ratio significantly (P<0.05) biased towards males. At higher length-classes (55 to less than 80 mm) males still outnumbered females, but there was no significant (P>0.05) deviation from the expected 1:1 ratio. In contrast, at sizes greater than 80 mm, there was a tendency for females to predominate, although the ratio, again, did not differ significantly from 1:1. Estimates of fecundity for ten Polymesoda erosa individuals within the shell length range of 65-85 mm of the above cohort showed that each clam contained 1.20 x 10 6 eggs (SD = 0.43 x 10 ( ’). Gonadosomatic index (GSI). The GSI of Polymesoda erosa varied significantly during the study period. Box and Whisker plots of gonadosomatic index for each sex showed large variability and strong asymmetry in all monthly samples (Fig. 5A, B). Each sample covered wide ranges of GSI values suggesting continuous breeding activity. However, the box plots showed differences in the positions of the medians of the data. In general, for both sexes, the medians increased from June 2001, reached a maximum in January 2002, then fell to low levels between February and May. After this period, the medians rose again and reached high values for the next three months (June to September 2002). These rises and falls in the medians were interpreted as difference in spawning intensities. The results of this exploratory data analysis were confirmed by ANCOVA which showed that the GSI values of males females differed significantly among months (males: F um = 6.6119, P<0. 05; females: F n 10 =6.0190,P<0.05). In males, post hoc comparisons among means using Tukey’s HSD showed significantly low values for male GSI values in February 2002 and May 2002. Similarly, in females, 40 Reproduction of Polymesoda erosa in northern Australia 140 - 120 100 E z 60 H 40 H 20 0 Female Male 46% 46% 37% n = n = 80 n = 76 62 — 47% n = 88 46% n = 118 54% n = 79 59% n = 78 40-45 46-50 51-55 56-60 61-65 66-70 71-75 76-80 81-85 86-90 >90 Size class (mm) F'g. 4. Size frequency and sex ratios of Polysoma erosa collected from Maningrida from June 2001 to September 2002. Figures above the columns indicate percentage of female and total number of individuals collected for that size class. significantly low GSI values occurred between February 2002 and May 2002. Two-way ANCOVA showed a not significant interaction (P n , |5 =l.0773, P>0.05) between sex and sampling month, indicating that males and females have similar patterns of gonad development. 1 lowever, both factors (i.e., month and sex) individually showed significant differences (month: F u , |5 =11.2941, P< 0.05; sex: F x 215 =84.1799, PC0.05). Females had significantly higher (P<0.05) GSI values than males. Analyses of reproductive condition by the gonad index (Fig. 6A, B) confirmed the evidence for two peaks in reproductive condition annually in both males and females, followed by troughs suggestive of spawning periods. Male reproductive cycle. The reproductive cycle of male Polymesoda erosa from Maningrida is presented in Figure 7A. Contingency chi-square test showed that the frequency of each gonadal stage was associated significantly with sampling month (X 2 4 4 = 82.84, P<0.05). In June 2001, about half the individuals were in early stages (primordial and developing), while maturing and ripe were 40% and 10%, respectively. In July the frequency of the ripe individuals increased to 40% and reached a maximum between July and September. Chi-square tests confirmed that there were significantly (P< 0.05) more individuals of primordial and developing stages than expected during June 2001. There was a short period of spawning during September 2001 but for the next few months until January 2002, only maturing and ripe animals were present. Seventy percent of individuals were ripe in January 2002. Starting in February, the numbers of spent or partly spawned animals increased from 20% to 50% in May 2002. During this period, the number of males with spent gonads was significantly higher than the expected values (P<0.05, chi-square test). Apparently, gonads developed rapidly after a long breeding period because in June 2002, 70% of the males were maturing and by August the sample was dominated by ripe males (50%). In September 2002, 30% of males were spent, suggesting another spawning event. The histological data for male confirm those obtained using the gonad index (Fig. 6A). Female reproductive cycle. As in males, the frequency of each female gonadal stage was associated significantly with sampling month {X : A = 115.98, P<0.05). The female reproductive cycle thus followed that of the males closely. All gametogenic stages occurred in most sampling months with a high proportion of maturing and ripe stages (Fig. 7B). Developing gonads, from primordial to mature, occurred during the period of June to July and, probably, through August 2001. Chi-square tests showed that these three stages had significantly higher values than expected (PcO.OS) during this period. Ripe females increased during the period, and in September 50% of individuals were ripe. Some individuals spawned in September 2001 as indicated by the presence of spent and primordial stages. The ripe individuals were most common (at least 60%) during October 2001 until January 2002. An intense spawning occurred in February 2002 after which 70% of the females were spent and 10% were partially spawned. As with males, spawning activity lasted until May 2002, although intensities decreased as the number of ripe animals decreased. The mass spawning events between February and May were confirmed by chi-square tests which showed a significantly higher number (/MJ.OS) of spent individuals during this period. There was no resting period for the female cycle because of rapid maturation during June to August 2002. By September 2002, some of the ripe females were ready to spawn again. The histological data for females confirm those obtained using the gonad index (Fig. 6B). 41 R. Gimin, T. Luong-Van, R. Mohan and A. D. Griffiths Month ? Median 25% - 75% Non-Outlier Range Outliers 5 Month O Median 25% - 75% Non-Outlier Range Outliers Fig. 5. Box and Whisker plots of gonadosomatic index in the monthly samples of A, male and B, female Polymesoda erosa individuals collected from Maningrida from June 2001 to September 2002. DISCUSSION Sex ratio. Polymesoda erosa is dioecious (gonochoristic) but shows no other morphological distinction between the sexes except for the gonads. This is in agreement with reports by Morton (1985) that P. erosa has separate sexes. Sexual differentiation in P. erosa is similar to other corbiculids, such as P. caroliniana (Bose) which inhabits saline marshes (Walker and Heffernan 1994), but is different from the freshwater P. solida (Philippi) which is monoecious (hermaphroditic) (Rueda and Urban 1998). Although the total number of each sex of Polymesoda erosa was equal, one sex predominated at certain shell lengths. Within the range of 50 to 80 mm shell length, the ratios between males and females were 1:1. However, in size classes less than 45 mm, there were significantly more males than females, whereas females were more common in the size classes greater than 85 mm. This pattern is very similar to that of the pearl oyster Pinctada imbricata (Roding) (O’Connor and Lawler 2004). The biased ratios in the population may be due to greater longevity in females than males, resulting in higher number of females than males in older (larger) animals, as in the case of Arctica islandica (Linnaeus) (Ropes el al. 1984; Fritz 1991; Thorarinsdottir and Steingrimsson 2000). There is no evidence that the higher number of 42 Reproduction of Polymesoda erosa in northern Australia females than males in the same class size is due to the faster growth rate in the former, as reported by Wells and Keesing (1989), Marsden (1999), Ward and Davis (2002) and Baghurst and Mitchell (2002). Hermaphroditism can also affect sex ratio, particularly beyond average size-classes (Littlewood and Gordon 1988; Fritz 1991; Thorarinsdottir and Steingrimsson 2000). In the present study no simultaneous hermaphroditism was observed, which is also the case for the population of P. erosa in Hong Kong (Morton 1985). Gonadosomatic index. Considerable overlaps in GS1 values among months indicate that Polymesoda erosa has asynchronous gonad development wherein both early developed and fully mature individuals are present in the population in each sampling month. The pattern of gonad development in P. erosa is typical for a continuous breeder (Grant and Tyler 1983). However, although mature individuals of P. erosa were in evidence year round, there were differences in spawning intensities. Month Month Fig. 6. Seasonal change in Gonad Index for male A, and female B, Polymesoda erosa from Maningrida over the period between June 2001 and September 2002. 43 R. Gimin, T. Luong-Van, R. Mohan and A. D. Griffiths A Spent jj] Partial spawned J Ripe Maturing Developing =1 Primordial 2001 2002 B 100 % 80% 60% S 40% 20 % 0 % Spent ]J] Partial spawned ^ Ripe Maturing Developing § Primordial 2001 2002 Fig. 7. Changes in histological rank for reproductive condition of A, male and B, female Polymesoda erosa from Maningrida between June 2001 and September 2002. * = no sampling during these months. GSI values showed that females had larger gonads than males. This shows that the pattern ofmaterial accumulation during development of the gonads is different between males and females (Grant and Tyler 1983). Nevertheless, the present study showed there was a synchronous pattern of gonad development and spawning between the sexes. Reproductive cycle. In general, the Polymesoda erosa population from Maningrida possessed mature individuals year round, as confirmed by the GSI, Gl and histological data. However, the population showed some seasonality in spawning intensities. Continuous breeding with different spawning intensities is typical of tropical bivalves (Urban 2001). The P. erosa population at Maningrida had two major spawning periods (i.e., a short season in September and an extended season occurring between February and May). This gametogenic pattern is different to that of the P. erosa population in Hong Kong, where the local seasonal cycle creates a single extended breeding period over summer (Morton 1985). Contrasting the two populations, there are clear differences in reproductive timing. In Hong Kong, Polymesoda erosa commences active gametogenesis between April and May with maturation during June and August. Spawning occurs at the end of summer and from September until the following February most individuals were still spent. Apparently this long recovery was affected by cold, winter water temperatures (Morton 1985). Conversely, at Maningrida, active gametogenesis starts in June, and by August most individuals have reached maturity. Rapid gametogenesis at Maningrida was proven by low percentages of early stages (primordial and developing) in the population throughout the year 44 Reproduction of Polymesoda erosa in northern Australia (Cardenas and Aranda 2000). At Maningrida, although a small proportion of the population spawns in September, mO s t ripe individuals retain their gametes and remain at u mature stage for a long period until January. This cohdition is an indication of ripe gamete accumulation (Cardenas and Aranda 2000). The period of ‘ripeness’ at Maningrida was extended by a second spawning season frOm February to May. In males, gonadal maturation takes place gradually to reach a maximum at the end of the season, whereas in females intense spawning occurs at the beginning of the season followed by spawning at reduced levels at the end. The resting period in Polymesoda erosa from Maningrida was short, if not absent. An absence of a resting period is common in tropical bivalves where gonads recover soon after spawning is complete (Pouvreau et al- 2000; Laureta and Marasigan 2000; Urban 2001). A rapid recovery rate and no observable resting period in a population could only happen when the production of a new cohort of germinal cells takes place continuously (Gonzalez et al. 2002). In the present study, even mature animals contained germinal cells at various early developmental stages. Also, during resorption of unreleased gametes, thp gonad continues to produce new germinal cells as can be seen from the presence of oogonia and lytic ova in the same gonad. These findings could explain the fast recovery rate of the P. erosa gonad. The ability to recover enables individuals to have multiple spawnings throughout the year as long as specific cues are available (Marsden 1999). Rapid redevelopment after spawning also happens when animals show partial spawning (Joll and Caputi 1995), or if spawning is incomplete (Rodriguez-Moscoso and Arnaiz 1998; Pouvreau et al 2000). Partially spawned specimens were recorded in Polymesoda erosa from Maningrida and Morton (1985) reported that for Hong Kong, even in ripe individuals, mature eggs never packed the follicles, suggesting a progressive release over a long period. This condition was also observed during this study. Such partial and incomplete spawning might contribute to the fast recovery of P. erosa. Newell et al. (1982) noted that ‘dribble spawning’ reduces the loss of potential recruits over the course of a year so that a catastrophic event kills few larvae. At Maningrida, Polymesoda erosa shows asynchronous gamete development. The presence of two or more stages confirms this asynchrony (Fournier 1992; Tirado and Salas 1998). However, because ripe individuals do not release their gametes, those individuals which develop later reach the same stage as those which mature earlier. Consequently, some months (October to January) are dominated by ripe individuals. Such a situation reinforces synchrony, at least at maturation, increasing the chances of successful fertilization. There was no evidence to link the spawning events of the Maningrida Polymesoda erosa population with changes in temperature, as suggested in subtropical Hong Kong (Morton 1985), or salinity. However, at Maningrida, seasonal temperature does not fluctuate as much as in Hong Kong, where temperatures rise above 35°C in summer but fall to near freezing in winter, and the fact that gametogenesis takes place rapidly in any month of the year suggests that temperature does not play a major role in directing gametogenesis in the tropics. Salinity can also regulate gametogenesis in marine bivalves (Giese 1959; Urban 2001), although how it affects gonadal maturation is unclear (Fournier 1992). In the present study, salinity measurements recorded a range of 5 to 40%o among samples inside the mantle cavity, suggesting that P. erosa at Maningrida was exposed to considerable fluctuations in salinity. I lowever, gametogenesis was not affected by rises or falls in salinity, as shown by the presence of various gamete developmental stages year round. The ability to retain mature gametes without resorption for long periods of time (up to five months) in Polymesoda erosa is unusual but, possibly, happens in other tropical bivalves, e.g. the pearl oyster Pinctada margaritifera (Linnaeus) from tropical regions (Pouvreau et al. 2000). In most bivalves, ripe gametes that are not released soon after maturation are reabsorbed. In the scallop, Argopecten purpuratus (Lamarck), mature oocytes undergo atresia (lysis) and are later reabsorbed when retained too long (Avendano and Le Pennec 1997). Low temperature and a lack of food are factors acting, independently or together, to cause atresia (Avendano and Le Pennec 1997). ACKNOWLEDGMENTS Thanks are due to AusAID for providing a PhD Scholarship to Ricky Gimin, to Ray Hall and the Djelk rangers from the Bawinanga Aboriginal Corporation, for proving the Polymesoda erosa specimens, to Ms Eleanor Hayward for assisting with the histology and to two anonymous reviewers for valuable comments to improve the manuscript. REFERENCES Avendano, M.and Le Pennec, M. 1997. Intraspccific variation on gametogenesis in two populations of the Chilean molluscan bivalve, Argopecten purpuratus (Lamarck). Aquaculture Research 28: 175-182. Bancroft, J.D. and Stevens, A. 1982. Theory and practice of histological techniques. Churchill Livingstone: Edinburgh. Baghurst. B.C. and Mitchell. 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Reproduction and fecundity o Wonax trunculus L., 1758 (Bivalvia: Donacidac) in the littoral of Malaga (Southern Spain). Journal of Shellfish Research 17: 169-176. Urban, H.J. 2001. Reproductive strategics in tropical bivalves (Pteria colymbus, Pinctada imbricata and Pinna cornea)'. Temporal coupling of gonad production and spat abundance related to environmental variability. Journal of Shellfish Research 20: 1127-1134. Walker, R. and Heffernan, P.B. 1994. Gametogcnic cycle of the Carolina marshclam, Polymesoda caroliniana (Bose, 1801), from coastal Georgia. American Malacological Bulletin 11: 57-66. Ward, DAV. and Davis, A.R. 2002. Reproduction of the turban shell Turbo torquatus Gmelin 1791 (Mollusca: Gastropoda), in New South Wales, Australia. Marine and Freshwater Research 53: 85-91. Wells, F.E. and Keesing, J.K. 1989. Reproduction and feeding in the abalone Haliotis roei G ray. Australian Journal of Marine and Freshwater Research 40: 187-197. Willan, R.C. and Dredge, M. 2004. Molluscs. Pp. 201-216. In: Description of key species groups in the Northern Planning Area. National Oceans Office: Hobart, Australia. Accepted 22 June 2005 46 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21 : 47-51 Short communication New distributional records for four species of Stromboidea (Mollusca: Gastropoda) from Australasia GiJS C. KRONENBERG 1 AND BUNJAMIN DHARMA 2 ' Milieu Educatie Centrum, P.O. Box 435, NL-5600AK, THE NETHERLANDS corresponding author: gijs. kronenberg@tisca!i. nl Hawakal VI/16, Grogol, Jakarta 11440, INDONESIA bdharma@dnet. net. id Keywords: Mollusca, Gastropoda, Stromboidea, Rostellariella, Mirabilistrombus, Euprotomus, Dolomena, Indo-Pacific, distribution. INTRODUCTION Recently the second author acquired some specimens of Mirabilistrombus listeri (T. Gray, 1852) and Rostellariella delicatula (Nevill, 1881) from the Arafura Sea, between the Aru Islands and Arnhem Land coast of the Northern Territory, Australia. As this locality is a long way from the presently known distribution of these species according to the literature and therefore quite unexpected, we put these new localities on record here. Kronenberg (2002) gave only Indian Ocean localities for Euprotomus aurora in the original description. Since then, some more localities have come to the attention of the authors, and we take the opportunity to record those as well. This is also the case for Dolomena hickeyi , originally described from off Queensland and Papua New Guinea (Willan 2000), but now realised as widely distributed in the south-western Pacific. Based on conchological characters, it has been advocated that the genus Strombus sensu Abbott (1960) is not monophylctic (e.g. Kronenberg and Vermeij 2002). Recently, this has been confirmed on the basis ofanatomical studies (Simone 2005) and molecular evidence (Latiolais 2003). The name Strombus should only be employed for a clade consisting of the American Recent (Neotropical) species and their fossil ancestors, and will probably also include the West African Strombus latus (Kronenberg and Vermeij 2002; Latiolais 2003; Kronenberg and Lee in prep.). To overcome this non-monophyletic use of Strombus for all Indo-Pacific species, we use the strombid taxa employed as subgenera by Abbott (1960) as full genera, but emphasising at the same time we do not advocate the elevation of all Abbott’s subgenera to generic status. For instance, some allocations by Abbott have already shown to be erroneous (Moolenbeek and Dekker 1993; Kronenberg 1998; Kronenberg and Vermeij 2002; Kronenberg and Lee in prep, and references in those papers). ABBREVIATIONS USED BDJI Bunjamin Dharma, Jakarta, Indonesia GKEN Gijs C. Kronenberg, Eindhoven, The Netherlands HDWN Henk Dekker, Winkel, The Netherlands MBWN Matthijs A. Bouwknegt, Wageningen, The Netherlands NTM Museum and Art Gallery of the Northern Territory, Darwin, Australia TBCM Tim Blackwood, Cohasset, Minnesota, USA VLFI Virgilio Livcrani, Faenza, Italy WBFT Winston Barney, Fort Worth, Texas, USA NEW DISTRIBUTIONAL RECORDS Family Rostellariidae Gabb, 1868 For recognition of Rostellariidae as a family, see Kronenberg and Burger (2002). Genus Rostellariella Thiele, 1929 For recognition of Rostellariella as a genus, see Kronenberg and Burger (2002). Rostellariella delicatula (Nevill, 1881) (Fig. 1A, B) Previously known distribution (taken from literature): northern part of the Indian Ocean, from Gulf of Aden to Sumatra, including Bay of Bengal (Walls 1980; Kronenberg and Berkhout 1984; DeTurck et at. 1999); Vietnam, off Khanh Hoa province (Thach 2005). 47 G. C. Kronenberg and B. Dharma New distributional record. Arafura Sea, between Aru and Australia, where three specimens were trawled alive at approximately 144 m (originally indicated as 80 fathoms) in black mud by Indonesian fishermen. Remarks. It is quite surprising to have this record of the well-known Rostellariella delicatula in the Arafura Sea so soon after the discovery of another species of this genus from roughly the same area (Morrison 2005). So far R. delicatula has not been collected together with R. lorenzi Morrison, 2005. The record from Vietnam (Thach 2005) concerns one single specimen illustrated in Thach (2005: pi. 16, fig. 3), and is the first record of this species from off Vietnam. This specimen was found at a depth of 80-100 m (Thach pers. comm, to GKEN, May 2005). As Melvill and Standen (1905) mention a mud substrate in their paper, we suspect that the habitat is the same for the Indian Ocean specimens and the Arafura Sea specimens reported herein. Melvill and Standen (1905) also report specimens from the Persian Gulf (24°49’N, 55°56’E) at a depth of 250 fathoms (approximately 450 m), but this locality appears to be erroneous because it plots out to be on the mainland of the Arabian Peninsula. Other localities given by Melvill and Standen, with depths varying from 140 to 175 fathoms (approximately 250-315 m), from the Gulf of Oman are more credible. Family Strombidae Rafinesque, 1815 Genus Mirabilistrombus Kronenberg, 1998 Mirabilistrombus listeri (T. Gray, 1852) (Fig. 1C, D) Previously known distribution (taken from literature): Gulf of Oman; northwest Indian Ocean; Bay of Bengal (Okutani 1965; Walls 1980; Kronenberg and Berkhout 1984; DeTurck et at. 1999). New distributional record. Arafura Sea, between Aru and Australia, where five dead specimens were trawled at approximately 144 m (originally indicated as 80 fathoms) by Indonesian fishermen. One specimen was still fresh with the typical colour pattern (Fig. 1C), but the pattern had faded in the other specimens, and they had a more or less uniform brown colour (Fig. ID). Remarks. The taxonomic placement of this species has been controversial. Abbott (1960) placed it in the subgenus Doxander Iredale. Iredale (1931), however, only gave a name and no description, and therefore Doxander Iredale, 1931 is a nomen nudum and not available (ICZN art. 13). We have found no earlier reference prior to that of Wenz (1940) wherein the name Doxander was made available, so the name Doxander should be attributed to Wenz, 1940. Okutani (1965) allocated this species to Euprotomus Gill, 1870, but is was re-allocated to Doxander by Walls (1980). Kronenberg and Berkhout (1984) followed Okutani. Based on conchological evidence, Kronenberg (1998) described Mirabilistrombus as a new genus to accomodate this species. The species has been reported as living at depths of 120 m (as 67 fathoms) (Abbott 1960) and 40-80 m (Kronenberg and Berkhout 1984). The Arafura Sea record is from slightly deeper water. Genus Euprotomus Gill, 1870 Euprotomus aurora Kronenberg, 2002 (Fig. IE, F) Previously known distribution (taken from literature): Indian Ocean localities listed by Kronenberg (2002). Additional records: Japan, Mikawa; Philippines, Bohol and Cebu; Australia, Northern Territory, Darwin (Monsecour 2004); Indonesia, Java, west coast (Dharma 2005). Nerv distributional records. Papua New Guinea, East Cape (10°15’S, I50°52’E), ITT. Williams, c. 1935, NTM P2357, 1 specimen; Indonesia, south of Sunda Strait, 2 specimens; Palabuhan Ratu, south-west Java, 1 specimen; Pangandaran, south-west Java, 13 specimens; east Java, 1 specimen, all coll. BDJI. New fossil record. KENYA, locality 2a: Quarry in reef-limestone, eastern shore of Mombassa I larbour, just north east of the Swimming Club at English Point, as Stromhus auris-dianae (sic) (Cox 1930: 137). Cox also mentioned the Pleistocene of Dar-es-Salaam (Tanzania). Remarks. There is no doubt about the authenticity of the Papua New Guinea specimen recorded here. In NTM there is an original, hand-written statement by Mrs Thea J. Williams on 24 October 1983 to the Museum’s Director donating the collection of her late husband. Rev. Henry T. Williams, to the Museum on file. Rev. Williams served as a missionary for 16 years on islands off Papua New Guinea between 1930 and 1946 (Richard C. Willan pers. comm, to GKEN May 2005). The specimens in the BDJI collection and the specimen from Darwin (Monsecour 2004) partially fill in the gap between the Indian Ocean distribution as originally reported by Kronenberg (2002) and the Papua New Guinea specimen reported herein. The Indonesian records in the BDJI collection probably represent viable populations, given the number of specimens encountered. The Darwin (Monsecour 2004) and Papua New Guinea (herein) records are probably incidental findings however, possibly from larvae being carried in some fashion to those localities, as Willan (2005) does not mention this species from Ashmore, Cartier and Hibernia reefs (Timor Sea), nor a population near Darwin, an area often visited by R.C. Willan (pers. comm, to GKEN). The Philippine and Japanese records by Monsecour (2004) cannot be accepted unreservedly. The Japanese record is especially questionable, as it is very far from the established distribution range. Both Euprotomus bulla (Roding, 1798) and E. aurisdianae (Linnaeus, 1758) occur in the Philippines (Springsteen and Leobrera 1986) 48 New distributional records for Australian Stromboidea big. I .A, 15, Rostellariella delicatula (Nevill, 1881). Arafura Sea, taken at approximately 144 m by Indonesian fishermen, length 51.2 mm BDJI collection (photo Bunjamin Dharma); C, D, Mirabilistrombus lisleri (T. Gray, 1852). Arafura Sea, taken at approximately 144 in by Indonesian fishermen; C, Length 113.8 mm, BDJI collection; D, length 114 mm, GKEN collection no. 6293 (photo Bunjamin Dharma); E, F, Euprotomus aurora Kronenberg, 2002. Papua New Guinea, East Cape, leg. H.T. Williams, c. 1935, length 63.5 mm NTM P2357 (photo NTM); G, H, Dolomena hickeyi (Willan, 2000). Philippines, Cebu, Punta Engaiio, length 48 mm (photo Douglas Tolrud). and southern Japan (Higo et al. 1999), and a mix up of samples at the hands of a shell dealers can easily happen (pers. obs. GKEN). According to Monsecour (pers. comm, to GKEN, May 2005) the specimen he reported from Cebu, collected by Mike Filmer, is a genuine record, as Filmeronly collected specimens he had found personally. More records from the Philippines however are needed to verify the existence of E. aurora from that area. Nevertheless, it is remarkable that so soon after the description of this species, at first believed as being endemic to the Indian Ocean, records from the Pacific Basin arc emerging, a matter that needs further investigation. 49 G. C. Kronenberg and B. Dharma Genus Dolomena Wenz, 1940 As mentioned above for Doxander, the genus Dolomena has long been attributed to Iredale (Abbott 1960; Walls 1980; Kronenberg and Bcrkhout 1984; DeTurck et al. 1999; Raven 2002). As for Doxander, Iredale (1931) only gave a name and no description, and therefore Dolomena Iredale, 1931 is a nomen nudum and not available (ICZN art. 13). We have found no earlier reference prior to that of Wenz (1940), wherein the name Dolomena was made available. Therefore the name Dolomena should be attributed to Wenz (1940). Dolomena hickeyi (Willan, 2000) (Fig. 1G-H) Previously known distribution (taken from literature): Papua New Guinea and eastern coast of Queensland, Australia (Willan 2000); Malaysia, Kota Kinabalu; Sabah, Malawali Island; and Brunei, Jerudong (Raven 2002); Indonesia, Karimata Strait (Dharma 2005). New distributional records. THAILAND: Andaman Sea, Kor Bon, WBFT, 6 specimens (see below); Gulf of Thailand, Pattani, taken from fishermen’s nets, September 1998, HDWN coll. no. 2011, 5 specimens; Ban Pak Nam Sakom, 60 km west of Pattani (6°57’N, 100°49’36”E); local fishery. January 2000, HDWN coll. no. 4285, 1 specimen; MALAYSIA: Johor, Sedili, ex Ng Hiong Eng, VLFI coll. no. 617-09, 1 specimen; Borneo, near wreck ‘Hajicki Marn\ ex H. Morrison, VLFI coll. no. 617-07, 1 specimen. SINGAPORE; WBFT collection, 2 specimens; VIETNAM: off Nha Trang, VLFI coll. no. 617-02, 1 specimen; INDONESIA: North West Sumatra, Sibolga, 7 specimens; West Borneo, Tanjung Batu, 7 specimens; West Borneo, Pajantan Island, 1 specimen; Karimata Strait, between Kalimantan and Sumatra, 6 specimens. East Java, 4 specimens, all BDJI collection; PHILIPPINES: Mindoro, Puerto Galera area, Sabang, on sandy substrate at 19 m using scuba, 1 empty shell, M. Bouknegt, 18 December 2004; Cebu, Punta Engano, TBCM collection; PAPUA NEW GUINEA: New Britain, Rabaul (ex Lamprell coll.), VLFI coll. no. 617-05, 2 specimens; SOLOMON ISLANDS: Guadalcanal, Honiara bay, by diver, VLFI coll, no. 617-03, 1 specimen. Remarks. Dolomena hickeyi was originally described in Labiostrombus Oostingh, 1925 (as a subgenus of Strombns) by Willan (2000). We do not concur with this view, and prefer to assign this species to Dolomena based on shell morphology, as indicated by Abbott (1960). More research however is needed, especially regarding the assignment of several species currently assigned to Dolomena, see also the remarks made by Kronenberg and Vermeij (2002) on shell morphology within Strombidae. After the publication by Raven (2002) it was predictable that more localities would become known for this species. Although a number of the samples mentioned above have minimal locality data, we are convinced that the species actually lives in, or near, the areas mentioned. However, the Kor Bon record and the Singapore record (both from WBFT collection) need confirmation (WBFT in e-mail to GKEN May 2005). The Singapore record may be referable to specimens actually collected in Karimata Strait. The specimens from both Ban Pak Nam Sakom and Pattani were probably collected within a range ol 100 km from those places, at a maximum depth of 50 m, but rather at 15—35 m (e-mail HDWN to GKEN May 2005). The Punta Engano specimen (TBCM collection) may actually be from another Philippine locality, as shell dealers use Punta Engano as a ‘catch all’ locality in the Philippines. Bouwknegt (e-mail to GKEN May 2005) provided detailed information on the specimen he found at Sabang, Mindoro: “1 found the shell on the evening of 18 December 2004 during a dive to the three small shipwrecks lying on a sandy bottom at 19 m depth, off the beach ol Sabang. Near the largest wreck, which stands upright, there is sometimes a very weak current at the bow. This results in some shell grit being deposited and 1 found the shell in this grit. Although empty, without any remains ol soft parts or smell, the shell is perfect, even the rim of the aperture has no scratches or dullness; this indicates, I think, that the species actually lives very near that spot (a gently sloping sandy bottom, with hardly any overgrowth). Only at 10 m there is some coverage and coral heads and only at depths below 30 m there is more coral gravel with coverings. The weak current can sometimes be detected, but appears to me as not being strong enough to make significant transport along the bottom plausible.” In the original description (Willan 2000), this species was compared with the (partly) sympatric Dolomena pulchella (Reeve); D. dilatata (Swainson); D. minima (Linnaeus) and the allopatric D. columba (Lamarck). It appears that it may also be confused with D. dilatata form orosmina (Duclos) sensu Abbott (1960) (pers. obs. authors; WBFT in e-mail). Based on Raven (2002), Dolomena hickeyi and Dolomena dilatata form orosmina (Duclos, 1844) sensu Abbott (1960) can be distinguished. The situation within this group of species (i.e. Dolomena dilatata sensu Abbott including Dolomena hickeyi ) is not completely resolved however, and shall be discussed elsewhere (Kronenberg and Raven in prep.). This species is probably rather common in Indonesia, as the second author has observed over a hundred specimens in a box at a shell dealer’s shop in Jakarta. ACKNOWLEDGMENTS We thank the Indonesian fishermen for giving us an opportunity to study the specimens of RostellarieUa delicatula and Mirabilistrombus listeri from the Arafura Sea. We also thank Dr Richard Willan, N I M, for making us aware of the Papua New Guinea record of Euprotomns aurora and photographing that specimen; Mr Matthijs A. Bouwknegt, Wageningen, The Netherlands, for detailed information about his discovery of Dolomena hickeyi, 50 New distributional records for Australian Stromboidea ar *d a photograph of his specimen; Mr Winston Barney, F< Vt Worth, Texas, USA ; Mr Uenk Dekker, Winkel, The ^therlands; Mr Tim Blackwood, Cohasset, Minnesota, ^A; and Mr Virgilio Liverani, Faenza, Italy, kindly tired the data of their specimens of Dolomena hickeyi w 'th us, Mr Douglas Tolrud, Cohasset, Minnesota, USA, photographing the specimen of D. hickeyi from the Tim ^ackwood collection. The first author wishes to thank Dr ^ Hilippe Bouchet, Museum Nationale d’l listoire Naturelle, ^ ; Vis, France, for drawing our attention to the Iredale (1931) Publication; and Ms Marianne Matthijssen, Eindhoven, TV , he Netherlands, for her continuing support. We also thank Richard Willan for kindly reading the manuscript and v;) luable suggestions for improvement. REFERENCES Abbott, R.T. I960. The genus Strombus in the Indo-Pacific. Indo- Pacific Mollusca 1 (2): 33-146. ^x, L.R. 1930. Reports on geological collections from the coastlands of Kenya Colony. VII. Post Pliocene mollusca. [in McKinnon Wood, M.] Monographs of the Geological department of the Hunterian Museum Glasgow University 4: 103-112. ° e Turck, K„ Kreipl, K„ Man in ‘t Veld, L.M. and Poppe.G.T. 1999. In Poppe, G.T. and Groh, K. (cds). A conchologicaliconography. Family Strombidae. ConchBooks: Hackenheim. D *Wma, B. 2005. Recent and fossil Indonesian shells. ConchBooks: Hackenheim. M'go, S., Callomon, P. and Goto, Y. 1999. Catalogue and bibliography of the marine shell-bearing Mollusca of Japan. Gastropoda. Bivalvia, Polyplacophora, Scaphopoda. Elle Scientific Publications: Osaka. International Commission on Zoological Nomenclature, 1999. International Code of Zoological Nomenclature , fourth edition. London. * re dale, T. 1931. Australian molluscan notes. No. 1. Records of the Australian Museum 18(1-5): 201-235, pis. 22-25. Kronenberg, G.C. 1998. Revision of Euprotomus Gill. 1870. 1. The systematic position of Strombus listen Gray. 1852. Vita Marina 45(3—4): 1-6. Kronenberg, G.C. 2002. Revision of Euprotomus Gill, 1870. 3. Description of Euprotomus aurora spec. nov. from the Indian Ocean (Gastropoda, Strombidae). Vita Malacologica 1: 55-60 Kronenberg, G.C. and Berkhout, .1. 1984. Strombidae. Vita Marina 31 (1-6): 263-326, pis 1-9. Kronenberg, G.C. and Burger, AAV. 2002. On the subdivision of Recent Tibia-Wkc gastropods (Gastropoda, Stromboidea), with the recognition ofthe family Rostellariidae Gabb, 1868, and a note on the type species of Tibia Roding, 1798. Vita Malacologica 1: 39-48. Kronenberg, G.C. and Vermeij, G.J. 2002. Terestrombus and Tridentarius, new genera of Indo-Pacific Strombidae (Gastropoda), with comments on included taxa and on shell characters in Strombidae. Vita Malacologica 1: 49-54. Latiolais, J.M. 2003. The phylogenetic underpinnings for spatial patterns of morphological disparity: analyses using strombid gastropods. Masters thesis. Available from http://etd. lsu.edu/docs/available/etd-1112103-135018/unrestricted/ JaredFinalThesis.pdf.pdf Melvill, J.C. and Standcn. R. 1905. Rostellaria delicatula Nevill. Notes upon its distribution and limits of variation. Journal of Conchology 11 (6): 161-163, I pi. Monsecour, D. 2004. Strombus (Euprotomus) aurora (Kronenberg, 2002) (Gastropoda: Stromboidea: Strombidae) op de Filippijnen. Neptunea 3 (4): 1-6. Moolenbeek, R.G. and Dekker, H. 1993. On the identity of Strombus decorus and Strombus persicus , with the description of Strombus decorus masirensis n. ssp. and a note on Strombus fasciatus. Vita Marina 42 (1): 3-10. Morrison, H.M. 2005. Description of Rostellariella lorenzi spec. nov. from the Arafura Sea area of eastern Indonesia (Gastropoda: Stromboidea: Rostellariidae). Visava I (4): 15-23. Okutani, T. 1965. Strombus (Euprotomus) listeri T. Gray - New Records; anatomy. Indo-Pacific Mollusca 1 (6): 399-400. Raven. J.G.M. 2002. Notes on molluscs from NW Borneo. I. Stromboidea (Gastropoda, Strombidae, Rostellariidae, Seraphidae). Vita Malacologica 1: 3-32. Simone, L.R.L. 2005. Comparative morphological study of representatives of the three families of Stromboidea and the Xenophoroidea (Mollusca, Caenogastropoda), with an assessment of their phylogeny. Arquivos de Zoologia 37 (2): 141-267. Springsteen, F.J. and Leobrera, F.M. 1986. Shells ofthe Philippines. Carfel Shell Museum: Malate. Thach, N.N. 2005. Shells of Vietnam. Conchbooks: Hackenheim. Walls, J.G. 1980. Conchs, tibias, and harps. T.F.H. Publications: Neptune. Wenz, W. 1938-1944. Handbuch der Paliizoologie. Band 6. Gastropoda Teil 1: AllgemeinerTeil und Prosobranchia. Part 1:1 -240 March, 1938; Part 2: 241-480 October. 1938; Part 3: 481-720 July, 1939; Part 4: 721-960 August, 1940; Part 5: 961-1200, October, 1941; Part 6: 1201-1506 October, 1943; Part 7: 1507-1539 + i-xii, November, 1944. Willan, R.C. 2000. Strombus hickeyi, a new species in the subgenus Labiostrombus (Gastropoda: Strombidae) from the tropical southwestern Pacific Ocean. Vita Marina 47 (1): 18-24. Willan, R.C. 2005. The molluscan fauna from the emergent reefs ofthe northernmost Sabul Shelf. Timor Sea - Ashmore, Cartier and Flibernia Reefs; biodiversity and zoogeography. The Beagle, Records of the Museums and Art Galleries of the Northern Territory. Supplement 1: 51-81. Accepted 6 September 2005 51 '■> . ' The heagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21: 53-66 Ascidians from the Solomon Islands PATRICIA KOTT Queensland Museum, PO Box 3300, South Brisbane QLD 4101, AUSTRALIA patricia. mather@qm.qld.gov. an ABSTRACT Twenty-eight ascidian species from the Solomon Islands (western Pacific Ocean) are recorded including two new species (one in each of the genera Trididemnum and Lissoclinum). Extensions of the known geographic range and/or species characteristics are documented for a further 19 species (of which ten were known previously from no more than five records). Seven well known and often encountered species are newly recorded, without discussion. In situ colour images constitute a unique source of information on the living organisms and contribute to their recognition in the field. The species list is dominated by the Didemnidae (15 species) reflecting the diversity of that family in shallow water, reefal habitats in the tropics. Other species recorded from all suborders are also colonial, most with habits convergent with the Didemnidae. Keywords: West Pacific, tropical, Didemnidae, in situ images, Trididemnum mellitum sp. nov., Lissoclinum karenae sp. nov. INTRODUCTION twenty-eight ascidian species were collected by S*“lJBA diving from depths of four to ten metres in ^ ie Solomon Islands (Florida and Russell Groups and Guadalcanal). The specimens were all collected in June, ZOOq Mature gonads and larvae were detected only in Did^mnum membranaceum. Although a general breeding season for western Pacific tropical species has not been detected, larvae being found at all times of the year for many species (see Kott 1990, 1992,2001), it is tempting to speculate that their absence from all but one of the present specimens coincides with the southern winter. However, there is no direct evidence of this. With the exception only of a new species in each of the genera Trididemnum and Lissoclinum , the species recorded here are not unexpected components of the Solomon Island fauna, although ten of these species were known previously from five or fewer records. All of the species, except Polysyncraton adelon F. and C. Monniot, 2001 and the new species have been recorded previously from Australian waters and the material generally confirms the hypothesis that the northern Australian ascidian fauna is part of the wider western Pacific tropical fauna, sometimes extending into the tropical Indian Ocean. In a large (187 species) collection from the tropical western Pacific, F. and C. Monniot (2001) recorded 21 species of Eudistoma (Polycitoridae), but only three species of that genus are in the present collection. Nevertheless, about 25% of the species recorded by the Monniots (loc. cit) are in the Didemnidae while 50% (15 species) of the present collection are didemnids. The dominance of the latter family confirms its high diversity in shallow water, tropical habitats. However, the large, colourful colonies occupying extensive areas are conspicuous and could cause some sampling bias. The availability of in situ photographic images makes the present work a valuable illustrated catalogue to the field identification of these organisms. Some of the living specimens are seen to look dramatically different from the preserved material (e.g. Polysyncraton cucuHiferum). Specimens examined are registered in the collection of the Queensland Museum (QM) and duplicates are in the Solomon Islands’ National Museum (SINM). Species reported in this study (♦records only) Diazonidae *Rhopalaea crassa Herdman, 1886 - QM G308764; SINM E004 Clavelinidae Clavelina arafurensis Tokioka, 1952 Clavelina oliva Kott, 1990 *Clavelina robusta Kott, 1990 - QM G308761, Fig. 7C this work Polycitoridae Eudistoma glaucum (Sluiter, 1909) Eudistoma laysani (Sluiter, 1900) lEudistoma inauratum F. and C. Monniot, 2001 Polycitor giganteus (Herdman, 1899) Didemnidae Leptoclinides cons tella tus Kott, 2001 Leptoclinides erinaceus Kott, 2001 Polysyncraton adelon F. and C. Monniot, 2001 Polysyncraton cucuHiferum (Sluiter, 1909) Polysyncraton dromide Kott, 2001 53 P. Kott Didemnum albopunctatum Sluiter, 1909 Didemnum arancium Kott, 2001 Didemnum lacertosum Monniot, 1995 Didemnum membranaceum Sluiter, 1909 *Didemmtm molle Herdman, 1886 - QM G308769, G308770; SINM E010E01I Didemnum rota Kott, 2004b Trididemnum mellitum sp. nov. Lissoclinum karenae sp. nov. Lissoclinum regmum Kott, 2001 Diplosoma versicolor Monniot, 1994 Perophoridae Perophora namei Hartmeyer and Michaelsen, 1928 *Ecteinascidia nexa Sluiter, 1904 - QM G308789; Fig. 9G this work Styelidae *Eusynstyela latericius Sluiter, 1904 - QM G308794, G308795; SINM E005 E032 *Botrylloides perspicuum Herdman, 1886 - QM G308796; SINM E015; Fig. 9H Pyuridae *Herdmania mofnus (Savigny, 1816) - QM G308792 TAXONOMY Family Clavelinidae Clavelina arafurensis Tokioka, 1952 (Figs 1A, B; 7A) Clavelina arafurensis Tokioka, 1952: 97. - Kott 1990: 38 and synonymy; F. and C. Monniot 2001: 229 and synonymy. Distribution. Previously recorded (see F. and C. Monniot 2001): Western Australia (Exmouth Gulf); Arafura Sea; Truk Atoll; Palau Islands; Philippines; Mozambique. New record: Solomon Islands (Russell Group, QM G308763, SINM 029). Description. Living zooids have the white granular patches each side of the branchial aperture extending dorsally to surround the atrial siphon as described previously for this species. These patches are granular and white in preservative. The branchial sac is transparent with black pigment in the haemocoele and around each of the apertures. Colonies have a ough basal common test with vertical more or less cylindrical lobes, each containing up to ten completely embedded zooids. Zooids have about 15 rows of branchial stigmata. In these specimens about 8 transverse muscles extend from the endostyle across the thorax where they merge with longitudinal bands from the branchial and atrial siphons and extend along each side of the abdomen, terminating posteriorly in the two horns, one each side of the vascular stolon. The stomach is about halfway down the relatively short abdomen. Larvae are in the posterior part of the atrial cavity. The larval trunk is 0.62 mm long and the tail is wound almost three-quarters of the way around it. Remarks. F. and C. Monniot (2001) have referred to partially embedded zooids, although in all other reported specimens, the zooids are entirely embedded in the separate lobes of the colonies. Also, as in most species in this genus, there are muscles emanating from the endostyle and curving to form the longitudinal bands from the branchial and atrial siphon. A specimen from the Philippines (F. and C. Monniot 2001: fig. 22) is a strongly contracted specimen in which the endostylar muscles appear to have become isolated from a band of dorsal longitudinal muscles. Apart from its strongly contracted condition, the musculature is not essentially different from that of other clavelinid zooids. Larvae have not previously been reported for this species. They are distinctive in the absence of a frontal plate and in their simple concave triradially arranged adhesive organs that resemble those of the temperate Clavelina baudinensis Kott. 1957. The latter species is similar to the present one in its embedded zooids and branched colony, although its zooid pigmentation is quite different and it lacks the endostylar muscles of other present species. Clavelina (diva Kott, 1990 (Figs 2; 7B) Clavelina oliva Kott, 1990: 55. - Kott 2002: 22. Distribution. Previously recorded (sec Kott 1990, 2002): Western Australia (Danipier Archipelago, Kendrew Island, Shark Bay, Houtman’s Abrolhos); Queensland (Lindeman Island, Lizard Island); Northern Territory (Darwin); Philippines. New record: Solomon Islands (Florida Group, QM G308762, SINM E025). Description. In life the small zooids have white thoraces with a blue band around the apertures and a blue abdomen showing clearly through the translucent, colourless test. Zooids appear to be solitary, the thoraces supported on a relatively narrow stalk containing the abdomen and the posterior abdominal vascular stolon. The stalk expands basally into thicker test by which it is attached to the substrate. The expanded base contains the terminal ampullae which detach from the vascular stolon and develop into replicates of the zooids. The separately opening apertures have smooth rims. About 10 rows of stigmata are in the branchial sac. The gut loop is short, the stomach about halfway down it, with a short duodenal area opening into a rounded posterior stomach. A short length of the cylindrical intestine is in the pole of the gut loop. The stomach is symmetrical, and its otherwise smooth wall is divided into four more or less equal longitudinal areas by straight narrow strips of what appears to be glandular epithelium that extend the full length of the stomach. Gonads are enclosed in the short gut loop. Remarks. The small separate zooids, relatively short gut loop and long vascular stolon with numerous ectodermal terminal ampullae in the base are characteristic of the species. The symmetrical stomach, with a smooth wall interrupted by four straight strips of glandular 54 Ascidians from the Solomon Islands cells, which create the characteristic square shape that Berrill (1950) refers to, appears to be characteristic of the Clavelinidae. Family Polycitoridae Polycitor giganteus (Herdman, 1899) (Fig. 7D) Polyclinum giganteus Herdman, 1899: 79. Polycitor giganteus. - Kott 1990: 171 and synonymy; F. and C. Monniot 2001: 249. Distribution. Previously recorded (see Kott 1990, F. and C. Monniot 2001): the species is recorded from locations all around Australia. Kott (2001) thought it was an indigenous species although records from Papua New Guinea (F. and C. Monniot 2001) and the newly recorded colony from the Florida Group (QM G308790, SINM E009) indicate that the species does have a wider range into the West Pacific. The type location is Port Jackson (not South Australia as in F. and C. Monniot 2001). Remarks. Three large, firm, gelatinous species are known in this genus, viz. Polycitor circes Michaelsen, 1930, P. giganteus and P. translucidus Kott. 1957. They all have long thoraces with numerous stigmata and their Figs 1-5. 