Records of the Museums and Art Galleries of the Northern Territory Volume 27 December 2011 The Beagle, Records of the Museums and Art Galleries of the Northern Territory (formerly ‘Records of the Northern Territory Museum of Arts and Sciences ’) ISSN 0811-3653 © Museums and Art Galleries of the Northern Territory, 2011. Printed by the Government Printing Office of the Northern Territory Volume 26 for 2010 was published on 17 December 2010 EDITORIAL COMMITTEE Richard C. Willan Editor Michael F. Braby Christopher J. Glasby Philip Short 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; Northern Territory and Oceanic history and archaeology. The Beagle is published once or twice a year, depending upon the material available. All contributions to The Beagle are reviewed by two referees and, where possible, at least one is internationally-based. Whilst articles for The Beagle w ill normally be 10-50 pages in length, shorter communications, notes and review articles may also be acceptable. Longer articles, significant works or substantial revisions, w'hich 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). All orders, subscriptions, back numbers and exchange enquiries should be addressed to the Library Manager, Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin NT 0801, AUSTRALIA, or e-mail library.dam@nt.gov.au 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 Gallery’ of the Northern Territoiy GPO Box 4646, Darwin NT0801, AUSTRALIA E-mail: richard. willan@nt.gov. au MUSEUM AND ART GALLERY NORTHERN TERRITORY Front cover: Helicteres serpens, a new species of creeping plant, is endemic to the northern part of the Northern Territory (see Cowie, pages 48^49). Drawing by Monika Osterkamp. Records of the Museums and Art Galleries of the Northern Territory Volume 27, December 2011 CONTENTS BROCKWELL, S., BOURKE, P., CLARKE, A., CRASSWELLER, C„ FAULKNER, P„ MEEHAN, B„ O’CONNOR, S., SIM, R. and WESLEY, D. - Holocene settlement of the northern coastal plains, Northern Territory, Australia.1 JACKES, B.R. - The vine Embelia tiwiensis (Angiospermae: Myrsinaceae), a new species from the Northern Territory, Australia.23 COWIE, I.D. - New taxa and notes on Helicteres L. (Malvaceae: Helicteroideae) from the Northern Territory, Australia.27 ALVAREZ, B. and HOOPER, J.N.A. Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of northern Australia. Family Halichondriidae.55 BRABY, M.F. - New larval food plant associations for some butterflies and diurnal moths (Lepidoptera) from the Northern Territory and eastern Kimberley, Australia.85 BROWN, G.R. - Dimorphothynmis (Hymenoptera: Thynnidae: Rhagigasterini) newly recorded from the Northern Territory, Australia. 107 BRUCE, A.J. - Periclimenaeus pulitzerfinali sp. nov., a new pontoniinc shrimp (Crustacea: Decapoda: Palaemonidae) from East Africa.113 DAVIE, P.J.F. - A new species and new records of Oncinopus (Crustacea: Brachyura: Inachidae) from northern Australia.123 REID, A. - Euprymna pardalota sp. nov. (Cephalopoda: Sepiolidae), a new dumpling squid from northern Australia.135 NIELSEN, J.G. and SCHWARZHANS, W. - A new genus and species of Bythitidae (Teleostei: Ophidiiformes) from northwestern Australia.143 SCHWARZHANS, W. and NIELSEN, J.G. - Revision of the genus Microbrotula (Teleostei: Bythitidae), with description of two new species and a related new genus...147 SCHWARZHANS, W. and M0LLER. P.R. - New Dinematichthyini (Teleostei: Bythitidae) from the indo-west Pacific, with the description of a new genus and five new species.161 SOMMER, J.A., LI, C„ BROZEK, J„ BESSERT, M.L., ORTI, G. and BERRA, T.M.- Low genetic diversity in Nurseryfish, Kurtus gulliveri (Perciformcs: Kurtidae), and an appraisal of its breeding system using microsatellite loci. 179 TAKITA, T„ LARSON, H.K. and ISHIMATSU, A. - The natural history of mudskippers in northern Australia, with field identification characters 189 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 21: 1-22 Holocene settlement of the northern coastal plains, Northern Territory, Australia SALLY BROCKWELL 1 , PATRICIA BOURKE 2 , ANNE CLARKE 3 , CHRISTINE CRASSWELLER 4 , PATRICK FAULKNER 5 , BETTY MEEHAN 1 , SUE O’CONNOR 1 , ROBIN SIM 1 and DARYL WESLEY 1 'Department of Archaeology and Natural Histoiy, School of Culture, History and Language College of Asia and the Pacific, Australian National University, Canberra ACT 0200, AUSTRALIA sally. brockwell@anu. edit, att 2 Heritage Branch, Department of Natural Resources, Environment, The Arts and Sport, PO Box 496, Palmerston NT0830, AUSTRALIA department of Archaeology, School of Philosophical and Historical Inquiry, University> of Sydney, Sydney NSW 2006, A USTRALIA 4 8 Wanguri Terrace, Wanguri NT0810, AUSTRALIA 5 School of Social Science, University of Queensland, St Lucia QLD 4072, AUSTRALIA ABSTRACT The northern Australian coastal plains are relatively recent landforms that have undergone dynamic evolution over the last 10,000 years. Over 300 radiocarbon dates have enabled archaeologists and geomorphologists to provide a more detailed interpretation of human settlement and resource use. This paper provides a synthesis of the archaeological evidence and integrates it within the palaeo-cnvironmental frameworks. It