1, Clavelina arafurensis (QM G308763): A, colony; B, larva; 2, Clavelina oliva (QM G308762), zooid; 3, Eudistoma laysani (QM G308798), colony; 4, Polysyncraton cuculliferum (QM G308780), surface of part of colony showing a common cloacal aperture; 5, Trididemnum mellitum (QM G308776, holotype): A, thorax; B, abdomen. Scale bars: 1A, 1.0 cm; IB, 5A, B. 0.1 mm; 2, 3. 4, 1.0 mm. 55 P. Kott long zooids have separately opening apertures and are not arranged in circles or rudimentary systems as so often they are in species in the Polycitoridae. Polycitor circes is recorded from the northern half of the Australian continent, Papua New Guinea, New Caledonia and the Philippines. The type location, said to be ‘Australia’ in F. and C. Monniot (2001), is Cockburn Sound on the central coast of western Australia. It has large larvae with triradially arranged adhesive organs, fine longitudinal stomach folds and black pigment in the zooids. Polycitor translucidus also is recorded from the Philippines, New Caledonia, French Polynesia and the northern half of the Australian continent, and it penetrates south to Cockburn Sound on the western coast of the continent and to Port Jackson and Wilson’s Promontory (Victoria) on the eastern coast. It has a transparent (rather than translucent) test, a stomach like the present species that has over four longitudinal folds, and small larvae (to 0.08 mm) with antero-median adhesive organs. Polycitorgiganteus has a translucent test, a larva about 1.25 mm long with antero-median adhesive organs, and the stomach has four longitudinal grooves. It is distinguished from P. translucidus by its translucent test and longer larva; and from P. circes by its stomach, lack of black pigment and the vertical arrangement of the antero-median adhesive organs. Eudistomaglaucum (Sluiter, 1909) (Fig. 7E) Polycitorglaucum Sluiter, 1909: 12. Eudistoma glaucutn. - Kott 1990: 208 and synonymy; ?F. and C. Monniot 2001: 240. ?Eudistoma viride. - F. and C. Monniot 2001: 247. Distribution. Previously recorded (see Kott 1990): Queensland Great Barrier Reef); Palau Islands, Okinawa, Indonesia, Fiji. New records : Solomon Islands (Florida Group, QM G308773, SINM E008). Remarks. The colony, consisting of firm, darkly pigmented, rounded heads on short stalks, with separately opening zooids arranged in circles, is characteristic of this species, as are the zooids with their long oesophageal necks. The white patches in the photographed specimen are small didemnid colonies. Tokioka (1955) compared this species with Eudistoma viride Tokioka, 1955, a junior synonym of Sigillina signifera (Sluiter, 1909), which has flat-topped colonies with dark bluish-green zooids (see Kott 1990). F. and C. Monniot (2001) assigned specimens with a long oesophageal neck to Tokioka’s species, despite the fact that the latter has shorter zooids with the pronounced muscular posterior abdominal stolon of Sigillina and the greenish-black colour of the preserved zooids of S. signifera. Eudistoma viride of F. and C. Monniot (2001) has a long oesophageal neck and a number of embryos being incubated in the branchial sac (rather than the single large embryo of Sigillina signifera, which is being incubated in a brood pouch constricted off from the thorax), and it may be a synonym of the present species. Eudistoma glaucum of F. and C. Monniot (2001) is recorded from Papua New Guinea but is not described and its identity is not confirmed. Eudistoma laysani (Sluiter, 1900) (Fig. 3; 7F) Distoma laysani Sluiter, 1900: 9. Eudistoma laysani. - Monniot 1988:212 and synonymy; Kott 1990: 214 and synonymy. Distribution. Previously recorded (see Monniot 1988, Kott 1990): New South Wales (Botany Bay, Lake Macquarie); Queensland (NSW-Queensland border, Noosa, Gladstone, Heron Island, Sarina); Lord Howe Island; New Caledonia. Palau Islands, '/Philippines, French Polynesia, Hawaii. New records: Solomon Islands (Russell Group, QM G308798-9, SINM E014, E022). Description. Small, well-spaced, stalked, fiat-topped vertical lobes arise from a sandy common basal mat. There is only a sprinkling of sand on the vertical lobes. Zooids are in circles of up to eight around the top of the lobes, their atrial apertures opening toward the centre of the upper surface. In life, the lobes are white and translucent with a conspicuous narrow black line around each aperture joined by a straight median line between the two apertures. In preservative, the zooids are pinkish-white, but the black line is not present. Zooids are small and very contracted in the newly recorded specimens. They have the usual strong transverse muscles and longitudinal bands, three rows of stigmata, long oesophageal neck, and gonads in the pole of the gut loop. Remarks. Zooids of the genus Eudistoma have a conservative morphology, and often the species are determined on the form of the colony (including the arrangement of zooids and their organisation into rudimentary cloacal systems), the nature of test inclusions and the form of the larvae. The present species, with its small, naked, vertical lobes on a common sandy base, is especially difficult to characterise and it is probable that specimens with a similar conservative morphology have been occasionally assigned erroneously to this taxon. Tokioka (1967) describes a large colony (from the Palau Islands) consisting of 13 vertical lobes. The zooids conform to those generally found in this genus. Their arrangement in the vertical lobes is not documented and their larvae have a trunk about 1.0 mm long. Kott (1990) did not regard these specimens as synonyms of the present species. Millar (1975) recorded similar colonies from the Philippines and Indonesia, although he found the zooids arranged in circular systems with atrial apertures in the centre of the lobe. Larvae in Millar’s (1975) specimen have a trunk of 0.54 mm long, similar to those recorded by Kott (1990) from a range of locations but about half the length reported by Tokioka (1967). Kott (1990) did not observe circular systems in a range of colony forms, from those with many small vertical lobes with relatively few zooids, to single lobes with many zooids. She may have overlooked these systems in the 56 Ascidians from the Solomon Islands preserved material. Nevertheless, it is possible that not all of the range of different colonies Kott (1990) assigned to this species are conspecific. Eudistoma album Monniot, 1988 from Tonga, New Caledonia and the Marianas Islands has similar zooids and similar (but smaller) larvae. However, it has cushion-like colonies rather than the small vertical lobes of the present species, and some specimens of E. laysani of Kott (1990) may belong to this species. Eudistoma laysani ofMiIlar(1975) from the Philippines has similar colonies to the present species and single systems in each lobe. However, it has yellow pigment spots each side of the dorsal ganglion, and may be conspecific with E. punctatum F. and C. Monniot, 2001, which also has a prominent orange/red spot on each side of the dorsal ganglion. Nevertheless E. laysani (Sluiter, 1900) has a wide range in the tropical Pacific. It is characterised by its colonies consisting of a sandy basal mat and vertical zooid-bearing lobes with rudimentary cloacal systems opening on the top of each lobe and a larval trunk 0.4-0.5 mm long with four median ampullae alternating with the antero-median adhesive organs. The black lines around the apertures and connecting them in the mid-dorsal line (observed in the in situ photographs of the newly recorded colonies) are recorded here for the first time. Eudistoma laysani of F. and C. Monniot (2001) from Papua New Guinea is a new record, but is not described and the identification is not confirmed. ?Eudistoma inauratum F. and C. Monniot, 2001 (Fig. 7G) Distribution. Previously recorded (see F. and C. Monniot 2001): Palau Islands. New record 4 . Solomon Islands (Russell Group, QM G308800, SINM E021). Description. In life, the colonies are small, upright translucent yellowish spherical heads on a short, cylindrical sandy stalk. Sand is not present in the upper part of the colony. Zooids open separately on the spherical head. They do not appear to be in circles. In the preserved colonies, the zooids are very contracted and their morphology could not be determined. Remarks. The newly recorded colonies resemble those of E. inauratum , although they are translucent yellow rather than orange. Detail of the zooid structure is obscured by contraction. Leptoclinides constellatus Kott, 2001 (Fig. 7H) Leptoclinides constellatus Kott, 2001: 51. - Kott 2004b: 49 and synonymy. Distribution. Previously recorded (see Kott 2004b): Queensland (Whitsunday Islands); Northern Territory (Darwin, Bynoe Harbour). New record 4 . Solomon Islands (Florida Group, QM G308774, SINM E016). Description. The newly recorded colony is a robust slab with black pigment cells crowded into bands in the roof of common cloacal canals to form wide, black lattice-like markings on the surface. Well-spaced branchial apertures are along each side of the common cloacal canals. Patches of beige/tan are also present in the surface of the living colony. The conspicuous superficial layer of bladder cells overlies the spicules, which are in the upper half of the colony surrounding the common cloacal cavities and in the zooid-free areas between and surrounded by the common cloacal canals. Spicules also are in a thin layer on the base of the colony. Spicules are stellate to 0.075 mm diameter with 9-11 chisel-tipped rays in optical transverse section. The lower half of the colony is transparent and aspiculate. Black, granular, spherical bodies are scattered through the test. Remarks. Although neither the black streaks in the surface nor the black pigment cells were detected in the type material, the aspiculate basal test and the shape and form of the relatively small spicules help to distinguish the species. Leptoclinides erinaceus Kott, 2001 (Figs 6A, 8A) Leptoclinides erinaceus Kott, 2001: 61. Distribution. Previously recorded (see Kott 2001): Western Australia (NNW of Port Hedland), Queensland (Hardy Reef). New record : Solomon Islands (Florida Group, QM G308768, SINM E001). Description. This fleshy looking slab has an aspiculate superficial layer of test that is particularly conspicuous as brownish marks where the surface is depressed over the radial canals that converge to the randomly spaced, large, sessile, common cloacal apertures, which have either blackish ortransparent rims. The colour in life is pink over the common cloacal canals with the branchial apertures showing as conspicuous black dots suggesting that the zooids are black in life. Spicules are in thin layers, one beneath the bladder cell layer and another in the base of the colony, and the remainder of the colony is aspiculate. Spicules are large (to 0.125 mm), stellate (with 13-15 and sometimes 11 conical rays in optical transverse section) with pointed or chisel-shaped tips. The zooids are of the usual form for this genus, with a posteriorly orientated atrial siphon. Remarks. This species has the same number of spicule rays as the temperate species L. magnistellus , but they are smaller. Further, the surface of/,, magnistellus is raspy to the touch and lacks the layer of bladder cells found in the present species. The spicules are identical to those previously described, although some have only 11 rays in optical transverse section. 57 P. Kott Fig. 6. Scanning electron micrographs ofcalcareous spicules from the test of: A, Leptoclinides erinaceus (QM G308768); B, Polysyncraton adelon (QM G308779); C, Polysyncraton cuculliferum (QM G308780); I), Didemnum arancium (QM G308777); E, Didemnum lacertosum (QM G308771); F, Didemnum rota (QM G308783); G, Trididemnum mellitum (QM G308776, holotype); H, Lissoclinum karenae (QM G308781, holotype). 58 Ascidians from the Solomon Islands Fig. 7. In situ images: A. Clavelina arafurensis (QM G308763); B. Clavelina oliva (QM G308762); C, Clavelina robusta (QM G308761); D, Polycitor giganteus (QM G308790); E, Eudistoma glaucum (QM G308773); F, ?Eudistoma laysani (QM G308799); G, lEudistoma inauratum (QM G308800); H, Leptoclinides conslellatus (QM G308774). 59 P. Kott Polysyncraton addon F. and C. Monniot, 2001 (Figs 6B, 8B) Polysyncraton addon F. and C. Monniot, 2001: 272. Distribution. Previously recorded (see F. and C. Monniot 2001): Palau Islands. New records: Solomon Islands (Russell Group, QM G308778-9, SINM E019-20). Description. The newly recorded specimen is a stiff, hard, thin, encrusting sheet, greenish-black in preservative. Black/green cells are mixed with the bladder cells in a thin superficial layer of test and black cells are scattered sparsely through the remainder of the test amongst the white spicules. Spicules are globular, to 0.05 mm diameter, with relatively crowded flat-tipped, rod-like rays. In life the colony is red with white on the base and around the sides. Zooids arc blackish-green, with a fine retractor from halfway down the oesophageal neck. Remarks. The thin colonies, globular spicules to 0.05 mm diameter and the greenish colour of the zooids appear to be characteristic of this readily identified species. In their variable size these globular spicules resemble those of Lissoclinum patella, although the species differ from one another in other respects, including their generic characters. Polysyncraton cuculliferum (Sluiter, 1909) (Figs 4, 6C, 8C) Diplosomoides cuculliferum Sluiter, 1909: 90. Polysyncraton cuculliferum. - Kott 2002: 30; Kott 2005: 2427 and synonymy. Distribution. Previously recorded (see Kott 2005): Northern Territory (Darwin, Bynoe Harbour). Queensland (Great Barrier Reef, Whitsunday Islands, Hinchinbrook Island, Lizard Island); Indonesia. New record : Solomon Islands (Russell Group, QM G308780). Description. The colony is a double cone with one large terminal common cloacal aperture on one of the cones, and another between the two cones. Both the common cloacal apertures are sessile with spicules in the rim of the opening. Branchial apertures are at the base of, and partially covered by, conspicuous pointed papillae which are directed toward the common cloacal apertures. In life the soft, green colony with its conspicuous surface papillae has a dramatic and unique appearance. Spicules are present in a thin layer in the surface test but are sparse elsewhere. They are stellate with 7-9 long, tapered and sharply pointed rays and are up to 0.86 mm diameter. The common cloacal cavity is vast and extends from a horizontal space posterior to the zooids that separates the surface zooid-bearing layer of test from the basal or central test. The posterior abdominal cavity is crossed by connectives between the basal or central test and the surface zooid-bearing layer of test and it opens into a large open space beneath the common cloacal apertures. Zooids are robust. The ventral branchial lobe is enlarged and projects into the surface papillum. A wide atrial aperture with a small atrial lip from its anterior riiy, exposes the branchial sac to the common cloacal cavity A retractor muscle was not detected in this specimen Oesophageal buds are present. Up to 6 or 7 immature testis follicles were detected iq some of the zooids. Remarks. The colony shape and texture of the newly recorded specimen resembles Didemnum mode (Herdmary 1886), however the surface papillae and the translucent green colour are both different from D. mode which i§ smooth and opaque, its green colour deriving from its symbiotic green symbionts. Also the distinctive stellate spicules of the present species are readily distinguishes from the globular spicules of D. mode. Specimens front Darwin Harbour (see Kott 2002, 2004a) are similar in appearance to the newly recorded specimen, although some specimens of the junior synonym, P. echinatum Kott, 200) from north-eastern Queensland, are larger colonies anq the one from Bowden Reef (Kott 2001: pi. 5A) appears to be a different colour. Polysyncraton dromide Kott, 2001 (Fig. 8D) Polysyncraton dromide Kott, 2001: 49. - Kott 2002; 32; Kott 2004a: 2477. Distribution. Previously recorded (see Kott 2004a); northern Australia (Torres Strait, Darwin); Western Australia (Cockburn Sound). New record'. Solomon Islands (Florida Group, QM G308772, SINM E007). Description. The colony is aspiculate, and forms an irregular, brown, encrusting sheet with what appear as a mosaic of brown blisters over the surface. These blisters are separated by narrow depressions of the surface test over the circular common cloacal canals. The zooids line the common cloacal canals, their branchial apertures being along each side of these depressions, and their broad atrial lips fan out across the roof of the common cloacal canals, Abdomina are embedded in the test. Zooids have a long fine retractor muscle. Remarks. The common cloacal systems, consisting of circular canals lined on each side by the zooids, surrounding firm stands of test that project like blisters on the surface of the colony, occur in Polysyncraton arvum Kott, 2004b and P. catillum Kott, 2004a (from Ashmore Reef), P. palliolum Kott, 2001 (from Rottnest Island, Western Australia), P. pseudorugosum Monniot, 1993 (from central Queensland, Darwin, the Northern Territory and the Coral Sea), P. purou C. and F. Monniot, 1987 (from the Great Barrier Reef, Darwin, the Philippines and French Polynesia), and P. robustum Kott, 2001 (from Cockburn Sound, Western Australia). Of these, P. purou (which is more darkly pigmented than the present species) is the only other species that is aspiculate. The present aspiculate specimen, like one front Darwin (see Kott 2004a), differs from the type material in lacking the burr-like spieules either scattered sparsely 60 Ascidians from the Solomon Islands in the surface test or crowded in the test over the common cloacal canals. In life the present specimen looks browner than previously photographed specimens, but it has the same metallic sheen in the surface (see Kott 2002: fig. 24g) and the specimens are otherwise identical. Didemnum albopunctatum Sluiter, 1909 (Fig. 8E) Didemnum albopunctatum Sluiter, 1909: 148. - Kott, 2004b: 52 and synonymy. Distribution. Previously recorded (see Kott 2004b): Western Australia (Ashmore Reef, Rowley Shoals, Houtmans Abrolhos); Queensland (Great Barrier Reef); Indonesia; Indian Ocean (Cocos Keeling). New record: Solomon Islands (Florida Group, QM G308775, SINM E012). Description. The colony is a soft, encrusting sheet with black pigment cells in a superficial bladder cell layer, and spherical black pigment cells mixed with crowded spicules fill the remainder of the colony. The soft, flexible rather than brittle colony may be the result of the small (to 0.038 mm diameter) globular spicules with flat-tipped crowded rays. In life, a ring of black pigment in the bladder cell layer sometimes surrounds each branchial aperture, although this often is absent and white patches surround the apertures. The horizontal common cloacal cavity is at thorax level and is crossed by the thoracic part of the zooids, each with a strip of test along the ventrum. Abdomina are embedded in the white (with crowded spicules) basal test where the pigment cells are only sparse. Zooids have a short retractor muscle from a short distance down the oesophageal neck. Remarks. This species has a characteristic aspiculate surface layer of test. Relatively few species in this genus have globular spicules like those in the present species. They are present in D. precocinum Kott, 2001, which is distinguished by its deeper common cloacal cavity and the absence of a black bladder cell layer. Didemnum jedanense Sluiter, 1909 has similar-sized spicules but they have irregular ray tips, the zooids are along each side of circular canals and spicules usually are less crowded and are missing from many parts of the colony. Didemnum arancium Kott, 2001 (Figs 6D, 8F) Didemnum arancium Kott, 2001: 150 and synonymy. Distribution. Previously recorded (see Kott 2001): Queensland (southern Great Barrier Reef); French Polynesia. New records: Solomon Islands (Russell Group, QM G308777, SINM E018). Description. The colony is a hard white slab in preservative, and looks the same in life. Spicules, crowded throughout, are burr-shaped to globular, the latter with tiny, short conical points in the fiat ends of the rod-like rays. Remarks. The material recorded from Australia (including the type colonies) are salmon-coloured to orange vermilion, or a deep orange colour. Despite the white colour of the newly recorded colonies, they are otherwise the same as previously recorded colonies. The spicules of this species are unique. Didemnum lacertosum Monniot, 1995 (Figs 6E, 8G) Didemnum lacertosum Monniot, 1995: 311. - Kott 2001: 199. Distribution. Previously recorded (see Kott 2001): Queensland (southern Great Barrier Reef); New Caledonia. New record : Solomon Islands (Florida Group, QM G308771). Description. In preservative, the newly recorded colony is a white cushion, with three or four large common cloacal apertures around the outer margin. Spicules, up to 0.05 mm diameter, are crowded throughout the colony. They are globular and burr-shaped (both with crowded rod¬ shaped fiat-tipped rays), and stellate (with up to 19 crowded fusiform rays in optical transverse section). Gonads were not detected in the colony. Remarks. The species is readily identified by its polymorphic spicules that resemble those in Didemnum moseleyi (Herdman, 1886) and D. poecilomorpha F. and C. Monniot, 1996 (with symbiotic plant cells in the colony), which are the same size but have fewer rays. Didemnum vesperi Kott, 2004b also has similar spicules to the present species but the stellate spicules have shorter rays and the species has characteristic small colonies with a central common cloacal aperture. Didemnum membranaceum Sluiter, 1909 (Fig. 8H) Didemnum membranaceum Sluiter, 1909: 58. - Kott 2004a: 2497 and synonymy; Kott 2004b: 56. Distribution: Previously recorded (sec Kott 2004a,b): Western Australia (Montebello Island to Dongara, Marmion Lagoon, Cockburn Sound); South Australia (Kangaroo Island, SAM); Queensland (Moreton Bay, Heron Island, Swain Reefs, Broadhurst Reef, Fantome Island, Davies Reef); Northern Territory (Darwin); Timor Sea, Indonesia, Mieronesica, French Polynesia, Flong Kong. New record: Solomon Islands (Florida Group, QM G308782, SINM E024). Description. In preservative, the newly recorded colony is a white cushion with the usual superficial dirty brown marks resulting from the brown pigment cells in the thin superficial test amongst the small pointed papillae that are often on the surface of colonies of this species. The crowded spicules are characteristic with relatively rare giant stellate spicules with few rays scattered amongst the ordinary stellate spicules with pointed conical rays. The horizontal common cloacal cavity with thoraces crossing it separately, each associated with a ventral strip of test is also characteristic as are the small comma-shaped zooids with a fine retractor muscle from halfway down the oesophageal neck and about six stigmata in the anterior row on one 61 P. Kott Fig. 8. In situ images: A, Leptoclinides erinaceus (QM G308768): 15, Polysyncraton adelon (QM G308778); C, Polysyncraton cuculliferum (QM G308780); D, Polysyncraton dromide (QM G308772); E, Didemnum albopunctatum (QM G308775): F, Didemnum aranciutn (QM G308777); G, Didemnum lacertosum (QM G308771); H, Didemnum membranaceum (QM G308782). 62 Ascidians from the Solomon Islands Fig. 9. In silu images: A, Didemnum rota (QM G308783); 15. Trididemnum mellilwn (QM G308776, holotypc); C, Lissoclinum karenae (QM G308781, holotypc); D, Lissoclinum reginum (QM G308784); E, Diplosoma versicolor (QM G308785); F, Perophora namei (QM G308787); G, Ecteinascidia nexa (QM G308789); H, Botrylloidesperspicuum (QM G308796). 63 P. Kott side of the branchial sac. Larvae are in the basal test of the newly recorded colony and are of the usual form with four pairs of ectodermal ampullae. Remarks. This is one of the most commonly recorded species in the shallow water ascidian fauna of the tropical western Pacific. Although generally its morphology is conservative and resembles many other Didemnum spp., it is readily identified by the giant spicules that occur amongst its otherwise unremarkable stellate spicules. The species also has a remarkable range around the Australian coast and supports the hypothesis that the continental shelf constitutes a bridge for species between the tropics and temperate waters (see Kott 1985). Didemnum rota Kott, 2004 (Figs 6F; 9A) Didemnum rota Kott, 2004b: 58. Distribution. Previously recorded (see Kott 2004b): Ashmore Reef. New record : Solomon Islands (Russell Group, QM G308783, SINM E031). Description. In preservative, the newly recorded colonies are fiat, hard, off-white sheets. The basal surface is white and wrinkled, possibly resulting from its removal from the substrate. The upper surface is yellowish and dimpled where the branchial apertures are withdrawn into the surface. The superficial test has a slightly fluffy look as if the otherwise crowded spicules are mixed with bladder cells. In life the colony appears to be a deeper yellow colour on the surface shading into a yellowish-red around the common cloacal apertures. Spicules (to 0.062 mm diameter) are stellate with 7-9 almost cylindrical rays in optical transverse section. The ray tips are relatively blunt. A thoracic cloacal cavity is shallow and horizontal and is crossed by the relatively robust thoraces with moderately long branchial siphons. The atrial aperture is sessile and lacks an anterior lip. A retractor muscle projects from about halfway down the oesophageal neck. Gonads were not detected. Remarks. Although the present colony is yellow in life, while the holotype of D. rota is brick-red (see Kott 2004b), the colonies appear to be similar and some traces of the red pigment are in the surface of the newly recorded specimen. Most aspects of the zooids and the spicules appear to be the same in these specimens. Spicules are similar though smaller and with longer, thinner rays than those of D. lillipution Kott, 2004b, which also is recorded from Ashmore Reef. However the colonies of D. lillipution are colourless, and thin and brittle, and its zooids are larger and less robust than those of the present species. Trididemnum mellitum sp. nov. (Figs 5A,B; 6G; 9B) Trididemnum savignii. - Kott 2004a: 2505. Distribution. Type locality. Solomon Islands (Florida Group, Nggele Side off Uru Point, 09°01.4"S 160°06.69’’E, 4-6m, coll. K. Gowlett-Flolmes 23 June 2004, holotype QM G308776; paratype SINM E013). Previously recorded'. Northern Territory (Darwin Harbour, Kott 2004a, QM G308618, G308626). Description. The holotype colony is a thin cushion to 2 cm diameter, although an in situ photograph shows a translucent yellow, irregular cushion. The colony is a pinkish colour in preservative with short, upright, almost cylindrical lobes on the upper surface. Large terminal common cloacal apertures are on the upright lobes of the colony. Large spicules (to 0.1 mm diameter) are evenly distributed (but not crowded) in a layer beneath an aspiculatc surface layer of bladder cells. Spicules are sparse in the remainder of the test. They are stellate with 11-13 spiky rays in optical transverse section. Flat-topped stubs sometimes are crowded between the base of the rays where, possibly, their conical tips have broken off. The common cloacal cavity is extensive and zooids arc suspended across it in test connectives that join the surface to the basal layer of test. A layer of faecal pellets is embedded in the basal test. Zooids are relatively small, with a conspicuous branchial siphon and posteriorly orientated atrial siphon, each with six very similar rounded lobes on the rim of the opening. An endostylar pigment cap was detected over the anterior end of the endostyle and the abdomen of the preserved zooids is covered with black squamous epithelium. Fine, parallel, longitudinal muscles are on the thorax and a short, thick retractor muscle extends from the posterior end of the branchial sac. An imperforate area is in front of the perforated part of the branchial sac. About 12 stigmata are present in each half row. The vas deferens coils eight times around the undivided testis follicle. Remarks. The specimen from Darwin assigned to Trididemnum savignii by Kott 2004a (QM G308626) has similar colony, spicules and is the same honey yellow colour as the present holotype and appears to be a synonym. Both specimens differ from T. savignii Herdman, 1886 in their yellow colour, lobed colonies, more numerous spicule rays, absence of spicules from the interior of the colony, and the presence of a layer of faecal pellets in the basal test. These colonies are irregular and lobed, but do not form the three-dimensional reticulae of Trididemnum sibogae. Trididemnum tomarahi C. and F. Monniot, 1987 has smaller spicules than the present species. The six rounded atrial lobes around the aperture are unusually conspicuous for this genus, resembling the branchial lobes, rather than the five pointed papillae around the rim of the atrial aperture in Leptoclinides spp. Despite its posteriorly orientated atrial siphon the species is readily distinguished from Leptoclinides spp. by the presence of the retractor muscle as well as its three rows of stigmata and its entire testis follicle. Etymology. The species name is derived from the colony colour in life, viz. mellitus, -a, -um, honey coloured. 64 Ascidians from the Solomon Islands Lissocliiutnt karenae sp. nov. (Figs 6H; 9C) Distribution. Type locality. Solomon Islands (Florida Group - small island E. of Sandfly Passage, under rock, 4-6m, SINM E023, coll. K. Gowlett Holmes, 28 June 2004, holotype QM G308781; paratype SINM E023). Description. The holotype is a tongue-shaped cushion about 2 cm long, salmon-pink and grey on the upper surface. It has a row of four common cloacal apertures ringed with grey owing to especially crowded pigment cells. These apertures are evenly spaced along the centre of the colony. The dark internal test is seen through the common cloacal openings. The colour of the upper surface of the colony is the result of red pigment cells mixed with white spicules in the upper half of the colony. In the preserved specimen, pigment is not present in the solid basal test which appears cream in colour. Otherwise the preserved specimen appears to be the same colour as it was in life. Stellate branchial apertures are evenly spaced on the surface. A vast horizontal common cloacal cavity is in the upper half of the colony and is crossed by the whole length of the zooids, each encased in a strip of test, exposing only almost the whole of the branchial sac. The spicules are the usual burr-like ones with many rod- like flat-tipped rays, although they are larger than those in most species of this genus, being to 0.065 mm diameter. Remarks. Although gonads were not detected, the species resembles Lissoclinum spp. in its large zooids with an almost rectangular thorax, large common cloacal cavity and burr-shaped spicules. The colony, with conspicuous common cloacal apertures on firm, fleshy rounded branches rather than flat sheets is most like Lissoclinum badium F. and C. Monniot, 1996, as is the dark internal test with pigment cells surrounding the zooids. However, it differs in having spicules almost twice the size of L. badium, and also it lacks the yellow pigment present in the latter species. Lissoclinum textile F. and C. Monniot, 2001 has similar but larger spicules (up to 0.08 mm diameter) and zooids are arranged along each side of circular common cloacal canals, which have brown pigment over them. Etymology. The species is named for Karen Gowlett- Holmes, who collected and photographed the specimens in this collection. Lissocliituin reginum Kott, 2001 (Fig. 9D) Lissoclinum reginum Kott, 2001: 319 and synonymy. - Kott 2004b: 65 and synonymy. Distribution. Previously recorded (see Kott 2004b): Western Australia (Ashmore Reef, Port Hedland); Queensland (Great Barrier Reef); Northern Territory (Darwin, Bynoe Harbour); Indian Ocean (Cocos Keeling Is). New record: Solomon Islands (Russell Group, QM G308784). Remarks. The in situ photograph of the newly recorded specimen is purple with white flecks around the common cloacal apertures (Fig. 9D) and closely resembles others previously recorded (Kott 2001: pi. 20c). The spicules are also identical with the burr-shaped spicules formerly described for this and other Lissoclinum spp. Diplosoma versicolor Monniot, 1994 (Fig. 9E) Diplosoma versicolor Monniot, 1994: 9. - Kott 2004b: 67 and synonymy. Distribution. Previously recorded (see Kott 2004b): Western Australia (Ashmore Reef, Montebello Island, Houtman’s Abrolhos); New South Wales (Lord Howe Island); Queensland (Southern Great Barrier Reef, Townsville, Lizard Island); Northern Territory (Darwin, Bynoe Harbour); Micronesia, New Caledonia, Philippines. New record: Solomon Islands (Florida Group, QM G308785, SINM E026). The species has also been recorded from Noosa on the south-eastern coast of Queensland. Remarks. The newly recorded colony of this commonly encountered species is soft, and has patches of whitish bodies (that do not appear to be calcareous) in the black- brown surface test. The yellowish patches present in most specimens of this species are not present, and the colony resembles the black and white specimens reported by F. and C. Monniot (2001). Family Perophoridae Peropltora namei Hartmeyer and Michaelsen, 1928 (Fig. 9F) Peropltora namei Hartmeyer and Michaelsen, 1928: 270. - Kott 1985: 108 and synonymy; F. and C. Monniot, 2001: 301 (records only). Distribution. Previously recorded (see Kott 1985, F. and. C. Monniot 2001): Coral Sea, Papua New Guinea, Philippines. New records: Solomon Islands (Russell Group, QM G308787-8, SINM E027). Remarks. Once known only from the Philippines, new records ol this fragile and beautiful species indicate that it has a wide range in the West Pacific. ACKNOWLEDGMENTS The collection was made under a permit from the Solomon Islands Ministry of Education and Training for the Solomon Islands National Museum. Additional information about this group of organisms is available as a result ot Karen Gowlett-Holmes' careful documentation ol the material, the high quality of her in situ photographic images and her careful indexing of the photographs to the fixed material. As a result, it now is possible to recognise many of the tropical West Pacific species in the field. I thank her for her efforts and the contribution she is making to the understanding of the Ascidiacea. 65 P. Kott I am grateful also to the Director and Board of the Queensland Museum for their continuing support of this project by providing laboratory accommodation and infrastructure. My research assistant Eileen Salisbury prepared and scanned the didemnid spicules, drew the black and white figures and maintained the database of specimens and helped to collate the final manuscript. This assistance is provided by a grant from the Australian Biological Resources Study (ABRS) and I am grateful for the continuing support of that agency. REFERENCES Berrill, N.J. 1950. The Tunicata. Ray Society Publications 133: 1-354. Hartmeyer, R. and Michaelsen, W. 1928. Ascidiae Diktyobranchiae und Ptychobranchiae. Fauna Sudwest-Australien 5: 251-460. Herdman. W.A. 1886. Report on the Tunicata collected during the voyage of HATS. ’Challenger' during the years 1873-76. Pt. II. Ascidiae conipositae. Zoology 14(38): 1-425, Herdman, W.A. 1899. Descriptive catalogue of the Tunicata in the Australian Museum. Australian Museum, Sydney, Catalogue 17: 1-139. Kott, P 1957. Ascidians of Australia II. Aplousobranchiata Lahille; Clavelinidae Forbes and Hanley and Polyclinidae Verrill. Australian Journal of Marine and Freshwater Research. 8(1): 64-110. Kott, P 1985. The Australian Ascidiaeea Pt I. Phlebobranchia and Stolidobranchia. Memoirs of the Queensland Museum 23: 1-440. Kott, P 1990. The Australian Ascidiaeea, Pt 2 Aplousobranchia(l). Memoirs of the Queensland Museum 29(1): 1-266. Kott, P. 1992. The Australian Ascidiaeea, Pi 3 Aplousobranchia (1). Memoirs of the Queensland Museum 32(2): 377-620. Kott, P. 2001. The Australian Ascidiaeea Pt 4, Didemnidae. Memoirs of the Queensland Museum 47(1): 1-410. Kott, P. 2002. Ascidiaeea (Tunicata) from Darwin, Northern Territory, Australia. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 18: 19-55. Kott, P. 2004a. New and little known species of Didemnidae (Ascidiaeea. Tunicata) from Australia (Part 2). Journal of Natural History 38( 19): 2455-2526. Kott, P. 2004b. Ascidiaeea (Tunicata) in Australian waters of the Timor and Arafura Seas. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 20: 37-81. Kott, P. 2005. New and little known species of Didemnidae (Ascidiaeea, Tunicata) from Australia (part 3). Journal of Natural History 39(26): 2409-2479. Michaelsen, W. 1930. Ascidae Krikobranchiae. Fauna Sudwest- Australiens. 5(7): 463-558. Millar, R.ll. 1975. Ascidians from the Indo-West Pacific region in the Zoological Museum, Copenhagen (Tunicata, Ascidiaeea). Steenstrupia 3(20): 205-336. Monniot, C. and Monniot, F. 1987. Les ascidies de Polynesie Frangaise. Memoires du Museum National d’Histoire Naturelle Paris 136: 1-155. Monniot, F. 1988. Ascidies de Nouvclle Caledonie V. Polycitoridae du lagon. Bulletin du Museum National d ’Histoire Naturelle. Paris , scr. 4 10A(2): 197-235. Monniot, F. 1993. Ascidies de Nouvellc-Caledonie XIII Le genre Polysyncraton (Didemnidae). Bulletin du Museum National d’Histoire Naturelle, Paris , ser. 4 15A (1-4): 3-17. Monniot, F. 1994. Ascidies de Nouvellc-Caledonie XIV. Le genre Diplosoma (Didemnidae). Bulletin du Museum National d'Histoire Naturelle, Paris, ser. 4 16A(1): 3-11. Monniot, F. 1995. Ascidies de Nouvclle-Caledonie XV. Le genre Didemnum. Bulletin du Museum National. d’Histoire Naturelle, Paris, ser. 4 I6A(2—4): 299-344. Monniot, F. and Monniot, C. 1996. New collections of ascidians from the western Pacific and southeastern Asia. Micronesica 29(2): 133-279. Monniot, F. and Monniot, C. 2001. Ascidians from the tropical western Pacific. Zoosystema 23(2): 201-383. Savigny, J.C. 1816. Memoires sur les Animauxsans Vertebres. Part 2. Paris: G. Dufour pp. 1-239. Sluiter, C.P. 1900. Tunicaten aus dem Stillen Ocean. Zoologische Jahrbiicher (Jena). (Systematik) 13: I 35. Sluiter, C.P. 1904. Die Tunicaten der Siboga-Expedition. Pt. I, Die socialen und holosomen Ascidien. Siboga Expeditie. 