characterises the timing, pattern and nature of human-environment interaction in this newly formed landscape over the last 10,000 years. Keywords: Northern Australia, coastal plains, landforms, anthropology, archaeology, radiocarbon dates, human settlement, resource use. INTRODUCTION Following the post-Pleistocene marine transgression, broad plains formed along coastal northern Australia and continued to develop throughout the Holocene through processes of sedimentation and coastal progradation. They contain large tidal flats and salt pans interspersed with cheniers and are drained by substantial rivers with extensive floodplains and estuaries (Lees & Clements 1987: 312). There has been considerable archaeological and geomorphological work undertaken on the Northern Territory coastal plains, although mostly independent of each other. The notable exceptions are joint studies on the South Alligator River (Hope et 'al, 1985; Woodroffe et al. 1988), the Blyth River by John Chappell and Rhys Jones (unpublished data), and the Sir Edward Pellew Group (Prebblc et al. 2005). The geomorphological evidence demonstrates that the coastal plains of northern Australia are relatively recent formations that have undergone dynamic evolution from the mid to late Holocene. Studies have been carried out on the Victoria and Daly Rivers in the west, Darwin Harbour, the Adelaide, Mary and South Alligator Rivers, Magela Creek, Blyth River, and Grootc Eylandt and the Edward Pellew islands in the east (Fig. 1). The evolutionary sequence of the coastal plains was similar right across the Northern Territory, the differences being caused by topography and timing. How did humans respond to these newly created landscapes? This question has been addressed by an extensive body of research over the past 40 years focused on the archaeology of the Northern Territory coastal plains. The early research effort was concentrated mainly in the Alligator Rivers region of what is today Kakadu National Park (Kamminga & Allen 1973;Schrire 1982; Jones 1985; Allen & Barton 1989), but the last 15 years has seen many other regions investigated. As a result of this research, many more radiocarbon dates have become available, providing an emerging pattern of Holocene settlement across the coastal plains of the Top End as a whole. This paper presents a synthesis of these new data to provide an overview of the chronology of occupation and the changing nature of the archaeological record, which includes several different site types - rockshelters, shell mounds, shell middens, earth mounds, artefact concentrations and contact sites. 1 S. Brockwell et al. Fig. 1. Location map, showing the Top End of the Northern Territory. In addition, information about palaeoclimates and the geomorphological evolution of the northern coastal plains is summarised to provide the environmental framework for the archaeological data. Visibility plays a major role in site location. Archaeological sites are difficult to observe in some coastal environments such as lateritc plains and swampy areas. During the marine transgression, former shorelines and associated archaeological evidence were submerged (Woodroffe et al. 1988). As well, the extensive nature of mangrove cover during the Big Swamp phase may have almost completely removed evidence of occupation, most of which for this period is found in rocksheltcrs. Open sites proliferated after sea level stabilisation approx. 6000 years ago, which is almost certainly a function of improved visibility. PALAEOCLIMATE It is widely acknowledged that there has been an increase in climatic variability in the Australasian region in the last few thousand years, in particular from approximately 1000 BP to the present (Kershaw 1983; Wasson 1986; Gagan et al. 1994; Kershaw 1995; Shulmeister 1999:82; Gagan & Chappell 2000; Prebble et al. 2005; Wasson & Bayliss 2010; Williams et al. 2010). Many of the longer- term trends in climate change that have occurred during the period spanning the mid Holocene to the present day are related to the El Nino/Southem Oscillation (ENSO) cycle, which has strongly influenced climatic patterns in Australia (McGlone et al. 1992; Shulmeister & Lees 1992; Jones et al. 1999; Shulmeister 1999; Gagan et al. 2004; Turney & Hobbs 2006), and at present represents the principal source of inter-annual climatic variability within the Indo-Pacific region (Glantz 1991; Diaz & Markgraf 1992; Allan et al. 1996; Rowland 1999) (Fig. 2). The pattern of climatic shifts portrayed in recent reviews of the climate history of the South Pacific region indicates a change from low seasonality in the early Holocene to increased seasonality in the late Holocene (Markgraf et al. 1992; Shulmeister 1999). There also appears to be a general trend toward increased aridity in the mid to late Holocene, as supported by data extracted from coral, foraminifera, and varve, lake and sea bottom sediments from sites in Australia and the circum-Pacific region (McPhail & Hope 1985; Brookfield & Allan 1989; Singh & Luly 1991; McGlone et al. 1992; Hope & Golson 1995; Kershaw 1995; Nott et al. 1999; Rodbell et al. 1999; McCarthy & Head 2001; Kim et al. 2002; Koutavas et al. 2002). Evidence from pollen records on Groote Eylandt are also indicative of this environmental change from continuously increasing rainfall (effective precipitation) during the early Holocene, to a period of reduced rainfall and increased climatic variability after 4000 BP (Shulmeister & Lees 1995). In the Groote Eylandt archipelago, effective precipitation declined sharply soon after 3700 BP, with evidence across Australia indicating that climate became more variable after this time (Shulmeister & Lees 1995). Gcomorphic data from cheniers and coastal dunefields (Lees & Clements 1987; Lees et al. 1990; Lees 1992; Lees et al. 1992) indicate that some of the observed changes in these systems were synchronous across northern Australia, and may represent coherent, broad-scale climatic signals (Shulmeister 1999; Prebble et al. 2005) (Fig. 2). 2 Holocene settlement of northern Australian coastal plains Age (Thousand years BP) 0 2 4 6 8 10 North Australian Climate B 80 Number 60 ENSO Events 40 20 C Progradation: Alligator Rivers Fig. 2. North Australian Holocene environmental and climatic phases (grey shading indicates main mounding period). - 1 - 1 - 1 -- —1 - V Increased Climatic Variability Sharp Decline in Effective Precip. Max. Effective Precip. Sea-Level Effects Dominant Gradual Increase in Effective Precipitation LANDSCAPE The dated archaeological sites are located roughly in five regions - (I) the West Coast represented by the Reynolds River (Guse & Majar 2000; Guse 2005); (2) the Darwin Region, including Darwin Harbour, Hope Inlet (Hiscock 1997; Bourke 2000; Hiscock & Hughes 2001; Crassweller 2002; Bourke 2004, 2005a, 2005b; Bourke & Crassweller 2006; Crassweller 2006;); theAdelaide River(Smith 1995; Crassweller 1996; Brockwell 2001,2006a, 2009) and Mary River (Baker 1981; Guse 1992); (3) western Arnhem Land, including the Alligator Rivers and Magela Creek (Schrire 1982; Meehan et al. 1985; Woodroffe et al. 1988; Allen & Barton 1989; Roberts et al. 1990; Mowat 1995; Roberts et al. 1998), Bald Rock and Malarrak in the Wellington Ranges (May et al. 2010) and the Cobourg Peninsula (Mitchell 1994a, 1994b, 1996); (4) central Arnhem Land represented by the Blyth River and Milingimbi (McCarthy & Setzler 1960; Mulvaney 1975; Meehan 1982, 1988, 1991; Roberts 1991, 1994; Brockwell et al. 2005); and (5) eastern Arnhem Land along the east coast of the Gulf of Carpentaria, including Cape Arnhem (Bourke 2001); Port Bradshaw (Schrire 1972), Blue Mud Bay (Faulkner & Clarke 2004; Faulkner 2006, 2008, 2009; Faulkner & Clarke 2009), Groote Eylandt (Clarke 1994; 2000a, 2000b; Clarke & Frederick 2006; 2008, 2011) and the Edward Pcllew islands (Sim 2002; Prebble et al. 2005; Sim 2005; Sim & Wallis 2008) (Fig. 1). The landscape covered by the study areas is diverse and includes: dissected sandstone and lateritic plateaux bordering the coastal plains; large rivers and creeks; immense floodplains containing vast freshwater wetlands; and saline mudflats further towards the coast. Mangrove forests fringe coastal cheniers and tidal mudflats, headlands protrude onto coastal salt fiats, mixed woodland dominates on slopes and plains. There arc large stretches of open mixed woodland, broad sandy beaches and extensive dune fields, pockets of monsoon rainforest, and grassland, broad sandy beaches, shallow bays and off-shore islands, some substantial in size that can sustain permanent human populations. ARCHAEOLOGY AND CHRONOLOGY This complex landscape is reflected in the archaeology, which consists of numerous occupied rockshelters dating back to the Pleistocene (Kamminga & Allen 1973; Schrire 1982; Jones & Johnson, 1985a, 1985b; Allen & Barton 1989, Roberts et al. 1990, 1998), thousands of panels of magnificent rock art in the escarpment and outliers on 3 S. Brockwell et al. 2 Sigma summed probability age ranges by location Reynolds River (n = 8) :■ on xi n n in 0 2000 4000 6000 8000 10000 Adelaide River (n = 14) m ii ■imi n an 0 2000 4000 6000 8000 10000 Cobourg Peninsula (r> = 18) ann □ mo 0 2000 4000 6000 8000 10000 Milingimbi (n = 11) hum in Dm 0 2000 4000 6000 8000 10000 Port Bradshaw (n = 3) i ■ cm 0 2000 4000 6000 8000 10000 Groote Eylandt (n = 33) ■O 1 0 0 2000 4000 6000 8000 10000 Fig. 3. C14 graphs by location. the coastal plains (Chaloupka 1985; Lewis 1988; Tason 1989; Chaloupka 1993; Tapon 1993; Taijon & Chippindale 1994; Ta?on & Brockwell 1995;TaQon<?7r//. 1996;Guse& Woolfe 2006; May et al. 2010; Ta<;on et al. 2010; Wesley in press), clusters of earth mounds and extensive artefact concentrations along the floodplain margins (Baker 1981; Smith 1981; Meehan et al. 1985; Brockwell 1989; Guse 1992, Hiscock et al. 1992; Brockwell 1996; Hiscock 1996; Guse & Majar 2000; Brockwell 2001, 2005; Guse 2005; Brockwell 2006a, 2006b, 2009), and middens and shell mounds on the lower floodplains and along the coast (McCarthy & Setzler 1960; Mulvaney 1975; Meehan 1982; Woodroffe et al. 1988; Hiscock & Mowat 1993; Clarke 1994; Roberts 1994; Mowat 1995; Hiscock 1997; Bourke 2000,2001,2004; Faulkner & Clarke 1994; Bourkc 2005b; Brockwell etal. 2005; Bourke & Crasswcllcr 2006; Faulkner 2006,2008; Sim & Wallis 2008; Faulkner 2009). Dates have been calibrated in order to ensure a robust comparison of the conventional radiocarbon