56A: 1-126. Sluiter, C.P. 1909. Die Tunicaten der Siboga Expedition Pt. 2. Die merosomen Ascidien. Siboga Expeditie 56B: 1-112. Tokioka, T. 1952. Ascidians collected by Messrs Renzi Wada and Seizi Wada from the Pearl Oyster bed in the Arafura Sea in 1940. Publications of the Seto Marine Biological Laboratory 2(2): 91-142. Tokioka, T. 1955. Contributions to Japanese ascidian fauna XL Sporadic memoranda (2). Publications of the Seto Marine Biological Laboratory 4(2-3): 205-22. Tokioka, T. 1967. Pacific Tunicata of the United States National Museum. Bulletin U.S. National Museum 251: 1-242. Accepted 9 October 2005 66 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21 : 67-72 A new species of the circumtropical goby genus Gnatholepis Bleeker (Teleostei: Gobiidae: Gobionellinae) from northern Australia HELEN K. LARSON 1 AND DUNCAN BUCKLE 2 'Museum and Art Gallery of the Northern Territory PO Box 4646, Darwin NT 0801, AUSTRALIA Email: helen.larson@nt.gov.au 2 Environmental Research Institute of the Supervising Scientist Department of the Environment and Heritage GPO Box 461, Darwin NT 0801, AUSTRALIA Email: Duncan.Buckle@deh.gov.au ABSTRACT A new species of Gnatholepis is described from northern Australia, where it is common on shallow intertidal rocky reefs. It is sexually dichromatic, with males having bright blue markings along the side of the body. The new species’ closest relative is G. gymnocara Randall and Greenfield, 2001, which is restricted to the east coast of Queensland. Both these species are small, lack scales on the side of the head and lack enlarged canine teeth in the lower jaw. The new species has shown to be basal to all other Gnatholepis species. Keywords: new species, Gnatholepis, Gobiidae, Gobionellinae, Indo-Pacific, northern Australia. INTRODUCTION The circumtropical marine gobiid genus Gnatholepis is unusual in that it is the only fully marine genus of the subfamily Gobionellinae, species of which are mostly estuarine to freshwater dwelling (Larson 2001; Thacker 2004). There has been little work done on the species of Gnatholepis, and attempts at distinguishing species have been few and not always in agreement (e.g. Iloese 1986; Kuiter and Tonozuka 2001; Nakabo 2002). Characters of the genus Gnatholepis were recently reviewed by Randall and Greenfield (2001) in their description of a new species from northern Australia ( G. gymnocara) and four new subspecies of G. cauerensis. They also created neotypes for Gobius anjerensis Bleeker, 1851 (the type species of the genus), and for Gnatholepis davaoensis Seale, 1910, and synonymised several species, without explanation, with Gnatholepis anjerensis. Randall and Greenfield’s work was the first attempt by anyone to review the genus. Randall and Greenfield (2001) illustrated an undescribed small species, known only from the Northern Territory (first collected there in 1981 by HKL) and off Cape York, and stated that it was most similar to their G. gymnocara, as both species lack scales on the cheek, opercle and predorsal region, unlike other species of the genus. In their description of G. gymnocara, Randall and Greenfield included two paratype specimens from the Northern Territory (AMS 1.23930-011). As no other specimens of G. gymnocara have so far been found in the Territory, these paratypes were examined and were found to be female specimens of the undescribed species (the larger specimen had a second dorsal fin count of 1,11, but anal ray count of 1,11, while the smaller specimen had 1,10 second dorsal rays and 1,11 anal rays; both have scaleless pectoral fin bases and typical female colouring for the new species). We describe this new species here, and briefly comment on some other species of the genus. MATERIAL AND METHODS Morphometries and measurements. Measurements were taken using electronic callipers and dissecting stereomicroscope. Counts and methods generally follow Hubbs and Lagler(1958), except for transverse scale counts (TRB), taken by counting the number of scale rows from the anal fin origin diagonally upward and back toward the second dorsal fin base, and head length is taken to the upper attachment of the opercular membrane. Morphometric values are expressed as a percentage of standard length (SL) or head length (HL). In the description, numbers in parentheses after counts indicate the number of specimens with that count, or the range of counts. Pterygiophore formula follows Birdsong etal. (1988). Vertebral counts and 67 H. K. Larson and D. Buckle other osteological information were obtained by clearing and double-staining. Terminology for lateral canals and sensory pores follows Larson (2001). Papillae rows are named based on Sanzo (1911). Abbreviations for institutions referred to are as in Leviton et al. (1985); but note that the name of the Museum and Art Gallery of the Northern Territory is no longer as it appears in Leviton et al. SYSTEMATICS Gnatholepis argus sp. nov. (Figs 1-5) Gnatholepisgymnocara Randall and Greenfield, 2001: 14 (in part). Gnatholepis sp. - Randall and Greenfield 2001: 16, pi. IIH; Thacker 2004: figs 1-2, tables 1-2. Diagnosis. A small Gnatholepis (up to 33 mm SL) lacking scales on side of head, predorsal midline and pectoral fin base, no distinct flap on end of lower lip (low fold may be present), and no distinct canine or enlarged teeth; sexually dimorphic in adult size and colour, with females averaging larger than males; sexually dichromatic: males with row of six dusky to blackish spots or blotches along mid-side of body, posteriormost 3-4 blackish spots darker and larger than anterior spots and surrounded by small iridescent blue spots in life, small dense black spots scattered on unpaired fins; females with broken grey line along mid-side of body and dusky to blackish spots or blotches indistinct and posterior blotches not darker than anterior ones, no small iridescent blue spots on body; second dorsal fin rays modally 1,10; anal fin rays modally 1,11; pectoral rays 15-18; longitudinal scales 24-28; predorsal scales absent from nape midline, scales on side of head may reach from just over opercle to nearly behind eyes. Material examined. 492 specimens, 6-33 mm SL. HOLOTYPE - NTM S.14965-032, 25.5 mm SL male, N side of Turtle Reef off Field Island, Northern Territory, 0-0.25 m, coll. II. Larson and party, 10 September 1999. PARATYPES - NORTHERN TERRITORY: NTM S.14965-008, 105(11-26), same data as holotype; QM 1.37222, 12(19-25.5), same data as holotype; USNM 384336, 12(19-25.5), same data as holotype; NTM S. 10005-034, 128(8-26), reef flat on E side Burford Island, off Aiton Bay, NT, 0-0.5 m, coll. II. Larson, 13 October 1981; BPBM 40168, 12(16-24.5), same data as preceding; NTM S.10006-027, 7(16-26), pools among Fig. 1.1 lolotype of Gnatholepis argus sp. nov., NTM S. 14965-032,25.5 mm SL male from north side of Field Island, Northern Territory. Fig. 2. Papillae pattern and sensory canal arrangement in 27.5 mm SL female Gnatholepis argus sp. nov.. East Point Reef, Darwin Harbour, Northern Territory (AMS 1.23930-011); also a paratype specimen of Gnatholepis gymnocara. A, lateral view; B, underside of head. 68 New goby from northern Australia mangrove roots along shoreline, E side Burford Island, off Aiton Bay, NT, 0-0.5 m, coll. H. Larson, 13 October 1981; NTM S.13237-038, 38(12-33), fringing reef flat on W side Rimbija Island, off Cape Wessel, NT, 0-0.5 m, coll. H. Larson and J. Hanley, 16 November 1990; NTM S.14966-022, 46(6-26), rocky reef on N side Field Island, NT, 0.05 m, coll. H. Larson, R. Williams and A. Pickering, 10 September 1999; NTM S.10411-026, 22(12-23), rocky reef off Bullocky Point, Fannie Bay, Darwin, NT, 0-0.25 in, coll. H. Larson, 11 December 1981; S.10033-005, 115 (9-22), reef Hat SE of Dudley Point, East Point area, Darwin, 0-0.25 m, coll. H. Larson and P. Horner, 13 November 1981. QUEENSLAND: AMS 1.19356-016, 61(22-33), N side Terry Beach, Prince of Wales Island, Torres Strait, 1 m, coll. D. Brown and W. Ponder, 2 July 1976. WESTERN AUSTRALIA: AMS 1.24713-005,1(29), No Name Bay, Dampier, coll. S. Blaber, 7 January 1989. Other material. NORTHERN TERRITORY: NTM S.13718-014, 2, Darwin Harbour; NTM S.12811-038, 3, Channel Island; NTM S.10429-033, 11, Channel Island; NTM S.12447-015, 3, East Arm Darwin Harbour; AMS 1.23948-013,2, East Arm; AMS 1.24676-014,12, East Arm; AMS 1.23930-011,2 paratypes of Gnatliolepis gymnocara. East Point; NTM S.10412-005, 3, Bullocky Point; AMS 1.24677-003, 12, Bullocky Point; NTM S.l5675-016, 8 , Dudley Point; NTM S.l 1813-004, 1, East Point; NTM S.l0417-001, 22, East Point; AMS 1.24678-007, 10, East Point; NTM S.15221-019, 1, Nightcl iff reef; AMS 1.24694- 004, 88 , Gunn Point; NTM S.12448-012, 21, Lee Point; AMS 1.24688-007, 2, Lee Point; NTM S.10415-019, 24, Lee Point; NTM S.10036-006, 12, Lee Point; NTM S.10690-005, 3, Vernon Islands; NTM S.10011-015, 10, Trepang Bay; NTM S.14961-007, 1, West Alligator Head; NTM S.l5532-018, 51, Field Island; NTM S.14471-007, 2, Field Island; NTM S.14469-004, 34, Field Island; NTM S. 14472-014, 22, Field Island; NTM S. 14665-024, 7, Field Island; NTM S.10004-034, 1, Sandy Island Number Two; NTM S.10603-030, 1, North Oxley Island. Description. Based on 80 specimens, 11.5-32.5 mm SL. An asterisk indicates the counts of the holotype (Fig. 1). First dorsal VI*; second dorsal 1,9-1,11 (modally 1,10*); anal 1,11-12 (modally 1,11*), pectoral rays 15-18 (modally 17*), segmented caudal rays always 17*; caudal ray pattern usually 9/8*; branched caudal rays 6/6 to 8/7* (modally 7/7); unsegmented (procurrent) rays 7/6 (I), 8/8 (1); longitudinal scale count 24-28 (mode 26, 27 in holotype); TRB 7 - 9/2 (mode 8 / 2 , IVz in holotype); circumpeduncular scales 11-12* (mode 12)(Table I). Gill rakers on outer face of first arch 0-1 + 2 3 (in 8 , modally 1+3). Pterygiophore formula 3-22110 (in 2). Vertebrae 10+16 (in 2), including urostyle. One (in 1) or two epurals (in 1). Two anal pterygiophores before haemal spine of first caudal vertebra (in 2 ). Body compressed, somewhat more rounded anteriorly. Head compressed, broader ventrally, slightly deeper than wide, ML 24.9-30.0% (mean 27.5%) of SL; head depth at Table 1. Meristics of specimens of Gnatliolepis argus sp. nov. Holotype Means Max. Min. Mode Second dorsal rays 1,10 1,10 1,11 1,9 1,10 Anal rays 1,11 fill 1,12 1,10 1,11 Pectoral rays right 17 17 18 16 17 Pectoral rays left 17 17 18 15 17 Caudal segmented 17 17 17 17 17 Caudal branched 15 14 16 12 14 Longitudinal scales 27 26 28 24 26 Transverse rows back 1'A 8/2 10 7 8/2 Transv. rows forward 7 9 12 7 9 Caudal peduncle scales 12 12 12 11 12 posterior preopercular margin 63.0-80.0% (mean 70.1%) of HL; head width at posterior preopercular margin 59.6-76.5% (mean 67.6%) of HL; head profile bluntly pointed to bluntly rounded. Mouth subterminal and small, almost horizontal; jaws generally reaching to below anterior half of eye; upper jaw length 29.3-39.0% (mean 34.2%) of HL. Lips smooth; upper lip broader than lower, lower lip with low twist or fold on posterior portion, but not forming triangular llap as in most Gnatliolepis, lip broadening anteriorly and interrupted at chin by narrow mental bump or short longitudinal fold on chin. Eye moderate, round, dorsolateral, top forming part of dorsal profile, 25.0-33.3% (mean 29.3%) of I1L. Snout blunt to rounded, 22.7-35.2% (mean 28.6%) of HL; posterior nostril small and round, adjacent to anterior margin of eye; anterior nostril in short tube, close to anterior nostril, at a level with ventral margin of eye. Interorbital very narrow, 4.4-8.6% (mean 6.3%) of HL. Body depth at anus 16.7-22.1% (mean 19.7%) of SL. Caudal peduncle compressed, length 12.0-18.2% (mean 16.0%) of SL; caudal peduncle depth 6.7-12.6% (mean 11.3%) of SL (Table 2). First dorsal fin rounded to roughly triangular or square, with no spines greatly elongate; third or fourth spine usually longest or subequal in length; when depressed, spine tips reach to first to third element of second dorsal fin or falling short of first fin element (usually in specimens less than 20 mm SL). Third dorsal spine 13.3-20.4% (mean 16.2%) of SL; fourth dorsal spine length 13.4-20.9% (mean 17.3%) of SL. Second dorsal fin almost as tall as first dorsal, rays longer posteriorly than anteriorly, fin pointed posteriorly. Anal fin somewhat lower than second dorsal, anteriormost rays shorter than posterior few rays; fin pointed to slightly rounded posteriorly. Second dorsal and anal fin rays, when depressed, reaching caudal fin in adults. Pectoral fin pointed, central rays longest, 22.6-30.4% (mean 26.1%) of SL; rays all branched but for upper and lowermost ray, fin reaches to above first few anal fin rays. Pelvic fins fused, frenurn with finely fimbriate margin, fins oval, reaching to above anus or nearly to anal fin origin, 22.9-28.7% (mean 26.5%) in SL. Caudal fin moderate, oval, 25.2-36.5% (mean 30.9%) of SL. Chin usually with small longitudinal bump or low fold. Gill opening restricted, extending anteriorly to lower edge of pectoral base. Gill rakers on outer face of first 69 H. K. Larson and D. Buckle Table 2. Measurements of specimens of Gnatholepis argus sp. nov., expressed as percentage of standard length (SL) or head length (HL). Holotype (male) Means (males) Max. (males) Min. (males) Means (females) Max. (females) Min. (females) Standard length 25.5 20.8 29.5 11.5 23.9 32.5 15.5 Head length in SL 26.3 27.3 29.4 24.9 27.9 30.0 25.9 Head depth in HL 74.6 69.7 80.0 66.1 70.2 80.0 63.0 Head width in 11L 73.1 67.5 73.7 59.6 67.5 76.5 61.1 Body depth in SL 20.0 19.1 20.9 16.7 20.4 22.2 18.6 Body width in SL 9.8 10.4 13.1 8.5 10.7 12.8 9.4 Caud. ped. length in SL 15.7 15.7 18.2 12.0 16.2 17.7 13.7 Caud. ped. length in SL 11.8 11.3 12.6 6.7 11.3 12.2 10.7 Snout length in HL 28.4 29.0 35.1 22.7 28.4 35.2 22.9 Eye width in HL 29.9 29.6 33.3 25.0 28.8 32.4 26.1 Jaw length in HL 37.3 35.0 39.0 29.5 33.3 38.7 29.3 Interorbital in HL 7.5 6.2 8.6 4.8 6.2 8.0 4.4 Pectoral fin in SL 26.3 26.0 30.4 22.6 26.1 28.3 23.7 Pelvic fin in SL 27.5 26.9 28.7 23.4 26.0 28.6 22.9 Caudal fin in SL 34.5 32.1 36.5 28.3 29.3 32.7 25.2 3 ,J D1 fin spine in SL - 16.9 20.4 14.7 15.5 17.6 13.3 4 lh D1 fin sp in SL 20.0 18.2 20.9 14.9 15.7 17.0 13.4 arch few, short and stubby, longest raker at angle of arch; few rudimentary rakers on inner face of first arch; outer rakers on second gill arch consisting of low pads of pointed papillae; outer rakers on remaining arches similar to those on first arch but longer. Inner face of upper limb of first gill arch, and to lesser extent, upper limbs of other arches, covered with low dense fleshy papillae which may form clumps or groups; dorsal portion of arch may have short fleshy protuberances ending in one or several papillae. Lower quarter to one-third (usually one-third) of first gill arch bound by membrane to inner face of opercle. Tongue short, tip blunt to concave. Teeth in upper jaw in two rows across front and one row at side ofjaw, outermost row teeth largest, curved and pointed, largest teeth at front on either side of symphysis; innermost row teeth quite small, sharp and evenly sized. Teeth in lower jaw in two rows, arranged similarly to upper jaw but outer row teeth smaller (especially in females); posteriormost outer row tooth may be slightly recurved. Predorsal scales smaller than body scales, cycloid, reaching on side of nape to variable extent, from just over opercle to nearly behind eyes; midline always naked. Ctenoid scales on side of body extending up to pectoral fin base. Opercle and preopercular area without scales. Breast naked (usually) or with small patch of embedded cycloid scales before pelvic fins (2-6 scales). Pectoral fin base naked. Belly scales usually cycloid; midline may be naked; ctenoid scales may be present posteriorly on midline in some specimens. Head pores with anterior nasal pore just anterior to anterior nostril tube, posterior nasal pore beside each posterior nostril, a pair of anterior interorbital pores, a single posterior interorbital pore, a postorbital pore, a terminal pore over opercle and an anterior and posterior temporal pore in short separate posterior portion of the oculoscapular canal over opercle; three preopercular pores present (Fig. 2A). Sensory papillae arranged in sparse transverse pattern (Fig. 2A); vertical row ot on opercle may be broken or continuous. Papillae on chin in two, short, longitudinal rows on either side of low bump (Fig. 2B). Coloration of fresh material. Randall and Greenfield (2001: pi. 11H) illustrated this species in colour. A photograph by Doug Floese, ofa fresh specimen from East Point (Fig. 3), shows the head and body pale pinkish orange with dark brown speckling and short vertical dark brown streaks and six rounded blackish spots along midside of body, side of head and body covered with dark-margined iridescent pale-blue oval spots, which cluster and partly coalesce around posteriormost four blackish round spots, forming ocellate dark spots. Blue spots on side of head fewer, may be larger and darker than those on body; cheek and opercle tend to be more orange than pinkish. Eye brown dorsally with dark brown to blackish marks around margin; iris pale gold, reddish dorsally; single short blockmark extends from ventral margin of eye partly onto cheek, several short black marks on cheek and near rictus. A broken line of dark brown dashes along midside of body, coinciding with midlateral round blackish spots. Unpaired fins pinkish, roughly banded with rows of white to bluish white spots; small dense black spots scattered over fins, rows of black spots present on first spine of each dorsal fin; anal fin darker pinkish brown than other fins, and oval blue (not bluish white) spots; pectoral fins with transparent membrane and rays pinkish orange, dotted with melanophores; pelvic fins brownish with white to orange mottling. The specimen in Randall’s photograph (Randall and Greenfield 2001: plate 111 I) has all the fins extended, so it can be seen that the black spots on the dorsal and caudal fins are arranged in approximate rows, and the black spots in the anal fin are coalesced into oblique streaks running dorsoposteriorly. 70 New goby from northern Australia The black marks on the cheek are joined to form an irregular blotchy line from the eye to the lower jaw. Coloration of preserved material. Head and body yellowish white, most scales with dusky margin or with brown to dark grey spot on rear margin; along mid-side of body a dark brown line broken into six sections, in males a rounded dusky to dark brown blotch around each section of brown line, posterior three round dark blotches darkest and surrounded with lighter brown pigment; in females, a faint dusky blotch or short bar around each section of mid-lateral brown line but no blotches particularly darker than others (Figs 1,4). Some specimens very pale, whitish, others show intense coloration. Dorsal midline with about 12 dusky to dark brown spots or small blotches, anteriormost blotch on nape above rear edge of opercle. Broken brownish to dark brown horizontal line from rear edge of eye extending back onto body, ending at point below third first dorsal fin spine or above pectoral fin base; broken line may be intensified over pectoral fin base into series of dark brown partly connected small spots; pectoral base with several dusky blotches or short horizontal dusky to brownish lines, which may extend partly on to base of fin. Side of head with few scattered dusky to dark brown small spots or blotches; narrow dusky to blackish vertical line crossing cheek from lower margin of eye, ending on cheek behind rictus, line may be irregular or broken into blotches; short horizontal dusky to blackish line from posterior end of upper lip extending back to fall short of, to meet, or to cross vertical cheek line, may extend back onto rear half of cheek. Dorsal margin of eye with dark brown or blackish blotch or short line, which does not extend into interorbital space. Dark brown to blackish short lines, spots or blotches around nostrils, anterior nostril tube dusky to dark brownish. Lips dusky with two to three short brown lines or spots crossing obliquely, angled toward, and fading out on, chin. Underside of head, breast and belly plain brownish or dusky. In males, first dorsal fin plain dusky to dark grey, with two to three irregular rows of oval dense black spots, three to four similar black spots evenly spaced along anterior face of first spine; oval transparent to whitish spots often scattered over fin. In females, first dorsal fin translucent to dusky grey, with four dark grey to blackish streaks or series of broken oblique dark lines; four blackish spots across anterior face of first dorsal spine; broad plain dusky grey band along distal margin of fin. Second dorsal fin, in males, similar to first dorsal fin but with narrow transparent margin to fin; dense black oval spots in three to four irregular rows, four black spots along anterior face of fin spine; oval translucent to whitish spots may be larger and more numerous than on first dorsal; fin usually darker posteriorly. Second dorsal fin in females very similar to first dorsal, but four to five rows of short dark streaks present, mostly on fin membranes; four to five dark spots on anterior face of fin spine. Anal fin in males plain dusky to dark grey with few scattered dense black spots, especially posteriorly; may be paler dusky band along fin margin. Anal fin plain dusky in females. Caudal fin translucent to dark grey (darkest in adult males), with irregular rows of small dense black to dark grey spots crossing fin; several black spots near upper and lower base of fin may be enlarged and intensely black; males with transparent oval spots on membrane, mostly toward central part of fin; fin Fig. 3. Gnatholepis argits sp. nov. Fresh male specimen (AMS 1.24677-003) from Bullocky Point reef. Darwin Harbour; photograph by Doug Hoese. Fig. 4. Gnatholepis argus sp. nov. Female paratype specimen (NTM S. 14965-008) from Field Island, Northern Territory (same locality as holotypc), showing difference in preserved colour pattern. 71 H. K. Larson and D. Buckle tends to be darkest ventrally. Pectoral and pelvic fins plain dusky to brownish grey. Distribution. Specimens are known only from northern Australia, where suitable intertidal coral/rocky reef habitat is present: from Dampier, Western Australia, across the Northern Territory to Cape Wessel, and from one locality in the Torres Strait (Prince of Wales Island). The apparent disjunct distribution is probably due to lack of collecting in suitable habitats. Ecology. This species has been collected from shallow intertidal pools (no deeper than 1 m) on rocky fringing reefs, where low corals, especially faviids, and sponges are present, with a substrate of sand to muddy rock; the surrounding waters are turbid. It can be locally abundant. Comparisons. This is a sister-species to Gnatholepis gymnocara, which appears to be restricted to the Queensland coast. Both species are small and differ from all other Gnatholepis in having a naked or mostly naked predorsal mid-line, no scales on the cheek and opercle, and all teeth in the lowerjaw are small and similar in size, with no enlarged or recurved canines. Gnatholepis argus differs from G. gymnocara in having 1,11 anal rays (versus 1,12), second dorsal rays 1,10, rarely 1,11 (versus 1,11, rarely 1.10 rays), no scales on the pectoral fin base (versus a few embedded scales), being smaller (greatest adult size 33 mm SL versus 36.6 mm SL), and in colour pattern differences: males being strongly marked, with small dense black spots on unpaired fins, and posteriormost 3-4 mid-lateral spots darker and larger than anterior spots (versus males without dense black spots on fins and no large dark spots on side of body). Etymology. In reference to the Greek god Argus, of the many watchful eyes, in reference to the eye-like blue- spotted pattern along the side of the body in males; a noun in apposition. DISCUSSION Thacker (2004) discussed the species of Gnatholepis and carried out a cladistic analysis using DNA data for what she considered to be six species, including G. argus (referred to as Gnatholepis sp. Randall and Greenfield), although she lacked DNA material of its sister species, G. gymnocara. She demonstrated that G. argus was basal to all other Gnatholepis species (Thacker 2004: fig. 2). However, the names Thacker assigned to each of the other Gnatholepis species did not coincide with those used by Randall and Greenfield (2001) and she did not examine type material of G. cauerensis or G. scapu/ostigma (while using the name G. scapu/ostigma for the species usually referred to as G. cauerensis). Material identified by Thacker (2004: 581-582, fig. 1) as G. anjerensis and G. davaoensiS from the Northern Territory are all G. argus. This is based upon direct examination, by the senior author, of the same AMS specimens that Thacker examined. Randall and Greenfield also synonymised several species of Gnatholepis without providing any reasons for doing so. For example, they placed Gobius ophthahnotaenia Bleeker as a synonym of Gnatholepis anjerensis, apparently without examination of type specimens. As a result, the two most recent reviews of Gnatholepis species are unsatisfactory, contradictory and confusing and thus in need of revision. We have commenced a revision of the genus Gnatholepis. The two basal small species, G. gymnocara and G. argus, appear to be the only taxa in the genus that do not require reviewing. ACKNOWLEDGMENTS Our thanks to Mark McGrouther (AMS) for rapid loans of specimens, to Doug Hoese (AMS) for drawing our attention to other material, and to Arnold Suzumoto (BPBM), Jeff Johnson (QM) and Jeff Clayton (USNM) for their very swift responses for catalogue numbers. The junior author would like to thank his partner Alicia for letting him take holidays to look at fish. REFERENCES Birdsong. R.S., Murdy, E.O. and Pezold, F.L. 1988. A study of the vertebral column and median (in osteology in gobioid fishes with comments on gobioid relationships. Bulletin of Marine Science 42(2): 174-214. Bleeker, P. 1851. Over eenige nieuwe soorten van Blcnnioi'den en Gobioi'dcn van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch-Indie 1: 236-258. Hoese, D.F. 1986. Family No. 240: Gobiidae. Pp. 774-807. In: Smith, M.M. and Heemstra, P.C. (eds) Smith’s sea fishes. Springer-Verlag: New York. Hubbs, C.L. and Lagler, K.F. 1958. Fishes of the Great Lakes Region. University of Michigan Press: Ann Arbor. Kuiter, R.H. and Tonozuka, T. 2001. Pictorial guide to Indonesian reef fishes Part 3 Jawfishes - sunfishes. Zoonetics: Melbourne. Larson. H.K. 2001. A revision of the gobiid fish genus Mugilogobius (Telcostei: Gobioidei), with discussion of its systematic placement. Records of the Western Australian Museum. Supplement No. 62: 1-233. Leviton, A.E., Gibbs, R.H., Heal, E. and Dawson, C.E. 1985. Standards in Herpetology and Ichthyology: Part I. Standard symbolic codes for institutional resource collections in I lerpetology and Ichthyology. Copeia 3: 802 -832. Nakabo, T. 2002. (ed.) Fishes of Japan with pictorial keys to the species, English edition II. Tokai University Press: Tokyo. Randall, J.E. and Greenfield, D.W. 2001. A preliminary review of the Indo-Pacific gobiid fishes of the genus Gnatholepis. Iclithyiological Bulletin 69: 1-17. Sanzo, L. 1911. Distribuzione delle papille cutanee (organi ciatiformi) e suo valore sistematico nei Gobi. Mitteilungen aus der Zoologischen Station zu Neapel 20: 249-328. Seale, A. 1910. New species of Philippine fishes. Philippine Journal of Science 4(6): 491-543. Thacker, C. 2004. Phylogeny and species boundaries in the gobiid genus Gnatholepis (Teleostei: Perciformes). Zoological Journal of the Linnean Society 142b: 573-582. Accepted 7 August 2005 72 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2005 21: 73-163 Review of the Dinematichthyini (Teleostei: Bythitidae) of the Indo-West Pacific. Part I. Diancistrus and two new genera with 26 new species WERNER SCHWARZHANS 1 , PETER RASK M0LLER 2 AND J0RGEN G. NIELSEN 2 'Ahrensburger Weg 103 D, 22359 Hamburg, GERMANY wwschwarz@aol. com 2 Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen 0, DENMARK pdrmoller@snm. ku. elk, JGNielsen@snm. ku.dk ABSTRACT An ongoing revision of the dinematichthyine fishes (Ophidiiformes, Bythitidae, Brosmophycinae) of the Indo-West Pacific based on ca. 5000 specimens will be published in parts. Part I includes 765 identified specimens in the genera Brotulinella (new genus with one new species), Diancistrus Ogilby. 1899 (with four described and 23 new species) and Paradiancistrus (new genus with two new species). The main distinguishing character in this group of presumably related genera is the male pseudoclasper pattern. Brotulinella is further characterized by the slender form and the generally higher number of precaudal vertebrae (12 versus 11 in the two other genera, except for one species with 12 in Diancistrus). Paradiancistrus is unique amongst Indo-West Pacific dinematichthyine genera in having only one instead of three lower preopercular pores. Brotulina Fowler, 1946, Calcarhrotula Fowler, 1946 and Parabrostnolus Machida, 1996, are regarded as junior synonyms of Diancistrus. The separating characters of the species are male pseudoclaspers, head squamation, head pores, otoliths, morphometric proportions, and fin ray and vertebral counts. Diancistrus . which has remained monotypic for more than 100 years, is here shown to be one of the most diverse genera of tropical reef fishes. The many unrecognised species have previously been confused with other genera such as Dinematichthys and Ogilbia. Pseudoclasper morphology is used to define three informal species groups in the genus Diancistrus. Keywords: viviparous brotulas, Indo-West Pacific, Australia, Indonesia, Micronesia, New Guinea, Philippines, Polynesia, Ryukyu Islands, coral reef fishes, Diancistrus, new genera, new species. CONTENTS INTRODUCTION.74 MATERIAL AND METHODS.75 DEFINITION OF CHARACTERS.75 COMPARATIVE MATERIAL.78 SYSTEMATICS.78 Tribe Dinematichthyini Cohen and Nielsen, 1978.78 Key to the genera of Dinematichthyini in the Indo-West Pacific.78 Brotulinella n. gen.79 Brotulinella taixvanensis n. sp.80 Diancistrus Ogilby, 1899.82 Key to the species of Diancistrus .88 Diancistrus alatus n. sp.89 Diancistrus alleni n. sp.91 Diancistrus altidorsalis n. sp.i.95 Diancistrus atollorum n. sp.97 Diancistrus beateae n. sp.100 Diancistrus brevirostris n. sp.103 Diancistrus eremitus n. sp.105 Diancistrus erythraeus (Fowler, 1946).108 Diancistrusfijiensis n. sp.110 Diancistrus fuscus (Fowler, 1946).113 73 W. Schwarzhans, P. R. Moller and J. G. Nielsen Diancistrus jackrandalli n. sp.115 Diancistrus jeffjohnsoni n. sp.117 Diancistrus karinae n. sp.119 Diancistrus katrineae n. sp. 121 Diancistrus leisi n. sp.125 Diancistrus longifilis Ogilby, 1899.126 Diancistrus mcgroutheri n. sp.129 Diancistrus machidai n. sp.131 Diancistrus manciporus n. sp.134 Diancistrus mennei n. sp.136 Diancistrus nigern. sp.138 Diancistrus novaeguineae (Machida, 1996).140 Diancistrus pohnpeiensis n. sp.143 Diancistrus robustus n. sp.146 Diancistrus springeri n. sp.147 Diancistrus tongaensis n. sp.149 Diancistrus vietnamensis n. sp.151 Diancistrus sp. 1 . 153 Diancistrus sp. 2 . 154 Diancistrus sp. 3 . 154 Paradiancistrus n. gen.154 Paradiancistrus acutirostris n. sp. 155 Paradiancistrus cuyoensis n. sp. 157 GEOGRAPHIC DISTRIBUTION. 159 ACKNOWLEDGMENTS. 162 REFERENCES. 162 INTRODUCTION In the latest comprehensive review of ophidiiform fishes (Nielsen et a!. 1999), the dinematichthyine fishes, a tribe within the subfamily Brosmophycinae of the viviparous family Bythitidae, were represented by 12-14 genera and ca. 25 species worldwide. For distinction from the other tribe within Brosmophycinae, the Brosmophycini, see the diagnosis to the Dinematichthyini below. After the recent review of the American Dinematichthyini (Moller et al. 2004a and 2005) the number now stands at 13 genera and 52 species. Following the ongoing review of the Indo-West Pacific Dinematichthyini currently under way, the number of species is likely to double. Some species may reach 100-120 mm in length, a few even 150 mm, others not more than 25-30 mm. When newly born, dinematichthyine fishes are 10-15 mm long. They live in shallow tropical to subtropical waters, hidden in holes and crevices of coral reefs, algae beds and rocky shores. Different reefal environments, such as reef drop-offs, surge channels, lagoons or tide pools can be dominated by different groups of the Dinematichthyini. The species of the genera Diancistrus and Dinematichthys for instance are typical of the main reef bodies, are less common in lagoons and rare in tide pools or rocky shores. The geographical distribution of most dinematichthyine species is very restricted, but there are also notable exceptions of wide distribution patterns (see for instance D. alleni n. sp., D. beateae n. sp. or D. katrineae n. sp. and section Geographic Distribution). Dinematichthyine species show relatively little variation in their general appearance, and their live coloration tends to be uniform, the prevailing colours being yellow, orange, red and brown. Moller et al. (2004a) used the apomorphic position of the copulatory organ below a covering fleshy hood in a cavity of the ventral body wall as the main diagnostic character for the bythitid tribe Dinematichthyini, thereby confirming Sedor's (1985) view. In all other viviparous bythitids (Brosmophycini and Bythitinae), the copulatory organ is an integrated part of the fleshy genital hood, with the penis present as a small, soft papilla. Pseudoclasper-like structures, if present, are indistinct papillae with or without a slight sclerification in the form of a superficial thickening of the epidermis. Until the introduction of scuba-diving and the use of fish poison, dinematichthyine fishes were rarely caught, but with these methods the catches have exploded. Today, many museums hold several thousand specimens of dinematichthyine fishes. Considering how common they are, dinematichthyine fishes must play an important role in the ecology of reefs. Due to the large amount of material with many new taxa we have decided to deal with the dinematichthyine fishes of the Indo-West Pacific in four volumes: the future 74 Dinematichthyine fishes of the Indo-West Pacific three parts will revise the genera Beaglichthys, Brosmolus, Monothrix and include several new genera, revise the genera Dermatopsis, Dermatopsoides and Dipulus and finally revise the most common genus Dinematichthys sensu lato. Prior to this review, Diancistrus was a monotypic genus, known only from the type species D. longifilis. It was described in 1899 by Ogilby based on a single type specimen caught at the isolated Lord Howe Island in the Tasman Sea. Since then, it has been recorded only as secondary citations in check lists or systematic reviews. The type specimen was reviewed and refigured for the first time 100 years later by Nielsen el al. (1999). Specimens caught off Lord I lowe Island were often identified as D. longifilis in various collections. A second dinematichthyine fish occurs at Lord Howe Island belonging to the genus Dinematichthys senstt lato, with which it was commonly confused. MATERIAL AND METHODS Examination of ca. 5000 specimens of Indo-West Pacific Dinematichthyini yielded 835 identified to the genera treated herein. Of these, 765 were identified to the species level whereas the remaining 70, mostly juvenile or female specimens, could only be identified to the genus level. The 765 identified specimens include a small number of specimens that are only tentatively assigned to species. Also included are specimens viewed and identified in the collections of the AMS and the USNM but not borrowed for detailed investigations. These are listed as additional specimens and are not referred to as type specimens for any of the new species. Despite the very large number of Indo-West Pacific specimens investigated, we expect that still more new species and more specimens of rare species will be found in collections or when regions now under-represented in collections are sampled for dinematichthyines. As for the latter we particularly expect more undescribed species from Indonesia, the Nicobar and Andaman Islands and the Indian subcontinent. The material and comparative material belongs to the following institutions: AMS (Australian Museum, Sydney), ANSP (Academy of Natural Sciences, Philadelphia), ASIZP (Academia Sinica Institute of Zoology, Taipei), BPBM (Bernice P. Bishop Museum, Honolulu), CAS (California Academy of Sciences, San Francisco), LACM (Los Angeles County Museum), MCZ (Museum of Comparative Zoology, Harvard Museum), MNHN (Museum Nationale d’Histoire Naturelle, Paris) NHM (= BMNH; The Natural History Museum, London), NMNZ (Museum of New Zealand Te Papa Tongarewa, Wellington), NSMT (National Science Museum, Tokyo), NMV (National Museum of Victoria, Melbourne), NTM (Museum and Art Gallery of the Northern Territory, Darwin), QM (Queensland Museum, Brisbane), ROM (Royal Ontario Museum, Toronto), SA1AB (South African Institute for Aquatic Biodiversity, formerly RUSI (JLB Smith Institute of Ichthyology), Grahamstown), SAM (South African Museum, Cape Town), SIO (Scripps Institute of Oceanography, La Jolla), SMNS (Staatliches Museum fur Naturkunde, Stuttgart), TAU (Tel Aviv University), UMMZ (University of Michigan, Museum of Zoology), USNM (United States National Museum, Washington). WAM (Western Australian Museum, Perth), YCM (Yokosuka City Museum), ZMUC (Zoological Museum, University of Copenhagen). Morphometric characters are given as percent of standard length (SL) throughout. In the descriptions holotype values are given first, followed by the range in paratypes in brackets. Size of eye is measured as horizontal diameter of pigmented eyeball. Meristic counts were made from radiographs, except for pectoral fin rays, gill rakers, teeth and scale rows. Abbreviations used in meristic counts are: D/V = anterior dorsal fin ray above vertebra number; D/A = anterior anal fin ray below dorsal fin ray number; V/A = anterior anal fin ray below vertebrae number. Otoliths were removed through the gill cavity by making a small cut above the gills on the right side. Size of body scales was measured on holotypes at mid-body above anal fin origin. Pseudoclaspers were observed by bending forward the fleshy hood covering the copulatory organ and thereafter by bending outwards the pseudoclaspers or spreading them and fixing them with a thin needle. It was not necessary to dissect pseudoclaspers for morphological analysis. In drawings, the pseudoclaspers and penis are shaded; other parts, such as the fleshy hood, isthmus or outline of the copulatory cavity are simple line drawings. In order to avoid excessive redundancy, the descriptions of the individual species of the genus Diancistrus concentrate on those characters depicting significant variance or difference from the general character states found within this genus. For further correlation of individual species, reference is made to Table 2 (pseudoclasper patterns, scales on operculum, eye size, total number of vertebrae), Table 3 (dorsal fin ray counts) and Table 4 (selected morphometric measurements). The ecology of most of the species is poorly known. From station data we have gathered information about habitat and depth range, but we have no, or very little, data about behaviour, live coloration and feeding. A number of females were examined for reproductive data, e.g., number and size of embryos. The distribution maps were created using Microsoft Encarta 2001 digital world atlas. DEFINITION OF CHARACTERS Male pseudoclaspers are the most important characters for separating the species. Juveniles and females can be difficult to identify for some species, even though the distribution pattern deduced from male occurrences in 75 W. Schwarzhans, P. R. Mnller and J. G. Nielsen many instances will allow at least a tentative specific assignment of females. Due to the somewhat inconsistent and confusing nomenclature and terminology of characters in the dinematichthyine literature, the more important characters have been re-defined in Moller el al. (2004a) and are here briefly summarized as follows: Head pore system (Fig. 1). The following pore rows are recognized: supraorbital row holds 3-4 pores; the posteriormost was previously termed the first lateral line pore (e.g. Machida 1994; Cohen and McCosker 1998). It is located above the opercular spine at the upper termination of the gill opening and is part of the supraorbital row. The infraorbital row holds 3 anterior and 2-3 posterior pores. The mandibular row holds 3 anterior and 3 posterior pores. The preopercular row generally holds 3 lower and 1 upper pore, but the number is reduced in some species and genera. The first and second lower preopercular pores are often placed in a common pore-like cavity. The latter two rows form a continuous system and are often referred to in literature as the preoperculo-mandibular pore row. Male copulatory organs (Fig. 2) are covered anteriorly by a thick fleshy genital hood that originates from the posterior margin of the anus. The organs consist of a penis flanked by one or two pairs of pseudoclaspers (three solely in Ogilbichthys Moller el al., 2004a, a genus from American waters) on each side, and a sometimes indistinct, accessory organ at the base of the penis. The pseudoclaspers may contain hardened, sclerified bodies that keep them in an upright position. These are anterior infraorbital pores 2 3 Fig. 1 . Head pore system terminology. Open pores shown as solid circles and pores hidden from lateral view as dashed circles. A, lateral view of head; B, ventral view of head. here termed supporters. A fleshy isthmus separates the pairs of pseudoclaspers, originating from the hood and joining the base of the penis. A ligament that passes around the posterior base of the penis connects the outer pseudoclaspers (o.p) distally. The inner pseudoclasper (i.p.) is free or joined to the outer pseudoclasper and/or to the isthmus between the pseudoclaspers by a ligament. For a detailed description of the reproductive ecology and functional anatomy of the copulatory apparatus see Turner (1946) and Suarez (1975). Pseudoclaspers are best studied in preserved fish, since the hood has been found to be very stiff in freshly dead specimens. In this paper one of the pseudoclaspers is shown usually from the left inside. Additionally, the entire copulatory apparatus is illustrated with the fleshy hood bent forward from an inclined lateral view, showing one pair of pseudoclaspers from the outside, the other from the inside. A schematic sketch of a ventral view with (or sometimes without) the hood bent forward is sometimes added, because at the first inspection of the organ this often gives valuable identification hints in the case of the genera reviewed here. The presence of one or two pairs of pseudoclaspers is, with very few exceptions, used for definition of genera, while details of the pseudoclasper morphology are used for species definition. In fact, there are only few instances where specific differences rely exclusively on other characters. The genera treated herein all have two pairs of pseudoclaspers, with the inner pseudoclasper placed at the anterior margin of the outer pseudoclasper. The inner pseudoclasper in these three genera is mostly much smaller than the outer pseudoclasper, often joined to it and sometimes atrophied to the extent of the complete loss of a supporter. Examination of the pseudoclaspers of more than 2000 mature male dinematichthyines has revealed that its morphology is very stable, both in terms of variability and ontogenetic changes (the latter after maturity has been reached). Within the Indo-West Pacific Dinematichthyini, limited allometric growth has been observed in few species, for instance of the genus Diancistrus. Sagittal otolith (Fig. 3). Terms of the sagitta follow Schwarzhans (1993). The sagittae (‘otolith’ in the following) of dinematichthyine fishes show little ontogenetic and intraspecific variation in contrast to certain other ophidiiform fishes. Sexual dimorphism of otoliths has been observed in several ophidiiform fishes (Schwarzhans 1994), but is found mostly negligible in the case of the Dinematichthyini. For a comprehensive review ofophidiiform otolith morphologies see Nolf (1980) and Schwarzhans (1981). The most easily recognizable character in dinematichthyine otoliths is the status of the sulcus, i.e. divided (separated colliculi) or undivided (fused colliculi) and the form of the ventral margin of the sulcus between the two. These characters are often used as additional characters for generic diagnoses. For instance 76 Dinematichthyine fishes of the Indo-West Pacific Fig. 2. Male copulatory organ terminology. A, ventral view of male copulatory organ ( Diancistrus karinae n. sp.); B-C, E, view of left pseudoclasper front inside, B, Diancistrus karinae n. sp., C, Diancistrus altidorsalis n. sp., E, Diancistrus beateae n. sp.); D, inclined lateral view of male copulatory organ (Diancistrus karinae n. sp.). Diancistrus can always be distinguished readily from Dinematichthys by the status of the eolliculi (fused versus separated). However, fusion of colliculi is a very common trend in Dinematichthyini and other ophidiiform fishes and has apparently occurred in multiple parallel lineages; i.e. there are several other dinematichthyine genera not closely related to Diancistrus with fused colliculi. In the case of Diancistrus the use of otolith morphology is restricted to selected cases for differentiation of the many species. Selected additional characters (general morphology) of important diagnostical value are: Head squamation usually requires removal of mucus, which often covers the head. Only a few species of Dinematichthys s.l. from the Indo-West Pacific show complete squamation of the head. In most other Dinematichthyini head squamation is restricted to the cheek and the operculum above (and sometimes also below) the opercular spine, or is missing entirely from the head. Such differences are used in generic as well as in species definitions. In the case of Diancistrus a thin skin may in addition cover those scales above and below the opercular spine. Their occurrence and number are often of important dorsal dorsal Fig. 3. Otolith (sagitta) terminology. Median view of right otolith: A, Dinematichthys ; B, Diancistrus. 77 W. Schwarzhans, P. R. Mnller and J. G. Nielsen diagnostic value in this genus. When the skin is removed and the scales are uncovered, they come loose and can then easily fall off, although scale pockets will give an indication that scales were there originally. Therefore, whenever possible, one side of a fish investigated for head squamation was left untouched. The anterior nostril is normally placed close to the tip of the snout in bythitids, but in Dinematichthys s.l. and a few related, yet undescribed Indo-West Pacific genera it is positioned mid-way between snout and posterior nostril, i.e., high above the upper lip. However, in the case of Diancistrus the anterior nostril is often positioned intermediate between that observed in Dinematichthys and the remainder of the Dinematichthyini and thus causes this character here to be less reliable than usual. Absolute body size of mature males and females is variable in the case of the genus Diancistrus, which contains species with mature males at a size of 30 mm SL that do not exceed 60 mm SL (for instance D. alatus and D. mcgroutheri) and others, which are only mature at sizes >55 mm SL and reach up to over 120 mm SL (D. beateae ). COMPARATIVE MATERIAL Indo-West Pacific Dinematichthyini. Beaglichthys macrophthalmus Machida, 1993: Holotype - NTM S.10395-001-1, non-type: WAM P. 28155-019. Brosmolus longicaudus Machida, 1993: Holotype-NTM S.10623-001. Dennatopsis macrodon Ogilby, 1896: Holotype - AMS I. 3505; non-types: WAM P. 27112-005 (3 specimens). Dennatopsis multiradiatus McCulloch and Waite, 1918: Paratype - AMS 1. 14614. Dennatopsoides kasougae (Smith, 1943): Holotype - RUS1 333, only radiograph examined; non-type - RUSI 8547. Dennatopsoides talboti Cohen, 1966: Holotype - RUSI 340, only radiograph examined; Paratypes - RUSI 339, only radiograph examined; non-types -SAM 21800; SAM 21963. Dipulus caecus Waite, 1905: non-types - WAM P. 29884-012; WAM P. 29886-007; ZMUC P 77485-87 (3 specimens). Dipulus norfolkanus Machida, 1993: non-types - NMNZ 11706; NMNZ 11742; NMNZ 27162 (3 specimens); NMNZ 27166 (4 specimens). Dinematichthys dasyrhynchus Cohen and Hutchins, 1982: Paratypes - AMS I. 20245-016 (3 specimens); non-types- WAM P. 27950-011 (2 specimens), WAM P. 27951-007, ZMUC P 77716-18. Dinematichthys iluocoeteoides Bleeker, 1855: BMNH 1862.2.28.65. Dinematichthys indicus Machida, 1994: Paratypes -ROM 37813-2 (14 specimens), ROM 58269 (8 specimens). Dinematichthys megasoma Machida, 1994: non-types - WAM P. 30308-001 (7 specimens), WAM P. 30909- 002 (9 specimens). Dinematichthys mizolepis Gunther, 1867: Holotype - BMNH 1867.5.13.17. Dinematichthys rytikyuensis Aoyagi, 1954: Holotype - YCM-P 30001(1); non-types -USNM 297347 (12 specimens), ROM 71850(5 specimens). Monotlirixpolylepis Ogilby, 1897: Holotype -AMS 1.3654. American Dinematichthyini. See Moller etal. 2004a and 2005. Brosmophycinae and Bythitinae. See Moller et al 2004b. SYSTEMATICS Tribe Dinematichthyini Cohen and Nielsen, 1978 (family Bythitidae Gill, 1861; subfamily Brosmophycinae Gill, 1862) Diagnosis. Male copulatory organ with a penis and 1-2 (rarely 3) pairs of pseudoclaspers in cavity of ventral body wall covered by a fleshy hood. First anal fin pterygiophore slightly to strongly elongated. Head pore system generally unreduced, 6 mandibular, 2-4 preopercular, 5-7 infraorbital and 3-4 supraorbital pores, including supraorbital pore above opercular spine. Posteriormost supraorbital head pore tubular. Comparison. The Dinematichthyini are best defined by the apomorphic position of the copulatory organ below a covering fleshy hood in a cavity of the ventral body wall (Sedor 1985; Moller et ah 2004a). In all other viviparous bythitids (Brosmophycini and Bythitinae), the copulatory organ is an integrated part of the fleshy genital hood, with the penis present as a small, soft papilla. Pseudoclasper- like structures, if present, are indistinct papillae with or without a slight sclerification in the form of a superficial thickening of the epidermis. The 3rd supraorbital head pore above the opercular spine is only known from one other bythitid genus outside the Dinematichthyini from Brosmophycis Gill, 1861. In Brosmophycis this pore is slit-like, not tubular as in the Dinematichthyini. Key to Indo-West Pacific genera of the Dinematichthyini la. Maxilla not vertically expanded postventrally; head without scales; lower lip with folded papillae; upper preopercular pore absent; one pair of (outer) pseudoclaspers; otolith with separate colliculi.2 lb. Maxilla vertically expanded postventrally; head with scales patches, at least on cheek, but commonly also on operculum above and below opercular spine; lower lip without folded papillae; upper preopercular pore present or absent; two (rarely one) pairs of pseudoclaspers; otolith with fused or separate colliculi. 4 2a. Opercular spine covered by skin, not exposed. . Dennatopsoides 2b. Opercular spine free, exposed.3 3a. Precaudal vertebrae 11-14; dorsal fin rays <85. . Dennatopsis 3b. Precaudal vertebrae 14-23; dorsal fin rays >85. . Dipulus 4a. Precaudal vertebrae 13-15; average number of dorsal fin rays per ray-bearing vertebra 2.4-2.7; 1 (rarely two) pairs of pseudoclaspers.5 78 Dinematichthyine fishes of the Indo-West Pacific 4b. Precaudal vertebrae 11-12 ( Dinematichthys dasyrhynchus 14); average number of dorsal fin rays per ray-bearing vertebra 1.9-2.3; two pairs of pseudoclaspcrs.8 5a. Upper preopercular pore present.6 5b. Upper preopercular pore absent.7 6a. One pair of (outer) pseudoclaspers; squamation only on cheek; one pair of supraorbital pores on occiput; canine teeth present; otolith with fused colliculi.... . Monothrix 6b. Two pairs of pseudoclaspers, the inner being stick-like and extending centrally over the outer pseudoclasper; squamation on head continuous across cheek and operculum; five pairs of supraorbital pores on occiput; no canine teeth; otolith with separate colliculi. . Dinematichthys dasyrhynchus 7a. One pair of (outer) pseudoclaspers; caudal vertebrae >42. Bros molus 7b. Two pairs of (centrally joined) pseudoclaspers; caudal vertebrae <38. Beaglichthys 8a. Squamation on head continuous across cheek and operculum. Dinematichthys (part) 8b. Squamation on head divided in separate patches on cheek and operculum.9 9a. Anterior nostril high on snout; otolith with separate colliculi; upper preopercular pore absent or present . Dinematichthys (part) 9b. Anterior nostril low on snout; otolith with fused colliculi; upper preopercular pore present (except absent in Diancistrus manciporus) . 10 10a. One lower preopercular pore; inner pseudoclasper joined anteriorly to outer pseudoclasper to form a u- shaped feature; otolith length to sulcus length usually >2.4. Paradiancistrus n. gen. 10b. Three lower preopercular pores (two of which are joined in a single opening); inner pseudoclasper joined anteriorly in various ways but not forming a u-shaped feature; otolith length to sulcus length usually <2.4.11 11a. Body robust, moderately slender to deep-necked (head height >15% SL except for Diancistrus jeffjohnsoni, body depth at anal >16% SL except for Diancistrus longi/ilis ); precaudal vertebrae 11 (except 12 in Diancistrus jeffjohnsoni)', maxilla with angular posteroventral widening close to its termination; body scales (1.2) 1.3-2.2% SL. Diancistrus lib. Body slender (head height <15% SL, depth at anal <16% SL); precaudal vertebrae predominantly 12 (rarely 11); maxilla rounded posteroventrally with weak knob in front of rear corner; body scales small, <1.2% SL. Brotulinella n. gen. Brotulinella n. gen. (Tables 2-4) Type species: Brotulinella taiwanensis n. sp. (type locality: southern tip of Taiwan Island, 21°N, 120°E). Diagnosis. Anterior nostril placed low on snout; male copulatory organ with two pairs of pseudoclaspers, the outer large, broad and blade-like, the inner a mere fleshy appendix fused to the outer pseudoclasper anteriorly and without a supporter; eyes moderately large (1.8- 2.6% SL); fishes small (maximum 60 mm SL, maturity reached at 35 mm SL), slender (head height <16% SL, depth at anal <16.5% SL); precaudal vertebrae 12 (rarely 11); head with scale patch only on cheek; otolith elongate, its sulcus inclined (5-10°), short (otolith length to sulcus length 2.2-2.4), with fused colliculi; maxilla rounded postero¬ ventrally with weak knob in front of rear corner; anterior anal fin pterygiophore long. Comparison. Brotulinella belongs among the dinematichthyine genera characterized by a combination of a low anterior nostril, two pairs of pseudoclaspers, a maxillary knob at the rear corner and an otolith with fused colliculi. Of the Indo-West Pacific genera so far described, Dinematichthys s.l. is distinguished by the high anterior nostril and an otolith with separate colliculi. The interrelated genera Dermatopsis, Dermatopsoides and Dipu/us differ in the absence of a vertically extended posterior maxilla, the lack of scales on the head, a single pair of pseudoclaspers and an otolith with separate colliculi. The remaining genera in the Indo-West Pacific share the four characters mentioned for Brotulinella above, except for the genera Brosmolus and Monothrix, which have a single pair of pseudoclaspers. Both genera, and Beaglichthys, further differ from Brotulinella in the higher number of precaudal vertebrae (13-15 versus 12). Brotulinella differs from Diancistrus and Paradiancistrus n. gen. in its slender shape (head height <16% SL versus >15% SL and body depth at anal <16.5% SL versus >16% SL) and the predominantly 12 precaudal vertebrae (versus 11, except 12 in Diancistrus jeffjohnsoni). Paradiancistrus n. gen. is further distinguished by the presence of only one lower preopercular pore (versus 3). The broad, blade¬ like outer pseudoclasper with the inner pseudoclasper anteriorly joined as a fleshy flap without a supporter further distinguishes Brotulinella from Diancistrus and Paradiancistrus n. gen. Of the American Dinematichthyini, Ogilbia, Pseudogilbia and Ogilbichthys are closest to Brotulinella. In these three genera, however, the inner pseudoclasper is free from the outer pseudoclasper and always contains a supporter. Otoliths of Ogilbia and some species of Ogilbichthys have separated colliculi, while some other species of Ogilbichthys and Pseudogilbia have fused colliculi like Brotulinella. Ogilbichthys is also readily recognized by the presence of two pairs of inner pseudoclaspers, a unique character in Dinematichthyini. Pseudogilbia finally shows a very different pseudoclasper arrangement (though with two pairs) and, like Paradiancistrus n. gen. only one lower preopercular pore. Species. The genus is monolypic. 79 W. Schvvarzhans, P. R. Moller and J. G. Nielsen Etymology. The name refers to the genus Brotulina, now regarded as a junior synonym of Diancistrus (see below; type-species; Brotulina fusca, now Diancistrus fuscus known from the northern Philippines, Taiwan and the Ryukyu Islands) and the small size of the fishes of this genus. Gender: feminine. Brotulinella taiwanensis n. sp. (Figs 4-6; Table 1) Material examined. (61 specimens, 23-60 mm SL). Holotype - USNM 221048, male, 43 mm SL, Taiwan, 21°N, 120°E (collector, method and date unknown). Paratypes- ANSP 163510,2 females, 35-49 mm SL and 1 juvenile, 31 mm SL, 20°24’N, 121°55’E, Batanes Province, white beach past Mahatae, Batan Island, surge channel at outer edge of reef flat, walls of channel rock, 0-6.5 m depth, Philippines, G.D. Johnson and W.F. Smith-Vaniz, 22 April 1987; ASIZP 59304, I male, 43 mm SL. Taiwan; BPBM 23339, 1 male, 36 mm SL and 1 female, 38 mm SL, Clf uan-Fan-Shih, Taiwan, J.E. Randall, G.W. Tribble and R.P.H. Rutherford, 16 July 1978; USNM 318065, 1 male, 36.5 mm SL, Batan Island, Batanes Province, Philippines, 18°N, 121°E; USNM 366695, 1 male, 34 mm SL and 1 female, 49 mm SL, SW tip of Taiwan, V.G. Springer, 5 May 1968; USNM 374176, 2 males, 37-40 mm SL, 1 female, 50 mm SL and 1 juvenile 23 mm SL, Babuyan Islands, Maybag Island, middle of S coast of island, along shore, Philippines, C.A. Ross and V. Samarita, 8 March 1990; USNM 374178, 1 male, 38 mm SL and 4 females, 32-60 mm SL, 20°25’N, 121°57'E, Batan Island, Batanes Province, Philippines, G.D. Johnson eta!., 22 April 1987; USNM 374188, 1 male, 51 mm SL and 4 females, 31-45 mm SL, S shore of Taiwan, just S of cut between large outstanding rock and Ch’uan-Fan-Shih, Taiwan, V.G. Springer et al., 24 April 1968; USNM 374189, 1 male, 51 mm SL, rocky headland NW of swimming beach of Sha Toa, Taiwan, J.H. Choat dal.. 5 May 1968; USNM 384606, 1 female, 43 mm SL, Taiwan, same location as holotype; ZMUC P 771468, 1 male, 52 mm SL, same data as USNM 374189; ZMUC P 771469, 1 female, 32 mm SL, same data as USNM 374178. Additional specimens. USNM 221051, 2 females, 25 and 53 mm SL, Taiwan; USNM 263711, 3 males and 8 females, 30-54 mm SL, Taiwan, 21°55'N, 120°49’E; USNM 263726, 1 male and 1 female, 25-34 mm SL, Taiwan; USNM 300087,3 females, 38-45 mm SL, Batanes Table 1. Meristic and morphometric characters of Brotulinella taiwanensis n. sp. Holotype „ . . USNM J 7 M0ty t pe 221048 +27 P arat yP es Mean (range) n Standard length in mm 43 41.0 (23-60) 28 Meristic characters Dorsal fin rays 78 73.3 (68-78) 26 Anal fin rays 57 56.7 (52-63) 26 Pectoral fin rays 17 18.8(17-21) 25 Precaudal vertebrae 12 11.7(11-12) 26 Caudal vertebrae 31 30.0 (29-31) 26 Total vertebrae 43 41.7 (40-43) 26 Rakers on anterior gill arch 12 14.0(11-17) 26 Pseudobranchial filaments 2 2 26 D/V 7 6.5 (6-7) 26 D/A 20 21.2(18-24) 26 V/A 15 14.2(12-15) 26 Morphometric characters in % of SL Head length 22.4 24.9 (22.4-27.0) 28 Head width 10.5 11.2 (9.5-13.0) 27 Head height 12.9 14.3 (12.9-15.9) 27 Upper jaw length 11.0 12.1 (10.9-13.3) 27 Maxillary height 3.3 3.8 (3.2-4.5) 27 Diameter of pigmented eye 2.0 2.2 (1.8-2.9)* 28 Interorbital width 5.4 5.8 (4.S-7.2) 27 Postorbital length 16.3 18.0(16.3-19.2) 27 Preanal length 49.3 48.1 (42.4-51.5) 27 Predorsal length 30.2 31.1 (28.4-33.2) 27 Body depth at origin of anal fin 13.8 14.5 (12.9-16.6) 27 Pectoral fin length 14.4 14.9(13.3-17.3) 27 Pelvic fin length broken 20.5 (16.9-25.3) 23 Base pelvic fin - anal fin origin 33.3 30.1 (25.0-34.3) 26 * Strong negative allometric change. Province, Philippines, 20°27’N, 121°57’E; USNM 300093, 2 females, 27 and 39 mm SL, Batanes Province, Philippines, 20°20’N, I21°49’E; USNM 300096, 1 female, 39 mm SL, Batanes Province, Philippines,20°25’N, I2I°56’E; USNM 366493, 2 males, 34-36 mm SL and 1 female, 30 mm SL, Taiwan, 2I°55’N, 120°49’E; USNM 366564, I male,40mm SL and 3 females, 35-54 mm SL, Taiwan: USNM 366565, 2 males and 2 females, 33-39 mm SL, Taiwan; USNM 367146, 1 male, 30 mm SL, Taiwan. Diagnosis. See generic diagnosis. Description. The principal meristic and morphometric characters are shown in Table 1. Body elongate, slender, head height <16% SL and body depth at anal <16.5% Fig. 4. Brotulinella taiwanensis n. sp. Holotype. USNM 221048, male, 43 mm SL, 80 Dinematichthyine fishes of the Indo-West Pacific SL. Eyes 1.8-2.9% SL. Head with scale patch on cheek containing up to 5 vertical rows of small scales. Maxillaries ending far behind eyes, dorsal margin of maxillaries covered by dermal lobe of upper lip, posterior end rounded, with weak knob on posterior ventral part but anterior to rear corner. Anterior nostril positioned low, 1 /4 the distance from tip of snout to anterior margin of eye. Posterior nostril large, about 1/3 the size of eye. Opercular spine pointed, with free tip. Anterior gill arch with 11-17 rakers, including 3 elongate rakers in the angle. Pseudobranchial filaments 2. Scales on body embedded in skin, horizontal diameter 1.1% SL. Predorsal length 28-33% SL, anal fin origin just before mid-body, distance pelvic fin base to anal fin base mostly 25-34% SL. Pectoral fins reaching halfway between verticals through anal and dorsal fin origins, 13-17% SL. pelvic short, 17-25% SL. Head sensory pores (Fig. 5 A-B). Supraorbital pores 3: 1st pore in front of 2nd anterior infraorbital pore, 2nd pore indistinct, above and behind eye, 3rd pore tubular, at upper termination of gill opening above opercular spine. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore behind anterior nostril, 2nd and 3rd anterior pores covered by dermal flap of upper lip, three posterior pores on rear part of upper lip. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large and tubular, with a single cirrus anteriorly, 2nd pore positioned in lateral skin fold, 3rd anterior at anterior termination of jugular isthmus, three posterior pores on rear part of lower jaw. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint opening; upper pore tubular. Lateral line configuration indistinct. Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 row of larger teeth anteriorly, blending into a few rows of granular teeth posteriorly. Anterior-most teeth length up to 1/2 diameter of pupil. Vomer horseshoe¬ shaped, with one row of large fangs up to 'A diameter of pupil. Palatine teeth in 2 rows, slightly larger in inner row. Dentary with 3 outer rows of granular teeth and 1 inner row of larger teeth anteriorly, blending into 1 row of 8 large fangs posteriorly, up to about 2/3 of pupil diameter. Otolith (Fig. 5 G-H). Elongate in shape, length to height about 2.2 (36-43 mm SL). Anterior tip pointed; posterior tip more robust. Dorsal rim straight, with obtuse predorsal and prominent postdorsal angle; ventral rim shallow and gently curved. Inner face slightly convex; outer face smooth, flat. Otolith length to sulcus length 2.2 to 2.4. Sulcus with undivided margin and fused colliculi. Sulcus inclined to otolith axis at 5° to 10°. Ventral furrow distinct, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, except for neural spines of vertebrae 3-4 (5) inclined and (5) 6-8 depressed, shorter in length than spines of 2nd-4th (5th) vertebrae. Bases of neural spines 5-9 enlarged. Parapophyses present from vertebra 6 to 11. Pleural ribs on vertebrae 2-11 (10), absent on last precaudal vertebra in all but one specimen. Epipleural ribs indistinct. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis in males and just below in females. Male copulatory organ (Fig. 5 C-F). Two pairs of pseudoclaspers; outer pseudoclasper large, broad, blade- Fig. 5. Brotulinella taiwanensis n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male copulatory organ, USNM 318065; D, ventral view of left pseudoclasper, USNM 318065; E, view of left pseudoclasper from inside, holotype; F, inclined lateral view of male copulatory organ, holotype; G, median view of right otolith, USNM 318065; H, ventral view of right otolith, USNM 318065. 81 W. Schwarzhans, P. R. Moller and J. G. Nielsen shaped with a broad supporter. Inner pseudoclasper a long fleshy flap anteriorly joined to the outer pseudoclasper, almost reduced to an appendix without supporter. Penis not much longer than pseudoclaspers, curved, broad at basis. Colour in alcohol. Live colour unknown. Light brown, head and upper 1/3 of body darker than flanks. Comparison. See comparison between Brotulinella and other genera. Distribution. Known from southern Taiwan and northern Philippines, chiefly Batanes Province (Fig. 6). 110"E 120°E 130“E 140”E Fig. 6. Sample sites of Brotulinella taiwanensis n. sp. One symbol represents several samples. Ecology. The species occurs in shallow water between rocks, mainly in surge channels. Etymology. The name taiwanensis refers to the type locality, Taiwan, Republic of China. Diancistrus Ogilby, 1899 (Tables 2-4) Diancistrus Ogilby, 1899: 743 (type species D. longifilis Ogilby, 1899 by monotypy) (see Eschmeyer 1998). Diancistrus - Cohen and Nielsen, 1978: 57; Paxton et al. 1989: 316; Nielsen etal. 1999: 128. Brotulina Fowler, 1946: 195, figs 59-60 (type species B. fusca Fowler, 1946 by monotypy), ‘new synonym’. Brotulina - Cohen and Nielsen, 1978: 56; Nielsen et al. 1999: 125. Calcarbrotula Fowler, 1946: 193, figs 57-58 (type species C. erythraea Fowler, 1946 by monotypy). Parabrosmolus Machida, 1996: 147, figs 1-4 (type species P. novaeguineae Machida, 1996 by monotypy), ‘new synonym’. Parabrosmolus - Nielsen et al. 1999: 124. Diagnosis. Anterior nostril placed closer to upper lip than to posterior nostril; tip of opercular spine free; male copulatory organ with two pairs of mostly highly diverse and specific pseudoclaspers, the outer pseudoclasper being the larger and more diverse; inner pseudoclasper small, positioned near anterior edge of outer pseudoclasper and joined to it in various degrees, rarely free (in D. alatus n. sp., D. megroutheri n. sp. and D. tongaensis n. sp.); otoliths with fused colliculi; otolith length to sulcus length <2.2; sulcus not incl ined; anterior anal fin ray pterygiophore long; lower preopercular pores 3 (2 lower and 1 upper); upper preopercular pore present (except absent in D. manciponis n. sp.); ventral maxillary knob at rear corner; precaudal vertebrae usually 11 (12 in D. jeffjohnsoni n. sp.); body robust, moderately slender to high-necked (head height >15% SL in adults, except for D. jeffjohnsoni n. sp. and depth at anal >16% SL except for D. longifilis ); dentary with large fang-like teeth posteriorly, of about 1/3 to 2/3 of pupil width. Description. Vertebrae 11-12+28-35 = 39-46, dorsal fin rays 66-91, anal fin rays 54-74, pectoral fin rays 16-21. ventral fin rays 2, caudal fin rays 15-17 (almost always 16), D/V = 5-7, D/A = 17-24, V/A = 12-15. Body robust, often deep-necked, head slender or massive, moderately to strongly compressed. Body covered with small scales, head with scale patch on cheek and many species also with scales on operculum. Maxillary ending far behind-eyes, dorsal margin covered by dermal lobe of upper lip, expanded posteriorly, angular or with knob at ventral rear corner. Anterior nostril placed low on snout, close to upper lip. Tip of opercular spine free, pointed. Anterior gill arch with 12-18 rakers, thereof 2-4 (usually 3) elongate rakers in an uninterrupted row at the angle. Pseudobranchial filaments 0-2 (almost always 2). Pectoral fin length 13-20% SL; pelvic fin length mostly 20-30% SL. Dorsal fin originates at about 1/3 from body length, predorsal mostly at 30-37% SL; anal fin origin at about mid-body, preanal mostly 43-53% SL; distance of pelvic fin base to anal fin base 21-32% SL. Caudal fin free, rounded. Positive allometric growth sometimes observed in body depth (depth at anal fin origin) and head length. Head pore system includes: Supraorbital pores 3: 1st at tip of snout, 2nd above and behind eye and 3rd above opercular spine; infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore behind anterior nostril, 2nd and 3rd anterior pores covered by dermal flap of upper lip, three posterior pores on rear part of upper lip; mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore at tip of lower jaw, 2nd anterior pore positioned in lateral skin fold, 3rd anterior at anterior termination of jugular isthmus, three posterior pores on rear part of lower jaw: preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint openings, 3rd pore just above, upper preopercular pore behind and above check scale patch (absent in D. manciponis n. sp.). Teeth pointed and granular, present on palatine, vomer, premaxilla and dentary. Lateral line system often indistinct. When observable, consisting of short dorsal row from behind opercular opening to vertical line through anal fin origin and 82 Dinematichthyine fishes of the Indo-West Pacific Table 2. Comparison of key characters of species of the genera Brotulinella n. gen, Diancistrus and Paradiancistrus n. gen. Diancistrus species groups refer to: I a = Diancistrus altidorsalis subgroup, lb = Diancistrus erythraeus subgroup, 2a = Diancistrus longifilis subgroup, 2b = Diancistrus springeri subgroup, 3a = Diancistrus fijiensis subgroup, 3b = Diancistrus katrineae subgroup. pseudoclaspers scales on operculum eye size vertebrae D. altidorsalis n. sp. D. niger n. sp. D. atollorum n. sp. D. erythraeus D. mcgroutheri n. sp. D. robustus n. sp. D. tongaensis n. sp. D. sp.3 rZ o 'c75 CD C. C — C <D S/J C ~ OJ. o — C -g'E. V C/3 CO X C.£- ■ Q. £ “ 6 CS-O C0T3 c c 3 co ~ zj ■ ^ C3 n d co co r- u. p u. CJ CO O CO v - X X X (x) X X (X) X X (X) X (X) X X (x) X (x) X X X X X X D. alleni n. sp. 2 a X X X (X) X D. beateae n. sp. 2 a X X X X D. fuscus 2 a X X X X X D. longifilis 2 a X (X) X (X) X (X) 1 X D. machidai n. sp. 2 a X (X) X X X X X D. manciporus n. sp. 2 a X X X X X D. novaeguineae 2 a X (X) X X X X X D. vietnamensis n. sp. 2 a X X X X (X) X D. leisi n. sp. 2 b X X X X D. springeri n. sp. 2 b X X X D. sp. 1 3 X X X D. alatus n. sp. 3a X X X X D. brevirostris n. sp. 3a X X X X X D. fijiensis n. sp. 3a X X (X) X X X X D.jackrandalli n. sp. 3a X X X X D. eremitus n. sp. 3b X X X X D. karinae n. sp. 3b X X (X) X (X) D.jeffjohnsoni n. sp. 3b X X X X X D. mennei n. sp. 3b X X X D. pohnpeiensis n. sp. 3b X X X D. katrineae n. sp. 3b X X X X D. sp. 2 3b X X X Paradiancistrus acu tiros Iris n. sp. X X X X Paradiancistrus cuyoensis n. sp. X X X X X Brotulinella taiwanensis n. sp. X X (X) X (X) X X X lo ^r 0> CO aj > lo o vo OJ CO 1) > X X X X X X X X X X X X X X X (x) X X X X X (x) X X 83 W. Schwarzhans, P. R. Mollerand J. G. Nielsen Table3. Frequency distribution of dorsal fin ray counts of the species of the genera Brotulinella n. gen., Diancistrus and Paradiancistrus n. gen. Diancistrus species groups refer to: la = Diancistrus altidorsalis subgroup, lb = Diancistrus erythraeus subgroup, 2a = Diancistrus longifilis subgroup, 2b = Diancistrus springeri subgroup, 3a = Diancistrus fijiensis subgroup, 3b = Diancistrus katrineae subgroup. o- =3 O CD C /3 .— O 1) D- C/J £2 C $ Q D. altidorsalis n. sp. la D. niger n. sp. la D. atollorum n. sp. lb D. erythraeus lb D. mcgroutheri n. sp. lb D. robustus n. sp. lb D. tongaensis n. sp. lb D. sp. 3 lb dorsal finrays 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 1 3 5 8 3 3 4 1 - 3 1 - 1 - - 2 2 1 4 5 3 4 2 1114 112 4 6 12 11 1 1334433321 1-1 D. alleni n. sp. 2 a D. beateae n. sp. 2 a D.fuscus 2 a D. longifilis 2 a D. machidai n. sp. 2 a D. manciporus n. sp. 2 a D. novaeguineae 2 a n. sp. D. vietnamensis 2 a n. sp. D. leisi n. sp. 2 b D. springeri n. sp. 2 b D. sp. 1 2 262 10 11 7834 1 77864 6 34 111444424111 241576722 2-1211 111-11 2-2132841-11 1 2 3 4 2 3 -11 11-3- 1 1 1 - - - - 1 1 D. alatus n. sp. 3a D. hrevirostris n. sp. 3a D. fijiensis n. sp. 3a D.jackrandalli n. sp. 3a D. eremitus n. sp. 3b D. karinae n. sp. 3b D.jefjjohnsoni n. sp. 3b D. mennei n. sp. 3b D. pohnpeiensis ,, n. sp. D. katrineae n. sp. 3b D. sp. 2 3b Paradiancistrus acutirostris n. sp. Paradiancistrus cuyoensis n. sp. Brotulinella taiwanensis n. sp. 1 - (1) - - 3 2 2 112 2 15 - - 1 1 -13 4 1 4 6 2 2 1 - 5 4 2 2 1 - 1 1 3 3 - 1 - 1 2 1 3 1 1 1 - 1 2 3 4 3 1 12 2 1 3 1 1 2 1 12 3 2 84 Dinematichthyine fishes of the Indo-West Pacific Table 4. Comparison of selected morphometric characters of the species of the genera Brotulinella n. gen., Diancistrus and Paradiancistrus n. gen. Diancistrus species groups refer to: la = Diancistrus altidorsalis subgroup, lb = Diancistrus erythraeus subgroup, 2a = Diancistrus longifilis subgroup, 2b = Diancistrusspringeri subgroup, 3a = Diancistrusfijiensis subgroup, 3b = Diancistrus katrineae subgroup. Diancistrus species group Head length Head height Depth at anal fin origin Body scales <25% SL 25 - 26% SL 26 - 27% SL 27 - 28% SL 28 - 29% SL >29% SL <14% SL 14 - 15% SL 15 - 16% SL 16 - 18% SL 18 - 20% SL 20 - 22% SL 22 - 24% SL >24% SL <14% SL 14 - 16% SL 16 - 18% SL 18 - 20% SL 20 - 22% SL >22% SL <1.2% SL 1.2- 1.4% SL 1.5- 1.6% SL 1.7 - 1.9% SL D. altidorsalis n. sp. la X X X X X X X X X X D. niger n. sp. la X X X X X X X D. atollorum n. sp. ib X X X X (X) X X D. erythraeus lb X X X X (X) X X X D. megroutheri n. sp. lb X X X X X X X X D. robustus n. sp. lb X X X X D. tongaensis n. sp. lb X X X X X X X D. sp.3 lb X X X X X D. alleni n. sp. 2a X X X X X X X X X X D. beateae n. sp. 2a (X) X X X X X X X X X X D.fuscus 2a X X (X) X X X X X X X D. longifilis 2a X X X X X X X X X D. machidai n. sp. 2 a X X X X X X X X X X X D. manciporus n. sp. 2a X X X X X X D. novaeguineae n. sp. 2a X X X X X X X X X X D. vietnamensis n. sp. 2a (x) X X X X X X X X X X D. leisi n. sp. 2b X X X X X X X X D. springeri n. sp. 2b X X X X (X) X X X D. sp. 1 2 X X X X X X D. alattts n. sp. 3a X X X X D. brevirostris n. sp. 3a X X X X X X X (X) X X X D. fijiensis n. sp. 3a X X X X X X (X) X X D.jackrandalli n. sp. 3a X X X X X D. eremitus n. sp. 3b X X X X X X X X X X D. karinae n. sp. 3b X X X X X X X X X D.jeffjohnsoni n. sp. 3b X X X X X X X X X D. mennei n. sp. 3b X X (X) X X X X D. pohnpeiensis n. sp. 3b X X X X X X D. katrineae n. sp. 3b X X X X X X X D. sp. 2 3b X X X Paradiancistrus acutirostr is n. sp X X X X Paradiancistrus cuyoensis n. sp. X X X X X X Brotulinella taiwanensis n sp. X X (X) X X X x X X S? as A 85 W. Schwarzhans, P. R. Moller and J. G. Nielsen mediolateral row from vertical line through anal fin origin to caudal fin origin. Otolith thin, moderately elongate to elongate. Sulcus with fused colliculi and no notch at ventral sulcus margin; otolith length to sulcus length <2.4; sulcus not inclined. Shape of otolith characterized by usually deep ventral rim and marked concavities at dorsal rim, one in front of predorsal, the other behind postdorsal angle. Anterior tip usually pointed; posterior tip pointed, but usually broader than anterior tip. Neural and haemal spines slender, except for neural spines of vertebrae 4 (and 5) inclined and (5) 6- 8 (9) depressed, shorter in length than spines of 2nd-4th vertebrae. Bases of neural spines 5-9 enlarged. Parapophyses present from vertebra (6) 7 to 10 (II). Pleural ribs on vertebrae 2-11 (10), present or absent on last precaudal vertebra. Epipleural ribs indistinct. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis or just below it, usually longer in males than in females. Male copulatory organ with two pairs of mostly highly diverse and specific pseudoclaspers (see section below). Live coloration uniformly yellow, greenish, brownish or reddish, without marks, but unknown for most species. Preserved colour usually light to dark brown. Comparison. Diancistrus is a common and the most speciose genus in the West Pacific. It is often confused with Dinematichthys, but pseudoclaspers of Diancistrus are generally much larger compared to Dinematichthys, so that in most instances both genera are easily distinguished. In Diancistrus the outer pseudoclasper is expanded in a wing, hook or ear-lobe shaped manner and the inner pseudoclasper is attached to the outer pseudoclasper anteriorly, whereas in Dinematichthys the outer and inner pseudoclaspers arc simple flap- or stick-like structures with the inner pseudoclasper inserted at about the middle of the base of the outer pseudoclasper. Another character, often found in keys to distinguish Dinematichthys from other Dinematichthyini, the high position of the anterior nostril, is not as reliable in respect to Diancistrus, which often shows intermediate positions of the anterior nostril, somewhat higher above the snout than usual in non-Dinematichthys genera. A very reliable and recommendable character to distinguish both genera in the case of females is the otoliths: those of Dinematichthys s.l. always have clearly separated colliculi, whereas those of Diancistrus always have completely fused colliculi. Also, the teeth in Diancistrus are generally longer than those found in Dinematichthys as observed already by Ogilby (1899: 730). The genera Dermatopsis, Dermatopsoides and Dipulus are readily distinguished from the other Indo-West Pacific genera so far described by the absence of a vertically expanded posterior maxilla, the lack of scales on the head, the single pair of pseudoclaspers and the otolith with separate colliculi. Beaglichthys, Brosmolus and Monothrix differ from Diancistrus (and Bratulinella and Paradiancistrus n. gen.) in the higher number of precaudal vertebrae (13-15 versus 11-12). Brosmolus and Monothrix have a single pair of pseudoclaspers. In respect to the differences of Diancistrus from Brotulinella see above. Paradiancistrus n. gen. is distinguished from Diancistrus by the presence of only 1 lower preopercular pore (versus 3) and the curved feature of the inner pseudoclasper (Figs. 72, 74). Remarks. Diancistrus was recognized as a valid genus in Cohen and Nielsen (1978) as well as Nielsen etal. (1999), but as monolypic and restricted to the Lord Howe Island in the South-West Pacific. Brotulina was regarded as valid from the Ryukyu Islands, while Calcarbrotula, established by Fowler from a single female from the Ryukyus, was considered a junior synonym of Brotulina. The monotypic Parabrosmolus was also regarded as valid and placed in the Brosmophycini following Machida’s (1996) statement of its lacking ossified elements in the male intromittant organ. As extensively discussed in Moller et al. (2004a), this character is no longer regarded as valid for distinguishing Brosmophycini and Dinematichthyini. Instead, the latter is now defined by their male copulatory organ having a penis and 1-2 (rarely 3) pairs of pseudoclaspers in a cavity of the ventral body wall covered by a fleshy hood (see above). A review of all involved type specimens clearly shows that all four nominal genera can conveniently be placed in a single genus, for which the hitherto little used genus name Diancistrus has priority. The three male holotypes of the type species of Diancistrus, Brotulina and Parabrosmolus exhibit pseudoclaspers with a hook- or stick-shaped outer pseudoclasper characteristic for one of the main species groups of Diancistrus (see below). For Calcarbrotula erythraea, which was based on a single female, males are now well documented in the material available and it clearly belongs to a second species group with ear-lobe shaped outer pseudoclaspers (see below). In fact, with the description of 23 new species the genus Diancistrus now comprises a total of 27 species and thus forms the most species-rich genus of the Dinematichthyini. Other very speciose genera are Ogilbia with 18 species (see Moller et al. 2005) from American waters and Dinematichthys s.l. with many undescribed species (under review). In abundance of specimens, the species of the genus Diancistrus are second in the Indo-West Pacific, surpassed only by those of the genus Dinematichthys s.l. Distribution. Diancistrus is widespread in the Indo- West Pacific from the Chagos Islands in the western Indian Ocean to Pitcairn and Ducie Atoll in the central Pacific. Only Dinematichthys s.l. is more widespread, reaching the eastern Indian Ocean and in being represented by a single species in the Caribbean. The high diversity observed in Diancistrus clearly has to do with geographical endemics in many areas and the (common) presence of several species per region (three to five species of Diancistrus occurring together is not rare), but there are also notable exceptions 86 Dinematichthyine fishes of the Indo-West Pacific of a few widespread species (for instance D. alleni n. sp., D. beateae n. sp. or D. katrineae n. sp.). Ecology. The species of Diancistrus are typically reef dwelling, most commonly associated with live coral reefs. They are less commonly observed in lagoonal environments. The only species and record from a non¬ reef related cave environment is Diancistrus katrineae n. sp. from Rapa, Tubuai Islands. Diancistrus specimens are generally caught in association with species of the genus Dinematichthys, which then usually constitute the larger proportion in terms of specimens with ratios ranging from about 3:1 to about 10:1. In various locations up to three, sometimes four, different species of Diancistrus have been caught simultaneously. This is the highest density of dinematichthyine species per genus observed at any location. Species groups. Diancistrus is the most speciose genus of the Dinematichthyini with 27 species described in the following and three further species left in open nomenclature because of lack of adequate material. The body proportions and sizes show more variation than found in most other dinematichthyine genera and so do the meristic values, except for the prccaudal vertebrae count, which is amazingly constant. The most universally useful character in Diancistrus is, like for all other dinematichthyine genera, the highly diagnostic and diverse male copulatory organ, except for a pair of species (D. longifilis - D. manciporus n. sp.) and a trio of species (D. fuscus - D. vietnamensis n. sp. - D. alleni n. sp.) with similar pseudoclaspers. Another very useful character to distinguish species of the genus Diancistrus is the head squamation, particularly the presence, number and orientation of scales on the operculum. Their exact evaluation, however, requires a certain treatment (see chapter ‘Definition of Characters’ above). The species of the genus Diancistrus can be arranged into species groups according to the pseudoclasper organization and the head squamation as follows: Pseudoclasper species grouping. 1) Diancistrus altidorsalis n. sp. / erythraeus species group: outer pseudoclasper ear-lobe shaped (Fig. 2C). This group can be subdivided into the Diancistrus altidorsalis n. sp. subgroup characterized by atrophied inner pseudoclaspers without supporter and large eyes (>2.5% SL) containing D. altidorsalis n. sp. and D. niger n. sp., and the Diancistrus erythraeus subgroup characterized by an inner pseudoclasper with supporter and small eyes (<2.0% SL) consisting of D. atollorum n. sp„ D. erythraeus, D. megroutheri n. sp., D. robustus n. sp. and D. tongaensis n. sp. 2) Diancistrus longifilis /springeri n. sp. species group: outer pseudoclasper hook- or stick- like (Fig. 2E). This group contains two subgroups, the Diancistrus longifilis subgroup with hook-like outer pseudoclaspers with D. alleni n. sp., D. beateae n. sp., D. fuscus, D. longifilis, D. machidai n. sp„ D. manciporus n. sp., D. novaeguineae and D. vietnamensis n. sp., and the Diancistrus springeri n. sp. subgroup with the species D. leisi n. sp. and D. springeri n. sp. 3) Diancistrusfijiensis n. sp. / katrineae n. sp. species group: outer pseudoclasper flat, wing-shaped (Fig. 2B). This group can be subdivided into the Diancistrus fijiensis n. sp. subgroup characterized by a bifurcate inner pseudoclasper comprising the species D. alatus n. sp., D. brevirostris n. sp„ D. fijiensis n. sp. and D. jackrandalli n. sp., and a second Diancistrus katrineae n. sp. subgroup characterized by a single-tipped inner pseudoclasper, which often extends anteriorly over the outer pseudoclasper containing D. eremitus n. sp., D. jeffjohnsoni n. sp., D. karinae n. sp., D. katrineae n. sp., D. mennei n. sp. and D. pohnpeiensis n. sp. Head squamation grouping. 1) Species with a continuous scale patch on the operculum extending above and below the opercular spine: D. beateae n. sp. and D. karinae n. sp. 2) Species with two separate scale patches on the operculum, one above and one below the opercular spine: D. altidorsalis n. sp., D. katrineae n. sp. and D. novaeguineae (large specimens only). 3) Species with scales on the operculum only above the opercular spine. This group can be subdivided into one subgroup with >4 scales above the opercular spine arranged in 2 to 3 rows comprising D. mennei n. sp., D. eremitus n. sp., D. fuscus, D. novaeguineae (most specimens), D. springeri n. sp. and D. vietnamensis n. sp., and another subgroup with <4 scales above the opercular spine arranged in a single row containing D. alleni n. sp., D. jackrandalli n. sp., D. jeffjohnsoni n. sp., D. machidai n. sp. (in part) and D. pohnpeiensis n. sp. 4) Species without scales on the operculum. This is the largest group containing D. alatus n. sp., D. atollorum n. sp., D. brevirostris n. sp.. D. erythraeus, D. fijiensis n. sp., D. leisi n. sp., D. longifilis, D. machidai n. sp. (in part), D. megroutheri n. sp., D. manciporus n. sp., D. niger n. sp., D. robustus n. sp. and D. tongaensis n. sp. Further research and confirmation is required to judge whether the pseudoclasper or head squamation groupings or a combination thereof are monophyletic. Therefore, in the following the species are described in alphabetical order and without grouping. Comparison to other species is restricted to species within a specific species group, defined by pseudoclasper morphology (see above) and sympatric species. The following descriptions of Diancistrus species takes into consideration all characters and methods investigated, but nevertheless makes significant use of pseudoclasper morphology. Obviously, without pseudoclasper information certain species can not definitely be distinguished. In these species females may be assigned to species when collected together with males of the respective species at the same location. Where this is not the case, such females will have to remain unidentified. 87 W. Schwarzhans, P. R. Muller and J. G. Nielsen Key to the species of Diancistrus la. Outer pseudoclasper ear-lobe shaped; no scales on operculum (except scales present on operculum in D. altidorsalis) .2 lb. Outer pseudoclasper hook, stick or wing shaped (the latter sometimes with concave inner face superficially resembling ear-lobe form); scales on operculum present or absent.8 2a. Eyes large (>2.5% SL); innerpseudoclasperatrophied without supporter.3 2b. Eyes small (<2.0% SL); inner pseudoclasper with supporter. 4 3a. Total vertebrae 43-45; scales present on operculum above opercular spine (6-9) and in separate patch below opercular spine (1-3); body scales >1.5% SL; colour light .. Diancistrus altidorsalis n. sp. 3b. Total vertebrae 41-42 (rarely 43); no scales on operculum; body scales <1.5% SL; colour black.... ..... Diancistrus niger n. sp. 4a. Total number of vertebrae 43; head massive (width >16% SL). Diancistrus robustus n. sp. 4b. Total number of vertebrae 39-42; head slender (width <16% SL) .5 5a. Inner pseudoclasper joined anteriorly to outer pseudoclasper; scale patch on lower cheek in 2 rows. 6 5b. Inner pseudoclasper free from anterior part of outer pseudoclasper; scale patch on lower cheek in 3-4 rows.7 6a. Inner pseudoclasper a small hook, ear-lobe shaped extension of outer pseudoclasper short and half-moon shaped; otolith length to height ratio 2.0; body scales <1.5% SL . Diancistrus erythraeus n. sp. 6b. Inner pseudoclasper forming a forward-inclined ear¬ lobe shaped extension, ear-lobe shaped extension of outer pseudoclasper complete; otolith length to height ratio >2.1; body scales >1.9% SL... . Diancistrus atollorum n. sp. 7a. Inner pseudoclasper forming a sharp, forward- inclined thorn or lobe, ear-lobe shaped extension of outer pseudoclasper short, confined to rear half.... . Diancistrus mcgroutheri n. sp. 7b. Inner pseudoclasper forming a broad, distalIy expanded flap, ear-lobe shaped extension of outer pseudoclasper well developed along entire length . . Diancistrus tongaensis n. sp. 8a. Outer pseudoclasper hook- or stick-shaped.9 8b. Outer pseudoclasper wing-shaped (Fig. 2B).20 9a. Outerpseudoclasperstick-shaped; inner pseudoclasper thin, almost as long as outer pseudoclasper. 10 9b. Outer pseudoclasper hook-shaped (Fig. 2E); inner pseudoclasper much smaller than outer pseudoclasper, usually about half the length.11 10a. Scale patch (7-8 scales) above opercular spine; eyes large (size >2.8% SL); total vertebrae 44-45; dorsal fin rays 78-84; body scales >1.7% SL. . Diancistrus springeri n. sp. 10b. No scales on operculum; eye small (size <2.0% SL); total vertebrae 41; dorsal fin rays 69-73; body scales <1.5% SL. Diancistrus leisi n. sp. 11a. Operculum with scales above and below opercular spine.12 1 lb. Operculum with scales only above opercular spine or without scales.13 12a. Continuous squamation on operculum above and below opercular spine; dorsal fin rays 84-91; anal fin rays 65-74; outer pseudoclasper broad, hook-shaped; inner pseudoclasper stout, fleshy, thick; body scales >1.7% SL. Diancistrus beateae n. sp. 12b. Squamation on operculum interrupted above and below opercular spine; dorsal fin rays 74-85; anal fin rays 60-69; outer pseudoclasper slender, hook- shaped; inner pseudoclasper stout, with broad fleshy appendix; body scales <1.5% SL...... Diancistrus novaeguineae [large specimens only] 13a. Scales present above opercular spine.14 13b. No scales above opercular spine (except rarely in D. machidai n. sp.).18 14a. Two or three rows of scales above opercular spine with >3 scales.15 14b. Single row with 1-2 (rarely 3) scales above opercular spine.17 15a. Three rows of mostly >7 (rarely 6) scales above opercular spine; inner pseudoclasper stout, with broad fleshy appendix .. . Diancistrus novaeguineae [most specimens] 15b. Two rows of scales above opercular spine with 3-7 scales; inner pseudoclasper stout, thick, with small fleshy appendix.16 16a. Dorsal fin rays usually >77 (74-85); otolith length to sulcus length <2.0, sulcus not inclined; dorsal head profile not elevated. Diancistrus fuscits 16b. Dorsal fin rays usually <77 (69-80); otolith length to sulcus length >2.3, sulcus inclined; dorsal head profile elevated (Figs. 66, 67A)... . Diancistrus vietnamensis n. sp. 17a. Dorsal fin rays 66-77; total vertebrae 40-42 (rare 43); inner pseudoclasper narrow, pointed; hook-like extension of outer pseudoclasper straight; body scales <1.5% SL. Diancistrus machidai n. sp. 17b. Dorsal fin rays 75-84; total vertebrae 43-45; inner pseudoclasper stout, fleshy; hook-like extension of outer pseudoclasper bent outwards; body scales >1.7% SL. Diancistrus alleni n. sp. 18a. Dorsal fin rays <77 (66-77); inner pseudoclasper narrow, pointed; hook-like extension of outer pseudoclasper about twice the length of anteriorly pointed inner pseudoclasper [specimens can have one or two scales above opercular spine]. . Diancistrus machidai n. sp. 18b. Dorsal fin rays>77 (77-85); inner pseudoclasper wide, concave; hook-like extension of outer pseudoclasper less than twice the length of inner pseudoclasper.. . 