ages from different regions and the different site types. Over 300 radiocarbon dates are now available. Details of calibration methods and a tabulated list of dated sites can be found Darwin Region (n = 81) I IILM_B_I 0 2000 4000 6000 8000 10000 Kakadu Rsgion (n = 34) nno ochmm an □ □1 0 2000 4000 6000 8000 10000 Blyth River (n = 22) i □ 0 2000 4000 6000 8000 10000 Cape Arnhem (n = 3) cm m 0 2000 4000 6000 8000 10000 Blue Mud Bay (n = 38) ■»rn ■Ti 0 2000 4000 6000 8000 10000 Edward Pollew Islands (n = ■OD [ZED (HDD 57) E □ □ 0 2000 4000 6000 8000 10000 in Brockwell el al. (2009). Evidence from rockshelters suggests that Aboriginal people began to occupy and use the resources of the coastal plains at the time of their formation over 10,000 years ago, but the majority of sites are 4000 years or younger. Fig. 3 illustrates chronology by location. On the basis of the environmental evidence, the archaeology can be divided into five temporal periods: 10,000-6000 BP. Evidence of occupation on the coastal plains has been dated from the early to mid Holocene period, encompassing the time of rising sea levels and the ‘Big Swamp Phase’. Effective precipitation and temperature were gradually increasing during this phase. During the post-Pleistocene transgressive phase, the down-cut river valleys of northern Australia were drowned. The various river systems responded differently to this event. Some, like Darwin Harbour, became deep-water embayments. Others, through processes of sedimentation, formed vast mangrove swamps. This has been described as the ‘Big Swamp Phase' and dates from about 7000 BP to approx. 4000 BP (Woodroffe et al. 1985, 1986; Chappell 1988; Woodroffe 1988; Woodroffe & Mulrcnnan 1993) (Fig. 4). 4 Holocene settlement of northern Australian coastal plains Fig. 4. Developmental phases of the northern river floodplains. Allen (1996: 198,201; Allen & Barton 1989: 104) made the point that the mangrove forests of the Big Swamp Phase would have presented a formidable barrier to movement and settlement. Access to the river channels and the coastal plains may have been impossibly difficult. The distribution of sites during the Big Swamp Phase in western Arnhem Land reflects this situation and is consistent with Allen and Barton’s prediction that “Seasonal movement and sites would have been restricted to the plateau valleys, the plateau margins, and the lowland corridors between the tidal floodplains” (Allen & Barton 1989: 104). This period is represented by cultural sequences mainly from rockshclters and two middens. In the west, Djagorda 2 shelter on the Reynolds River contained a low density of stone artefacts (Guse 2005: 113). At the edge of floodplains in sub-coastal western Arnhem Land, the lower levels of Nawamoyn and Ngarradj Warde Djobkeng shelters in the Alligator Rivers region were dominated by estuarine shellfish soon after the establishment of the vast mangroves of the Big Swamp Phase approx. 7000 BP (Kamminga & Allen 1973: 45-47; Schrire 1982: 85-97, 118-30; Allen & Barton 1989: 89). At Nawamoyn and Malakunanja II fragments of mammals (open woodland), fish (unidentified) and reptiles (woodland) occurred throughout the midden levels, as did stone artefacts and bone tools (mostly spatulate points at Nawamoyn). There were human burials in Nawamoyn and Malakunanja II. Malarrak and Bald Rock in the Wellington Ranges were first occupied in the Pleistocene and appear to have been occupied throughout the Holocene (May et al. 2010). Although not dated, the base of the deposit at Bomgolo Shelter in eastern Arnhem Land contains marine fauna, which Schrire (1972: 662) surmises is consistent with occupation after sea level stabilisation 6000 -7000 years ago. Also in eastern Arnhem Land, the lower levels of Mushroom Rock (VB17) and Wobuya Shelter (WS) on Vanderlin Island in the Edward Pellcw islands contain stone artefacts and the remains of shellfish from mangroves and open mudflats (Sim & Wallis 2008: 102) (Fig. 5). During this period, the Edward Pellew islands were still attached to the mainland. Following the post-glacial marine transgression, the islands appear to have been abandoned approx. 6700 years BP and not reoccupied until 4200 years ago (Sim & Wallis 2008: 102). In western Arnhem Land, one date was taken from a midden (Kapalga P) buried on the lower South Alligator River floodplains, which is dominated by the mangrove shellfish Telescopium telescopium. The substrate contains mangrove wood from the Big Swamp Phase (Woodroffe et al. 1988: 97). Muyu-ajirrapa midden on the Blyth River in central Arnhem Land was also dated to this period. However, there may be a problem with this charcoal date, as its shell pair was much younger. The midden is dominated by Dosinia sp„ an open beach species that proliferates in sandy mud flats. Although direct correlation of art styles, climatic phases and Aboriginal settlement is problematic, it has been argued that the pattern of settlement and economic organisation described above is reflected in the rock art, and can suggest 5 S. Brockwell et al. Fig. 5. Vanderlin Island rockshelter. Photograph: Robin Sim. insight into the underlying social systems. However, it must be pointed out that these conclusions are highly interpretive and are not closely tied to the radiocarbon chronology. For example, it is argued for western Arnhem Land that the so- called ‘Dynamic Figures’ depicted in hunting scenes with macropods and other terrestrial fauna are associated with the arid, inland environment that existed approx. 10,000 years BP (Chaloupka 1993; Chippindale & Taqon 1998) (Fig. 6). The mostly male figures wear elaborate head dresses, pubic fringes and bustles. They use weapons that include single pronged spears, boomerangs and hafted axes, and carry dilly bags (Chaloupka 1993: 106-110; Ta 9 on&Chippindale 1994;Ta9on& Brockwell 1995: 684). The head dresses and bustles have been compared with those commonly found today in Papua New Guinea and West Papua, and may refer to a time when the land formed one continent with Australia (Chaloupka 1993:110).Ta9on(1994:113)argues that the rate of art production varied over time and that this period is one of the periods when rock art was produced prolifically. The rock art of this period is also regionally consistent between central and western Arnhem Land, which implies cultural uniformity over a wide area (Tagon 1993: 118). Layton (quoted in Taqon & Chippindale 1994: 216, 225) suggests that this is the result of environmental conditions that required collective hunting strategies to capture large terrestrial fauna. The implied later art of this period, the so-called ‘Post-Dynamic Figures’ (Chaloupka 1985; Chippindale & Taqon 1998: 107) are shown in more static poses and their relationship to the Dynamic Figures is not known. 6000-4000 BP. The period 6000-4000 BP encompassed sea level stablilisation and the decline of the ‘Big Swamp Phase’. Following the cessation of sea-level rise approx. 6000 BP, sedimentation and coastal progradation resulted in vertical accretion of the floodplains, and caused the development of extensive coastal plains. Effective precipitation and temperature continued to increase until approximately 5000 BP. With the decline of tidal influence, mangroves retreated seawards and towards the banks of the rivers and had mainly disappeared from the floodplains by 4000 years BP (Woodroffe et al. 1988: 98; Woodroffe & Mulrennan 1993) (Fig. 4). There are dates from rockshelters, shell mounds and earth mounds during this period. On the Reynolds River, three Tabletop Range rockshelters, Djagorda 2 and Walker Creek 6, were excavated. Although only Majar was dated to this period, dates and rates of deposition from these sites imply an increasing site usage in the region after 4500 BP. There was also a large increase of stone artefacts deposited at Majar approximately 4000 years ago (Guse 2005: 97). In western Arnhem Land, there is a marked decrease in the mangrove-associated shellfish Polymesoda erosa and Teleseopium telescopium and an increase in Cerithidea sp. [probably C. anticipate! ] between the lower and upper midden levels at Malangangerr and Nawamoyn (Schrire 1982: 88-89, 120-22, 233-34; Hiscock 1999: 94-95), linking these sites to the decline of the Big Swamp Phase. Telescopium telescopium prefers a Rhizopliora mangrove habitat, which dominated in the Big Swamp Phase. Similarly, Polymesoda erosa is likely to be more abundant in Rhizopliora forests where it is found along small streams. Cerithidea sp. can be gathered in large numbers from the trunks of Avicennia and Bruguiera mangrove trees, which survive in drier conditions, and replaced Rhizopliora at the end of the Big Swamp Phase in archaeological sites in western Arnhem Land (Hiscock 1999: 95-96). Malangangerr and Nawamoyn contain many bone tools, with an increase in bipoints in the upper midden levels (Schrire 1982: 85-97). Based on ethnographic evidence, which associates bone uni and bipoints with multi-pronged spears used for fishing in western Arnhem Land, Schrire (1982: 63, 95, 249) speculates that this pattern indicated a shift from exploitation of mangrove/mudflat shellfish to a greater reliance on fishing towards the end of this period. Malakunanja II contains a similar range of fauna to Malangangerr. It also contains burials (Kamminga & Allen 1973: 45-46). Allen & Barton (1989: 90) argue that the dating of the top spit of Malakununja II to the latter part of this period indicates that this site was mostly abandoned coincident with the retreat of the mangroves. Uni facial and bifacial stone points first appeared in the sands below the midden at Ngarradj between 4862 - 3837 cal. BP (Allen & Barton 1989: 93). As Allen and Barton said of the rockshelters in western Amhem Land, “The fact that every date that can be reliably associated with the mangrove/ mudflat shell middens at these sites falls between 3000 and 7000 years BP suggests a high degree of correlation between the archaeological record and the geomorphological evidence for the local occurrence of mangroves” (1989:90). In eastern Amhem Land the lower level at Bomgolo, the top of which is dated to this period, contains stone and bone tools, and a wide variety of fauna that reflects exploitation of the open sea (dugong and fish), sandy shore (turtles and turtle eggs), rocky shore (oysters, fish, crabs), mudflats (estuarine shellfish) and land (mammals, reptiles, echidnas, 6 Holocene