19 Dinematichthyine fishes of the lndo-West Pacific 19a. Upper preopercular pore present . . Diancistrus longifilis n. sp. 19b. Upper preopercular pore absent (Fig. 50A). . Diancistrus manciporus n. sp. 20a. Inner pseudoclasper free, broad-based, bifurcate, usually not extending over anterior part of outer pseudoclasper.21 20b. Inner pseudoclasper joined to outer pseudoclasper anteriorly, narrow-based, thin, with single pointed tip, usually extending over anterior part of outer pseudoclasper.24 21a. Two scales above opercular spine; eye small (<2.1% SL); total vertebrae 40-41; inner pseudoclasper short, without spines; body scales <1.5% SL . . Diancistrus jackrandalli n. sp. 21b. No scales above opercular spine; eyes large ( >2.1% SL); total vertebrae 42-44; inner pseudoclasper with long bifurcate portions; body scales >1.7% SL... 22 22a. Anterior branch of inner pseudoclasper twice as long as posterior branch, both with rounded tip . . Diancistrus alatus n. sp. 22b. Both branches of inner pseudoclasper sharp and of about equal length .23 23a. Branches of inner pseudoclasper broad based, flap¬ like. Diancistrus brevirostris n. sp. 23b. Branches of inner pseudoclasper thin, thorn-like... . Diancistrusfijiensis n. sp. 24a. Scales on operculum present above and below opercular spine; body scales >2.0% SL.25 24b. Scales on operculum present only above opercular spine; body scales <1.7% SL.26 25a. Continuous squamation on operculum above and below opercular spine; head slender; outer pseudoclasper llat. Diancistrus karinae n. sp. 25b. Squamation on operculum interrupted, above opercular spine 13-21 scales, below opercular spine 3-7 scales; head massive; outer pseudoclasper with concave inner face. Diancistrus katrineae n. sp. 26a. Precaudal vertebrae 12 (rarely 11); dorsal fin rays 70- 77; anal fin rays 55-63; single scale above opercular spine; outer pseudoclasper flat; otolith with regularly curved dorsal rim .. Diancistrus jeffjohnsoni n. sp. 26b. Precaudal vertebrae 11; dorsal fin rays >77; anal fin rays >62; number of scales above opercular spine >1; otoliths of males with postdorsal angle.27 27a. Scale patch above opercular spine with 3 rows and 7-9 scales; outer pseudoclasper flat, short (about 1 1/2 the length of inner pseudoclasper) . . Diancistrus mennei n. sp. 27b. Scale patch above opercular spine with 1-2 rows and 2-5 scales; outer pseudoclasper with concave inner face...28 28a. Scales above opercular spine 4-5 in 2 rows; total vertebrae 42-44; D/A 22-24; outer pseudoclasper without fleshy bulge posteriorly. . Diancistrus eremitus n. sp. 28b. Scales above opercular spine 2-3 in 1 row; total vertebrae 45-46; D/A 20-22; outer pseudoclasper with fleshy bulge posteriorly. . Diancistrus polinpeiensis n. sp. Diancistrus alatus n. sp. (Figs 7, 8, 29; Table 5) Material examined. (1 specimen, 32 mm SL). Holotype - USNM 372953, male, 32 mm SL, 19°16’S, 174°22’W, small reefs between Ha’apai and Vava’u Island groups, Tonga, caves and surge channels into reef front, rubble at base of reef, 2-7 m, J. T. Williams et al., 10 Nov. 1993. Diagnosis. Body slender, small; vertebrae 11+31 =42, dorsal fin rays 79, anal fin rays 59; outer pseudoclasper large, wing-shaped, bent outwards, with flat, broad supporter; inner pseudoclasper free, broad-based, bifurcate, its anterior branch twice as long as posterior branch extending over anterior part of outer pseudoclasper, both with rounded tip; scale patch only on cheek, no scales on operculum; otolith length to sulcus length 2.2. Description. The principal meristic and morphometric characters are shown in Table 5. Body and head profile slender, fish small, mature at 32 mm SL. Head with narrow scale patch on cheek containing up to 4 vertical rows of small scales. Horizontal diameter of scales on body about Table 5. Meristic and morphometric characters of Diancistrus alatus n. sp. Holotype USNM 372953 Standard length in mm 32 Meristic characters Dorsal fin rays 79 Anal fin rays 59 Pectoral fin rays 20 Precaudal vertebrae 11 Caudal vertebrae 31 Total vertebrae 42 Rakers on anterior gill arch 17 Pseudobranchial filaments - D/V 6 D/A 19 V/A 13 Morphometric characters in % of SL Head length 27.2 Head width 9.4 Head height 17.2 Upper jaw length 12.8 Maxillary height - Diameter of pigmented eye 2.5 Interorbital width 4.1 Postorbital length 18.1 Preanal length 49.4 Predorsal length 32.8 Body depth at origin of anal fin 15.0 Pectoral fin length 14.4 Pelvic tin length 23.8 Base pelvic fin - anal fin origin 28.8 89 W. Schwarzhans, P. R. M 0 ller and J. G. Nielsen 1.7% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner and knob in front of rear angle. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril small, the size of anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 8 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 row of larger teeth at symphysis, followed by about 4 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 1 tooth row, the posterior teeth the larger. Palatine teeth in 1 row with about 10 teeth. Dentary with 3 outer rows of granular teeth and 1 inner row of larger teeth anteriorly, blending into 1 row of 8 fangs posteriorly, fang length up to about 1/3 of pupil diameter. Otolith (Fig. 8 F-G). Moderately elongate, length to height 2.0 (32 mm SL); otolith length to sulcus length 2.2; sulcus inclined at an angle of 5°. Anterior tip of otolith pointed; posterior tip less pointed; dorsal rim with moderate concavities anteriorly and posteriorly. Axial skeleton (of holotype). Neural spine of vertebrae 4 inclined and 5-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 8 C-E). Two pairs of large pseudoclaspers. Outer pseudoclasper large, wing-shaped, bent outwards, extending well outside even in resting position; its supporter flat, broad; inner pseudoclasper free, broad-based, bifurcate, its anterior branch twice as long as posterior branch extending over anterior part of outer pseudoclasper, both with rounded tip. Isthmus between pseudoclaspers narrow. Penis long, curved, uniformly thin. Coloration. Live colour unknown. Uniformly light grey-brown when preserved. Comparison. Diancistnis alatus belongs to the group of species with wing-shaped outer pseudoclaspers and bifurcate inner pseudoclaspers, which further contains 1 cm Fig. 7. Diancistnis alatus n. sp. Holotype. USNM 372953, male, 32 mm SL. Fig. 8. Diancistnis alatus n. sp. Holotype. USNM 372953. A, lateral view of head; B, ventral view of head; C, ventral view of male copulatory organ; D, view of left pseudoclasper from inside; E, inclined lateral view of male copulatory organ; F, median view of right otolith; G, ventral view of right otolith. 90 Dinematichthyine fishes of the Indo-West Pacific the species D. brevirostris n. sp., D. fijiensis n. sp. and D. jackrandalli n. sp. Diancistrus alalus is distinguished from all of them in the anterior branch of the inner pseudoclasper being longer than the posterior branch. From D. jackrandalli n. sp. it is further distinguished by the higher number of vertebrae (42 or more versus 41 or less), the larger eye (>2.0% SL versus <2.0% SL) and the absence of scales on the operculum above the opercular spine (versus present). Compared to D. brevirostris n. sp. and D. fijiensis n. sp., D. alalus usually has a lower number of vertebrae (42 versus 42-45). In D. fijiensis n. sp. the sulcus on the otolith is more anteriorly positioned than in D. alalus and with the dorsal rim of the otolith very gently curved without distinct angles or concavities. In Tonga, Diancistrus tongaensis n. sp. and D. manciporus n. sp. occur in addition to D. alalus. Diancistrus tongaensis n. sp. belongs to the species group with ear-lobe shaped outer pseudoclaspers and small eyes and thus is readily distinguished. Diancistrus manciporus n. sp. belongs to the species group with hook shaped outer pseudoclaspers and is the only species of the genus lacking a upper preopercular pore. Diancistrus alalus probably represents the species with the largest pseudoclaspers (in comparison with the size of the fish) within the genus (extrudes extensively from below the fleshy hood in the resting position). Distribution (Fig. 29). Known from the holotype caught off the northern Tonga Islands. Etymology. Alatus (Latin = winged), referring to the unusually large, protruding wing-shaped outer pseudoclaspers. Diancistrus alleni n. sp. (Figs 9-12; Table 6) Material examined. (102 specimens, 29-70 mm SL). Holotype- AMS 1. 19108-140, male, 52 mm SL, 14°40’S, 145°28’E, Lizard Island, Queensland, Great Barrier Reef, 1-10 m, D.F. Hoese and party, 17 Nov. 1975. Paratypes -AMS IA.7-91, I female, 62 mm SL, 16°37’S, 168°09’E, Vanuatu, Epi Island, A.R. McCulloch and party; AMS 1. 18469-162, I embryo-bearing female, 44 mm SL and 1 female, 46 mm 03°0TS, 128°03’E, Ceram, Maluku, Indonesia, J.R. Paxton, 1 April 1975; AMS 1. 18739-037,4 males, 43-61 mm SL and 2 females, 47-50 mm SL, 14°42’S, 145°27’E, Lizard Island, Great Barrier Reef, J.R. Paxton and party, 21 Nov. 1975; AMS I. 19108-141, 1 female, 47 mm SL, same data as holotype; AMS 1.20779-232, 1 male, 45 mm SL, 13°05’S, 143°57’E, Cape York, N end ofTijou Reef, Great Barrier Reef, AMS and AIMS team, 22 Feb. 1979; AMS 1.22581-025,2 males, 54 56 mm SL, 15°49'S, I45°50’E, Escape Reef, back reef slope. Great Barrier Reef, Queensland, 10 14 nr depth, AMS party, 29 Oct. 1981; AMS I. 22582-078, 1 female, 54 mm SL, 15°49’S, 145°50’E, Escape Reef, outer barrier reef. Great Barrier Reef, 14 17 m depth, AMS party. 29 Oct. 1981; AMS I. 33693-007, 1 male, 51 mm SL, 11°42’S, 144°04’E, Great Table 6. Meristic and morphometric characters of Diancistrus alleni n. sp. Holotype AMS I. 19108-140 Holotype + 64 paratypes Mean (range) N Standard length in mm 52 47.3 (38-70) 64* Meristic characters Dorsal fin rays 81 79.1 (75-84) 39 Anal fin rays 66 64.5 (60-70) 40 Pectoral fin rays 19 18.9(18-20) 31 Precaudal vertebrae 11 11 42 Caudal vertebrae 33 32.9 (31-35) 40 Total vertebrae 44 43.9 (42-45) 40 Rakers on anterior gill arch 16 15.4(13-18) 41 Pseudobranchial filaments 2 2 41 D/V 7 6.4 (6-7) 39 D/A 20 19.6(16-23) 39 V/A 14 13.2(12-14) 39 Morphometric characters in % of SL Head length 25.8 27.2 (25.1-29.1) 41 Head width 11.0 13.1 (11.0-16.1) 37 Head height 19.2 18.8(15.9-22.0) 38 Upper jaw length 11.2 13.2(11.2-14.7) 41 Maxillary height 4.6 4.6 (3.8-5.4) 40 Diameter of pigmented eye 2.5 2.8 (2.4-3.5) 41 Interorbital width 5.6 5.5 (3.6-7.6) 38 Postorbital length 17.3 19.5 (17.3-23.1) 39 Preanal length 47.5 48.1 (43.1-56.6) 41 Predorsal length 31.9 33.4 (28.9-35.9) 40 Body depth at origin of anal fin 18.3 18.4(16.6-20.9) 41 Pectoral fin length 17.5 16.2 (13.7-19.7) 34 Pelvic fin length 20.6 23.3 (17.8-27.7) 32 Base pelvic fin - anal fin oriuin 30.4 26.5 (20.3-31.1) 40 * One paratype with broken tail. Barrier Reef, 11 Jan. 1993; BPBM 17545,2 females, 37-39 mm SL, Tutuila Island, American Samoa, J.E. Randall, R. C. Wass and McGuire, II May 1974; BPBM 24123, 1 male, 58 mm SL and 1 female, 59 mm SL, Tutuila Island, American Samoa, R.C. Wass, 1976-77; CAS 222743, 1 male, 70 mm SL, Bagabag Island, Papua New Guinea, S. G. Poss, D.G. Catania and party, 18 May 1987; NSMT P. 49905, 1 female, 51 mm SL, Meno Island, Lombok Strait, Indonesia, K. Matsuura, 24 July 1996; ROM 42713, 2 females, 36-51 mm SL, 09°S, 159°E, Chagos Archipelago, R. Winterbottom, Oct. 1983; ROM 78114, 1 male, 49 mm SL and I female, 34 mm SL, 05°S, 071°E, Chagos Archipelago; USNM 209549, 3 females, 36-53 mm SL, Haruku Island, point east of Tandjung Naira, 5 m depth, Maluku Province, Indonesia, V.G. Springer and M.F. Gomon, 15 Jan. 1973; USNM 222482,1 male, 65 mm SL, Tutuila Island, American Samoa, R. C. Wass; USNM 247276, 3 males, 43 -48 mm SL and 2 females, 45 mm SL, 05°21’S, 072°12’E, Salomon Atoll, Chagos Archipelago, A.R. Emery etal., 14 March 1979; USNM 362752, 13°52’S, 167°33’E, 1 male, 49 mm SL and 2 females, 47-51 mm SL, Vanua Lava, Banks Islands, Vanuatu, J.T. Williams et al., 16 May 1997; USNM 366509, 1 female, 42 mm SL, 91 W. Schwarzhans, P. R. Moller and J. G. Nielsen 07°15’S, 072°22’E, Chagos Archipelago; USNM 384597, 4 males, 33-52 mm SL and 2 females, 33-52 mm SL, 05°52’S, 110°25’E, Karimundjawa Archipelago, Java Sea, Indonesia, V.G. Springer et ai, 29 March 1974; WAM P. 27469-009, 1 male, 48 mm SL, 15°50’S, 145°50’E, Queensland, Great Barrier Reef, Escape Reef, G.R. Allen etal., 31 Oct. 1981; WAM P. 28554-003, 3 females, 44-59 mm SL. Chagos Archipelago; WAM P. 30340-010, 1 male, 60 mm SL and 1 female, 45 mm SL, 05 o ll’S, 145°50’E, Madang, Papua New Guinea, G.R. Allen, 1991; WAM P. 30844-049, I male, 29 mm SL, 11°58’S, 123°21’E, Hibernia Reef, Timor Sea, G.R. Allen, 19 Sept. 1994; WAM P. 31144-024, 2 females, 33-49 mm SL, 05°18’S, 150°08’E, Kimbe Bay, Bismarck Archipelago, G.R. Allen, 20 April 1996; WAM P. 31202-002, 1 male, 50 mm SL and 1 female, 34 mm SL, I4°15’S, 125°18’E, Bonaparte Archipelago, East Montalivet Island, Western Australia, G.R. Allen, 24 Sept. 1996; WAM P. 31437-037, 1 male, 58 mm SL and 1 female, 52 mm SL, 12°15’S, 122°58’E, Timor Sea, Ashmore Reef, Western Australia, J.B. Hutchins, I Oct. 1997; WAM P. 31438-083, 1 male, 45 mm SL and 2 females, 30-54 mm SL, 12°15’S, 122°58’E, Ashmore Reef, Timor Sea, Western Australia, J.B. Hutchins, 3 Oct. 1997; WAM P.31654-002, 1 male,51 mmSLand I female, 51 mm SL, 14°07’S, 125°16’E, Bonaparte Archipelago, Ingram Shoals, Western Australia, G.R. Allen, 20 Sept. 1998; ZMUC P 771486, I female, 52 mm SL, same data as WAM P. 28554-003; ZMUC P 771487, 1 male, 45 mm SL, same data as WAM P. 30844-049. Additional specimens. AMS I. 17094-070, 2 males, 08°S, 151°E; AMS I. 17472-056, 1 female, 56 mm SL, Vanuatu; AMS I. 19456-036, 1 male, 45 mm SL, Queensland, Great Barrier Reef, Lizard Island; AMS I. 22586-009, 1 male, 60 mm SL and 3 females, 39-45 mm SL, Queensland, Great Barrier Reef, Escape Reef; AMS I. 22611-017, 2 males, 29-51 mm SL, Queensland, Great Barrier Reef, Escape Reef; USNM 263685, 3 males and 10 females, 30-68 mm SL, 01°33’S, 144°59’E, Papua New Guinea; USNM 263747, 7 males, 38-64 mm SL, Solomon Islands, Bougainville Island; USNM 356170, 1 male, 44 mm SL, I7°03’S, 168 0 21’E, Vanuatu, Emae Island; USNM 376192, 3 females, 51-53 mm SL, 05°52’S, I10°25’E, Karimundjawa Archipelago, Java Sea, Indonesia, V.G. Springer and party, 29 March 1974; USNM BBC 1488-90, 1 male and 1 female, 40-58 mm SL, Papua New Guinea, Madang. Tentatively assigned specimens. BPBM 33057, 1 female, 45 mm SL, North Male Atoll, Maidive Islands, J.E. Randall, R.C. Anderson, Adam and Shareef, 24 March 1988. Diagnosis. Vertebrae 11+32-34=43-45, dorsal fin rays 75-84, anal fin rays 60-70; outer pseudoclasper hook¬ shaped, hook-like extension bent outwards, with massive supporter; inner pseudoclasper anteriorly connected to outer pseudoclasper, stout, fleshy; 1 or 2, rarely 3 large scales on operculum above opercular spine aligned in single row and usually covered by mucus and skin. Description. The principal meristic and morphometric characters are shown in Table 6. Body compact with blunt, fleshy snout; fishes mature at about 40 to 45 mm SL. Head with narrow scale patch on cheek containing up to 4 or 5 vertical rows of small scales on upper half; I or 2, rarely 3 large scales on operculum above opercular spine aligned in single row and usually covered by mucus and skin. Horizontal diameter of scales on body about 1.9% SL. Maxillaries expanded posteriorly with prominent knob at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril small, the size of anterior nostril and about 1/5 the size of eye. Head sensorypores (Fig. 10 A-C). Supraorbital pores 3- Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 row of larger teeth at symphysis, followed by about 5 rows of granular teeth posteriorly- Largest teeth anteriorly about 'A pupil diameter. Vomer horseshoe-shaped, with 2 tooth rows, with the larger teeth posteriorly on inner row. Palatine with 2 tooth rows, with teeth on inner row slightly larger than on outer row. Dentary with 3 outer rows of granular teeth and 1 inner row of larger teeth anteriorly, blending into 1 row of 6 very large fangs posteriorly, fang length up to about 1/1 of pupil diameter. Otolith (Fig. 10 H). Moderately elongate, length to height 1.9-2.0 (40-60 mm SL); otolith length to sulcus length 2.1-2.2; sulcus inclined at an angle of 5°. Anterior tip of otolith pointed; posterior tip abrupt. Fig. 9. Diancistrus cilleni n. sp. Holotype. AMS 1. 19108-140, male, 52 mm SL. 92 Dinematichtliyine fishes of the Indo-West Pacific Fig. 10. Diancistrus alleni n. sp. A, lateral view of head, AMS I. 18739-037, male, 61 mm SL; B, ventral view of head, AMS I. 18739-037, male, 61 mm SL; C, lateral view of head, USNM 384597, female, 53 mm SL; D, ventral view of male copulatory organ, WAM P. 31438- 083,43 mm SL; F. view of left pscudoclasper from inside, USNM 362752, 50 mm SL; F, view of left pseudoclasper from inside, USNM 247276,48 mm SL; G, inclined lateral view of male copulatory organ, USNM 362752, 50 mm SL; H. median view ot right otolith, WAM P. 30844-049, male, 45 mm SL. Axial skeleton (ofholotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10 (11). First anal fin pterygiophore elongated, close to but not reaching tip of last prccaudal parapophysis in males and shorter in females. Male copulatory organ (Fig. 10 D-G). Two pairs of large pseudoclaspers. Outer pseudoclasper large, hook¬ shaped, its tip bent outwards, with massive supporter; inner pseudoclasper anteriorly con nected to outer pseudoclasper, stout, fleshy. Isthmus between pseudoclaspers moderately wide. Penis long, curved, thin. Coloration. Live colour unknown. Uniformly light to medium brown when preserved. Comparison. Diancistrus alleni belongs to the group of species with hook- or stick-shaped outer pseudoclaspers, which further contains the species D. beateae n. sp„ D. fuscus, D. leisi n. sp., D. longifilis, D. machidai n. sp., D. manciporus n. sp., D. novaeguineae, D. springeri n. sp. and D. vietnamensis n. sp. Of these, D. beateae n. sp. is readily distinguished by the complete scale coverage on the operculum, whereas D. leisi n. sp., D. longifilis and D. manciporus n. sp. lack scales on the operculum. Diancistrus alleni is distinguished from D. longifilis, with which it may overlap in distribution along the southern reaches of the Great Barrier Reef, and D. manciporus n. sp. by the inner pseudoclasper being thick, stout and fleshy versus wide and thin and concave in shape towards the isthmus. This is important to notice, since from New Caledonia two undisputable male specimens ofD. longifilis have been studied, which were bearing a single scale above the opercular spine just like in D. alleni. Fig. 11. Diancistrus alleni n. sp. Embryo 5 mm TL from AMS 1. 18469-162, 44 mm SL. 93 W. Schwarzhans, P. R. Moller and J. G. Nielsen 70°E 80°E 90°E 100°E IIO'E 120°E 130°E 140°E 150°E 160°E 170°E 180° 170°W O D. alleni © D. longifilis © D. manciporus * D. sp.1 Fig. 12. Sample sites of Diancistrus alleni n. sp„ D. longifilis, D. manciporus n. sp. and Diancistrus sp. 1. One symbol may represent several samples. Similar observations have been made on D. machidai n. sp., which also has specimens without, or with only 1 or 2 scales, above the opercular spine. Diancistrus alleni is distinguished from D. machidai n. sp. by the higher number of dorsal fin rays (75-84 versus 66-77), the higher number of vertebrae (43-45 versus 40-43), the outer pseudoclasper with a hook-like extension bent outwards (versus straight) and the inner pseudoclasper being stout and fleshy (versus narrow and pointed). Diancistrus alleni differs from D. leisi n. sp. and D. springeri n. sp. in the latter having a straight, stick-like outer pseudoclasper. Diancistrus alleni differs from D. novaeguineae in having 1-3 scales above the opercular spine (versus large scale patch above the opercular spine containing 6 to 11 scales) and the pseudoclasper pattern composed of an outer pseudoclasper with a hook-like extension bent outwards (versus blunt hook-shaped) and the inner pseudoclasper being stout and fleshy (versus wide with a little lobe anteriorly and an extended flap posteriorly). The closest species to D. alleni are D.fuscus and D. vietnamensis n. sp., which share the same pseudoclasper pattern, but it differs from both in the smaller number of scales above the opercular spine (1-2, rarely 3 versus 3-7), although there seems to be a marginal overlap. From D. vietnamensis n. sp. it further differs in the low neck (versus high) and the usually higher number of dorsal fin rays (75-84 versus 69-80). Also in geographical distribution D. alleni (chiefly south of the Equator) seems to be disjunct in distribution from D. fuscus (northern Philippines and Ryukyus) and D. vietnamensis n. sp. (northern Vietnam and Hainan Island). Biology. A 44 mm SL female (AMS I. 18469-162) contains embryos, 4-5 mm TL, with a few pigment spots above the vent and three short rows of pigment spots posteriorly on the body (Fig. 11). Distribution (Fig. 12). Diancistrus alleni is one ol the most common and widespread species of the genus. Its distribution ranges from the Chagos and Maidive Archipelagos, where it represents the western-most occurrence of the genus, to Java, along the northern Australian coast from 14°S, 125°E southwards to the central Great Barrier Reef at 15°S, 145°E, and east to New Guinea, Solomon Islands, Vanuatu and Samoa. The occurrence in the Chagos and Maldives appears to be disjunct from the other occurrences, but neither pseudoclaspers, head squamation, meristics or any other character investigated indicate morphological differences. In the east, D. alleni occurs commonly associated with D. novaeguineae and, occasionally, D. machidai n. sp. Ecology. The species was found in coral reef habitats at 5-17 m depth. 94 Dinematichthyine fishes of the Indo-West Pacific Etymology. Named in honour of Gerald R. Allen, Perth, Western Australia, in recognition of his many contributions to the ichthyology of the West Pacific. Diancistrus altidorsalis n. sp. (Figs 13, 14, 20; Table 7) Material examined. (88 specimens, 23-102 mm SL). Holotype- WAM P. 30410-001, male, 66 mm SL, 04°35’N, 118°45’E, Malaysia, Sabah, Bodgaya Island, sand and coral bottom at 15-20 m, G.R. Allen, 1992. Paratypes - BPBM 40210, 1 female, 43 mm SL, 01°N, 124°E, Manado, Bunaken, Sulawesi, Indonesia, J.E. Randall, Severns and R.C. Pyle, 29 Oct. 1991; USNM 263697, 1 male, 63 mm SL, 2 females, 55-67 mm SL and 1 juvenile, 26 mm SL, 00°49’S, 130°56’E, Batanta Island, Papua, Indonesia, B.B. Collette eta /., 2 July 1979; USNM 263701,2 males, 49-62 mm SL and 3 females, 23-39 mm SL, 05°17’S, 122°04’E, Kabaena Island, SE Sulawesi, Indonesia V.G. Springer. 24 Feb. 1974; USNM 263703, 2 males and 1 female, 52-86 mm SL, 05°52’S, 110°24’E, Karimundjawa Archipelago, Java Sea, Indonesia; USNM 263705, 9 males, 47-70 mm SL and 13 females, 60-85 mm SL, 05°51’S, 106°35’E; Seribu Islands, Java Sea, Indonesia, V.G. Springer et al., 5 April 1974; USNM 374157, 1 male, 46 mm SL, 10°42’S, 165°50’E, Ndendo Island, Graciosa Bay, Santa Cruz Islands, Solomon Islands, J. T. Williams et al., 26 Sept. 1998; WAM P. 31213-007, 1 male, 63 mm SL, 05°10’S, I45°49’E, Madang, lagoon, Papua New Guinea, G.R. Allen and A. Jenkins, 21 Oct. 1996; WAM P. 31355-026, I male, 61 mm SL and 2 females, 63-79 mm SL, 10°46’S, I51°42’E, Irai Island, Conflict Group, Papua New Guinea, G.R. Allen, 1 Oct. 1997; WAM P. 31491-001, 1 male, 91 mm SL and 3 females, 38-95 mm SL, 00°29’S, 122°04’E, Togean Islands, Sulawesi, Indonesia, G.R. Allen, 31 Oct. 1998; WAM P. 31736-004, 1 female, 35 mm SL, U°15’S, 152°08’E, Panarairai, Louisiade Islands, Papua New Guinea, G.R. Allen, 22 June 2000; ZMUC P 771372-3, 1 male, 93 mm SL and 1 female, 61 mm SL, Bali, Indonesia (aquarium fish); ZMUC P 771450, 1 male, 69 mm SL, Bali, Indonesia (aquarium fish); ZMUC P 771457-8, 1 male, 75 mm SL and 1 female, 66 mm SL, Bali, Indonesia (aquarium fish); ZMUC P 771464-65, 1 male, 68 mm SL, I female, 59 mm SL, Bali, Indonesia (aquarium fish). Additional specimens. AMS I. 39013-013, 1 female, 67 mm SL, Santa Cruz Islands, Solomon Islands; AMS I. 39029-003, 1 male, 78 mm SL and 1 female, 54 mm SL, 09 ° 47 ’S, 167°05’E, Santa Cruz Islands, Solomon Islands; USNM 99173, 1 male, 81 mm SL, Tomahu Island, Maluku, Indonesia; USNM 263684,3 males and 8 females, 33-102 mm SL, 01°14’S, 144°22’E, Ami Island, Papua New Guinea; USNM 263706, 1 male, 67 mm SL, 01°33’S, 144°59’E, Papua New Guinea; USNM 376189, I male, 84 mm SL, 05°52’S, 11°25’E, Karimundjawa Archipelago, Java Sea, Indonesia; USNM 376215,4 males and 14 female, 32-65 mm SL, Madang, Papua New Guinea. Tabic 7. Meristic and morphometric characters of Diancistrus altidorsalis n. sp. Holotype WAM P. 30410-001 Holotype + 40 paratypes Mean (range) N Standard length in mm 66 58.8 (23-95) 41 Meristic characters Dorsal fin rays 77 79.9 (76-85) 29 Anal fin rays 61 64.2 (59-68) 29 Pectoral fin rays 18 17.7(17-19) 27 Precaudal vertebrae 11 11 31 Caudal vertebrae 32 32.6 (32-34) 31 Total vertebrae 43 43.6 (43-45) 31 Rakers on anterior gill arch 14 15.9(14-18) 30 Pseudobranchial filaments 2 2 29 D/V 6 6.4 (6-7) 29 D/A 19 19.6 (18-22) 29 V/A 13 13.1 (12-14) 29 Morphometric characters in % of SL Head length 27.1 27.7 (25.3-29.0) 30 Head width 16.1 15.1 (12.6-18.1) 30 Head height 18.5 18.6(15.7-21.7) 30 Upper jaw length 14.0 13.9(12.9-14.9) 29 Maxillary height 4.5 4.5 (3.6-5.1) 30 Diameter of pigmented eye 3.5 3.3 (2.7-4.3) 32 Interorbital width 6.7 6.8 (5.6-7.9) 28 Postorbital length 19.6 20.0(18.2-21.5) 29 Preanal length 48.2 47.6 (44.4-50.7) 30 Predorsal length 34.8 34.8 (32.3-38.5) 30 Body depth at origin of anal fin 22.6 20.5 (16.5-23.2) 30 Pectoral fin length 17.2 17.0(12.9-19.8) 29 Pelvic fin length - 23.2 (20.6-25.6) 23 Base pelvic fin - anal fin origin 27.4 26.8 (22.2-30.9) 29 Diagnosis. Vertebrae 11+32-34=43-45, dorsal fin rays 76-85, anal fin rays 59-68; eyes large (>2.5% SL); outer pseudoclasper large, broad, ear-lobe shaped; inner pseudoclasper a fleshy appendix at the anterior-inner rim ol the outer pseudoclasper, without supporter; large scale patch on operculum above opercular spine (6-9 scales) and 1 to 3 isolated scales below opercular spine; head profile strongly ‘high-necked / hump-backed’; life colour yellow. Description. The principal meristic and morphometric characters are shown in Table 7. Body compact, high- necked, highest just in front of dorsal fin origin, snout slender, often somewhat pointed; fishes large, mature at about 50 to 55 mm SL. Head with broad scale patch on cheek, two separate scale patches on operculum, one large, above opercular spine, with 6 to 9 scales in 2 to 3 rows and 1 to 3 isolated scales below opercular spine. Horizontal diameter of scales on body about 1.9% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril small, the size of anterior nostril and about I/7th to l/10th the size of eye. 95 W. Schwarzhans, P. R. Moller and J. G. Nielsen Fig. 13. Diancistrus altidorsalis n. sp. Holotype. WAM P. 30410-001, male, 66 mm SL. Fig. 14. Diancistrus altidorsalis n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male copulatory organ, WAM P. 31491-001, 91 mm SL; D. view of left pseudoclasper from inside, WAM P. 31491-001, 91 mm SL; E, view ol left pseudoclasper from inside, holotype; F, inclined lateral view of male copulatory organ, WAM P. 31491-001, 91 mm SL; G, median view of right otolith, holotype; II, ventral view of right otolith, holotype. 96 Dinematichthyine fishes of the Indo-West Pacific Head sensory pores (Fig. 14 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of liolotype). Premaxilla with 6 rows of granular teeth and 2 rows of larger teeth at symphysis, followed by about 9 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 3 rows of teeth, the posterior inner teeth enlarged. Palatine teeth in 3 rows, largest on inner row. Dentary with 4 outer rows of granular teeth and 2 inner rows of anteriorly larger teeth, blending into 1 row of 15 large fangs posteriorly, which are up to about 1/3 of pupil diameter. Otolith (Fig. 14 G-H). Moderately elongate, length to height 1.9-2.0 (35-95 mm SL); otolith length to sulcus length 2.0-2.2; sulcus slightly inclined at an angle <5°. Anterior tip of otolith pointed; posterior tip broader; rims gently curved except sometimes an obtuse postdorsal angle. Axial skeleton (of liolotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, but not reaching tip of lastprecaudal parapophysis. Male copulatory organ (Fig. 14 C-F). Two pairs of large pseudoclaspers. Outer pseudoclasper large, broad, ear-lobe shaped, with a massive supporter; inner pseudoclasper a fleshy appendix of somewhat variable shape at anterior- inner rim of the outer pseudoclasper, atrophied without supporter. Isthmus between pseudoclaspers moderately wide. Penis long, curved, with broad base and sharp tip. Coloration. Live colour bright yellow. Uniformly light brown when preserved. Comparison. Diancistrus altidorsalis belongs to the group of species with ear-lobe shaped outer pseudoclaspers. It is easily distinguished from all other species of the group in being the only one with scales on the operculum. Together with D. niger n. sp., it is further distinguished by the large eye (>2.5% SL) from the Diancistrus erythraeus subgroup with small eyes (<2.0% SL) comprising D. atollorum n. sp., D. erythraeus, D. mcgrouthen n. sp., D. robustus n. sp. and D. tongaensis n. sp. The small-eyed species always have a supporter in the inner pseudoclasper, which is lacking both in D. altidorsalis and D. niger n. sp. Diancistrus niger n. sp. is unique in the dark, almost black, colour in preservation. Diancistrus altidorsalis differs from D. niger n. sp. in the higher number of dorsal fin rays (76-85 versus 69-76), anal fin rays (59- 68 versus 52-59) and the less elongate otolith (otolith length to height ratio 1.9-2.0 versus 2.2-23). Diancistrus altidorsalis further differs from the other specimens of the group by the high number of vertebrae (43 and more), shared only by D. robustus n. sp. Furthermore it seems to be the largest species in this group. Diancistrus altidorsalis is a very characteristic species of the genus Diancistrus occurring over a large area and (partly) co-occurring with a variety of other species such as D. alleni, D. beateae n. sp., D. karinae n. sp., D. machidai n. sp., D. novaeguineae and D. springeri n. sp., all of which do not share the ear-lobe shaped outer pseudoclasper. Diancistrus beateae n. sp. and D. karinae n. sp. differ in the complete coverage of the operculum with scales above and below the opercular spine, whereas D. alleni and D. machidai n. sp. show only few scales (or none, as the case may be with D. machidai n. sp.) above the opercular spine. Diancistrus springeri n. sp. and D. novaeguineae come closest of those with a large scale patch above the opercular spine, but no scales below, except for a very few large specimens of D. novaeguineae , where even a few scales below the opercular spines can be present. In those rare instances distinction would largely depend on pseudoclaspers, but D. altidorsalis appears more high- necked than D. novaeguineae. Biology. Diancistrus altidorsalis is one of the few species that has been observed alive (in the ZMUC aquarium). Several specimens were obtained from Bali by the Danish Aquarium in Charlottenlund under the name “Brotulina fusca”. Larger dominating specimens were observed to make attacks towards smaller specimens, which were hiding below rocks or any other shelter provided. Otherwise, the fishes swam actively forward and backward at similar ease with undulating movements of their vertical fins. One male was observed to flip the cover of the copulatory organ backward and forward so that the organ was exposed or covered while swimming. Distribution (Fig. 20). Diancistrus altidorsalis is widespread and fairly common in the central part of the West Pacific. It is known from the Java, Bali and Sabah in the west to Sulawesi and New Guinea and reaches eastward to the Solomon Islands (including Santa Cruz Islands). Etymology. Named after the characteristic high- necked profile - alius (Latin) = high, and dorsalis (Latin) = neck. Diancistrus atollorum n. sp. (Figs 15, 16, 17; Table 8) Material examined. (26 specimens, 25-58 mm SL). Holotype — CAS 222744, male, 47 mm SL, Falalap Islet, I laluk Atoll, Yap Slate, Federated States of Micronesia, off outer reef at 1-4 m, Pakal, Tachim, Yarofoma and R.R. Rolen, 22 Oct. 1953. Paratypes- BPBM 15366, I male, tail broken, Onoloa Atoll, Gilbert Islands, Kiribati; CAS 81438, 1 male, 38 mm SL and 1 female, 46 mm SL, Kapingamarangi Atoll, Pohnpei State, Federated States of Micronesia, lagoon coral heads on inner margin of reef flat, Atta, Kindaro and R.R. Rofen, 14 July 1954; CAS 222745,3 males, 39-47 mm SL and 3 females, 45-58 mm SL, same data as holotype; MCZ 158558, 1 male, 45 mm SL, 04°28’S, 172°10’W, Orona Atoll, Gilbert Islands, Kiribati, R.M. Bailey, 3 July 2000; USNM 263668, 1 male, 35 mm SL, Kwajalein Atoll, Marshall Islands, 1 Nov. 1964; USNM 372948, 2 males, 32 mm SL and 3 females, 25-33 mm SL, Rongerik Atoll, 97 W. Schwarzhans, P. R. Moller and J. G. Nielsen Table 8. Meristic and morphometric characters of Diancistrus atollorum n. sp. Holotype CAS 222744 Holotype + 25 paratypes n Mean (range) Standard length in mm Meristic characters 47 41.1 (25-58) 25* Dorsal fin rays 76 73.7 (72-76) 18 Anal fin rays 60 57.5 (56-61) 18 Pectoral fin rays 21 21.0 (20-22) 21 Precaudal vertebrae 11 10.9(10-11) 19 Caudal vertebrae 30 29.7 (29-31) 18 Total vertebrae 41 40.7 (40-42) 18 Rakers on anterior gill arch 12 15.5(12-18) 18 Pseudobranchial filaments 2 2 12 D/V 6 6.1 (6-7) 19 D/A 20 20.3 (17-22) 19 V/A 13 12.9(12-14) 19 Morphometric characters in % of SL Head length 27.8 27.5 (25.7-29.4) 19 Head width 14.3 13.5(10.7-16.6) 17 Head height 17.6 17.0(15.7-19.0) 18 Upper jaw length 15.2 14.0(12.9-15.5) 17 Maxillary height 4.5 4.3 (3.6-5.2J 19 Diameter of pigmented eye 1.8 17(1.2-2.0) 21 Interorbital width 7.3 7.1 (6.2-8.4) 21 Postorbital length 20.6 20.8(19.8-22.4) 18 Preanal length 49.1 48.1 (44.2-53.0) 20 Predorsal length 32.9 32.1 (29.4-34.8) 20 Body depth at origin of anal fin 18.6 18.5(16.8-20.6) 21 Pectoral fin length 16.3 16.0(13.3-17.3) 21 Pelvic fin length 28.4 27.2 (24.8-29.2) 14 Base pelvic fin - anal fin origin 28.3 27.4 (24.6-30.6) 20 Nail broken in one paratype Marshall Islands. L.P. Schultz and E.S. Herald, 28 June 1946; USNM 372949, 1 male, 40 mm SL and 1 female, 40 mm SL, Rongelap Atoll, Marshall Islands, L.P. Schultz and E.S. Herald, 19 June 1946; USNM 372951, 1 male, 45 mm SL, Bikini Atoll, Marshall Islands, V.P. Brocket ai ,21 July 1947; USNM 372952, 1 male, 50 mm SL, Bikini Atoll, Marshall Islands, V.P. Brock and E.S. Herald, 7 Aug. 1946; USNM 372971, 1 male, 33 mm SL, Bikini Atoll, Marshall Islands, V.P. Brock and E.S. Herald, 7 Aug. 1946; USNM 376184, 2 females, 41-44 mm, Kwajalein Atoll, Marshall Islands, A.B. Amerson, 15 Oct. 1964; ZMUC P 771482, 1 male, 39 mm SL and ZMUC P 771483 1 female, 47 mm SL, same data as CAS 222745. Diagnosis. Vertebrae 11+29-31=40-42, dorsal fin rays 72-76, anal fin rays 56-61; eyes small (<2.0% SL); outer pseudoclasper deep, ear-lobe shaped, opened towards ventral; inner pseudoclasper firmly joined to outer pseudoclasper anteriorly, with supporter, forming a sharp, forward-inclined lobe; narrow scale patch on cheek, no scales on operculum; head profile slender; otolith length to height ratio >2.1. Description. The principal meristic and morphometric characters are shown in Table 8. Body moderately compact, with slender fleshy snout; fishes mature at about 35 to 40 mm SL. Head with narrow scale patch on cheek, no scales on operculum. Horizontal diameter of body scales 2.0% SL. Maxiliaries slightly expanded posteriorly with angle at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril small, the size of anterior nostril and about l/4th the size of eye. Head sensory pores (Fig. 16 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 5 rows of granular teeth and 1 row of larger teeth at symphysis, followed by about 8 rows of granular teeth posteriorly. Anterior-most teeth on inner row the largest, about the size of pupil diameter. Vomer horseshoe-shaped, with 2 tooth rows of granular teeth. Palatine teeth in 2 rows, largest on inner row. Dentary with 5 outer rows of granular teeth and 1 inner row of larger teeth anteriorly. Totally, about 6 large teeth on inner row, about the size of pupil diameter. Otolith (Fig. 16 F-G). Elongate with a much expanded slender posterior tip resulting in a length to height relation of 2.15 (40 mm SL); otolith length to sulcus length 2.3; sulcus not inclined. Anterior tip of otolith pointed; dorsal rim gently curved without angles. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, but not reaching tip ot last precaudal parapophysis. Fig. 15. Diancistrus atollorum n. sp. Holotype. CAS 222744, male, 47 min SL. 98 Dinematichthyine fishes of'the Indo-West Pacific Fig. 16. Diancistrus atollorum n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype; C, view of left pseudoclasper from inside, holotype; D, view of left pseudoclasper from inside, CAS 222745, 47 mm SL; E, inclined lateral view of male copulatory organ, CAS 222745, 47 mm SL; F, median view of right otolith, MCZ 158558, male, 45 mm SL; G, ventral view of right otolith, MCZ 158558, male, 45 mm SL. Male copulatory organ (Fig. 16 C-E). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper deep, ear-lobe shaped, opened towards ventral; inner pseudoclasper firmly joined to outer pseudoclasper anteriorly, with supporter, forming sharp, forward-inclined lobe. Isthmus between pseudoclaspers moderately wide. Penis short, curved, slender. Coloration. Live colour. Uniformly light brown when preserved. Comparison. Diancistrus atollorum belongs to the species group with ear-lobe shaped outer pseudoclaspers and small eyes (Diancistrus erythraeus subgroup) further containing D. erythraeus, D. mdgroutheri n. sp., D. robustus n. sp. and D. tongaensis n. sp. Except for O D. atollorum © D. erythraeus © D. mergroutheri O D. robustus © D. tongaensis * D. sp.3 Fig. 17. Sample sites of Diancistrus atollorum n. sp.. D. erythraeus, D. megroutheri n. sp., D. robustus n. sp., D. tongaensis n. sp. and Diancistrus sp. 3. One symbol may represent several samples. 99 W. Schwarzhans, P. R. Moller and J. G. Nielsen D. robustus n. sp. the species of this subgroup are further characterized by the low number of vertebrae (39-41, rarely 42 versus 43 in D. robustus n. sp.) and the slender head (head width <16.5% SL versus >16.5% SL in D. robustus n. sp.). Of the remaining four species, D. atollorum is closest to D. erythraeus, being distinguished by the complete ear-lobe shaped outer pseudoclasper (versus incomplete, half-moon shaped), the large-lobed inner pseudoclasper (versus reduced, small, hook-like) and the more elongate otolith (otolith length to height >2.1 versus 2.0). Diancistrus atollorum is the only small-eyed Diancistrus species and also the only one with ear-lobe shaped outer pseudoclaspers within its area of distribution. It does not seem to overlap with any other species of the group. It may occur locally with D. beateae n. sp., D. mennei n. sp., D. eremitus n. sp., D. karinae n. sp. or D. pohnpeiensis n. sp., all of which are readily distinguished not only by the pseudoclasper pattern, but also by the presence of scales on the operculum, the large eye (>2.5% SL versus <2.0% SL) and the higher number of vertebrae (generally 43 and more versus <42). Finally, these are all species growing to larger body sizes (up to 80 mm SL at least versus <60 mm SL). Distribution (Fig. 17). Diancistrus atollorum is widespread and fairly common along the shores of the Micronesian atolls, from the Marshall Islands (Bikini, Rongelap, Rongerik, Kwajalein) in the north to Ifaluk and Kapingamarangi in the south and eastwards to Kiribati (Onotoa and Orona). To the north - Ryukyu Islands and northern Philippines D. atollorum is replaced by D. erythraeus, to the south-east - Fiji, Tonga, Samoa - by D. tongaensis n. sp. and to the south-west - New Guinea, Vanuatu, Great Barrier Reef- by D. mcgroutheri n. sp. Etymology. Named after its habitat among the Micronesian atolls. Diancistrus beateae n. sp. (Figs 18, 19, 20; Table 9) Material examined. (50 specimens, 39-132 mm SL). Holotype - CAS 222746, male, 87 mm SL, 01°N, 154°E, Kapingamarangi Atoll, Pohnpei State, Micronesia, lagoon coral heads on inner margin of reef fiat, Atta, Kindaroand R.R. Rofen, 14 July 1954. Paratypes-AMS 1.20641-001, 1 female, 43 mmSL, 11°N, 166°E, Rongelap Island, Marshall Islands, V.P. Brock and party, 25 July 1946; AMS I. 20779-034, 1 female, 99 mm SL, 13°05’S, 143°57’E, Cape York, north end of Tijou Reef, Great Barrier Reef, 25 m depth, AMS and AIMS party, 22 Feb. 1979; AMS 1. 33731-069, 1 male, 71 mm SL, 10°13’S, I44°24’E, Ashmore Reef, Coral Sea, 22 Jan. 1993; ANSP 135375, I male, 89 mm SL, 14°35’S, 145°36’E, Yonge Reef, Great Barrier Reef, 26 Nov. 1975; BPBM 13660, 1 female, 132 mm SL, Pixi Reef off Cairns, Great Barrier Reef, G.R. Allen, 2 July 1972; BPBM 40211, 1 female, 76 mm SL, Tutuila Island, American Samoa, R.C. Wass, 1976-77; USNM 222481, 1 male, 73 mm SL, Tutuila Bay, Table 9. Meristic and morphometric characters of Diancistrus beateae n, sp. Holotype CAS 222746 Holotype + 42 paratypes Mean (range) n Standard length in nun 87 73.0(39-132) 43 Meristic characters Dorsal fin rays 84 86.7 (84-91) 40 Anal fin rays 67 70.3 (65-74) 40 Pectoral fin rays 20 19.8(19-21) 20 Precaudal vertebrae 11 11 40 Total vertebrae 46 45.0 (44-46) 40 Rakers on anterior gill arch 16 16.1 (14-19) 31 Pseudobranchial filaments 2 2 30 D/V) 6 6.0 (5-7) 40 D/A 20 21.5(19-24) 40 V/A 13 13.0(12-14) 40 Morphometric characters in % of SL Head length 29.3 28.7 (26.8-30.3) 31 Head width 13.9 14.5(12.0-18.1) 31 Head height 21.6 23.0(18.6-27.3) 31 Upper jaw length 14.0 14.2(12.8-15.1) 31 Maxillary height 5.4 5.4 (4.6-6.0) 31 Diameter of pigmented eye 3.1 3.0 (2.6-3.5) 31 Interorbital width 6.4 6.3 (5.0-7.2) 31 Postorbilal length 21.3 20.7 (19.2-22.2) 31 Preanal length 47.1 47.8 (44.3-52.5) 31 Predorsal length 34.6 34.7 (32.1-37.9) 31 Body depth at origin of anal fin 22.4 21.1 (17.5-23.6) 30 Pectoral fin length 19.2 17.7(14.9-20.4) 20 Pelvic tin length 24.6 23.8 (21.2-26.3) 25 Base pelvic fin - anal fin origin 25.6 25.9 (22.1-29.8) 31 American Samoa, R.C. Wass; USNM 263664, 1 male, 95 mm SL, I6°45’S, 179°07’E, Wailangilala Island, Fiji, R. Bolin et al., 26 May 1965; USNM 366841, 1 male, 58 mm SL and 1 female, 60 mm SL, 18°58’S, 179°52’.,W. Totoya Island, Fiji, K. Bruwelheide et at., 27 April 1982: USNM 372950, 1 female, 45 mm SL,01°33’S, 144°59’E, Hermit Island, Papua New Guinea, V.G. Springer el at ., 30 Oct. 1978; USNM 372959,1 male, 75 mm SL, Kiriwina Island, Trobriand Islands, Papua New Guinea, T. Roberts, 19 Sept. 1975; USNM 372961, 2 males, 60-80 mm SL, 2 females, 59-68 mm SL and 2 juveniles, 46-53 mm SL, 01°33’S, 144°59’E, Hermit Island, Papua New Guinea, V.G. Springer et al., 30 Oct. 1978; USNM 372963, 2 males, 59-72 mm SL, 18°58’S, I79°52’W, Totoya Island, Fiji, V.G. Springer et al., 27 April 1982; USNM 372964, 1 female, 123 mm SL, 13°38’S, 167°30’E, Banks Islands, Vanuatu, J. T. Williams and R. Mooi, 20 May 1997; USNM 372965, 2 males, 52-72 mm SL and 1 female, 77 mm SL, 01°31’S, 145 o 0TE, Hermit Island, Papua New Guinea, V.G. Springer et al., 1 Nov. 1978; USNM 372967, 2 males, 88-96 mm SL and 1 female, 87 mm SL, 18°52’S, !78°30’E, Vuro Island, Fiji, R. Bolin et al., 8 May 1965; USNM 372968, 5 females, 41-112 mm SL, 17°44’S, 177°17TE, Malamala Island, Fiji, V.G. Springer 100 Dinematichthyine fishes of the Indo-West Pacific etal., 25 May 1982; USNM 374162, 1 male, 73 mm SL, Tutuila Bay, Samoa, R.C. Wass, 2 Oct. 2003; USNM 374163, 1 male, 59 mm SL, 18°08’S, 178°24’E, Fiji, V.G. Springer et al., 19 April 1982; USNM 374191, 1 male, 53 mm SL, 05°52’S, 110°25’E, Karimundjawa Islands, Java, V.G. Springer et ah, 29 March 1974; WAM P. 30618-031, 1 male, 82 mm SL and 1 female, 53 mm SL, 00°S, 122°E, Togean Islands, Sulawesi. Indonesia; WAM P. 31144-025, 2 females, 39-74 mm SL, 05°18’S, 150°08’E, Kimbe Bay, Bismarck Archipelago, Papua New Guinea, G.R. Allen, 20 April 1996; WAM P.31438-006, I female, 81 mm SL, 12° 15’S, 122°58’E, Ashmore Reef, Timor Sea, Western Australia, J.B. Hutchins, 3 Oct. 1997; ZMUC P 771472, 1 male, 78 mm SL and ZMUC P 771473, 1 female, 72 mm SL, same data as USNM 372961. Additional specimens. AMS I. 17499-003, 1 male, 85 mm SL, Solomon Islands, Florida Island; AMS I. 22611 - 017, 1 female, 74 mm SL, Escape Reef, Great Barrier Reef, Queensland, Australia; AMS 1. 