settlement of northern Australian coastal plains Fig. 6. Dynamic figure. Photograph: Daryl Wesley. birds) (Schrire 1972: 662, 665). There is no evidence for occupation during this period at Vanderlin Island in the Edward Pellew islands, which appears to have been abandoned just before it was truncated from the mainland by rising sea levels (Sim & Wallis 2008: 102). /O/r/f/t/ra-dominated shell mounds begin to appear during this period in the west at Werat 1 on chenicr ridges of the coastal plains of Reynolds River (Gusc 2005: 114), and on Field Island 1 and 2 in western Arnhem Land (Hiscock & Mowat 1993; Mowat 1995: 154-55). This pattern reflects exploitation of beach species from open sandy mud flats, the preferred habitat of Anadara granosa. On the West Alligator River some undated shell mounds were dominated by Anadara granosa, and others by open beach species such as Marcia hiantina and Circe sp., which suggests that the occupants were exploiting open mud and sandy beaches extant in the mid-Holocene (Mowat 1995: 154-55). Variability in species composition between these mounds is linked to the formation of mounds at different times and reflects changing environmental conditions along the coast in the mid to late Holocene (Mowat 1995: 154, 163). Mowat (1995: 163) concludes that the coastal areas were not abandoned as conditions changed, but rather the inhabitants adapted their foraging strategics accordingly. Hiscock (1999:96) observes that the existence of shell mounds from different regions indicates widespread foraging along the coast at this time. The base of earth mound HD1 on the now sub-coastal portion of the Adelaide River floodplains contains estuarine shell associated with the Big Swamp Phase (Brockwell 2006a); and while data about shell species were not colleeted, Woodroffe et ah (1988: 97) record grindstones, hammerstones and manuports from the surface mound Kapalga H2 on the black soil plains of the lower South Alligator River in western Arnhem Land, which elsewhere has been classified as an earth mound (Brockwell 2006b). At Point Stuart on the mouth of the Mary River, Baker (1981: 79) infers a date of 4500 years BP for a midden located on a chenier ridge on the basis of geomorphological investigations. In the lower levels Telescopium telescopium and Crassostrea sp. dominate the midden. These estuarine species inhabit stands of Rhizophora mangoves, which were extant on the floodplains during the Big Swamp Phase. Rock art during this period, such as ‘Simple Figures with boomerangs’ and ‘Yam Style Figures’ (Chaloupka 1985), shows increasing diversity and regionalisation (Ta ? on 1993: 114; Ta?on & Chippindale 1994:217). Early X-ray art also appears. It shows simple internal features and is dated to a minimum age of 4000 years BP, based on radiocarbon dating of beeswax from a rockshelter in western Arnhem Land (Nelson et al. 1995; Watchman & Jones 2002). A greater diversity of animal species is depicted, including those from the estuarine floodplains of the Big Swamp, such as barramundi (Lates calcarifer), catfish (Arias sp.) and estuarine crocodile (Crocodylus porosus) (Chaloupka 1985: 277; Lewis 1988: 50-55; Chaloupka 1993: 162; Ta?on 1993; Ta$on & Chippindale 1994: 217; Chippindale & Tapon 1998: 107). There are changes in 7 S. Brockwell el al. material culture too. Boomerangs, associated with hunting large animals on open plains, disappear and new weapons make their appearance, including spear-throwers, spears with hafted spear points and three-pronged spears associated with fishing (Chaloupka 1985: 277; Lewis 1988: 55, 96; Chaloupka 1993: 146; Tagon & Chippindale 1994: 224; Chippindale & Tagon 1998; Brockwell & Akerman 2007). It is suggested that large battle scenes depicted in the art of this period may reflect more tightly organised societies in conflict over access to resource bases (Tagon & Chippindale 1994: 224-225). Lewis (1988: 90) suggested that conflict might have been created by the drowning of coastal land by rising seas and forced migration inland onto other territorial land, although there is no direct evidence for this. It is also suggested by some authors that the Yam Figures might be associated with rising seas (Chaloupka 1985; Lewis 1988; Chaloupka 1993: 138-145; Tagon & Chippindale 1994; Tagon el al. 1996). Chaloupka (1985: 276) argued that yams need over 120 cm of rainfall to thrive and may have been an important staple replacing grass seeds in the preceding drier period. He also comments that the break from both previous and subsequent styles was such that it implies major ideological and cultural shifts (1993: 143). Likewise, Tagon el al. (1996) argue that the appearance of the Rainbow Serpent motif, a hugely important feature of current mythology, was associated with increased rainfall and sea-level rise in the mid-Holocene. Tagon & Brockwell (1995: 25) point out that land lost to rising water was probably observable in a lifetime. Tagon el al. (1996: 117) argue that “Ultimately, the most unusual aspects of change — snakes slithering away from drowning landscapes, rainbows overhead and strange ‘new’ creatures such as pipefish washed ashore —were combined into a powerful metaphor and symbol of change itself.” Lewis (1988: 91) suggests that the Rainbow Snake, which became a common image in many different accessible locations at or near campsites in this period, acted as a unifying symbol in times of change. Drawing an association between the mid-Holocene establishment of the monsoon pattern of rain and rock art, Chaloupka (1985: 277) speculates that the Lightning Man ( Namarrgon ) may have appeared in the art at this time. 4000-2000 BP. Sites dated to 4000-2000 BP encompass a Transition Phase between estuarine conditions and the beginnings of the Freshwater Phase on the floodplains of the north. Higher effective precipitation continued into this period until approximately 3700 BP, followed by increasing patterns of aridity. The rivers underwent a so- called Sinuous Phase’, which describes their meandering nature across the coastal plains. The following ‘Cuspate Phase’ of river development refers to the formation of wide reaches and pointed bends, which occurred after 2500 years ago. Continued sedimentation and the slowing of coastal progradation during this phase led to a cut-off of the tidal influence. Palaeochannels were created and the floodplains contained a mosaic of estuarine, freshwater and mudflat areas (Woodroffe el al. 1986; Chappell 1988; Woodroffe 1988; Woodroffe & Mulrennan 1993) (Fig. 4). Increasing numbers of shell mounds, earth mounds and middens fall within this time period. Initial observations suggest established settlement patterns developed in the Reynolds River area after 4000 BP. Another indicator is perhaps the uniformity of the artefact assemblages across the different environmental zones after 4000 BP (Guse 2005: 112). In western Arnhem Land, Allen & Barton (1989: 90-91) concluded that all four rockshcltcr sites, Malakunanja II, Malangangerr, Nawamoyn and Ngarradj Warde Jobkeng, were mostly abandoned by the end of the Transition Phase and only sporadically occupied afterwards. However, Fliscock (1999: 96) argued that the timing for this model of abandonment may be too rigid and that at the rockshelter of Paribari accumulation of midden deposit continued for sometime afterwards. The existence of an intermediate saline mudflat phase in the development of the floodplain is consistent with and confirmed by the compositional changes in the middens in the nearby archaeological sites. At Malangangerr, Nawamoyn and Paribari, there is a shift over time from exploitation of the bivalve species Polymesocla erosa in the lower midden levels to the mud whelk Cerithidea sp. in the upper levels (Schrire 1982: 233). This event is dated to about 3000 years BP at Paribari (Schrire 1982: 51). The abandonment of Malangerrangerr, Nawamoyn, Ngarradj Warde Jobkeng and Malakanunja II at the same time suggests local shifts in settlement and subsistence according to the increasingly mosaic environment on the floodplains during the Transition Phase (Hiscock 1999: 96). According to Allen and Barton (1989: 93-94), the earliest secure date for the appearance of stone unipoints and bipoints in western Arnhem Land plains rockshelters is 4000 BP and they were not widespread in the region until after 3500 BP, although Jones & Johnson (1985a: 206) argue tor an older date for their first appearance. The fact that these tools appear for the first time only in the upper levels of Nawamoyn and Malanangangcrr supports the abandonment theory approx. 3000 BP. Stone unipoints and bipoints appear only in the lower levels of Paribari. Surface mound (Kapalga N) on the lower South Alligator River coastal plains located above freshwater black soil (floodplain clay) contain Nerita balteata , which lives mangrove trunks, and Certhidea sp., found in the substrates of stands of the drier Avicennia and Bruguiera, suggesting exploitation of mangroves in the Transition Phase (Woodroffe el al. 1988: 96-97). There is an increase in numbers of open sites toward the end of this period around 2000 BP. Earth mounds I ID 1 and MP2 on the Adelaide River floodplains contain increasing quantities of stone artefacts, fish bone and bone points that have been associated with multi-pronged spears for fishing (Brockwell 2005, 2009; Brockwell & Akerman 2007) (Fig. 7). Nearby is Scotch Creek 1, an open site located next to a deep and permanent freshwater billabong Holocene settlement of northern Australian coastal plains Fig. 7. Bone points from the Adelaide River. Photograph: Darren Boyd. in open woodland two to three kilometres southeast of the main floodplains of the Adelaide River. In 1980, Smith (1981; Smith & Brockwell 1994) excavated a test pit and located cultural remains in a layer of black soil between 60 to 70 cm in depth above a clay substrate. A horizon of stone points located 10 to 15 cm above the clay substrate is dated to approx. 