22612-010, 1 female 59 mm SL, Escape Reef, Great Barrier Reef, Queensland, Australia; AMS 1.22619-006, 1 female, 62 mm SL, Escape Reef, Great Barrier Reef, Queensland, Australia; AMS I. 39048-021, 1 female, 73 mm SL, Solomon Islands, Stewart Island; USNM 358482, 1 female, 58 mm SL, 16°36’S, 168°09’E, Vanuatu; USNM 366850, 1 female, 85 mm SL, I7°06’S, 177°13’E, Fiji. Diagnosis. Vertebrae 11+33-35=44-46, dorsal fin rays 84-91, anal fin rays 65-74; eyes large (2.6—3.5% SL); outer pseudoclasper massive, hook-shaped, very broad in ventral view; inner pseudoclasper stout, fleshy, not extending over anterior rim of outer pseudoclasper; broad scale patch on cheek, large scale patch on operculum continuous above and below opercular spine. Description. The principal meristic and morphometric characters are shown in Table 9. Body moderately compact, often high-necked, highest in front of origin of dorsal fin, snout fleshy; fishes large, mature at about 55 to 60 mm SL. Head with broad scale patch on cheek and continuous and broad scale patch on operculum above and below opercular spine, sometimes almost the size of the scale patch on the cheek. Horizontal diameter of scales on body about 1.7% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril partly covered by large flap anteriorly, about twice the size of anterior nostril and about 1/4 the size of eye. I lead sensory pores (Fig. 19 A-C). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 7 rows of granular teeth and two rows of larger teeth at symphysis, followed by about 11 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 2 rows ofteeth, the posterior with 2-3 fang-like teeth. Palatine teeth in 3 rows of 15, 10 and 12 teeth, largest on inner row. Dentary with 5 outer rows of granular teeth and 2 inner rows of anteriorly larger teeth, blending into 1 row of 15 large fangs posteriorly, fang size up to about 2/3 of pupil diameter. Otolith (Fig. 19 H-l). Moderately elongate with a pointed, dorsally shifted posterior tip, length to height 1.9- 2.0 (52-81 mm SL); otolith length to sulcus length 1.9- 2.0; sulcus not inclined. Anterior tip of otolith pointed; dorsal rim rather fiat posteriorly without or with weak postdorsal angle in otolith of females, with more pronounced postdorsal angle in otoliths of males. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore slightly elongated, not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 19 D-G). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper broad, hook-shaped and massive; inner pseudoclasper stout, fleshy, massive, sometimes enveloped from the outside by fleshy forward extension at the base of the outer pseudoclasper. Isthmus between pseudoclaspers wide. Penis short, curved, thick. Coloration. Live colour unknown. Uniformly light brown when preserved. Comparison. Diancistrus heateae belongs to the large group with hook- or stick-like outer pseudoclaspers. It differs from all other species in the group by the very Fig. 18. Diancistrus beateae n. sp. Holotype. CAS 222746, male, 87 mm SL. 101 W. Schwarzhans, P. R. Moller and J. G. Nielsen Fig. 19. Diancistrus beateae n. sp. A, lateral view of head, holotype; B, lateral view of head, WAM P. 30618-031, male, 81 mm SL; C. ventral view of head, holotype; D, ventral view of male copulatory organ, USNM 372967,96 mm SL; E, view of left pseudoclasper from inside, holotype; F, view of left pseudoclasper from inside, USNM 372967,96 nun SL; G, inclined lateral view of male copulatory organ. USNM 372967,96 mm SL; H, median view of right otolith, WAM P. 31438-006, female, 81 mm SL; 1. median view of right otolith, AMS I. 33731-069, male, 71 mm SL. 102 Dinematichthyine fishes of the Indo-West Pacific O D. beateae © D. machidai © D. springeri ® D. altidorsalis © D. niger Fig. 20. Sample sites of Diancistrus beateae n. sp„ D. machidai n. sp„ D. springeri n. sp., D. altidorsalis n. sp. and D. niger n. sp. One symbol may represent several samples. broad and massive outer pseudoclasper, which is best seen in ventral view after bending forward the hood cover. Together with D. karinae n. sp. it is the only species ol the genus Diancistrus with a complete scale cover on the operculum above and below the opercular spine and joined around it, which makes it an easily recognizable species despite its wide geographical range and co-occurrence with several other species of the genus. From D. karinae n. sp. it differs in the pseudoclasper morphology (broad hook-shaped versus flat wing-shaped) and also in the larger number of dorsal fin rays (84-91 versus 78-83) and the mostly lower D/V (6 versus 7). Diancistrus beateae is one of the largest species of Dinematichthyini with a maximum recorded size of 132 mm SL. Distribution (Fig. 20). Diancistrus beateae is amongst the more widespread Diancistrus species, with a single record each from the Marshall Islands (Rongelap) and Micronesia (Kapingamarangi) in the north, to Sulawesi and New Guinea and further to the Solomon and Vanuatu Islands, Fiji and Samoa in the south-east. It further occurs along the Great Barrier Reef and is also known from a few isolated localities to the south-west, off Java and in the Timor Sea. In the central part of its distribution area (Sulawesi, New Guinea) D. beateae overlaps with D. karinae with which it may be confused most easily in the absence of males. Etymology. Named after Beate Schwarzhans for her most valuable support of her husband during his many hours invested in this study. Diancistrus brevirostris n. sp. (Figs 21, 22, 29; Table 10) Material examined. (11 specimens, 20-59 mm SL). Holotype - USNM 338982, male, 41 mm SL, 20°29’S, 166°19’E, Ouvea Atoll, lie Haute, Loyalty Islands, deep surge channel on ocean side of reef at 3-10 m, J. T. Williams, J.-L. Menou and P. Tirard, 19 Nov. 1991. Paratypes - SMNS 21614, 1 female, 59 mm SL, 21°35’37”S, 167°50’51”E, Cap Wabao, 6 km SW Tadin, Mare Island, Loyalty Islands, coralline rock on side of exposed reef channel and cave, 1-5 m depth, R. Fricke, 16 July 1999; USNM 338982, 1 male, 33 mm SL and 1 female, 58 mm SL, same data as holotype; USNM 356170, 1 male, 35 mm SL, 17°03’S, 168°21’E, Shepherd Islands, Emae Island, Vanuatu, J. T. Williams et al. , 28 April 1999; USNM 361739, 3 females, 39-44 mm SL, 17°31’S, 168°19’E, Efate Island, Vanuatu, J. T. Williams et al., 4 May 1997. Tentatively assigned specimens. USNM 338982, 1 juvenile, 21 mm SL, same location as holotype; USNM 356170, 2 females, 20 mm SL, 17°03’S, 168°21’E, Emae Island, Vanuatu. Diagnosis. Vertebrae 11+32-34=43-45, dorsal fin rays 71-82, anal fin rays 58-69; eyes large (2.3-2.8% SL); outer 103 W. Schwarzhans, P. R. Mailer and J. G. Nielsen Table 10. Meristic and morphometric characters of Diancistrus brevirostris n. sp. Holotype USNM 338982 Holotype + 7 paratypes Mean (range) n Standard length in mm Meristic characters 41 43.9 (33-58) 8 Dorsal fin rays 79 77.0 (71-82) 5 Anal fin rays 63 63.2 (58-69) 5 Pectoral fin rays 20 19.9(18-21) 8 Precaudal vertebrae 11 11 5 Caudal vertebrae 33 32.8 (32-34) 5 Total vertebrae 44 43.8 (43-45) 5 Rakers on anterior gill arch 14 15.5(13-19) 4 Pseudobranchial filaments - 2 5 D/V 7 6.6 (6-7) 5 D/A 21 19.8 (17-22) 5 V/A 14 Morphometric characters in % ofSL 13.6(13-14) 5 Head length 24.8 25.9 (24.8-26.7) 8 Head width 13.3 12.3(10.3-14.0) 8 Head height 14.3 18.1 (14.3-21.1) 8 Upper jaw length 11.9 12.5(11.9-13.3) 8 Maxillary height 4.3 4.0 (3.8-4.3) 8 Diameter of pigmented eye 2.4 2.5 (2.3-2.8) 8 Interorbital width 6.4 5.4 (4.1-7.3) 8 Postorbital length 18.1 18.5(18.1-19.3) 8 Preanal length 48.4 46.6(43.6-51.4) 8 Predorsal length 30.4 32.2(30.4-34.1) 8 Body depth at origin of anal fin 16.9 18.1 (16.2-20.1) 8 Pectoral fin length 15.4 16.7(14.1-19.6) 7 Pelvic fin length 26.1 23.7 (20.9-26.1) 8 Base pelvic fin - anal fin origin 29.7 27.3 (24.9-29.7) 8 pseudoclasper moderately large, broad, wing-shaped, its tip slightly bent outwards, with a characteristic kink at the base seen from ventral view; inner pseudoclasper bifurcate, both branches of about equal length, broad based, flap-like; narrow scale patch on cheek with 4-5 scale rows, no scales on operculum; head short, with blunt snout; otolith thick with gently curving dorsal rim. Description. The principal meristic and morphometric characters are shown in Table 10. Body slender, but with short massive head and blunt snout, fishes small, mature at about 35 mm SL. Head with 4-5 scale rows on upper cheek, no scales on operculum. Horizontal diameter of scales on body about 1.9% SL. Maxillaries slightly expanded posteriorly with knob near rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril slightly larger than anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 22 A-C). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior); the 1st anterior mandibular pore not tubular and without cirri. Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 5 rows of granular teeth and 1 row of larger teeth at symphysis, followed by about 5 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 3 tooth rows, the larger teeth on inner row posteriorly. Palatine teeth in 2 rows of equally long teeth. Dentary with 5 outer rows of granular teeth and 1 inner row of larger teeth anteriorly, blending into 1 row of 5 large fangs posteriorly, up to about 3/4 of pupil diameter. Otolith (Fig. 22 11-1). Moderately elongate with sharply pointed anterior and rounded posterior tip, length to height 2.0 (33-41 mm SL); otolith length to sulcus length 2.0-2.2; sulcus not inclined; dorsal rim regularly curving, almost symmetrical to ventral rim. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, sometimes reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 22 D-G). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper broad, wing-shaped, its tip slightly bent outwards, with characteristic kink at the outer side and base when seen from ventral view, resulting in square-shaped appearance; inner pseudoclasper bifurcate, both branches of about equal length, broad-based, flap-like. Isthmus between pseudoclaspers narrow. Penis short, curved, stout. Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Diancistrus brevirostris belongs to the small group with wing-shaped outer pseudoclaspers and bifurcate inner pseudoclaspers, which further contains D. alatus, D. fijiensis n. sp. and D. jackrandalli n. sp. Ol those, D. fijiensis n. sp. is closest in meristics, with the large eye, the absence of scales on the operculum (all of which is also shared with D. alatus) and the two branches Fig. 21. Diancistrus brevirostris n. sp. Holotype. USNM 338982, male, 41 mm SL. 104 Dinematichthyine fishes of the Indo-West Pacific Fig. 22. Diancistrus brevirostris n. sp. A, lateral view of head, holotype; B, ventral view ol head, holotype; C, lateral view of head, USNM 356170, male, 35 mm SL; D, ventral view of male copulatory organ, USNM 356170, male, 35 mm SL; E. view of left pseudoclasper from inside, USNM 356170, male, 35 mm SL; F, view ofleft pseudoclasper from inside, holotype; G, inclined lateral view ofr.iale copulatory organ, holotype; H, median view of right otolith, holotype; 1, ventral view of right otolith, holotype. ol'the bifurcate inner pseudoclasper being equally long (the anterior branch being twice as long as the posterior in D. alatus). Diancistrus brevirostris differs from D.fijiensis n. sp. in the broad based flap-like nature of the bifurcate inner pseudoclasper (versus narrow-based and sharply pointed), the characteristic kink at the outer side of the outer pseudoclasper base seen in ventral view and the more massive head and blunt snout. Also D. fijiensis n. sp. generally has fewer and larger scales on the cheek than D. brevirostris. In Vanuatu, D. brevirostris may occur associated with D. alleni, D. beateae, D. megroutheri n. sp. and D. novaeguineae. Diancistrus megroutheri n. sp. belongs to the group with ear-lobe shaped outer pseudoclaspers and small eyes; the other species are readily distinguished by the presence of scales on the operculum. A further similar looking species occurring in Vanuatu is Paradiancistrus acutirostris n. sp. This species is recognized by the curved structure of the fused inner and outer pseudoclaspers, the presence of only one lower preopercular pore (versus 3) and the very narrow scale patch on the cheek (2 scale rows on upper cheek versus 4-5). At the Loyalty Islands, D. brevirostris may co-occur with D. longifilis and D. tongaensis n. sp., both species without scales on the operculum. While Diancistrus tongaensis n. sp. belongs to the group with ear-lobe shaped outer pseudoclaspers and small eyes, D. longifilis would probably be the most easiest to confuse it with, in the absence of males. Diancistrus longifilis does exhibit a more slender, less blunt snout and has otoliths with marked pre- and postdorsal angles. Distribution (Fig. 29). Diancistrus brevirostris is only known from the Loyalty Islands and some islands of Vanuatu. Etymology. Named after short, blunt snout - brevis (Latin) = short and derivation of rostrum ( rostris - Latin) = snouted. Diancistrus eremitus n. sp. (Figs 23,24, 40; Table 11) Material examined. (6 specimens, 43-83 mm SL). Holotype - USNM 372957, male, 71 mm SL, 01°33’S, 144°59’E, Manus Province, Bismarck Sea, Bismarck Archipelago, Hermit Islands, Arnot Island, ocean side of reef at drop off, Papua New Guinea, ocean side of reef 105 W. Schvvarzhans, P. R. Moller and J. G. Nielsen at drop off, 0-16 m, V.G. Springer et al., 30 Oct. 1978. Paratypes — CAS 2223206, 1 male, 43 mm SL, 1 female, 49 mm SL, 17°25’S, 179°10’W. Fiji, Lau Group, Mago Island NW side, sand, rubble and hard coral rock, D.W. Greenfield, K.. Longenecker, and R. Langston, 3 Jan. 2003; USNM 372958, 2 males, 47-83 mm SL, 1 female, 49 mm SL, 08°23’S, 162°51’E, Stewart Island, Solomon Islands, J. T. Williams et al., 3 Oct. 1998. Diagnosis. Vertebrae 11+31-33=42-44, dorsal fin rays 83-86, anal fin rays 66-71; eyes very large (>3.0% SL), D/V 5-6; outer pseudoclasper large, more than 2 times the length of inner pseudoclasper, wing-shaped, with broad base and slightly concave pointed tip without fleshy bulge; inner pseudoclasper joined to outer pseudoclasper anteriorly, thin, with single pointed tip, short, not extending over anterior part of outer pseudoclasper; moderately broad scale patch on cheek with 5-6 scale rows on upper cheek, small scale patch on operculum above opercular spine (4-5 scales in 2 rows); head massive; otolith moderately slender, length to height 1.9-2.0. Description. The principal meristic and morphometric characters are shown in Table 11. Body moderately compact, highest behind origin of dorsal fin at about 10th dorsal fin ray, with massive head, fishes mature at about 50 mm SL. Mead with moderately broad scale patch on cheek with 5-6 scale rows on upper cheek, small scale patch on operculum above opercular spine (4-5 scales) in 2 rows and no scales below opercular spine. Horizontal diameter of scales on body about 1.6% SL. Maxillaries expanded posteriorly with prominent knob at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril very small, size of anterior nostril and about l/10th size of eye. Head sensory pores (Fig. 24 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and two rows of larger teeth at symphysis, followed by 8 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 3 rows of teeth, the posterior with enlarged teeth. Palatine teeth in two rows of 15 and 8 teeth, Table II. Meristic and morphometric characters of Diancistrus eremitus n. sp. Holotype USNM 372957 Holotype + 3 paratypes Mean (range) n Standard length in mm Meristic characters 71 62.5 (47-83) 4 Dorsal fin rays 85 84.5 (83-86) 4 Anal fin rays 67 68.3 (66-71) 4 Pectoral fin rays 20 20 2 Precaudal vertebrae 11 11 4 Caudal vertebrae 31 32.0 (31-33) 4 Total vertebrae 42 43.0 (42-44) 4 Rakers on anterior gill arch 15 15 2 Pseudobranchial filaments 2 2 4 D/V 5 5.8 (5-6) 4 D/A 24 22.7 (22-24) 4 V/A Morphometric characters in % 13 of SL 13.3 (13-14) 4 1 lead length 25.8 27.6 (25.8-29.5) 4 Head width 13.2 12.9(11.6-14.8) 4 Head height 22.1 20.6(19.0-22.1) 4 Upper jaw length 13.7 13.5(13.0-14.2) 4 Maxillary height 5.2 4.8 (4.3-5.2) 4 Diameter of pigmented eye 3.2 3.1 (3.0-3.3) 4 Interorbital width 6.5 5.7 (5.1-6.5) 4 Postorbital length 18.6 19.2(18.6-20.5) 4 Preanal length 49.0 49.6 (47.7-52.0) 4 Predorsal length 32.8 33.3 (32.8-33.7) 4 Body depth at origin of anal fin 19.3 19.7(18.3-21.8) 4 Pectoral fin length - 17.1 (16.1-18.0) 2 Pelvic fin length 23.2 22.1 (19.2-23.2) 4 Base pelvic fin - anal fin origin 28.3 27.6 (26.1-28.9) 4 the inner row with fewer and larger teeth. Dentary with 5 outer rows of granular teeth and 2 inner rows of larger teeth anteriorly blending into 1 row of 3-4 large fangs posteriorly, up to more than 1/2 of pupil diameter. Otolith (Fig. 24 H-K). Moderately elongate with moderately pointed anterior and posterior tips, length to height 1.9-2.0 (50-85 mm SL); otolith length to sulcus length 1.85-2.0; sulcus slightly inclined at 5°. Dorsal angle weak or absent in otoliths of females, more pronounced in otoliths of males. Fig. 23. Diancistrus eremitus n. sp. Holotype. USNM 372957, male, 71 mm SL. Dinematichthyine fishes of the Indo-West Pacific Fig. 24. Diancistrus eremitus n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype: C, ventral view of male copulatory organ, USNM 372958, male, 83 mm SL; D, view of left pseudoclasper from inside, USNM 372958, male, 83 mm SL; E, ventral view of male copulatory organ, holotype; F, view of left pseudoclasper from inside, USNM 372958, male, 47 mm SL; G, inclined lateral view of male copulatory organ, USNM 372958, male, 83 mm SL; H, median view of right otolith, USNM 372958, male, 47 mm SL; I, median view of right otolith, USNM 372958, male, 83 mm SL; K, ventral view of right otolith. USNM 372958, male, 83 mm SL. 107 W. Schwarzhans, P. R. Moller and J. G. Nielsen Axial skeleton (of holotype). Neural spine of vertebrae 4 inclined and 5-8 depressed. Parapophyses present from vertebra 6 to 11. Pleural ribs on vertebrae 2-10 (11). First anal fin pterygiophore slightly elongated, not reaching tip of last precaudal parapophysis in males, short in females. Male copulatory organ (Fig. 24 C-G). Two pairs of large pseudoclaspers. Outer pseudoclasper large, more than 2 times the length of inner pseudoclasper, wing¬ shaped, with broad base and slightly concave pointed tip, not bending outwards, with broad supporter; inner pseudoclasper joined to outer pseudoclasper anteriorly, thin, with single pointed tip, short, not extending over anterior part of outer pseudoclasper. Isthmus between pseudoclaspers wide. Penis short, curved, with thin pointed tip. An ontogenetic succession of pseudoclaspers from three males of different sizes of D. eremitus shows a reduced morphological expression in the smallest subadult specimen. Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Diancistrus eremitus belongs to the group with wing-shaped outer pseudoclasper and a single-tipped inner pseudoclasper, which further contains D. jeffjohnsoni n. sp„ D. karinae n. sp., D. katrineae n. sp„ D. mennei n. sp. and D. pohnpeiensis n. sp. Of these, D. mennei n. sp. and D. pohnpeiensis n. sp. are closest. It differs from D. mennei n. sp. in the fewer scales above the opercular spine (4 versus 8) and the large outer pseudoclasper of more than 2 times the length of the inner pseudoclasper (versus \'A length of inner pseudoclasper). From D. pohnpeiensis n. sp. it differs in the absence of a fleshy bulge posteriorly at the outer pseudoclasper, 4 scales above the opercular spine in 2 rows (versus 2-3 in a single row) and the number of vertebrae (42-44 versus 45-46). These three species do not seem to overlap geographically, i.e. they presumably replace each other in their respective areas of occurrence. At Flermit Island, New Guinea, Diancistrus eremitus is associated with D. alleni, D. altidorsalis, D. atollorum, D. beateae and D. karinae n. sp.. Of these, D. eremitus is easiest to confuse with D. alleni in females. Diancistrus eremitus has 4 scales in 2 rows above the opercular spine in a single row, whereas D. alleni has 1 or 2, rarely 3 scales. No other species are known from the Stewart Islands, but from the nearby main Solomon Islands, where D. eremitus seems to be missing, D. alleni, D. mcgroutheri n. sp. and D. novaeguineae have been recorded. Distribution (Fig. 40). Diancistrus eremitus is known only from three oceanic islands off the north coast of the New Guinea-Solomon Island chain respectively, namely the Flermit Island and Stewart Islands, and from one location off western Fiji in the Lau Group. Etymology. Named after the Latin translation (= eremitus) of the type locality - Hermit Island. Diancistrus erythraeus (Fowler, 1946) (Figs 17,25, 26; Table 12) Calcarbrotula erythraea Fowler, 1946: 193, fig. 57-58 (type locality: Aguni Shima, Ryukyu Islands, Japan). Brotulina erythraea - Machida in Masuda et al. 1984: 101; Nielsen et al. 1999: 126. Material examined. (11 specimens, 30-70 mm SL). Holotype - ANSP 72069, female, 64 mm SL, Aguni Shima, Ryukyu Islands, Japan, E.R. Tinkham. Additional specimens. BP13M 8710, 1 male, 38 mm SL and 3 females, 30-55 mm SL, Ishigaki-jima, Ryukyu Islands, J.E. Randall and ATI Banner, 22 May 1968; USNM 365838, 1 female, 70 mm SL, I6°26'N, 119°56’E, Pangasian Island, Philippines, E.O. Murdy, 16-17 April 1980; USNM 372955, 1 male, 34 mm SL and 1 female, 47 mm SL, 10°52'30”N 120°56’00”E, Palawan Province, Bararin Island, W side (Cuyo Island), Philippines, 0-14 m depth, 23 May 1978; USNM 374197, 2 males, 33-44 mm SL, Philippines; USNM 374199, 1 male, 35 mm SL, 24°27’N, 124°12’E, Ishigaki-jima, Ryukyu Islands, J. T. Williams et al., 23 Feb. 1998. Diagnosis. Vertebrae 11+28-30=39-41, dorsal fin rays 70-76, anal fin rays 55-61; eyes very small (<2.0% SL); outer pseudoclasper ear-lobe shaped, the half-moon shaped Table 12. Meristic and morphometric characters of Diancistrus erythraeus (Fowler, 1946). Holotype ANSP 72069 Holotype + 10 specimens Mean (range) n Standard length in mm 64 44.1 (30-70) ii Meristic characters Dorsal fin rays 73 73.3 (70-76) ii Anal fin rays 61 58.5 (55-61) ii Pectoral fin rays - 20.4 (20-21) 9 Precaudal vertebrae 10 10.9(10-11) 11 Caudal vertebrae 30 29.3 (28-30) 11 Total vertebrae 40 40.3 (39-41) 11 Rakers on anterior gill arch 15 15.7(14-18) 11 Pseudobranchial filaments 2 1.5(0, 2) 11 D/V 7 6.3 (6-7) 11 D/A 18 19.2(18-21) 11 V/A 13 13 11 Morphometric characters in % of SI. Head length 28.2 27.6 (26.2-29.0) 11 Head width 12.9 13.1 (11.6-15.6) 11 Head height 16.6 17.0(15.9-18.7) 11 Upper jaw length 14.9 13.9(13.0-14.9) 11 Maxillary height 5.2 4.3 (3.8-5.2) 11 Diameter of pigmented eye 1.6 1.6 (1.2-1.9) 11 Interorbital width 7.5 6.9 (6.2-7.5) 11 Postorbital length 21.7 20.8(19.7-21.7) 10 Preanal length 48.1 47.2 (45.1-49.9) 11 Predorsal length 33.8 32.8 (30.7-34.8) 1) Body depth at origin of anal fin 18.1 18.0(15.9-19.8) 11 Pectoral fin length 15.7 15.7(13.5-17.6) 11 Pelvic fin length 21.1 25.6(21.1-28.6) 9 Base pelvic fin - anal fin origin 30.1 28.2 (25.3-31.0) 11 108 Dinematichthyine fishes of the Indo-West Pacific ear-lobe extension inwardly directed; inner pseudoclasper firmly joined to outer pseudoclasper anteriorly, with supporter, reduced small hook-like, often with a fleshy flap at its base; 5-7 rows of small scales on upper cheek, no scales on operculum; head profile slender; otolith length to height ratio <2.0. Description (Fig. 25). The principal meristic and morphometric characters are shown in Table 12. Body moderately compact, with slender fleshy snout; fishes small, mature at about 30 to 35 mm SL. Head with narrow scale patch on cheek, no scales on operculum. Horizontal diameter of scales on body about 1.4% SL. Maxillaries expanded posteriorly with prominent angle or knob at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril about twice the size of anterior nostril and about 1/4 the size of eye. Head sensory pores (Fig. 26 A-C). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of a 35 mm SL non-type - USNM 374199). Premaxilla with 5 rows of granular teeth and 1 row of larger teeth at symphysis, followed by 5 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 2 tooth rows, posterior row with larger teeth. Palatine teeth in two rows, inner row with larger teeth. Dentary with 4 outer granular tooth rows and 1 inner row of larger teeth anteriorly, blending into 1 row of 6 large fangs posteriorly, fangs up to 3/4 of pupil diameter. Otolith ( Fig. 26 G-H). Moderately elongate with rather blunt anterior and posterior tips resulting in a length to height relation of 2.0 (35 mm SL); otolith length to sulcus length 2.3; sulcus slightly inclined at 5°. Dorsal rim with rounded, obtuse pre- and postdorsal angles. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, but mostly not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 26 D-F). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper ear¬ lobe shaped, but incomplete like a half-moon extension, which is inwardly directed; inner pseudoclasper firmly joined to outer pseudoclasper anteriorly, with supporter. reduced to a small hook, often with a fleshy flap at its base. Isthmus between pseudoclaspers narrow. Penis short, curved, moderately slender. Coloration. Live colour probably red according to Fowler (1946) and Machida (1984). Uniformly light brown when preserved. Comparison. Diancistrus erythraeus belongs to the group of species with ear-lobe shaped outer pseudoclaspers and small eyes (Diancistrus erythraeus subgroup), further containing D. atollorum, D. mcgroutheri n. sp., D. rohustus n. sp. and D. tongaensis n. sp. Except for D. robustus n. sp., the species of this subgroup are further characterized by the low number of vertebrae (39-41, rarely 42 versus 43 in D. robustus n. sp.) and the slender head (head width <16% SL versus >16% SL in D. robustus n. sp.). The remaining four species are best distinguished by means of pseudoclaspers. In D. mcgroutheri n. sp. and D. tongaensis n. sp. the inner pseudoclasper is free from the outer pseudoclasper, whereas in D. atollorum and D. erythraeus (and D. robustus n. sp.) it is firmly joined to the outer pseudoclasper. Apparently, D. erythraeus is closest to D. atollorum. Diancistrus erythraeus co-occurs with a number of Diancistrus species but is the only species with ear-lobe shaped outer pseudoclaspers. The only other small-eyed species with which it occurs along the Ryukyu Islands is D. jackrandalli n. sp., which readily differs in the form of the pseudoclaspers (wing-shaped outer pseudoclasper) and the presence of scales on the operculum above the opercular spine. Diancistrus erythraeus does not seem to overlap in distribution with any other species of the Diancistrus erythraeus subgroup. Remarks. Specimens from the Ryukyu Islands seem to have consistently smaller eyes than those from the northern Philippines, and they also seem to consistently lack the fleshy flap at the base of the inner pseudoclasper that is usually observed in specimens from the Philippines. More material needs to be investigated before it can be judged whether these subtle differences reflect the presence of two separate species. Distribution (Fig. 17). Diancistrus erythraeus is distributed along the Ryukyu Islands and along the northern Philippines, but is not known from Taiwan. To the south and east it is replaced by D. atollorum. Fig. 25. Diancistrus erythraeus (Fowler, 1946). Holotype. ANSP 72069, female, 64 mm SL. 109 W. Schwarzhans, P. R. Moller and J. G. Nielsen Fig. 26. Diancistrus erythraeus (Fowler, 1946). A, lateral view of head. BPBM 8710, female, 55 mm SL; B, lateral view of head, USNM 374197, male, 44 mm SL; C, ventral view of head, USNM 374197, male, 44 mm SL; D, view of left pseudoclasper from inside, BPBM 8710, 38 mm SL; E, view of left pseudoclasper from inside, USNM 374197, male, 44 mm SL; F, inclined lateral view of male copulatory organ, BPBM 8710, 38 mm SL; G, median view of right otolith, USNM 374199, male, 35 mm SL; H, ventral view of right otolith, USNM 374199, male, 35 mm SL. Diancistrus fijiensis n. sp. (Figs 27, 28, 29; Table 13) Material examined. (20 specimens, 26-49 mm SL). Holotype - USNM 374161, male, 41 mm SL, 18°52’S, 178°30’E, Vuro Island, Great Astrolabe Reef, Fiji, inner reef, R. Bolin etal ., 8 May 1965. Paratypes- ROM 78115, 1 male, 31 mm SL, 18°S, 178°E, Fiji; USNM 366849, 1 male, 26 mm SL, 17°12’S, 176°54’E, Viwa Isl„ Fiji, V.G. Springer et al., 27 May 1982; USNM 372970, 1 male, 30 mm SL and 4 females, 28-42 mm SL, same data as the holotype; USNM 372972, 1 male, 39 mm SL, Mamanutha Group, NW end of Malolo Island, S side of channel, Fiji, 17°45’S, 177°04’W, 0-20 m depth, V.G. Springer et al., 25 May 1982; USNM 374158,1 male, 32 mm SL and 1 female, 32 mm SL, 18°50’S, 178°32’E, Mbulio Isl., Fiji. R. Bolin et al., 28 Aug. 1963; USNM 374160, 1 male, 49 mm SL, 21°38’S, 178°45’W, Southern Lau Group, Ono-Ilau Island, outside barrier reef on NW side of island, Fiji, 15-18 m depth. V.G. Springer et al., 1 May 1982; USNM 384591, 1 female, 30 mm SL, same data as holotype; WAM P. 30791- 001, 1 male, 43 mm SL, 18°44’S, 178°29’E, Dravuni Isl, Fiji, A. R. Emery etal., 31 March 1983; ZMUC P 771474, 1 male, 40 mm SL, same data as for USNM 374160; ZMUC P 771475, 1 female, 30 mm SL, same data as for USNM 384591. Additional specimens. AMS I. 18438-015, 1 female, 49 mm SL, Fiji; USNM 259451,3 females, 42-47 mm SL, 19°49’S, 178°I5’W, Fiji. Diagnosis. Vertebrae 1 14-31-33=42-44, dorsal fin rays 70-81, anal fin rays 58-63; eyes large (2.4-3.2% SL); outer pseudoclasper moderately large, broad-based, wing-shaped, its tip strongly bent outwards; inner pseudoclasper bifurcate, both branches of about equal length, narrow, thin, forming sharp spines; scale patch on cheek with 3-4 rows of large scales, no scales on operculum; head slender; 1st anterior mandibular pore large, tubular, forwardly directed; otolith thick with irregularly curved dorsal rim. Description. The principal meristic and morphometric characters are shown in Table 13. Body slender, but with massive head and short snout; fishes small, mature at about 110 Dinematichthyine fishes of the Indo-West Pacific Table 13. Meristic and morphometric characters of Diancistrus fijiensis n. sp. Holotype USNM 374161 Holotype + 15 paratypes Mean (range) n Standard length in mm Meristic characters 41 36.0 (26-48) 16 Dorsal fin rays 77 76.7 (70-81) 16 Anal fin rays 60 61.1 (58-63) 16 Pectoral fin rays 19 19.7(18-21) 12 Precaudal vertebrae 11 11 16 Caudal vertebrae 32 32.0(31-33) 16 Total vertebrae 43 43.0 (42-44) 16 Rakers on anterior gill arch 16 16.1 (14-18) 14 Pseudobranchial filaments 2 2 13 D/V 6 6.2 (6-7) 14 D/A 20 20.3 (18-22) 14 V/A 13 Morphometric characters in % of SL 13.1 (13-14) 15 Head length 26.0 25.7 (24.5-27.0) 15 Head width 11.9 12.1 (10.0-13.8) 15 Head height 15.7 17.2(15.4-21.5) 15 Upper jaw length 13.1 12.8(11.9-13.9) 14 Maxillary height 4.4 4.1 (3.5-4.6) 13 Diameter of pigmented eye 2.7 2.8 (2.2-3.2) 15 Interorbital width 6.6 5.8 (4.0-7.3) 14 Postorbital length 17.9 18.1 (16.9-19.2) 15 Prcanal length 48.1 46.9(41.4-50.5) 15 Prcdorsal length 31.9 31.5 (29.5-34.1) 15 Body depth at origin of anal fin 16.4 16.9(16.3-18.1) 14 Pectoral fin length 15.4 15.0(12.8-16.7) 14 Pelvic fin length 26.8 25.3 (22.3-28.1) 9 Base pelvic fin - anal fin origin 29.5 27.1 (22.4-29.5) 14 30 mm SL. Mead with 3-4 rows of large scales on cheek, no scales on operculum. Horizontal diameter of scales on body about 1.8% SL Max diaries expanded posteriorly with angle at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril small, the size of anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 28 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior): the 1st anterior mandibular pore tubular, large, pointing towards anterior and with single cirrus. Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 5 rows of granular teeth and 1 row of larger teeth at symphysis, followed by 5 rows of granular teeth posteriorly. Largest teeth up to Zi pupil diameter. Vomer horseshoe-shaped, with 2 tooth rows, posterior row with larger teeth. Palatine teeth in two rows, slightly larger on inner row. Dentary with 4 outer granular tooth rows and 1 inner row of larger teeth anteriorly, blending into 1 row of about 13 large fangs posteriorly, fangs up to more than 2/3 of pupil diameter. Otolith (Fig. 28 G-H). Moderately elongate with broad anterior and rounded posterior tip, length to height 1.9 (49 mm SL); otolith length to sulcus length 2.0; sulcus markedly inclined at almost 10°; dorsal rim irregularly curving. Axial skeleton (of holotype). Neural spine of vertebrae 4 inclined and 5-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10 (11). First anal fin ptcrygiophore elongated, sometimes reaching tip of last precaudal parapophysis. Male copulatorv organ (Fig. 28 C-F). Two pairs of moderately large pseudoclaspers. Outer pscudoclasper with broad base, wing-shaped, its tip strongly bent outwards, broad supporter; inner pseudoclasper bifurcate, both branches of about equal length, narrow, very thin and spiny. Isthmus between pseudoclaspers narrow. Penis short, curved, with broad base and thin tip. Coloration. Live colour not known. Uniformly medium to dark brown when preserved. Comparison. Diancistrus fijiensis belongs to the small group with wing-shaped outer pseudoclaspers and bifurcate inner pseudoclaspers, which further contains D. a/atus, D. brevirostris and D. jackrcmdalli n. sp. Of these, D. brevirostris is the closest. For differentiation from D. alatus and D. jackrandalli n. sp. see their species accounts. Other species of Diancistrus occurring in Fiji are D. beateae, with hook-shaped outer pseudoclaspers and extensive squamation on the operculum, D. eremitus, with wing-like outer peudoclaspers and scales on the operculum above the opercular spine, and the two small-eyed species with ear-lobe outer pseudoclasper D. robustus n. sp. and D. tongaensis n. sp. Distribution. Diancistrus fijiensis is only known from the Fiji Islands. Etymology. Named after the type locality - Fiji. Fig. 27. Diancistrus fijiensis n. sp. Holotype. USNM 374161, male, 41 mm SL. W. Schwarzhans, P. R. Moller and J. G. Nielsen Fig. 28. Diancistrus Jijiensis n. sp. A, lateral view of head, holotype; B. ventral view of head, holotype; C, view of left pseudoclasper from inside, USNM 374160, 48 mm SL; D, view of left pseudoclasper from inside, USNM 374160, 44 mm SL; E, inclined lateral view of male copulatory organ, USNM 374160,44 mm SL; F, ventral view of male copulatory organ, USNM 374160, 48 mm SL; G, median view of right otolith, USNM 374160, male, 44 mm SL; H. ventral view of right otolith, USNM 374160, male, 44 mm SL. E 110*E 120*E 130’E 140*E 150*E 160*E WE 180* 170'W O D. alatus © D. brevirostris © D. fijiensis 0 D. jackrandalli © D. /e/s/ Fig. 29. Sample sites of Diancistrus alatus n. sp., D. brevirostris n. sp., D. fijiensis n. sp., D. jackrandalli n. sp. and D. leisi n. sp. One symbol may represent several samples. Dinematichthyine fishes of the Indo-West Pacific Diancistrus fuscus (Fowler, 1946) (Figs 30, 31, 57; Table 14) Brotulina fusca Fowler, 1946: 195, fig. 59-60 (type locality: Aguni Shima, Ryukyu Islands, Japan). Brotulina fusca - Machida in Masuda elal. 1984: 101; Chen and Shao 1991: 14; Nielsen et al. 1999: 126. Material examined. (28 specimens, 27-67 mm SL). Holotype- ANSP 72070, male, 60 mm SL, Aguni Shima, Ryukyu Islands, Japan. Additional specimens. BPBM 40212, 1 male, 50 mm SL, Ishigaki-jima, Ryukyu Islands, J.E. Randall and A.H Banner, 22 May 1968; NSMT P. 49441, I female, 62 mm SL, 24°19’N, 124°H’E, Ishigaki-jima, Ryukyu Islands, K. Matsuura, 7 Dec. 1995; USNM 372940, I male, 58 mm SL, 10°35’N, 122°08’E, Palawan, Philippines, J. T. Williams et al., 25 Sept. 1995; USNM 372954, 2 males, 37-47 mm SL, Ch’uan-Fan-Shi, Taiwan, V.G. Springer etai, 23 April 1968; USNM 372960, 1 male, 67 mm SL, 12°50’N, 120°45’E, Mindoro, Philippines, J. T. Williams, 10 Sept. 2003; USNM 374177, 1 male, 60 mm SL and 5 females, 47-58 mm SL, Ch’uan-Fan-Shi, Taiwan, V.G. Springer et al., 23 April 1968; USNM 374180, 1 male, 43 mm SL, 24°27’N, 124°12’E, Ishigaki-jima, Ryukyu Islands, J. T. Williams et al., 23 Feb. 1998; USNM 384594, I female, 28 mm SL, 10°53’N, 121°1TE, Cuyo Island NE of Palawan, Philippines, 0—21 m, 26 May 1978; USNM 384595, 1 male, 47 mm SL, 20°24’N, 121°55’E, Batanes Islands, Philippines, G.D. Johnson et al., 1 May 1987; USNM 384596, 3 males, 41-56 mm SL and 3 females, 35-56 mm SL, 10°52’N, 120°56’E, Cuyo Islands, Palawan, Philippines, 23 May 1978; USNM 384598, 1 male, 52 mm SL and 1 female, 48 mm SL, 20°24’N, 121°55’E, Batanes Province, Philippines, G.D. Johnson et al., 2 May 1987; YCM-P 36436, 1 female, 62 mm SL, Kakeroma Island, Amami Islands, northern Ryukyu Islands; YCM-P 36596, 1 female, 48 mm SL, Kakeroma Island, Amami Islands, northern Ryukyu Islands; ZMUC P 771484, 1 male, 48 mm SL and ZMUC P 771485, 1 female, 43 mm SL, same data as for USNM 384596. Diagnosis. Vertebrae 1 1+32-34=43-45, dorsal fin rays 74-85 (usually >77), anal fin rays 58-69, D/V 6 (rarely 7); eyes moderately large (2.4 -3.4% SL); outer pseudoclasper with hook-shaped extension oriented backward at 90° angle and slightly bent outwards and with rounded tip; inner pseudoclasper short, joined to outer pseudoclasper Table 14. Meristic and morphometric characters of Diancistrus fuscus (Fowler, 1946). Holotype ANSP 72070 Holotype + 27 specimens Mean (range) n Standard length in mm 60 50.4 (27-66) 28 Meristic characters Dorsal fin rays 82 79.4 (74-85) 28 Anal fin rays 65 64.7 (58-69) 28 Pectoral fin rays - 18.3(17-20) 19 Precaudal vertebrae 11 11 28 Caudal vertebrae 33 32.8 (32-34) 28 Total vertebrae 44 43.8 (43-45) 28 Rakers on anterior gill arch 16 15.2(13-17) 22 Pseudobranchial filaments 1 1.7 (0-2) 21 D/V 7 6.3 (6-7) 28 D/A 20 19.4(18-22) 28 V/A 14 13.2(12-14) 28 Morphometric characters in % of'SL 1 lead length 25.3 26.9 (25.3-28.9) 24 Head width 13.9 12.7(11.0-15.7) 24 Head height 15.8 18.1 (15.8-21.2) 22 Upper jaw length 13.2 13.1 (11.8-14.4) 25 Maxillary height 4.1 4.4 (3.9-5.0) 24 Diameter of pigmented eye 2.7 2.9 (2.4-3.4) 24 Interorbital width 7.0 6.0 (4.1-8.5) 25 Postorbital length 17.6 19.3(17.6-21.4) 25 Prcanal length 47.4 47.8 (43.9-57.3) 25 Predorsal length 27.7 33.3 (27.7-35.8) 24 Body depth at origin of anal fin 17.6 18.5(16.7-20.6) 25 Pectoral fin length 14.3 16.0(14.0-21.8) 22 Pelvic fin length 21.3 23.7(19.1-28.1) 20 Base pelvic fin - anal fin origin 28.0 26.9 (23.7-33.2) 25 anteriorly, with supporter, with small hook and wide thin flap at its base; scale patch on cheek with 5-6 scale rows on upper cheek, scales on operculum only above opercular spine, 3-7 scales in 2 rows; head profile slender, broad; otolith length to sulcus length <2.0, sulcus not inclined. Description. The principal meristic and morphometric characters are shown in Table 14. Body compact with blunt, fleshy snout; fishes mature at about 45 mm SL. 1 lead with scale patch on cheek containing 5-6 scale rows on upper cheek, 4-7 (rarely 3) scales on operculum above opercular spine in two rows. Horizontal diameter of scales on body about 1.2% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin Fig. 30. Diancistrus fuscus (Fowler, 1946). Holotype. ANSP 72070, male, 60 mm SL. 113 W. Schwarzhans, P. R. Mollerand J. G. Nielsen Fig. 31. Diancistrus fuscus (Fowler, 1946). A, lateral view of head. YCM-P 36436, female, 62 mm SL; B, ventral view of head, YCM-P 36596, female, 48 mm SL; C, Lateral view of head, USNM 372960, male, 67 mm SL: I). ventral view of male copulatory organ, USNM 384596, 56 mm SL; E, view of left pseudoclasper from inside, USNM 384596, 56 mm SL; F, view of left pseudoclasper from inside, USNM 384595,47 mm SL; G. inclined lateral view of male copulatory organ, USNM 384596,56ntm SL; U, median view of right otolith, USNM 372960, male, 66 mm SL; I, ventral view of right otolith, USNM 372960, male, 66 mm SL; K, median view of right otolith, NSM1 P. 49441, female, 62 mm SL; L, ventral view of right otolith, NSMT P. 49441, female, 62 mm SL. of eye. Posterior nostril small, the size of anterior nostril and about 1/6—1/8 the size of eye. Head sensory pores (Fig. 31 A-C). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of a 60 mm SL, male non-type - USNM 374177). Premaxilla with 4 rows of granular and 1 row of larger teeth at symphysis, followed by 5 rows of granular teeth posteriorly. Largest teeth up to 1/4 pupil diameter. Vomer horseshoe-shaped, with 3 tooth rows, posterior row with larger teeth. Palatine teeth in 3 rows anteriorly and 2 rows posteriorly, slightly larger on inner row. Dentary with 5 outer granular tooth rows and 1 inner row of larger teeth anteriorly, blending into 1 row of about 6 large fangs posteriorly, which arc up to about 2/3 of pupil diameter. Otolith (Fig. 31 H-L). Elongate, length to height 2.0-2.1 (47-67 mm SL); otolith length to sulcus length 1.8-2.0; sulcus not inclined. Posterior tip of otolith broad. Dorsal rim flat in otoliths of males, gently curved without postdorsal angle in otoliths of females. Otoliths of females more thickset than those of males. This subtle sexual dimorphism is opposite to the trend observed in most other species of the genus Diancistrus. 114 Dinematichthyine fishes of the Indo-West Pacific Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 (9) depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10 (11). First anal fin pterygiophore elongated, almost reaching tip of last precaudal parapophysis in males, shorter in females. Male copulatory organ { Fig. 31 D-G). Two pairs of large pseudoclaspers. Outer pseudoclasper with hook-shaped extension oriented backward at 90° angle and slightly bent outwards and with rounded tip: inner pseudoclasper short, joined to outer pseudoclasper anteriorly, with supporter, with small hook and wide thin flap at its base. Isthmus between pseudoclaspers narrow. Penis long, rather straight, with broad base and thin tip. Coloration. Live colour brown (according to Fowler 1946); specimen photographed by J. T. Williams (USNM 372960) dark brown with translucent fins, light yellow at base. Uniformly medium to dark brown when preserved. Comparison. Diancistrus fuscus belongs to the large group of species with hook- or stick-shaped outer pseudoclaspers, which further contains the species D. alleni, D. beateae, D. leisi n. sp., D. longifilis, D. machidai n. sp., D. manciporus n. sp., D. novaeguineae, D. springeri n. sp. and D. vietnamensis n. sp. Of those, D. beateae is readily distinguished by the complete scale coverage on the operculum, whereas D. leisi n. sp., D. longifilis and D. manciporus n. sp. lack scales on the operculum. Diancistrus machidai n. sp. has specimens without, or with only 1 or 2, scales above the opercular spine, but is distinguished by the lower number of dorsal fin rays (66-77 versus 74-85, usually 77-84), the lower number of vertebrae (40-43 versus 43-45), the straight outer pseudoclasper (versus slightly bent outwards) and the inner pseudoclasper being narrow and pointed (versus short and with small hook). Diancistrus leisi n. sp. and D. springeri n. sp. differ in the straight, stick-like outer pseudoclasper. Diancistrus alleni, D. novaeguineae and D. vietnamensis n. sp. are the species closest to D. fuscus. Of these D. novaeguineae is recognized by the broad, fleshy appendix of the inner pseudoclasper and generally larger number of scales above the opercular spine (7-16 versus 3-7). Diancistrus fuscus, D. alleni and D. vietnamensis n. sp. all share the same pseudoclasper pattern. Diancistrus alleni differs from the other two in the smaller number of scales above the opercular spine (1-2, rarely 3 versus 3-7). Finally, D. fuscus differs from D. vietnamensis n. sp. in the usually higher number of dorsal fin rays (74-85 versus 69-80), the D/V (6, rarely 7 versus 7), the head profile (low versus high neck) and the relative sulcus length (otolith length to sulcus length <2.0 versus >2.3). At the Ryukyu Islands and northern Philippines D. fuscus co-occurs with two small-eyed species, D. erythraeus and D.jackrandalli n. sp., and at the northern Philippines also with D. karinae n. sp., which is readily distinguished by its continuous scale patch above and below the opercular spine. Possibly there is yet another Diancistrus species at the Ryukyu Islands (mentioned later as Diancistrus sp. 1). Distribution (Fig. 57). Diancistrus fuscus is known from the Ryukyu Islands in the north and Taiwan to the northern Philippines southward to approximately 10°N. Further to the south, across the equator, D. fuscus is replaced by D. alleni. In the west along Flainan and northern Vietnam it is replaced by D. vietnamensis n. sp. Remarks. ‘Brotulina fusca’ is used as a trade name in the aquarium industry. Specimens purchased from vendors and said to be from Bali have proven to belong to D. altidorsalis (see respective account). Diancistrus jackrandalli n. sp. (Figs 29, 32, 33; Table 15) Material examined. (3 specimens, 51-62 mm SL). FIolotype - BPBM 40213, male, 51 mm SL, Ishigaki- jima, Ryukyu Islands, reef about 1/2 mile off harbour of Ishigaki City; 20-35 feet, J.E. Randall and A.H Banner, 22 May 1968. Paratypes - BPBM 40214, 1 female, 51 mm SL, same data as holotype; YMC-P 34215, 1 female, 62 mm SL, Kakeroma Island, Amami Islands, northern Ryukyu Islands. Table 15. Meristic and morphometric characters of Diancistrus jackrandalli n. sp. Holotype BPBM 40213 Holotype + 2 paratypes Mean (range) n Standard length in mm Meristic characters 51 54.7 (51-62) 3 Dorsal fin rays 77 77.3 (76-79) 3 Anal fin rays 64 63.7 (60-67) 3 Pectoral fin rays 21 21 3 Precaudal vertebrae 11 11 3 Caudal vertebrae 30 29.7 (29-30) 3 Total vertebrae 41 40.7 (40-41) 3 Rakers on anterior gill arch 15 15.5(15-16) 2 Pseudobranchial filaments 0 0.7 (0,2) 3 D/V 6 6.3 (6-7) 3 D/A 21 21.3(21-22) 3 V/A 13 Morphometric characters in % of SL 13.7(13-14) 3 Head length 27.8 27.5 (26.9-28.0) 3 1 lead width 14.5 15.3(14.5-16.1) 2 Head height 19.5 19.0(18.6-19.5) 2 Upper jaw length 13.9 14.0(13.4-14.7) 3 Maxillary height 4.9 4.7 (4.4-4.9) 2 Diameter of pigmented eye 1.6 1.8 (1.6-2.1) 3 Interorbital width 7.8 7.0 (5.5-7.8) 3 Postorbital length 20.5 20.0(19.0-20.6) 3 Preanal length 49.1 50.7 (48.3-54.5) 3 Predorsal length 31.6 32.2 (31.5-33.4) 3 Body depth at origin of anal fin 20.5 20.2(19.2-20.8) 3 Pectoral fin length 15.0 15.8(14.9-17.4) 3 Pelvic fin length 26.6 25.1 (23.6-26.6) 2 Base pelvic fin - anal fin origin 28.8 27.5 (24.8-28.9) 3 115 W. Schwarzhans, P. R. Mollerand J. G. Nielsen Fig. 32. Diancistrus jackrandalli n. sp. Holotype. BPBM 40213, male, 51 mm SL. Diagnosis. Vertebrae 11+29-30=40-41, dorsal fin rays 76-79, anal fin rays 60-67; eyes small (1.6-2.1% SL); outer pseudoclasper moderately large, wing- shaped, slightly bent outwards; inner pseudoclasper almost free, bifurcate, with stout hook-like anterior branch and broad flap-like posterior branch; scale patch on cheek with 6-7 rows of small scales on upper cheek, 2 large scales on operculum above opercular spine; head massive, broad; otolith length to height ratio 2.1. Diancistrus jackrandalli is the only species of the genus Diancistrus, which combines small eyes (<2.1% SL) with presence of scales on the operculum. Description. The principal meristic and morphometric characters are shown in Table 15. Body moderately compact, with massive head; fishes mature at more than 50 mm SL. Head with scale patch on cheek with 6-7 rows on the upper cheek, 2 large scales on operculum above opercular spine. Horizontal diameter of scales on body about 1.3% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner. Anterior nostril low on snout, 1/3 distance from lip of snout to anterior margin of eye. Posterior nostril very small, the size of anterior nostril and about 1/8 the size of eye. Head sensory pores (Fig. 33 A). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 row of larger teeth at symphysis, followed by 4 rows of granular teeth posteriorly. Anterior- most large teeth about 1/3 of pupil diameter. Vomer horseshoe-shaped, with 4 tooth rows anteriorly, and 1 row posteriorly. Teeth of equal size. Palatine teeth in 3 rows anteriorly and 2 rows posteriorly. Teeth of equal size. Dentary with 2 outer rows of granular teeth and 2 inner rows of larger teeth anteriorly, blending into 1 row of 11 large fangs posteriorly, fangs up to about 1/2 pupil diameter. Otolith (Fig. 33 B). Moderately elongate with pointed anterior and expanded posterior tips, length to height 2.1 (51 mm SL); otolith length to sulcus length 2.1; sulcus Fig. 33. Diancistrus jackrandalli n. sp. A, lateral view of head, holotype; B, median view of right otolith, YMC 34215, female, 62 mm SL; C, view of left pseudoclasper from inside, holotype; D, inclined lateral view of male copulatory organ, holotype. 116 Dinematichthyine fishes of the Indo-West Pacific slightly inclined at 5°. Dorsal rim with shallow postdorsal area. Axial skeleton (of holotype). Neural spine of vertebrae 4 inclined and 5-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10 (11). First anal fin pterygiophore slightly elongated, not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 33 C-D). Two pairs of moderately large pseudoclaspers. Outer wing-shaped, slightly bent outwards, with broad supporter; inner pseudoclasper almost free, bifurcate, with stout hook¬ like anterior branch and broad flap-like posterior branch. Isthmus between pseudoclaspers wide. Penis short, curved, broad at base with slender tip. Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Diancistrus jackrandalli belongs to the small group of species with wing-shaped outer pseudoclaspers and bifurcate inner pseudoclaspers, which include D. alatus, D. brevirostris and D. fijiensis. Prom these it is readily distinguished by the small eye (<2.1% versus >2.2% SL), the presence of scales above the opercular spine (versus absent) and the bifurcate but not spiny inner pseudoclaspers (versus spiny). In the Ryukyu Islands, D.jackrandalli co-occurs with D- erythraeus , a small-eyed species with ear-lobe shaped outer pseudoclaspers, D. fnscus and an undescribed species, Diancistrus sp.l, which all belong to different species groups and are easily distinguished. Distribution (Fig. 29). Diancistrus jackrandalli is only known from the Ryukyu Islands. Etymology. Named in honor of John (Jack) E. Randall, Honolulu, BPBM, Hawaii, in recognition of his many contributions to ichthyology. Diancistrus jeffjolmsoni n. sp. (Figs 34, 35, 40; Tabic 16) Material examined. (24 specimens, 37-79 mm SL). Holotype - QM 1. 33769, male, 50 mm SL, 17°08’S, 139°36’E, Sweers Island, Gulf of Carpentaria, Australia, rocky reef with slabs of dead coral, large caves, brown algae at 2-3.5 m, J. Johnson and A. Gill, 15 Nov. 2002. Paratypes - AMS I. 24678-010, 3 males, 43-51 mm SL and 2 females, 48-59 mm SL, 12°25’S, 130°49’E, Darwin Harbour, Northern Territory, Australia, D.F. Floese and S. Reader, 31 Aug. 1984; QM I. 33742, 1 male, 41 mm SL and 2 females, 49-79 mm SL, 16°38’S, 139°53’E. Mornington Island, Gulf of Carpentaria, Queensland, Australia, J. Johnson and A.C. Gill, 22 Nov. 2002; WAM P. 27967-005, 2 males, 43-60 mm SL and 1 female, 61 mm SL, 24°29’S, H3°25’E, Carnarvon, Western Australia, J.B. Hutchins et < 25 April 1983; WAM P. 27980-064, 1 male, 53 mm SL, 20°26’S, 115°35’E, Montebello Islands, Western Australia, G.R. Allen, 25 May 1983; WAM P. 30920-002, 1 male, 46 tom SL and 3 females, 41-49 mm SL, I5°59’S, 124°17’E, Heywood Islands, Western Australia, J.B. Hutchins, 22 Nov. 1994; WAM P. 31251-055, 1 male, 51 mm SL and 3 Table 16. Meristic and morphometric characters of Diancistrus jeffjohnsoni n. sp. Holotype QM 33769 Holotype + 23 paratypes Mean (range) n Standard length in mm 50 51.6 (37-79) 24 Meristic characters Dorsal fin rays 70 73.5 (70-77) 24 Anal tin rays 55 58.2 (55-63) 24 Pectoral fin rays 17 17.5 (17-18) 22 Precaudal vertebrae 12 11.9(11-12) 24 Caudal vertebrae 31 31.3 (30-32) 24 Total vertebrae 43 43.2 (42-44) 24 Rakers on anterior gill arch 16 15.2(14-17) 23 Pseudobranchial filaments 2 1.9 (1-2) 23 D/V 7 6.7 (6-7) 24 D/A 21 20.4(19-23) 24 V/A 15 14.4(14-15) 24 Morphometric characters in % of SL Head length 25.4 25.8 (24.6-26.9) 22 1 lead width 12.0 12.7(11.4-14.7) 21 Head height 15.4 15.4(14.3-17.3) 21 Upper jaw length 12.2 12.7(12.2-13.4) 22 Maxillary height 4.1 4.1 (3.7-4.6) 22 Diameter of pigmented eye 3.0 2.7 (2.4-3.0) 23 Interorbital width 6.4 6.3 (5.6-7.1) 23 Postorbital length 18.7 18.8(17.8-20.2) 21 Preanal length 50.6 49.9 (46.6-55.2) 23 Predorsal length 32.4 32.7 (29.1-35.0) 23 Body depth at origin of anal fin 17.4 17.0(15.4-18.6) 23 Pectoral fin length 15.7 15.4(13.3-16.7) 23 Pelvic fin length 24.5 24.2 (21.0-27.4) 21 Base pelvic fin - anal fin origin - 30.0 (27.5-34.0) 21 females, 57—59 mm SL, 15°55’S, 124°03’E, Montgomery Reef, Kimberley, Western Australia, S. Morrison, 3 Dec. 1996; WAM P. 31389-001, 1 female, 37 mm SL, 16°5TS, 122°40’E, Beagle Bay, Lacepede Islands, Western Australia, J.B. Hutchins, 27 Aug. 1997; ZMUC P 771478, 1 male, 44 mm SL and ZMUC P 771479,1 female, 60 mm SL, same data as WAM P. 27967-005. Diagnosis. Vertebrae (11-) 12+ 30-32=42-44, dorsal fin rays 70-77, anal fin rays 55-63; eyes moderately large (2.4-3.0% SL), D/V 6-7; outer pseudoclasper large, twice the length of the inner pseudoclasper, wing-shaped, with narrow base, flat and its posterior part strongly bent backward; inner pseudoclasper almost completely joined to outer pseudoclasper anteriorly, thin, small, with single tip, not extending over anterior part of outer pseudoclasper; narrow scale patch on cheek with 3—5 scale rows on upper cheek, single large scale on operculum above opercular spine, usually covered by mucus and skin; head slender, elongate (head depth 14-17% SL); 1st and 2nd lower preopercular pore with separate opening (not fused as in all other species of the genus); otolith slender, length to height ratio 2.2-2.3, otolith length to sulcus length 2.2-2.4, dorsal rim regularly curved without angles. 117 W. Schvvarzhans, P. R. Mollerand J. G. Nielsen Fig. 34. Diancislrus jeffjohnsoni n. sp. Holotype. QM 33769, male, 50 mm SL. Description. The principal meristic and morphometric characters are shown in Table 16. Body and head slender, snout pointed; fishes mature at about 40-45 mm SL. Head with narrow scale patch on cheek with 3-5 scale rows on upper cheek, single large scale on operculum above opercular spine, usually covered by mucus and skin (seemingly a consistent character since no specimens with 0 or 2 such scales has been found). Horizontal diameter of body scales 1.3% SL. Maxillaries expanded posteriorly with angle at rear ventral corner and sometimes a knob in front of rear angle. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril large, larger than anterior nostril and about 1/4 size of eye. Head sensory pores (Fig. 35 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior), the 2nd anterior mandibular pore unusually large, about twice the size of posterior mandibular pores. Preopercular pores 4 (3 lower and 1 upper); 1st and 2nd lower preopercular pores separated. Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 tooth row anteriorly, including a few fang-like teeth closest to symphysis, up to 1/2 pupil diameter. Five rows of granular teeth posteriorly. Vomer teeth in 2 rows, with a few fangs on inner row posteriorly. Palatine teeth in 2 rows, teeth on inner row about 1/3 of pupil diameter, more than twice as long as teeth on outer row. Dentary with 4 rows of granular teeth and 1 tooth row anteriorly, blending into one row of about 10 large, curved, fang-like teeth posteriorly, fangs more than 1/2 pupil diameter. Otolith (Fig. 35 H). Elongate with pointed anterior and rounded posterior tip, length to height 2.2-2.3 (51-59 Fig. 35. Diancistrus jeffjohnsoni n. sp. A, lateral view ofliead, AMS I. 24678-010. male, 51 mm SL; B, ventral view of head, AMS I 24678-010, male, 47 mm SL; C, view of left pseudoclasper from outside, WAM R 31251-055, 51 mm SL; D, view of left pseudoclasper from ventral, WAM P. 31251-055, 51 mm SL; E, view of left pseudoclasper from inside, WAM P. 31251-055, 51 mm SL; F, view of left pseudoclasper from inside, holotype; G, inclined lateral view of male copulatory organ, WAM P. 31251-055, 51 mm SL; H, median view of right otolith, WAM P 31251-055, 51 mm SL. Dinematichthyine fishes of the Indo-West Pacific mm SL); otolith length to sulcus length 2.2-2.4; sulcus not inclined; dorsal rim regularly curved without angles, almost symmetrical to ventral rim. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to (11) 12. Pleural ribs on vertebrae 2-11. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis in males, slightly shorter in females. Male copulatory organ (Fig. 35 C-G). Two pairs of large pseudoclaspers. Outer pseudoclasper long, twice the length of inner pseudoclasper, wing-shaped, with narrow base, flat, with posterior part strongly bent backward, its supporter slender and strongly curved following curvature of the outer pseudoclasper; inner pseudoclasper almost completely joined to outer pseudoclasper anteriorly, thin, small, with single tip, not extending over anterior part of outer pseudoclasper, with small, slender supporter. Isthmus between pseudoclaspers narrow. Penis long, rather straight, thin. Coloration. Live colour unknown. Uniformly light brown or greyish when preserved. Comparison. Diancistrus jeffjohnsoni belongs to the group with a wing-shaped outer pseudoclasper and a single-tipped inner pseudoclasper, which includes D. eremitus, D. karinae n. sp., D. katrineae n. sp., D. mennei n. sp. and D. pohnpeiensis n. sp. It is easily distinguished from these species by the characters listed in the diagnosis, particularly the single scale above the opercular spine or the low number of dorsal fin rays (<77 versus >78). Its pseudoclasper pattern resembles best that of D. karinae n. sp. (see respective account). In fact, D. jeffjohnsoni is readily distinguished from all other Diancistrus species as being the only one with predominantly 12 precaudal vertebrae (versus 11, which also occurs, though rarely, in D. jeffjohnsoni). It is furthermore one of the few Diancistrus species that can be easily recognized by means ol otoliths with their elongate shape and the regularly curved dorsal rim. Distribution (Fig. 40). Diancistrus jeffjohnsoni is distributed along the northern coast of Australia, from 24°S, 113°E, just north of Shark Bay, Western Australia, to 17°S, 139°E, in the Gulf of Carpentaria, Queensland. Etymology. Named in honour of Jeff Johnson, Queensland Museum, Brisbane, who has collected the holotype specimen. Diancistrus karinae n. sp. (Figs 36, 37, 40; Table 17) Material examined. (16 specimens, 35-82 mm SL). Holotype - BPBM 36712, male, 64 mm SL, 01°N, 124°E, Manado, Bunaken, Sulawesi, Indonesia, base of drop off at 16 m, J.E. Randall, M. Severns and R.C. Pyle, 29 Oct. 1991. Paratypes - AMS I. 18677-019, 1 female, 82 mm SL, Philippines, J.D. Lewis, 1974; BPBM 9938,2 females, 41-55 mm SL, 07°N, 134°E, Palau, J.E. Randall, A.R. Emery and P. Helfman, 22 April 1970; BPBM 28527, 1 female, 67 mm SL, Sumilon Isl. SE of Cebu, Philippines, J.E. Randall, M. Gawel and K.E. Carpenter 3 June 1981; BPBM 31454,1 female, 62 mm SL, 07°N, 134°E, Palau, J.E. Randall and R.C. Pyle, 15 July 1986; ROM 55147, 1 female, 48 mm SL, 09°N, 123°E, Visayan Islands, Philippines, June 1988; USNM 224328, 1 male, 79 mm SL and 1 female, 35 mm SL, 06°56’N, 158°06’E, Pohnpei Island, Pohnpei State, Federated States of Micronesia, V.G. Springer et al. , 11 Sept. 1980; USNM 263661, 1 male, 75 mm SL and 1 female, 35 mm SL, 09°N, 118°E, Puerto Princesa, Palawan, Sulu Sea, Philippines, R.E. Schroeder and G. Hendler, 7 Aug. 1979; USNM 263682, 1 male, 56 mm SL, 01°33’S, 144°59’E, Hermit Island, Papua New Guinea, V.G. Springer et al., 2 Nov. 1978; ZMUC P 771476, 1 male, 59 mm SL, same data as USNM 263682; ZMUC P 771477, 1 female, 75 mm SL, same data as USNM 263661. Additional specimens. USNM 366603, 1 female, 54 mm SL, 09°31’N, 123°40’E, Bohol Island, Philippines; USNM 366680, 1 female, 47 mm SL, 09°N, 118°E, Puerto Princesa, Palawan, Philippines. Diagnosis. Vertebrae 11-12+32-35=43-46, dorsal fin rays 78-83, anal fin rays 61-68; eyes very large (2.7-3.8% SL), D/V 7; outer pseudoclasper large, about 1 1/2 length of inner pseudoclasper, wing-shaped, flat and its posterior Table 17. Meristic and morphometric characters of Diancistrus karinae n. sp. Holotype BPBM 36712 Holotype + 13 paratypes Mean (range) n Standard length in mm 61 58.1 (35-82) 14 Meristic characters Dorsal fin rays 80 80.6 (78-83) 13 Anal fin rays 64 64.3 (61-68) 13 Pectoral fin rays 19 18.9(18-21) 14 Precaudal vertebrae 11 11.1 (11-12) 13 Caudal vertebrae 34 33.4 (31-35) 13 Total vertebrae 45 44.5 (43-46) 13 Rakers on anterior gill arch 13 15.3(13-17) 14 Pseudobranchial filaments 2 2 13 D/V 7 7 13 D/A 19 19.6(18.0-21.0) 13 V/A 14 13.2(13.0-14.0) 13 Morphometric characters in % of SL Head length 27.8 27.0 (25.4-29.1) 14 Head width 15.8 14.1 (11.1-16.8) 13 Head height 16.7 16.9(15.7-18.8) 13 Upper jaw length 14.2 13.2(12.3-14.2) 14 Maxillary height 5.0 4.6 (4.0-5.1) 13 Diameter of pigmented eye 3.1 3.1 (2.7-3.8) 14 Interorbital width 7.4 6.9 (6.0-7.5) 13 Postorbital length 19.6 19.3(18.0-21.2) 13 Preanal length 48.9 47.4(43.3-51.4) 14 Predorsal length 35.5 34.3 (31.9-35.6) 14 Body depth at origin of anal fin 20.3 18.7(16.4-20.4) 13 Pectoral fin length 17.9 16.7(13.3-19.6) 14 Pelvic fin length 22.9 23.3 (21.6-25.1) 11 Base pelvic fin - anal fin origin 28.6 27.7(23.2-31.3) 14 119 W. Schvvarzhans, P. R. Moller and J. G. Nielsen Fig. 36. Diancistrus karinae n. sp. Holotype. BPBM 36712, male, 64 mm SL. part strongly bent backward; inner pseudoclasper joined to outer pseudoclasper anteriorly, long, single-pointed tip strongly extending over anterior part of inner pseudoclasper; scale patch on cheek with 5-6 scale rows on upper cheek, large scale patch on operculum continuous above and below opercular spine; head high, usually high- necked; otolith length to height ratio 2.0-2.1. Description. The principal meristic and morphometric characters are shown in Table 17. Body moderately compact, often high-necked, highest near origin of dorsal fin; fishes large, mature at about 50 to 55 mm SL. Head with moderately broad scale patch on cheek with 5-6 scale rows on upper cheek, large scale patch on operculum forming a continuous patch above and below opercular spine. Horizontal diameter of body scales 2.2% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril small, the size of anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 37 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and I upper). Dentition (of holotype). Premaxilla with 5 rows of granular teeth and 2 tooth rows anteriorly, including a few fang-like teeth closest to symphysis, up to 1/2 pupil diameter. Five granular rows of teeth posteriorly. Vomer teeth in 2 rows, with 2-3 fangs on inner row posteriorly. Palatine teeth in 2 rows, teeth on inner row about 1/3 of pupil diameter, more than twice that of teeth on outer row. Dentary with 4 granular rows and 2 tooth rows anterior, blending into one row of about 10 large, curved, fang-like teeth posteriorly, fang size more than 1/2 pupil diameter. Otolith (Fig. 37 G-I). Moderately elongate with rounded anterior and expanded posterior tips, length to height 2.0-2.1 (64-79 mm SL); otolith length to sulcus length 2.0-2.1; sulcus slightly inclined at 5°. Broad and rounded postdorsal angle in otoliths of males, no postdorsal angle and gently curved dorsal rim otoliths of females. Otoliths of males are slightly thicker than those of females. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-11. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis in males, slightly shorter in females. Male copulatory organ (Fig. 37 C-F). Two pairs of large pseudoclaspers. Outer pseudoclasper long, about 1 'A length of inner pseudoclasper, wing-shaped, with narrow base, flat and its posterior part strongly bent backward, its supporter slender and strongly curved with curvature of outer pseudoclasper; inner pseudoclasper joined to outer pseudoclasper anteriorly, long, single-pointed tip strongly extending over anterior part of outer pseudoclasper, with large supporter. Isthmus between pseudoclaspers wide. Penis long, curved, thin. Coloration. Live colourreddish brown. Uniformly light brown when preserved. Comparison. Diancistrus karinae belongs to the group with wing-shaped outer pseudoclasper and a single-tipped inner pseudoclasper, which includes D. jeffjohnsoni, D. katrineae n. sp., D. eremitus, D. mennei n. sp. and D. pohnpeiensis n. sp. In Pohnpei it co-occurs with D. pohnpeiensis n. sp. and in Hermit Island with D. eremitus , but is distinguishable by its complete coverage of scales on the operculum above and below the opercular spine. The shape of the flat outer pseudoclasper with the slender, strongly backward curved supporter resembles best that of D. jeffjohnsoni, but the pseudoclaspers of D. karinae are characterized by a rather strong and long inner pseudoclasper. Together with D. beateae, D. karinae is the only species of the genus Diancistrus with a continuous scale patch on the operculum above and below the opercular spine. Diancistrus karinae differs in the pseudoclasper morphology (flat wing-shaped versus broad hook-shaped), the smaller number of dorsal fin rays (78-83 versus 84-91) and the higher D/V (7 versus 6). It is possible that the distribution ranges of the two species overlap marginally, but even then female specimens of the two species can be distinguished by the dorsal fin ray counts. Distribution (Fig. 40). Diancistrus karinae is widely distributed in the northern West Pacific, from Palawan and the Visayan Islands of the northern Philippines to the northern tip of Sulawesi in the west and to the oceanic islands Palau, Pohnpei and Hermit Island in the east. 120 Dinematichthyine fishes of the Indo-West Pacific Fig. 37. Diancistrus karinae n. sp. A, lateral view of head, USNM 263682, male, 69 mm SL; B, ventral view of head, USNM 263682, male, 69 mm SL; C, ventral view of malecopulatory organ, USNM 224328, 79 mm SL; D, view of left pseudoclasper from inside, USNM 224328, 79 mm SL; E, view of left pseudoclasper from inside, holotype; F, inclined lateral view of male copulatory organ, holotype; G, median view of right otolith, USNM 224328, male, 79 mm SL; II, ventral view of right otolith, USNM 224328, male, 79 mm SL; I, median view of right otolith, AMS 1. 18677-019, female, 82 mm SL. Etymology, Named after Karin Bloch for her most valuable support of her husband Jorgen Nielsen during his many hours invested in this study. Diancistrus katrineae n. sp. (Figs 38, 39, 40; Table 18) Material examined. (16 specimens, 36-102 mm SL). Holotype - BPBM 40215, male, 71 mm SL, 27°S, 144°W, Katrineae, SW side of Kakatrineaeo lti, Tubuai Islands, French Polynesia, 20 m in cave, J.E. Randall and D. Bryant, 16 Feb. 1971. Paratypes - AMS 1. 253780-008. 1 female, 59 mm SL, I4°53’S, 148°43’E, Matahiva, Huahine, Tubuai Islands, French Polynesia, J. Bell, 1983; AMS I. 28950-064, 1 male, 49 mm SL and 1 female, 49 mm SL, 17°29’S, 149°5FW, Moorea, Tahiti, Society Islands, French Polynesia, T. Trnski, A. Lefevre and R. Galzin, 22 Feb. 121 W. Schwarzhans, P. R. Moller and J. G. Nielsen 1989; BPBM 6093, 1 male, 76 mm SL, 17°S, 149°W, Tahiti, Society Islands, French Polynesia, J.E.Randall, Walters, D. Devaney and Richert, 4 Sept. 1967; BPBM 8358, 1 female, 46 mm SL, Papote Bay, Tahiti, Society Islands, French Polynesia, J.E.Randall and Adam, 6 March 1969; BPBM 9935, 2 females, 36-41 mm SL, Tahiti, J.E. Randall, 11 March 1964;BPBM 13934,2females,42-78mm SL,21°S, 159°W, Rarotonga, Cook Islands, J.E. Randall and D.B. Cannoy, 10 March 1971; BPBM 17291, 1 male, 102 mm SL and 2 females, 51-72 mm SL, same data as holotype; ZMUC P 771480, 1 female, 56 mm SL, same data as BPBM 13934. Tentatively assigned specimens: BPBM 12274, 1 female, 81 mm SL, 24°S, 124°W, Ducie Atoll, Pitcairn Islands, J.E. Randall, Costello, D.B. Cannoy and S. Christian, 15 Jan. 1971; BPBM 40216,1 female, 48 mm SL, 24°S, 124°W, Ducie Atoll, Pitcairn Islands, J.E. Randall, Costello, D.B. Cannoy and S. Christian, 14 Jan. 1971. Diagnosis. Vertebrae 11-12+33-35=44-46, dorsal fin rays 78-86, anal fin rays 63-69; eyes large (2.0-3.1% SL), D/V 6; outer pseudoclasper large, broad, wing-shaped, with concave inner face; inner pseudoclasper joined to outer pseudoclasper anteriorly, thin, with narrow base and single-pointed tip extending over anterior part of outer pseudoclasper; moderately broad scale patch on cheek with 5-8 scale rows, large scale patch on operculum above opercular spine (13-21 scales) and separated smaller scale patch below opercular spine (3-7 scales); head massive, thick. Description. The principal meristic and morphometric characters are shown in Table 18. Body moderately compact, with massive fleshy snout; fishes large, mature at about 50 mm SL. Head with moderately broad scale patch on cheek with 5-8 scales rows on upper cheek, large scale patch on operculum above opercular spine (13-21 scales) and separated smaller scale patch below opercular spine (3-7 scales). Horizontal diameter of body scales 2.0% SL. Maxillaries expanded posteriorly with distinct knob at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril small, slightly larger than anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 39 A-D). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular Table 18. Meristic and morphometric characters of Diancistrus katrineae n. sp. Holotype BPBM 40215 Holotype + 13 paratypes Mean (range) N Standard length in mm 71 59.1 (36-102) Meristic characters Dorsal fin rays 85 82.6 (78-86) 14 Anal fin rays 67 66.4 (63-69) 14 Pectoral fin rays 22 20.8 (20-22) 11 Precaudal vertebrae 11 11.1 (11-12) 14 Caudal vertebrae 34 33.7 (33-35) 14 Total vertebrae 45 44.8 (44-46) 14 Rakers on anterior gill arch 17 16.0(14-17) 13 Pseudobranchial filaments 2 2 14 D/V 6 6.1 (6-7) 14 D/A 22 21.1 (18-24) 14 V/A 14 13.4(13-14) 14 Morphometric characters in % of SL Head length 27.7 27.1 (24.9-28.2) 14 Head width 13.8 14.2(11.8-18.6) 12 1 lead height 17.8 19.3 (16.7-21.8) 13 Upper jaw length 14.3 13.1 (12.0-14.3) 14 Maxillary height 5.3 4.8 (3.9-5.3) 12 Diameter of pigmented eye 2.6 2.7 (2.0-3.1) 14 Interorbital width 7.2 5.7 (4.3-7.2) 14 Poslorbital length 20.0 19.5 (16.7-20.4) 13 Preanal length 46.3 47.6 (44.9-51.2) 14 Prcdorsal length 33.4 32.3 (28.0-34.2) 11 Body depth at origin of anal fin 20.2 18.7(15.5-21.4) 13 Pectoral fin length 16.0 16.1 (12.9-18.6) 12 Pelvic fin length 26.2 23.8(19.5-27.6) 10 Base pelvic fin - anal fin origin 26.5 26.2 (20.7-31.2) 14 pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 6 rows of granular teeth and 2 rows of larger teeth at symphysis, followed by 7 rows of granular teeth posteriorly. Vomer horseshoe-shaped, with 3 rows of teeth, with teeth in inner row only slightly larger than outer rows. Palatine teeth in 3 rows of similar sized teeth. Dentary with 5 outer rows of granular teeth and 1 inner row of larger teeth anteriorly blending into 1 row of about 10 large fangs posteriorly, fang size up to about 1/3 of pupil diameter. Otolith (Fig. 39 G-I). Moderately elongate with almost symmetrically pointed anterior and posterior tip, length to height 2.0 (49-81 mm SL); otolith length to sulcus length 2.1; sulcus slightly inclined at 5°. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 5-8 depressed. Parapophyses present Fig. 38. Diancistrus katrineae n. sp. Holotype. BPBM 40215, male, 71 mm SL. 122 Dinematichthyine fishes of the Indo-West Pacific Fig. 39. Dicincistrus katrineae n. sp. A, lateral view of head, BPBM 6093, male, 76 mm SI.; B, ventral view of head, BPBM 6093, male, 76 mm SL; C, lateral view of head, holotype; I), lateral view of head, BPBM 12274, tentatively assigned female, 81 mm SL; E, view of left pseudoclasper from inside, BPBM 17291, male, 102 mm SL; F, inclined lateral view of male copulatory organ, BPBM 17291, male, 102 mm SL; G, median view of right otolith, BPBM 12274, tentatively assigned female, 81 mm SL; II. median view of right otolith, AMS I. 28950-064, male, 49 mm SL; I, ventral view of right otolith, AMS I. 28950-064, male, 49 mm SL. 123 W. Schwarzhans, P. R. Moller and J. G. Nielsen from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, mostly reaching near to tip of last precaudal parapophysis. Male copulatory organ (Fig. 39 E-F). Two pairs of pseudoclaspers. Outer pseudoclasper large, wing-shaped, with concave inner face, with broad base and broad supporter; inner pseudoclasper small joined to outer pseudoclasper anteriorly, thin, with narrow base and single pointed tip extending over anterior part of outer pseudoclasper, with large supporter. Isthmus between pseudoclaspers moderately wide. Penis short, curved, thin. Coloration. Live colour bright yellow to yellow-orange, yellowish grey in small specimens. Uniformly light brown when preserved. Comparison. Diancistrus katrineae belongs to the subgroup characterized by a wing-shaped outer pseudoclasper and a single-tipped inner pseudoclasper. This subgroup includes D. eremitus, D. jeffjohnsoni, D. karinae, D. mennei n. sp. and D. pohnpeiensis n. sp. Diancistrus katrineae differs from these by having a scale patch above and one below the opercular spine on the operculum, and the concave inner face of the outer pseudoclasper. Diancistrus karinae has a continuous scale patch above and below the opercular spine, whereas the other species only have a single scale patch above the opercular spine. Diancistrus katrineae differs further from D. karinae in the more massive head and the lower D/V (6 versus 7). The only other species with separate scale patches above and below the opercular spine arc D. altidorsalis with the high-necked head profile and the distinctly ear-lobe outer pseudoclasper, and, in rare instances, D. novaeguineae, which has hook-shaped outer pseudoclaspers, and, from the genus Dinematichthys, Dinematichthys megasoma Machida, 1994, occasionally in very large specimens. In Tahiti, D. katrineae co-occurs with the small-eyed D. tongaensis n. sp. of the group with ear-lobe shaped outer pseudoclaspers. Also, there is evidence of a possible further species of the same (D. katrineae) subgroup from Rurutu, Tubuai Islands, mentioned later as Diancistrus sp. 2. Remarks. Diancistrus katrineae shows certain regional differences, most obviously in head squamation: the 4 specimens from Tahiti and the 3 from the Cook Islands seem to have consistently more scales on the lower opercular scale patch (Fig. 39 A) than those from the Tubuai Islands (5 specimens. Fig. 39 C) and Ducie Atoll (2 specimens, Fig. 39 D) (6-7 versus 2-3). Specimens from Tahiti have smaller eyes (2.0-2.2% SL versus generally 2.5-3.1% SL). No other characters support these differences. The specimens from Ducie Atoll are further characterized by lesser and larger scales on the cheek (5-6 versus 7-8 scale rows on upper cheek) (Fig. 39 D) and a much more compressed otolith (length to height ratio 1.8 versus 2.0) (Fig. 39 G). Since only two females are known from Ducie Atoll they are only tentatively included. Distribution (Fig. 40). Diancistrus katrineae is the most eastwardly distributed species of the genus, known from the Society Islands, the Cook and Tubuai Islands and reaching even as far as Ducie Atoll, Pitcairn Islands, the most isolated atoll in the south-eastern Pacific. O D. katrineae © D. eremitus © D. karinae O D. jeffjohnsoni © D. mennei Q D. pohnpeiensis 4 D. sp.2 Fig. 40. Sample sites of Diancistrus katrineae n. sp.. D. eremitus n. sp., D. karinae n. sp., D. jeffjohnsoni n. sp., D. mennei n. sp., D. pohnpeiensis n. sp. and Diancistrus sp. 2. One symbol may represent several samples. 124 Dinematichthyine fishes of the Indo-West Pacific Etymology. Named after Katrine Worsaae for her most valuable support of her husband Peter Rask Moller during his many hours invested in this study. Diancistrus leisi n. sp. (Figs 29, 41, 42; Table 19) Material examined. (4 specimens, 33-46 mm SL). Holotype - AMS I. 33740-061, male, 33 mm SL, 10°S, 144°E, Ashmore Reef, NE end, 10°09.58’S 144°34.94’E, 1-2 in depth, FNQ party, 25 Jan. 1993. Paratypes - AMS 1. 33740-062, 1 female, 46 mm SL, same data as holotype; AMS I. 33708-132, 1 female, 34 mm SL, Reef 10-418, Coral Sea 10°59.98’S 144°01.22’E, 2-9 m depth, FNQ team, 15 Jan. 1993; AMS I. 22582-020, 1 female, 52 mm SL, 15°49’S, 145°50’E, Cooktown area, Escape Reef North, outer barrier reef. Great Barrier Reef, Queensland, 14-17 m depth, AMS party, 29 Oct. 1981. Diagnosis. Vertebrae 11+30=41, dorsal fin rays 69-73, anal fin rays 55-57; eyes small (1.7—1.9% SL), D/V 6; outer pseudoclasper short, only slightly longer than inner pseudoclasper, stick-shaped; inner pseudoclasper anteriorly joined to outer pseudoclasper, thin, spiny, strongly forward inclined; narrow scale patch on cheek with 3-4 scale rows on upper cheek and only 1-2 scale rows on lower cheek, no scales on operculum; head high, laterally strongly compressed, eye located high; otolith length to height ratio 2.0-2.1. Description. The principal meristic and morphometric characters are shown in Table 19. Body slender, laterally compressed, with dorsally depressed head profile; fishes small, mature at about 30 mm SL. Head with narrow scale patch on cheek with 3-4 scale rows on upper cheek and only 1-2 scale rows on lower cheek, no scales on operculum. Horizontal diameter of body scales 1.2% SL. Maxillaries expanded posteriorly with angle at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril large, half-moon shaped due to large anterior flap, twice the size of anterior nostril and about 1/3 the size of eye. Head sensory pores (Fig. 42 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior), the 3rd posterior mandibular pore twice the size of the two other posterior mandibular pores. Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 tooth row anteriorly, including a few fang-like teeth closest to symphysis, up to 'A pupil diameter. Four rows of granular teeth posteriorly. Vomer teeth in 2 rows of equally sized teeth. Palatine teeth in 2 rows, teeth on inner row about 1/3 of pupil diameter, more than twice as long as teeth in outer row. Dentary with 4 rows of granular teeth and 2 tooth rows anteriorly, blending into one row of about 10 large, curved fang-like teeth posteriorly, size more than 1/2 pupil diameter. Table 19. Meristic and morphometric characters of Diancistrus leisi n. sp. Holotype AMS 1. 33740- 061 Holotype + 3 paratypes Mean (range) n Standard length in mm Meristic characters 33 39.5 (33-46) 4 Dorsal fin rays 70 70.7 (69-73) 3 Anal fin rays 57 56.0 (55-57) 3 Pectoral fin rays 19 19.8(19-21) 4 Precaudal vertebrae 11 11 3 Caudal vertebrae 30 30 3 Total vertebrae 41 41 3 Rakers on anterior gill arch 16 16.5 (16-18) 4 Pseudobranchial filaments 2 2 4 D/V 6 6 3 D/A 19 20.7 (19-22) 3 V/A 13 Morphometric characters in % of SL 13.3(13-14) 3 Head length 28.3 28.0 (26.5-29.8) 4 Head width 11.1 12.2(11.1-13.8) 4 Head height 17.3 16.3(15.1-17.3) 4 Upper jaw length 13.7 13.8(13.3-14.6) 4 Maxillary height 4.2 4.2 (3.8-4.8) 4 Diameter of pigmented eye 1.7 1.8 (1.7-1.9) 4 Interorbital width 7.3 7.3 (6.9-7.6) 4 Postorbital length 21.1 21.2 (20.8-22.2) 4 Preanal length 46.5 46.1 (43.9-48.4) 4 Predorsal length 33.3 33.1,(31.4-34.0) 4 Body depth at origin of anal fin 16.9 17.1 (16.5-17.6) 4 Pectoral fin length 16.0 16.0(14.4-18.0) 4 Pelvic fin length 27.1 27.2 (27.0-27.5) 4 Base pelvic fin - anal fin origin 27.1 27.1 (26.5-28.3) 4 Otolith (Fig. 42 E-F). Elongate with pointed anterior and posterior tips, length to height 2.0-2.1 (34-46 mm SL); otolith length to sulcus length 2.3-2.4; sulcus slightly inclined at 5°. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore slightly elongated, not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 42 C-D). Two pairs of rather small pseudoclaspers. Outer pseudoclasper short, only slightly longer than inner pseudoclasper, stick¬ shaped; inner pseudoclasper anteriorly joined to outer pseudoclasper, thin, spiny, strongly forward inclined; both inner and outer pseudoclasper nearly reduced to their respective supporters. Isthmus between pseudoclaspers narrow. Penis moderately long, curved, thin. Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Within the large group of species with hook- or stick-shaped outer pseudoclaspers, D. leisi belongs to the small group with the stick-shaped ones, together with D. springeri n. sp., from which it differs in the smaller eyes 125 W. Schwarzhans, P. R. Moller and J. G. Nielsen Fig. 41. Diancistrus leisi n. sp. Holotype. AMS 1. 33740-061, male, 33 mm SL. (<2.0% versus >2.5% SL), the absence of scales above the opercular spine (versus present), the narrow scale patch on the cheek (versus broad), the lower number of vertebrae (41 versus 44-45) and dorsal fin rays (69-73 versus 78-84) and the laterally strongly compressed head (versus not compressed). Besides the pseudoclasper pattern, the characteristic head squamation pattern, the laterally strongly compressed head and the dorsally shifted small eyes distinguish D. leisi from the many other Diancistrus species occurring along the reefs of the Coral Sea and the northern Great Barrier Reef. Distribution (Fig. 29). Diancistrus leisi is known from a few specimens from offshore reefs in the Coral Sea and outer reefs of the northern part of the Great Barrier Reef, Queensland, Australia. Ecology. The specimens were collected in the outer barrier reef at 2-17 m depth. Etymology. Named in honour of Jeff Leis, Sydney, Australia, in recognition of his many contributions to the study of fish larvae. Diancistrus iongifilis Ogilby, 1899 (Figs 12, 43, 44; Table 20) Diancistrus iongifilis Ogilby, 1899: 744, figs 59-60 (type locality: Lord Howe Island, off eastern Australia). Diancistrus Iongifilis - Paxton et al. 1989: 316; Nielsen etal. 1999: 129. Material examined. (120 specimens, 19-77 mm SL). Holotype -QM-1785, male, 76 mm SL, 31°S, 159°E, Lord Howe Island. Further specimens. AMS 1A. 2044, 1 male, 71 mm SL, 20°S, 148°E, Hayman Island, Great Barrier Reef, Queensland, Australia; AMS I. 17377-018, 2 males, 40-44 mm SL, 31°32’S, 159°04’E, Lord Howe Island, J.E. Randall, 24 Feb. 1973; AMS I. 17445-148, 3 males, 29-48 mm SL and 4 females, 42-63 mm SL, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia, 23°30’S 152°05’E, F.H. Talbot and party, 19 Sep. 1968; AMS I. 19108-142, 1 female, 36 mm SL, I4°40’S, 145°28’E, Lizard Island, Queensland, Great Barrier Reef, 1-10 m depth, D.F. Hoese and party, 17 Nov. 1975; AMS 1. Fig. 42. Diancistrus leisi n. sp. A, lateral view of head. AMS I. 33740-062, female, 46 mm SL; B, ventral view of head, AMS I. 33740- 062, female, 46 mm SL; C, view of left pseudoclasper from inside, holotype; D, inclined lateral view of male copulatory organ, holotype; E, median view of right otolith. AMS 1. 33740-062, female, 46 mm SL; F, ventral view of right otolith, AMS I. 33740-062, female, 46 mm SL. 126 Dinematichthyine fishes of the Indo-West Pacific 20201-043, 12 males, 45-76 mm SL and 3 females, 49-61 mm SL, 23°30’S, 152°05’E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia, 2 m depth, D.F. Hoese, 29 Sep. 1971; AMS I. 20208-004, 3 males, 37-58 mm SL and 1 female, 60 mm SL, 23°30’S, 152°05’E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia, D.F. Hoese, 27 Sept. 1971; AMS 1. 20210-004, 3 males, 44-49 mm SL and 2 females, 48-53 mm SL, 23°30’S, I52°04’E, Capricorn Group, One Tree Island, lagoon. Great Barrier Reef, Queensland, Australia, F.H. Talbot and party, 5 Oct. 1971; AMS I. 20211-004, 2 females, 49 mm SL, 23°S, 152°E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia, F.H. Talbot and party, 5 Oct. 1971; AMS I. 20561-004, 5 specimens, 23°S, 152°E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia; AMS I. 21540-065, 1 male, 50 mm SL, 14°S, 145°E, Lizard Island, Eagle Cay, Great Barrier Reef, Queensland, Australia, 14°39’S 145°27’E, D.F. Hoese and H. Larson, 12 Feb. 1975; AMS 1.22583-019,1 male, 52 mm SL and 1 female, 43 mm SL, 15°S, 145°E, Escape Reef, Great Barrier Reef; AMS 1. 22633-103, 1 male, 40 mm SL and 2 females, 27-38 mm SL, 15°49’S, 145°50’E, Escape Reef, outer barrier reef, Great Barrier Reef, Queensland, Australia, 6 Nov. 1981; AMS I. 27138-048, 1 female, 52 mm SL, 29°27’S, 159°05’E, Middleton Reef, A.C. Gill and S. Reader, 5 Dec. 1987; AMS 1.27155-013,2 males, 70-72 mm SL and 3 females, 57-65 mm SL, 29°56’S, 159°02’E, Elizabeth Reef, A.C. Gill, Cordall and Ledbitter, 11 Dec. 1987; AMS I. 34311-015, 1 male, 45 mm SL and 5 females, 37-54 mm SL, 22°14’S, 150°19’E, Collins Island, Cannibal Group, Great Barrier Reef, Queensland, Australia, 15 Sept. 1993; AMS I. 34311-016, 1 male, 41 mm SL and 3 females, 28-37 mm SL, 22°14'S, 150°19’E, Collins Island, Cannibal Group, Great Barrier Reef, Queensland, Australia, 15 Sept. 1993; BPBM 11419, 1 male, 69 mm SL, Noumea, outside of barrier reef. New Caledonia, J.E. Randall. P. Fourmanoirand R. Grandperrin, 14 Aug. 1971; BPBM 14943,2 females, 63-69mm SL, 31°S, I59°E, Lord Howe Island, J.E. Randall, B.C. Russell and B. Goldman, 26 Feb. 1973; BPBM 27168, 1 female, 50 mm SL, New Caledonia, Jan. 1979; MNHN 1980-0243, 1 male, 77 mm SL and 2 females, 35-66 mm SL, New Caledonia; MNHN 1980-0244, 2 males, 59-64 mm SL and 2 females, 27-37 mm SL, New Caledonia; MNHN 1980-0634, 1 female. Table 20. Meristic and morphometric characters of Diancistrus longifilis Ogilby, 1899 Holotype QM-I 785 Holotype + 119 specimens Mean (range) n Standard length in mm 76 53.6 (27-77) 69 Meristic characters Dorsal fin rays 83 81.2 (77-85) 36 Anal fin rays 65 64.8 (60-69) 36 Pectoral fin rays 20 19.1 (18-21) 22 Preeaudal vertebrae 11 11 37 Caudal vertebrae 33 33.0 (32-35) 37 Total vertebrae 44 44.0 (43-46) 37 Rakers on anterior gill arch 14 15.2(13.0-18.0) 30 Pseudobranchial filaments 2 2 (1-2) 26 D/V 7 6.4 (6-7) 37 D/A 22 21.6(18.0-25.0) 37 V/A 14 13.6(13.0-14.0) 38 Morphometric characters in % of SL Head length 26.3 26.5 (25.1-28.4) 30 Head width 17.1 13.2(10.1-17.1) 31 1 lead height 17.1 16.0(14.3-18.6) 31 Upper jaw length 13.7 13.1 (12.0-14.3) 30 Maxillary height 5.1 4.0 (3.5-5.1) 30 Diameter of pigmented eye 2.4 2.7 (2.4-3.2) 31 Interorbital width 7.3 6.7 (5.6-7.4) 31 Postorbital length 19.0 19.3(18.0-20.5) 23 Preanal length 42.8 47.5 (42.4-52.2) 31 Prcdorsal length 33.6 32.0 (30.4-33.6) 31 Body depth at origin of anal fin 18.6 17.0(14.0-18.6) 31 Pectoral fin length 16.2 15.5(12.9-17.7) 31 Pelvic fin length 23.2 23.1 (21.0-25.5) 21 Base pelvic fin - anal fin origin 27.9 28.1 (22.0-33.2) 30 52 mm SL, New Caledonia; MNHN 1980-0695, 1 female, 62 mm SL, New Caledonia; SMNS 19829, 2 females, 63-64 mm SL, 21°35’45”S, I67°50’06”E, Cap Wabao, 150 m NNE of Cape, Baie de Tadin, 6 km SW Tadin, W coast, Mare Island, Loyalty Islands, crevices of coralline rock adjacent to a surge channel [high energy area] of a seaward fringing reef, 2-3.8 m depth , R. Fricke, 12 Nov. 1997; USNM 263714, 1 male, 57 mm SL and 1 female, 57 mm SL, 23°S, 152°E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia; USNM 263724, 2 males, 55-56 mm SL, 23°S, I52°E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia; USNM 366562, 3 males and 4 females, 29-55 mm SL, 23°S, 152°E, Capricorn Group, One Tree Island, Fig. 43. Diancistrus longifilis Ogilby, 1899. Holotypc. QM-I 785, male, 76 mm SL. 127 W. Schwarzhans, P. R. Muller and J. G. Nielsen F op Fig. 44. Diancislrus longifilis Ogilby, 1899. A, lateral view of head, AMS I. 20201-043, male, 71 mm SL; B, ventral view of head, AMS 1. 20201-043, male, 71 mm SL; C, view of left pseudoclasper from inside, MNHN 1980-0243, 77 mm SL; D, view of left pseudoclasper from inside, MNHN 1980-0244, 64 mm SL; E, inclined lateral view of male copulatory organ, MNHN 1980-0244, 59 mm SL; F, view of left pseudoclasper from inside, AMS I. 20201-043, male, 45 mm SL; G, median view of right otolith, WAM P. 29637-023, male, 46 mm SL. Great Barrier Reef, Queensland, Australia; USNM 366586,4 females, 42-55 mm SL, 23°S, 152°E, Capricorn Group, One Tree Island, Great Barrier Reef. Queensland, Australia; USNM 366588, 1 male and 3 females, 54-63 mm SL, 23°S, 152°E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia; USNM VGS 66-8,2 males and 4 females, 50-63 mm SL, 23°S, 152°E, Capricorn Group, One Tree Island, Great Barrier Reef, Queensland, Australia; WAM R 27065-001, 7 females, 19-59 mm SL, 24°46’S, 152°24’E, Bagara, Great Barrier Reef, Queensland, Australia, J.B. Hutchins, 3 Dec. 1980; WAM P. 29627-018, 1 male, 59 mm SL and 2 females, 31 mm SL, 16°31’S, 147°50’E, Holmes Reef, Coral Sea, G.R. Allen and M. Allen, 4 Nov. 1987; WAM P. 29637-023, 1 male, 46 mm SL and 2 females, 52-65 mm SL, 17°37’S, 151°26’E, Lihou Reef, Coral Sea, G.R. Allen etal ., 12 Nov. 1987; ZMUC P 771493, 1 male, 62 mm SL, same data as AMS I. 17377-018; ZMUC P 771492, 1 female, 49 mm SL, same data as AMS 1. 19108-142. Diagnosis. Vertebrae 11+32-35=43-46, dorsal fin rays 77-85, anal fin rays 60-69; outer pseudoclasper with long, sharply pointed hook-like extension, slightly bent outwards, and with thin supporter; inner pseudoclasper anteriorly connected to outer pseudoclasper, wide and concave due to broad thin flap; no scales on operculum (rarely a single scale above the opercular spine); upper preopercular pore present. Description. The principal meristic and morphometric characters are shown in Table 20. Body slender with pointed snout; fishes mature at about 40 to 45 mm SL. Head with scale patch on cheek containing 5-9 scale rows on the upper cheek; no scales on operculum, except for very few specimens with a single scale above the opercular spine. Horizontal diameter of body scales 1.4% SL. Maxillaries expanded posteriorly with prominent angle at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril small, slightly larger than anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 44 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). 128 Dinematichthyine fishes of the Indo-West Pacific Dentition (of liolotype). Premaxilla with 6 rows of granular teeth and 2 tooth rows anteriorly, including a few fang-like teeth closest to symphysis, up to 1/2 pupil diameter. Eight rows of granular teeth posteriorly. Vomer with 5 anterior rows of granular teeth and 1 inner row of larger teeth. Palatine teeth in 4 rows anteriorly and 2 rows posteriorly, largest on inner row. Dentary with 6 rows of granular teeth and 1 tooth row anteriorly, blending into one row of about 15 large, curved, fang-like teeth posteriorly, size more than 1/2 pupil diameter. Otolith (Fig. 44 G). Moderately elongate, length to height 2.0-2.1 (46-65 mm SL); otolith length to sulcus length 2.0-2.1; sulcus inclined at an angle of 5°. Anterior tip of otolith pointed; posterior tip less pointed. Axial skeleton (of liolotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10 (11). First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis in males, somewhat shorter in females. Mate copulatory organ (Fig. 44 C-F). Two pairs of large pseudociaspers. Outer pseudoclasper with long, sharply pointed and slightly bent outwards hook-like extension, with thin supporter; inner pseudoclasper anteriorly connected to outer pseudoclasper, wide and concave due to broad thin flap, sometimes with indication of furcation distally. Isthmus between pseudociaspers narrow. Penis long, curved, thin. Coloration. Live colour reported yellow or red. Uniformly light to medium brown when preserved. Comparison. Diancistrns longifilis belongs to the group of species with hook- or stick-shaped outer pseudociaspers, which includes D. alleni, D. beateae, D.fuscus, D. leisi, D. machidai n. sp., D. manciporus n. sp., D. novaeguineae, D. springeri n. sp. and D. vietnamensis n. sp. Diancistrns longifilis shares the absence of scales on the operculum with D. leisi , D. machidai n. sp. and D. manciporus n. sp. There are, however, very few undisputable male records of D. longifilis with a single scale above the opercular spine, an observation more regularly made also in D. machidai n. sp. Then, distinction ofD. longifilis from D. alleni relies on the pseudoclasper pattern. Both species overlap in their geographical distribution along the northern Great Barrier Reef. While/), machidai n. sp. differs in several characters (see account), does D. manciporus n. sp. differs mainly in the lack of the upper preopercular pore (versus present). Distribution (Fig. 12). Diancistrns longifilis is a common species along the southern Great Barrier Reef, Queensland, Australia, reaching southwards to 24°S and northwards to 14°S, where it is sympatric with D. alleni , which reaches southwards to about 15°S. Diancistrns longifilis is the only species of the genus Diancistrns along the isolated Lord Flowe Ridge - Lord Howe Island, Middleton and Elizabeth Reefs which is also the southern-most occurrence of the genus (31°S). To the east, D. longifilis is known from New Caledonia and the Loyalty Islands; further to the east, at the Tonga Islands, it is replaced by D. manciporus n. sp. Diancistrns mcgroutheri n. sp. (Figs 17,45, 46; Table 21) Material examined. (17 specimens, 30-60 mm SL). Holotype - WAM P. 29627-047, male, 36 mm SL, 16°3TS, 147°50’E, Holmes Reef, Coral Sea, 30-60 m, G.R. Allen and M. Allen, 4 Nov. 1987. Paratypes - AMS I. 18739- 121, 3 females, 32-45 mm SL, 14°42’S, 145°27’E, Lizard Island, Great Barrier Reef, J.R. Paxton and party, 21 Nov. 1975; AMS I. 33752-023, 1 male, 45 mm SL, 09°34’S, 144°46’E, Port lock Reef, Coral Sea, 29 Jan. 1993; USNM 366525,1 male, 47 mm SL, 05°S, 145°’E, Madang Harbour, Papua New Guinea, B. Collette, I June 1970; USNM 374182, 1 female, 43 mm SL, Lemus Island, New Ireland, Papua New Guinea, T. Roberts, 20-25 Jan. 1976; WAM P. 27469-010, 1 male, 43 mm SL, 15°50’S, I45°50’E, Escape Reef, Queensland, Great Barrier Reef, G.R. Allen et al„ 31 Oct. 19081. Additional specimens. AMS I. 19483-083, 1 male, 30 mm SL, 14°S, 145°E, Lizard Island, Great Barrier Reef, Queensland, Australia; AMS I. 20757-112, 1 female, 60 mm SL, Raine Island, Great Barrier Reef, Queensland, Australia; AMS I. 20774-016, 1 female, 38 mm SL, 14°S, 144°E, Cape Melville, Great Barrier Reef, Queensland, Australia; AMS I. 20775-092, 1 male, 40 mm SL, Raine Island, Great Barrier Reef, Queensland, Australia; AMS I. 22573-014, 1 male, 39 mm SL and 1 female, 45 mm SL, 15°S, 145°E, Escape Reef. Great Barrier Reef. Queensland, Australia; AMS 1.23848-004, 1 female, 40 mm SL, 14°S, I45°E, Lizard Island, Great Barrier Reef, Queensland. Australia; AMS I. 25109-062, 1 male, 42 mm SL, 13°S, I46°E, Osprey Reef, Coral Sea; USNM 366503, 1 female, 55 mm SL, 05°S, 145°’E, Madang, Papua New Guinea. Diagnosis. Vertebrae 11+30=41, dorsal fin rays 73-76, anal fin rays 56-61; eyes small (1.5-2.1% SL); outer pseudoclasper with small ear-lobe extension opening inwards; inner pseudoclasper free from outer pseudoclasper, with supporter, forming a sharp, forward- inclined thorn; narrow scale patch on check, no scales on operculum; head profile slender; otolith length to height ratio 2.0-2.2. Description. The principal meristic and morphometric characters are shown in Table 21. Body slender, with slender snout; fishes small, mature at about 30 to 35 mm SL. Head with narrow scale patch on cheek with 4-5 scale rows on upper cheek, no scales on operculum. Horizontal diameter of body scales 1.3% SL. Maxillaries expanded posteriorly with angle at rear ventral corner. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril about twice the size of anterior nostril, surrounded by funnel-shaped circular flap, about 1/4 the size of eye. Head sensory pores (Fig. Fig. 46 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). 129 W. Schwarzhans, P. R. Moller and J. G. Nielsen Table 21. Meristic and morphometric characters of Diancistrus mcgroutheri n. sp. Holotype WAM P. 29627- 047 Holotype + 7 paratypes Mean (ranee) N Standard length in mm Meristic characters 35 40.3 (31-48) 8 Dorsal fin rays 75 73.9 (73-76) 8 Anal fin rays 61 58.8 (56-61) 8 Pectoral fin rays 22 20.8 (20-22) 6 Precaudal vertebrae 11 11 8 Caudal vertebrae 30 30 8 Total vertebrae 41 41 8 Rakers on anterior gill arch 16 16.1 (14-18) 8 Pseudobranchial filaments - 2 6 D/V 6 6 8 D/A 20 20.1 (19-22) 8 V/A 13 Morphometric characters in % of SL 13 8 Head length 28.1 27.3 (26.0-29.9) 8 Head width 12.5 12.9 (11.7-14.5) 8 Head height 16.5 16.8(15.8-18.5) 8 Upper jaw length 13.8 13.6(12.8-14.1) 8 Maxillary height 4.2 4.0 (3.5-4.4) 8 Diameter of pigmented eye 1.6 1.7(1.5-2.1) 8 Interorbital width 7.0 7.2 (6.4-8.0) 8 Postorbital length 20.2 20.5(19.6-21.4) 8 Preanal length 49.1 46.7 (44.8-49.1) 8 Predorsal length 33.3 32.4 (31.4-33.5) 8 Body depth at origin of anal fin 18.7 18.0(17.3-18.7) 8 Pectoral fin length - 15.7(15.1-16.4) 7 Pelvic fin length 25.1 26.0 (21.5-29.4) 7 Base pelvic fin - anal fin origin 30.7 28.5 (25.3-33.3) 8 Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 tooth row anteriorly, including a few fang-like teeth closest to symphysis, up to Vi pupil diameter. Six rows of granular teeth posteriorly. Vomer with 1 row of granular teeth. Palatine with 2 irregular rows of equally sized teeth. Dentary with 5 rows of granular teeth and 1 tooth row anteriorly, blending into one row of about 8 large, curved, fang-like teeth posteriorly, about the size of pupil diameter. Otolith (Fig. 46 G-H). Elongate, with much expanded slender posterior tip resulting in length to height relation of 2.0-2.2 (36-45 mm SL); otolith length to sulcus length 2.2; sulcus not inclined. Anterior and posterior tips of otolith moderately pointed. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, but not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 46 C-F). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper ear-lobe shaped, the small ear-lobe extension confined to rear half and opening inwards; inner pseudoclasper free from outer pseudoclasper, with supporter, forming sharp, forward-inclined thorn, anteriorly connected to isthmus by ligament, posteriorly with small fleshy flap. Isthmus between pseudoclaspers moderately wide. Penis short, curved, slender. Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Diancistrus mcgroutheri belongs to the group of species with ear-lobe shaped outer pseudoclaspers and small eyes ( Diancistrus erythraeus subgroup) also containing/?, atollorum, D. erythraeus, D. rohustus n. sp. and D. tongaensis n. sp. Except for D. rohustus n. sp., the species of this subgroup are further characterized by the low number of vertebrae (39-41, rarely 42 versus 43 in D. rohustus n. sp.) and the slender head (head width <16% SL versus >16% SL in D. rohustus n. sp.). The remaining four species are best distinguished by means of pseudoclaspers. In D. mcgroutheri and D. tongaensis n. sp. the inner pseudoclasper is free from the outer pseudoclasper, whereas in D. atollorum and D. erythraeus (and D. rohustus n. sp.) it is firmly joined to the outer pseudoclasper. Apparently, D. mcgroutheri is closest to D. tongaensis n. sp., differing in the small, inwardly directed ear-lobe extension of the outer pseudoclasper (versus large and ventrally opened) and the thorn-like anteriorly inclined inner pseudoclasper (versus broad, distally expanded flap). Within its area of distribution D. mcgroutheri could potentially be confused with D. leisi in the absence of males, being the only other small-eyed Diancistrus species, and also in lacking scales on the operculum. However, D. Fig. 45. Diancistrus mcgroutheri n. sp. Holotype. WAM R 29627-047, male, 36 mm SL. 130 Dinematichthyine fishes of the Indo-West Pacific Fig. 46. Diancistrus mcgroutheri n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype; C, view ot left pseudoclasper from ventral, holotype; D, view of left pseudoclasper from inside, holotype; E, view of left pseudoclasper from inside, WAM P. 27469- 010,43 mm SL; F, inclined lateral view of male copulatory organ, WAM P. 27469-010,43 mm SL; G, ventral view of right otolith, AMS 1. 33752-023, male 45 mm SL; II, median view of right otolith, AMS I. 33752-023, male 45 mm SL. leisi is also distinguished by the narrow scale patch on the lower check and the laterally very compressed head with the dorsally positioned eyes. Distribution (Fig. 17). Diancistrus mcgroutheri is mainly known from the Coral Sea, i.e., the northern Great Barrier Reef and along the southern part of New Guinea (Papua New Guinea) including Madang and the Trobriand Islands. Much further to the west, south of Sumatra, possibly related small-eyed female specimens are mentioned as Diancistrus sp. 3. Etymology. Named in honour of Mark McGrouther, Sydney, Australia, in recognition of his great help with our revision of the Dinematichthyini. Diancistrus machidai n. sp. (Figs 20, 47, 48; Table 22) Material examined. (10 specimens, 31-60 mm SL). Holotype - USNM 372962, male, 42 mm SL, 05°17’20”S, 122°04’00”E, Tallabassi Bay, just off NE tip of Big Damalawa Islet, Kabaena Island, Sulawesi, Sulawesi Tenggara Province, Indonesia, 2-15 m, V.G. Springer and M.F. Gomon, 25 Feb. 1974. Paratypes - AMS I. 18469-183, 1 male, 45 mm SL, 03°0FS, 128°03’E, Ceram, Moluccas, Indonesia, J.R. Paxton, 1 April 1975; BPBM 26739, 1 male, 31 mm SL, 05°S, II9°E, Ujung Padang, Sulawesi, Indonesia, J.E. Randall, GAV. Tribble and R.P.H. Rutherford, 8 Sept. 1978; USNM 99224, 1 male, 41 mm SL, 09°N, 125°E, Surigao, Mindanao, Philippines, RV Albatross, 8 May 1908; USNM 263686,1 male, 42 mm SL, I0°52’N, 120°56’E, Cuyo Islands, Palawan, Philippines, 23 May 1978; USNM 300088, 1 male, 60 mm SL, 20°24’N, 121°55’E, Batanes Islands, Philippines, G.D. Johnson etal., 1 May 1987; USNM 366498, 1 male, 35 mm SL, 05°52’S, 110°25’E, Karimundjawa Archipelago, Java Sea, Indonesia, V.G. Springer et at., 29 March 1974; USNM 374159, 1 male, 43 mm, 05°51’S, 106°34’E, Seribu Islands, Java Sea, Indonesia, V.G. Springer et a /., 5 April 1974; USNM 374181, 1 male, 34 mm SL, 20°24’N, 121°55’E, Batanes Province, Philippines, G.D. Johnson et a /., 2 May 1987; USNM 374193, 1 female, 47 mm SL, 09°10’N, 123°26’E, Siquijor Island, Visayan Islands, Philippines, V.G. Springer etal , 10 May 1978. Diagnosis. Vertebrae 11+29-31 (32)=40-42 (43), dorsal fin rays 66-77, anal fin rays 50-64; eyes large (2.4-3.3% SL), D/V 6-7; outer pseudoclasper long, thick, with almost straight hook-like extension in ventral view; inner pseudoclasper anteriorly joined to outer pseudoclasper, broad, its supporter extended as distinct thorn to nearly length of extension of outer pseudoclasper; moderately wide scale patch on cheek with 5-6 scale rows on upper cheek, 0-2 scales on operculum above opercular spine; head slender, broad; otolith length to height ratio 2.0-2.1; sulcus short, ratio otolith length to sulcus length 2.3-2.5. 131 W. Schwarzhans, P. R. Moller and J. G. Nielsen Table 22. Meristic and morphometric characters of Diancistrus machidai n. sp. Holotype USNM 372962 Holotype + 9 paratypes Mean (range) N Standard length in mm 42 42.1 (31-60) 10 Meristic characters Dorsal fin rays 75 72.5 (66-77) 10 Anal fin rays 59 57.7 (50-64) 10 Pectoral fin rays - 18.3(17-19) 8 Precaudal vertebrae 11 11 10 Caudal vertebrae 31 30.7 (29-32) 10 Total vertebrae 42 41.7(40-43) 10 Rakers on anterior gill arch 17 16.8(14-21) 10 Pseudobranchial filaments 2 2 8 D/V 6 6.5 (6-7) 10 D/A 19 19.0(17-21) 10 V/A 13 13.2(13-14) 10 Morphometric characters in % of SL Head length 26.2 27.0 (25.6-29.8) 9 Head width 12.1 11.6(10.3-13.3) 8 Head height 20.7 17.6(14.3-20.7) 9 Upper jaw length 13.1 13.3(12.3-14.9) 9 Maxillary height - 4.3 (3.8-4.9) 7 Diameter of pigmented eye 3.1 2.8 (2.4-3.3) 9 Interorbital width 6.2 5.7 (3.9-7.6) 9 Postorbital length 18.3 19.3 (17.7-22.5) 9 Preanal length 50.0 49.2 (47.5-52.2) 9 Predorsal length 36.4 35.1 (32.1-38.6) 9 Body depth at origin of anal fin 20.5 17.6(14.4-20.5) 9 Pectoral fin length - 15.7(12.9-17.8) 8 Pelvic fin length 24.3 24.9 (22.7-28.1) 4 Base pelvic fin - anal fin origin 29.0 28.8(25.8-31.7) 9 Description. The principal meristic and morphometric characters are shown in Table 22. Body and head slender; fishes mature at about 35 to 40 mm SL. Head with scale patch on cheek with 5-6 scale rows on upper cheek, 0-1, rarely 2 scales above the opercular spine. Horizontal diameter of body scales 1.3% SL. Maxillaries expanded posteriorly with angle at rear ventral corner and indistinct knob in front of rear angle. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril located very close to eye, usually opening towards the eye, slightly larger than anterior nostril, about 1/5 the size of eye. Head sensory pores (Fig. 48 A-C). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior), the 3rd posterior mandibular pore twice the size of the two other posterior mandibular pores. Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 6 rows of granular teeth and 1 tooth row anteriorly, including a few fang-like teeth closest to symphysis, up to 1/3 pupil diameter. Six granular rows posteriorly. Vomer with 2 tooth rows, with teeth on posterior inner row the largest. Palatine with 2 rows of equally sized teeth. Dentary with 5 rows of granular teeth and 1 tooth rows anteriorly, blending into one row of about 7 large, curved, fang-like teeth posteriorly, fang size about 1/2 the size of pupil diameter. Otolith (Fig. 48 M). Moderately elongate with pointed anterior and more rounded posterior tips, length to height 2.0-2.1 (34-62 mm SL); otolith length to sulcus length 2.3-2.5; sulcus slightly inclined at 5°. Otoliths from males with gently curved dorsal rim and shallow or absent postdorsal angle, otoliths from females with marked postdorsal angle and flat middle part of dorsal rim. As in D.fuscus, this subtle sexual dimorphism is opposite to the trend observed in most other species of the genus. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis in males, slightly shorter in females. Mate copulatory organ (Fig. 48 D-L). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper long, thick, with almost straight hook-like extension in ventral view, but bent at a 90° angle in lateral view, with massive supporter; inner pseudoclasper anteriorly joined to outer pseudoclasper, broad, its strong supporter extended as a distinct thorn to nearly the length of the extension of the outer pseudoclasper, with fleshy flap along posterior margin. Isthmus between pseudoclaspers moderately wide. Penis moderately long, curved, tapering with thin tip. Fig. 47. Diancistrus machidai n. sp. Holotype. USNM 372962, male, 42 min SL. 132 Dinematichthyine fishes of the Indo-West Pacific Fig. 48. Diancistms machidai n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype; C, lateral view of head, USNM 374159, male, 43 mm SL; I), ventral view of male copulatory organ, holotype; E, view of left pseudoclasper from inside, holotype; F, inclined lateral view of male copulatory organ, holotype; G, ventral view of male copulatory organ, USNM 374159, 43 mm SL; H, view of left pseudoclasper from inside, USNM 374159, 43 mm SL; I, view of left pseudoclasper from inside, USNM 374181,34 mm SL; K, inclined lateral view of male copulatory organ, USNM 374181, 34 mm SL; L, inclined lateral view of male copulatory organ, USNM 374159, 43 mm SL; M, median view of right otolith, USNM 300088, male, 60 mm SL. 133 W. Schwarzhans, P. R. Moller and J. G. Nielsen Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Diancistrus machidai belongs to the group of species with hook- or stick-shaped outer pseudoclaspers, which includes D. alleni, D. beateae, D. fuscus, D. leisi, D. longifilis, D. manciponts n. sp., D. novaeguineae, D. springeri n. sp. and D. vietnamensis n. sp. Diancistrus machidai has either no scales on the operculum like D. leisi (which, however, has stick-like outer pseudoclaspers), D. longifilis and D. manciponts n. sp., or 1 or 2 scales above the opercular spine similar to D. alleni. Diancistrus machidai is easily recognized by the long spiny inner pseudoclasper, whereas D. leisi differs in the stick-like outer pseudoclasper. Diancistrus machidai is further distinguished from D. alleni, D. longifilis and D. manciponts n. sp., by its low number of vertebrae (40-43 versus 43-45) and the low number of dorsal fin rays (66-77 versus 75-85). Remarks. Diancistrus machidai shows an unusual large degree of variability, e.g., in the presence or absence of scales above the opercular spine and the wide range in dorsal fin ray counts. At the southern limit of its distribution, off Java, a specimen is recorded with 2 scales above the opercular spine and a more straight and long outer pseudoclasper (Fig. 46 C, G, H, L). At the northern limit, Batanes Islands of the Philippines, a specimen was found to have a shorter outer pseudoclasper, which is also more flat and less bent than the typical hook-shape (Fig. 46 I, K). It is possible that with more material available, a separation into two species may become advisable. Distribution (Fig. 20). Diancistrus machidai is widely distributed in the West Pacific but not common. It ranges from Java and Sulawesi in the south-west to the Batanes Islands of the Philippines in the north-east. Etymology. Named in honour of Yoshihiko Machida of Japan for his many contributions to the knowledge of ophidiiform fishes. Diancistrus manciponts n. sp. (Figs 12,49, 50; Table 23) Material examined. (5 specimens, 23-67 mm SL). Holotype - USNM 374179, male, 53 mm SL, 21°02°S, 175°12’W, Atata Island, reef near ship channel, Tongatapu, Tonga, vertical wall to about 20 m with sand and rubble at base, 15-20 m, J. T. Williams et ai, 22 Oct. 1993. Paratypes- USNM 384599, 1 female, 67 mm SL, 1 sub¬ adult male, 38 mm SL and 2 juveniles, 23-27 mm SL, same data as holotype. Diagnosis. Vertebrae 11+32-34=43-45, dorsal fin rays 79-84, anal fin rays 64-67; outer pseudoclasper with hook- like short and pointed extension and slightly bent distally and with thin supporter; inner pseudoclasper anteriorly connected to outer pseudoclasper, wide and concave due to broad thin flap, strongly anteriorly inclined; no scales on operculum; upper prcopercular pore absent. Description. The principal meristic and morphometric characters are shown in Table 23. Body slender with pointed snout; fishes mature at more than 50 mm SL. Head with moderately wide scale patch on cheek containing 5-6 scale rows on the upper cheek; no scales on operculum. Horizontal diameter of body scales 1.6% SL. Maxillaries expanded posteriorly with angle at rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril about the size of anterior nostril and about 1/6 the size of eye. Head sensory pores (Fig. 50 A-B). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 5 rows of granular teeth and I tooth row anteriorly, including a few fang-like teeth closest to symphysis, up to 1/3 pupil diameter. Eight rows of granular teeth posteriorly. Vomer with 3 rows of teeth, largest posteriorly on the inner row. Palatine teeth in 3 irregular rows, teeth on inner row slightly larger than on outer rows. Dentary with 5 rows of granular teeth and 1 tooth row anteriorly, blending into one row of about 10 large, curved, fang-like teeth posteriorly, fang size up to 1/2 pupil diameter. Table 23. Meristic and morphometric characters of Diancistrus manciponts n. sp. Holotype USNM 374179 Holotype + 4 paratypes Mean (range) N Standard length in mm 53 41.0(22-65) 5 Meristic characters Dorsal tin rays 79 81.4 (79-84) 5 Anal fin rays 64 65.8 (64-67) 5 Pectoral fin rays 20 20 4 Precaudal vertebrae 11 11 5 Caudal vertebrae 33 32.8 (32-34) 5 Total vertebrae 44 43.8 (43-45) 5 Rakers on anterior gill arch 16 16.0(15-17) 4 Pseudobranchial filaments 2 2 4 D/V 6 6 5 D/A 19 20.6 (19-22) 5 V/A 13 13 5 Morphometric characters in % of SL Head length 26.6 27.6 (26.6-29.0) 5 Head width 12.6 12.9(12.0-13.6) 4 Head height 16.5 16.9(16.5-17.7) 4 Upper jaw length 12.5 12.8(12.4-13.5) 3 Maxillary height 4.2 4.1 (3.6-4.6) 4 Diameter of pigmented eye 2.8 3.1 (2.8-3.5) 5 Interorbital width 5.8 6.6 (5.8-79) 4 Postorbital length 18.6 19.4 (18.6-20.6) 3 Preanal length 49.0 47.6 (46.7-49.0) 3 Prcdorsal length 32.1 33.3 (32.1-34.0) 4 Body depth at origin of anal fin 16.3 17.2(16.3-18.4) 4 Pectoral fin length 15.5 15.7(14.4-16.6) 4 Pelvic fin length 24.8 24.1 (21.8-25.7) 3 Base pelvic fin - anal fin origin 28.0 26.3 (24.7-28.0) 4 134 Dinematichthyine fishes of the Indo-West Pacific Fig. 49. Diancistrus manciponts n. sp. Holotype. USNM 374179, male, 53 mm SL. Otolith (Fig. 50 F-G). Moderately elongate, length to height 1.9-2.0 (55-67 mm SL); otolith length to sulcus length 2.1-2.2; sulcus inclined at an angle of 5°. Anterior tip of otolith pointed; posterior tip less pointed. Strong postdorsal angle in males, more rounded in females. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-9 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2- (10) 11. First anal fin pterygiophore elongated, but not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 50 C-E). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper hook-shaped, hook-like extension short, pointed and slightly bent at its termination, with thin supporter; innei pseudoclasper anteriorly connected to outer pseudoclasper, wide and concave due to broad thin flap, strongly anteriorly inclined. Isthmus between pseudoclaspers narrow. Penis long, curved, moderately thin. Coloration. Live colour not known. Uniformly light brown when preserved. Fig. 50. Diancistrus manciponts n. sp. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male copulatory organ, holotype; D, view of left pseudoclasper from inside, holotype; E, inclined lateral view of male copulatory organ, holotype; F, median view of right otolith, holotype; G, median view of right otolith, USNM 384599, female, 67 mm SL. 135 W. Schwarzhans, P. R. Moller and J. G. Nielsen Comparison. Diancistrus manciporus belongs to the group of species with hook- or stick-shaped outer pseudoclaspers like D. alleni, D. beateae, D. fuscus, D. leisi, D. longifilis, D. machidai, D. novaeguineae, D. springeri n. sp. and D. vietnamensis n. sp. It is most similar to D. longifilis , from which it is probably derived. The main differences to D. longifilis are the lack of an upper preopercular pore (versus present) and the strongly forward inclined inner pseudoclasper (versus not forward inclined). Distribution (Fig. 12). Diancistrus manciporus has so far only been found at the southern part of Tongatapu. Etymology. The name refers to the lack of an upper preopercular pore - manats (Latin) = missing and poms (Latin) = pore. Diancistrus ntennei n. sp. (Figs 40, 51, 52; Table 24) Material examined. (10 specimens, 25-76 mm SL). FIolotype - BPBM 22384, male, 48 mm SL, Eniwetok Atoll, Marshall Islands, J.E. Randall et al., April-May 1978. Tentatively assigned specimens: AMS I. 18051-048, 1 female, 65 mm SL, 0I°44’N, 172°59’E, Abaiang Atoll, Gilbert Islands, Kiribati, D.F. Flocse and B. Goldman, 10 Nov. 1973; MCZ 158555, 1 female, 31 mm SL, 02°50’S, 171°42’W, Canton Island, Kiribati, S. Bailey et al., 4 July 2000; MCZ 158556, 1 juvenile, 25 mm SL, 03°43’S, 170°42’W, Phoenix Island. Kiribati, S. Bailey et al., 6 July 2000; MCZ 162574, 4 females, 42-76 mm SL, 02°46’S, 171°43’W, Canton Island, Kiribati, G.R. Allen et al., 24 June 2002; MCZ 162575, 2 females, 39-45 mm SL, 04°31’S, 172°13’W,Orona Atoll, Gilbert Islands, Kiribati, G.R. Allen, 2 July 2002. Diagnosis. Vertebrae 11+34=45, dorsal fin rays 88, anal fin rays 69; eye large (3.4% SL), D/V 6; outer pseudoclasper moderately large, short (about 1 A length of inner pseudoclasper), wing-shaped, flat with broad base; inner pseudoclasper joined to outer pseudoclasper anteriorly, with thin, sharp, slightly anteriorly directed single-pointed tip extending over anterior part of outer pseudoclasper; moderately broad scale patch on cheek with 5 scale rows on upper cheek, large scale patch on operculum above opercular spine (8 scales); head broad, rather fiat dorsally; otolith slender, length to height ratio 2.2. Table 24. Mcristic and morphometric characters of Diancistrus rnennei n. sp. Holotype BPBM 22384 9 tentatively assigned spec. Mean (range) N Standard length in mm Meristic characters 47 46.4 (24-76) 10 Dorsal fin rays 88 83.3 (80-86) 10 Anal fin rays 69 65.3 (62-68) 10 Pectoral fin rays 21 18 3 Precaudal vertebrae 11 11 10 Caudal vertebrae 34 34.0 (33-35) 10 Total vertebrae 45 45.0 (44-46) 10 Rakers on anterior gill arch 21 16.0(15-18) 4 Pseudobranchial filaments 2 2 4 D/V 6 6 10 D/A 23 20.7 (19-22) 10 V/A 13 Morphometric characters in % of SL 12.9(12-14) 10 Head length 28.1 27.1 (26.2-27.8) 4 Head width 13.0 13.3 (13.1-13.7) 4 I lead height 20.0 18.1 (15.6-23.0) 4 Upper jaw length 15.1 13.5(12.9-14.2) 4 Maxillary height 4.7 4.4 (3.9-5.2) 4 Diameter of pigmented eye 3.4 3.2 (3.2-3.3) 4 Interorbital width 6.4 6.5 (6.3-6.8) 4 Postorbital length 19.8 19.6(18.8-20.3) 4 Preanal length 49.1 46.2 (44.4-48.8) 4 Predorsal length 35.1 32.0 (30.3-34.5) 4 Body depth at origin of anal fin 18.9 17.8(16.1-19.2) 4 Pectoral fin length 19.1 17.9(17.5-18.3) 3 Pelvic fin length 22.6 24.0 (23.6-24.6) 4 Base pelvic fin - anal fin origin 28.1 26.3 (25.2-26.9) 4 Description. The principal meristic and morphometric characters are shown in Table 24. Body moderately compact, with broad, dorsally compressed head and pointed snout, mature at about more than 45 mm SL. Flead with moderately broad scale patch (5 scale rows) on upper cheek, large scale patch on operculum above opercular spine (8 scales) in 3 rows and no scales below opercular spine. Horizontal diameter of body scales 1.6% SL. Maxillaries expanded posteriorly with prominent angle al rear ventral corner. Anterior nostril low on snout, 1/3 distance from tip of snout to anterior margin of eye. Posterior nostril about twice the size of anterior nostril and about 1/5 the size of eye. Fig. 51. Diancistrus rnennei n. sp. Holotype. BPBM 22384, male, 48 mm SL. 136 Dinematichthyine fishes of the Indo-West Pacific Fig. 52. Diancistnis mennei n. sp. A. lateral view of head, holotype; B, ventral view of head, holotype; C, lateral view of head, MCZ 162574, tentatively assigned female, 76 mm SL; D, view of left pseudoclasper from inside, holotype; E, inclined lateral view of male copulatory organ, holotype; F. median view of right otolith, MCZ 162574, tentatively assigned female, 76 mm SL; C, median view of right otolith, holotype; 11 , ventral view of right otolith, holotype. Head sensory pores (Fig. 52 A-C). Supraorbital pores 3. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 4 rows of granular teeth and 1 tooth row anteriorly, with teeth up to 1/4 of pupil diameter. Seven granular rows posteriorly. Vomer teeth in 2 rows, teeth on inner row slightly larger than on outer row. Palatine teeth in 2 rows of similar sized teeth. Dentary with 3 granular rows and 1 tooth row anterior, blending into one row of about 5 large fang-like teeth posteriorly. Otolith (Fig. 52 F-ll). Elongate with pointed anterior and posterior tips, length to height 2.2 (48 mm SL); otolith length to sulcus length 1.9; sulcus slightly inclined at 5°. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 52 D-E). Two pairs of moderately large pseudoclaspers. Outer pseudoclasper short (about I 1/2 length of inner pseudoclasper), wing- shaped, with broad base, broad supporter and fiat inner face, not expanded distally; inner pseudoclasper joined to outer pseudoclasper anteriorly, thin, sharp, slightly anteriorly directed, with single-pointed tip extending over anterior part of outer pseudoclasper. Isthmus between pseudoclaspers moderately wide. Penis long, curved, thin. 137 W. Schwarzhans, P. R. Moller and J. G. Nielsen Coloration. Live colour not known. Uniformly light brown when preserved. Comparison. Diancistrus mennei belongs to the group with wing-shaped outer pseudoclasper and a single-tipped inner pseudoclasper. which includes D. katrineae, D. eremitus, D. karinae, D. jeffjohnsoni and D. polmpeiensis n. sp. It differs from all these species in the relatively short outer pseudoclasper and the anteriorly directed inner pseudoclasper. Diancistrus katrineae and D. karinae both have scales on the operculum above and below the opercular spine (versus above opercular spine only). From the remainder of species D. mennei differs in the higher number of scales above the opercular spine (8 versus 1-4) and the more slender otoliths (length to height ratio 2.2 versus <2.1). Diancistrus mennei occurs associated with D. beateae, from which it is easily distinguished by the lack of a continuous scale patch on the operculum with scales below the opercular spine (versus present), and D. atollorum, which belongs to the group with ear-lobe shaped outer pseudoclaspers, small eyes and no scales on the operculum. Remarks. The holotype from the Eniwetok Atoll, Marshall Islands, represents the only male known. The 9 females from Kiribati closely resemble the holotype in all characters. Due to the geographic distance between the two localities and the lack of a male these specimens are tentatively assigned and not designated as paratypes. Distribution (Fig. 40). Diancistrus mennei is known from the Marshall Islands and probably from Kiribati. Etymology. Named afterTammes Menne, Copenhagen, Denmark, for his great help during this revisionary work. Diancistrus niger n. sp. (Figs 20, 53, 54; Table 25) Material examined. (8 specimens, 30-45 mm SL). Holotype - WAM P. 31543-005, male, 40 mm SL, 00°37S, 130°33E, Raja Ampat Islands, Mansuar Island, Waigeo, Papua, Indonesia, 0-4 m, G.R. Allen and S. Morrison, April 1999. Paratypes - WAM P. 31543-004, 2 males, 30-42 mm SL and 2 females, 38-44 mm SL, same data as holotype; WAM P.31541-011, 1 male, 45 mm SL, 00°48’S, 130°30’E, Raja Ampat Islands, Batanta Island, Waigeo, Papua, Indonesia, G.R. Allen, 14 April 1999; ZMUC P 771470, 1 male, 32 mm SL and ZMUC P 771471, 1 female, 43 mm SL, same data as WAM P. 31543-004. Diagnosis. Vertebrae 11+30-32=41-43, dorsal fin rays 69-76, anal fin rays 52 59; eyes large (3.0-3.6% SL); outer pseudoclasper large, broad, ear-lobe shaped, opening ventrally; inner pseudoclasper a fleshy appendix at anterior-inner rim of outer pseudoclasper, without supporter; no scales on operculum; head compressed, blunt; otoliths elongate, ratio otolith length: height 2.3, anterior and posterior tips of otolith almost equally pointed; colour dark grey to black in preservation. Description. The principal meristic and morphometric characters are shown in Table 25. Body moderately compact, but not high-necked, snout pointed; fishes small, mature at about 30 mm SL. Head with narrow scale patch on cheek with 5 scale rows on upper cheek, no scales on operculum. Horizontal diameter of body scales 1.4% SL. Maxillaries slightly expanded posteriorly, broadly rounded. Anterior nostril low on snout, 1/4 distance from tip of snout to anterior margin of eye. Posterior nostril large, slightly larger than anterior nostril and about 1/5 the size of eye. Head sensory pores (Fig. 54 A-B). Supraorbital pores 3. Additional small pore often visible below eye. Infraorbital pores 6 (3 anterior and 3 posterior). Mandibular pores 6 (3 anterior and 3 posterior). Preopercular pores 4 (3 lower and 1 upper). Dentition (of holotype). Premaxilla with 3 rows of granular teeth and 2 tooth rows anteriorly, including one fang-like tooth closest to symphysis, about 'A pupil diameter. Six rows of granular teeth posteriorly. Vomer teeth in 3 rows, teeth on inner row slightly larger than on outer rows. Palatine teeth in 3 rows of similar sized teeth. Dentary with 2 rows of granular teeth and 2 tooth rows anteriorly, blending into one row of about 10 large fang-like teeth posteriorly, fang size about 1/2 pupil diameter. Table 25. Meristic and morphometric characters of Diancistrus niger n. sp. Holotype WAM P. 31543-005 Holotype + 7 paratypes Mean (ranee) N Standard length in mm Meristic characters 40 39.0 (30-45) 8 Dorsal fin rays 75 73.4 (69-76) 7 Anal fin rays 59 56.6 (52-59) 7 Pectoral fin rays 17 17.2(17-18) 6 Precaudal vertebrae 11 11 7 Caudal vertebrae 31 30.9 (30-32) 7 Total vertebrae 42 41.9(41-43) 7 Rakers on anterior gill arch 15 14.6(13-17) 8 Pseudobranchial filaments 2 2 8 D/V 7 6.9 (6-7) 7 D/A 20 19.9(19-21) 7 V/A Morphometric characters in 1 14 /, of SL 13.6(13-14) 7 Head length 30.7 29.0(28.1-30.7) 8 Head width 15.4 15.3(14.1-16.6) 8 Head height 18.3 18.5(17.2-19.2) 8 Upper jaw length 14.7 14.2(13.7-14.7) 8 Maxillary height 4.6 4.8 (4.6-5.8) 8 Diameter of pigmented eye 3.2 3.3 (3.0-3.6) 8 Interorbital width 6.8 6.8 (64-7.2) 8 Postorbital length 21.1 20.3(19.4-21.1) 8 Preanal length 53.1 50.6 (46.1-53.1) 8 Predorsal length 37.2 35.7 (34.6-37.2) 8 Body depth at origin of anal fin 22.8 21.0(20.0-22.8) 8 Pectoral fin length 20.1 17.5(15.4-20.1) 8 Pelvic fin length 24.6 24.0 (22.3-25.4) 5 Base pelvic fin - anal fin origin 31.4 28.5(25.5-31.5) 8 138 Dinematichthyine fishes of the Indo-West Pacific Fig. 53. Diancistras niger n. sp. Holotype. WAM P. 31543-005, male, 40 mm SL. Otolith (Fig. 54 G-H). Very slender, elongate, length to height 2.3 (32-44 mm SL); otolith length to sulcus length 2.2-2.3; sulcus not inclined. Anterior tip of otolith sharply pointed; posterior tip nearly as pointed; dorsal rim shallow, with obtuse pre- and postdorsal angles, no postdorsal concavity. Axial skeleton (of holotype). Neural spine of vertebrae 4-5 inclined and 6-8 depressed. Parapophyses present from vertebra 7 to 11. Pleural ribs on vertebrae 2-10. First anal fin pterygiophore elongated, but not reaching tip of last precaudal parapophysis. Male copulatory organ (Fig. 54 C-F). Two pairs of large pseudoclaspers. Outer pseudoclasper large, broad, ear-lobe shaped, opening ventrally and anteriorly continuously connected to inner pseudoclasper, with large but poorly defined supporter; inner pseudoclasper a fleshy appendix at anterior-inner rim of outer pseudoclasper, without supporter, forming a folded double flap. Isthmus between pseudoclaspers narrow. Penis long, curved, thin. Fig. 54. Diancistrus niger n. sp. A. lateral view of head, WAM P. 31543-004, male 42 mm SL; B. ventral view of head, WAM P. 31543- 004, male 42 mm SL; C, ventral view of male copulatory organ, holotype; D, view of left pseudoclasper from inside, WAM P. 31541-011, 45 mm SL; E, inclined lateral view of male copulatory organ, holotype; F, view of left pseudoclasper from outside, WAM P. 31541-011. 45 mm SL; G. median view of right otolith, WAM P. 31543-004. male 42 mm SL; H, ventral view of right otolith, WAM P. 31543-004, male 42 mm SL. 139 W. Schwarzhans, P. R. Mollerand J. G. Nielsen Coloration. Live colour not known. Uniformly dark grey to black when preserved. Comparison. Diancistrus niger belongs to the group of species with ear-lobe shaped outer pseudoclaspers and to the subgroup with large eyes and atrophied inner pseudoclaspers without supporter, which contains two species, D. altidorsalis and D. niger. Diancistrus niger is easily distinguished from D. altidorsalis by the specific shape of the pseudoclaspers, the dark grey to black colour in preservation (versus light brown), the low number of vertebrae (41-32 versus 43-44), the low number of dorsal (69-76 versus 76-85) and anal fin ray (52-59 versus 61-68), the lack of scales on the operculum (versus two scale patches on the operculum, one above and one below the opercular spine) and finally the very slender otolith (otolith length to height 2.3 versus 1.9-2.0). Remarks. Three of the 8 specimens known show deformation of the vertebral column. Distribution (Fig. 20). Diancistrus niger is known only from the Raja Ampat Islands in the strait between the Waigeo Island and the Vogelkop Peninsula of northern New Guinea, Papua, Indonesia. Etymology. The name refers to the black colour- niger (Latin) = black. Diancistrus novaeguineae (Machida, 1996) (Figs 55, 56, 57; Table 26) Parabrosmolus novaeguineae Machida, 1996: 148, fig. 1-4 (type locality: Kranket Island, Madang Flarbour, Papua New Guinea). Parabrosmolus novaeguineae - Nielsen et al. 1999: 125. Material examined. (58 specimens, 28-103 mm SL). Holotype - BMNH 1974.5.25.3586, male, 37 mm SL, Kranket Island, Madang Harbour, Papua New Guinea, 30 May 1970. Further specimens. AMS I. 17487-018, 1 female, 59 mm SL, Doma Reef, Guadalcanal, Solomon Islands; AMS I. 18739-120, 1 male, 76 mm SL, 14°42’S, 145°27’E, Lizard Island, Great Barrier Reef, Queensland, Australia, J.R. Paxton and party, 21 Nov. 1975; AMS I. 19108-149, 1 male, 57 mm SL and 2 females, 39-48 mm SL, 14°40’S, 145°28'E, Lizard Island, Queensland, Great Barrier Reef, Queensland, Australia, D.F. Hoese and party, 17 Nov. 1975; AMS I. 20547-087, 1 male, 103 mm SL, Panasesa reef, Louisiade Archipelago, Papua New Guinea, B. Goldman. 24 March 1969; AMS I. 20937-034, 1 female, 63 mm SL, I2°02’S, 143°17’E, Haggerstone Island, Cape York, Great Barrier Reef, Queensland, Australia, 20 March 1979; AMS 1.20956-099,2 females, 28-70 mm SL, 17°04’S, 143°57’E, Tijou reef, Cape York, Great Barrier Reef, Queensland, Australia, 23 Feb. 1979; AMS I. 21540-065, 1 male, 14°S, 145°E, Lizard Island, Great Barrier Reef, Queensland, Australia; AMS I. 22612-010,4 males, 50-55 mm SL and 2 females, 45-50 mm SL, I5°49’S, 145°50’E, Escape Reef, Great Barrier Reef, Queensland, Australia, G.R. Allen Table 26. Meristic and morphometric characters of Diancistrus novaeguineae (Machida, 1996) Holotype BMNH 1974.5.25.3586 Holotype + 57 specimens N Mean (ramie) Standard length in mm Meristic characters 36.5 54.2 (27-103) 45 Dorsal fin rays 76 79.3 (74-85) 24 Anal fin rays 63 64.1 (60-69) 24 Pectoral fin rays 18 19.0(17-21) 27 Precaudal vertebrae 11 11 15 Caudal vertebrae 33 32.9 (32-34) 24 Total vertebrae 44 43.9 (43-45) 24 Rakers on anterior gill arch 15 15.1 (13-17) 34 Pseudobranch ial filaments 2 2.0(2-3) 32 D/V 6 6.3 (6-7) 24 D/A 19 19.7(18-23) 24 V/A 13 13.2(13-14) 24 Morphometric characters in % of SL Head length 28.2 27.2 (25.7-29.2) 35 Head width 14.4 13.6(10.9-17.2) 34 Head height 18.2 17.2(14.7-20.0) 34 Upper jaw length 14.0 13.4(11.8-14.6) 34 Maxillary height 4.4 4.3 (3.7-5.0) 35 Diameter of pigmented eye 3.4 2.8 (2.1-3.4) 36 Interorbital width 7.3 6.9 (5.7-7.8) 35 Postorbital length 20.1 19.7(17.9-21.5) 28 Preanal length - 46.1 (39.3-49.1) 35 Predorsal length 34.9 32.9 (29.2-35.2) 35 Body depth at origin of anal fin 19.3 18.4(15.1-20.8) 34 Pectoral fin length - 16.5(11.9-18.9) 32 Pelvic fin length - 24.3 (21.4-27.6) 19 Base pelvic fin - anal fin origin 23.7 25.8 (22.8-30.1) 34 and W.A. Starck, 1 Nov. 1981; AMS 1. 37229-010, 1 male, 66 mm SL, Vanuatu; AMS I. 37920-031, I male, 56 mm SL, 2 females, 50-58 mm SL and 1 juvenile, 32 mm SL, 13°S, 167°E Vanua Lava, Banks Islands, Vanuatu; AMS I. 39013-013, 1 female, 67 mm SL, 10°S, 166°E Reef Islands, Santa Cruz Islands, Solomon Islands; AMS I. 39033-003,1 male, 46 mm SL, 10°44’S, 166°49’E, Graciosa Bay, Nendo Islands, Santa Cruz Islands, Solomon Islands, 26 Sept. 1998; BPBM 32584, 1 female, 49 mm SL, 05°S, 145°E, Kranket Island, Madang, Papua New Guinea, J.E. Randall, 12 Nov. 1987; CAS 65661,2 females, 47-54 mm SL, 05°S, 145°E, Madang, Papua New Guinea, S.G. Poss, D.G. Catania and party, 12 May 1987; ROM 78116, 1 female, 42 mm SL, 09°S, 159°E, Guadalcanal, Solomon Islands; USNM 99061, 1 female, 100 mm SL, 03°S, 126°E, Uki, Buru Island, Maluku Province, Indonesia, RV Albatross Expedition, 9 Dec. 1909; USNM 99169, I female, 77 mm SL. 03°S, 126°E, Tifu Bay, Buru Island, Maluku Province, Indonesia, RV Albatross Expedition, 10 Dec. 1909; USNM 210252, I male, 37 mm SL and I female, 37 mm SL, 03°S, 128°E, Ambon, Maluku Province, Indonesia; USNM 377206, 1 female, 50 mm SL, 05°S, 145°E, Madang, Papua New Guinea; USNM 361548,1 female,45 mm SLJ3°32’S, 140 Dinematichthyine fishes of the Indo-West Pacific Fig. 55. Diancistrus novaeguineae (Machida, 1996). Holotype. BMNH 1974.5.25.3586, male, 37 mm SL. 167°20’E, Banks Islands, Vanuatu; USNM 366473, 4 females, 36-58 mm SL, 04°20’S, 129°54'E, Banda Islands, Banda Sea, Maluku Province, Indonesia; USNM 366481, 3 females, 39-60 mm SL, 05°S, 145°E, Madang, Papua New Guinea; USNM 366497, 3 females, 30-59 mm SL, 03°47’S, 128°06’E, Ambon, Maluku Province, Indonesia; USNM 366502, 2 females, 34-62 mm SL, 05°S, 145°E, Madang, Papua New Guinea; USNM 372966, 1 male, 82 mm SL, 23°25’S, 151°55’E, Heron Island, Great Barrier Reef, Queensland, Australia, J.H. Choat, 23 Feb. 1967; USNM 384600, 1 male, 45 mm SL and 1 juvenile, 28 mm SL, 16°47’S, 168°21’E, Epi Island, Vanuatu, J. T. Williams et ai, 16 June 1996; USNM 384601, 1 male, 48 mm SL and 3 females, 49-65 mm SL, 19°31’S, I69°29’E, Tanna Island, Vanuatu, J.T. Williams etui, 2 June 1996; ZMUC P 771490, 1 male, 61 mm SL, same data as AMS I. 18739- 120; ZMUC P 771491, 1 female, 56 mm