3000 BP and therefore to the Transition Phase (Smith & Brockwell 1994: 91; Smith 1995). On the basis of ethnographic association of stone points with light weight dueling spears, Smith (1995) speculates that the point horizon might represent a period of conflict associated with the changing and unpredictable nature of the resource base on the floodplains during this period. It is argued that the age of the site is no older than 4000 years as there are no mangrove shellfish present (Smith & Brockwell 1994: 93). The site has been occupied repeatedly since then (Crassweller 1996: 1). There are stone artefacts and faunal remains throughout the excavation (Smith & Brockwell 1994:91). The animal taxa present include macropods, bird, freshwater turtles and fish (the only identified species being Arius sp.). The faunal analysis indicates that macropods (open woodland) arc always a relatively more important taxon than at the main Adelaide River floodplain sites. Crassweller (1996: 78) concludes that the site was used as a base camp to exploit the billabong and open woodland and that manufacture and repair of stone artefacts took place there. Hiscock (1999: 97) suggests that settlement may have shifted away from the escarpment and floodplains to the coast in this period, which was rapidly prograding and probably productive during the Transition Phase. This is supported by the increasing number of coastal sites. Sites dating from this period include Wcrat 14 in the west, which is located on a chenier ridge of the coastal plains of Reynolds River. It is dominated by the open beach species Anadarci granosa (Guse 2005), as is MAI in the Darwin region (Bourkc 2000; 2005b). The shell mound Mari- Maramay 1 on Croker Island is dominated by Gafrariiim tumidum, another open beach species (Mitchell 1994a: 281). Macassar Well shell mound at Milingimbi (McCarthy & Setzler 1960; Mulvaney 1975) and the shell mounds at Blue Mud Bay (BMB/29, BMB/82) are also dominated by Anadara granosa (Faulkner & Clarke 2004; Faulkner 2006: 189; Hiscock & Faulkner 2006; Faulkner 2008,2009) (Fig. 9). In central Arnhem Land on the Blyth River, Maganbal midden dates to early in this period and is dominated by Anadara granosa. In eastern Arnhem Land, Gaynada midden on Cape Arnhem is dominated by Marcia luantina, another open beach species (Bourke 2001). At Blue Mud Bay, BMB/018 is the oldest dated midden in the study area and is dominated by Pinctada sp. (pearl oyster) found in shallow water and Melo sp. (baler shell) found on sandflats or mudflats (Faulkner 2006: 112; 2008). BMB/033 and BMB/067 are dominated by species from shallow water and sandy mudflats (Faulkner 2006: 169). On Grootc Eylandt, the base of Malmudinga midden is dominated by marine shells (Clarke 1994). Sub-coastally, several shell middens are recorded on the lower South Alligator River, western Arnhem Land. Kapalga D is located on a salt mudflat surface and dominated by mangrove shellfish species Terebralia palustris and Telescopiwn telescopium. Palaeochannel middens (concentrated shell) Kapalga F and K are located on freshwater black soil (floodplain clay) and on the bank of a palaeochannel, respectively. Shell from Kapalga F was not collected but Kapalga K contains the mangrove species Polymesoda erosa and Cerithidea sp. (Woodroffe et al. 1988:97). The mixed composition of the soil substrates and shell species and the concentrated location of sites suggest varying and mobile foraging strategies in response to the mosaic and dynamic nature of the floodplains during the Transition Phase. On the other hand, the pattern of occupation in rockshelters on Vanderlin Island in the Edward Pellew islands follows a somewhat different trajectory. While (Wobuya Shelter) was reoccupied or occupied for the first 9 S. Brockwell et al. Fi 8- 8 ' Depiction of fish (Malarrak, western Arnhem Land). Photograph: Daryl Wesley. time during this period (Wommgulumba, Babangi, Walala III, Victoria Bay III, Scissibar Creek, Boinmamda), this was followed by a low intensity or abandonment phase of these sites between 2500 and 1700 years BP (Sim & Wallis 2008). More complex and diverse rock art forms emerge towards the end of this period, including stick figures, abstract designs, beeswax figures and detailed X-ray art (Lewis 1988: 95; Tagon 1993: 114; Chippindale & Tagon 1998: 107). The X-ray art depicts a mixture of estuarine and freshwater species, especially fishes (Tagon 1987) (Fig. 8). It is argued that multi-pronged spears portrayed in the art arc associated almost exclusively with fishing as a major economic activity on the floodplains during the Transition Phase (Brockwell & Akerman 2007). Some rock art complexes in western Arnhem Land contain scenes of large numbers of human figures and spears in a melee that suggests fighting, perhaps over changing resource availability, during this Transition Phase. Most of this change occurs in the latter half of this period and accelerates throughout the next period, after 2000 BP (Lewis 1988: 309; Tagon 1993: 114). 2000-500 BP. This period encompasses the early Freshwater Phase, when freshwater from the annual monsoons ponded behind cheniers and in palaeochannels. By 2000 BP this process had created the vast freshwater floodplains and wetlands that are a major feature of the northern coastal plains today (Chappell 1988; Woodroffe et al. 1988) (Fig. 4). With a continuing decline in effective precipitation, climatic variability increased in the region from approx. 1000 BP. There are dates from increasing numbers of sites including rocksheltcrs (all but one located on the east coast), shell mounds, earth mounds and especially shell middens. 1 he midden dates are predominantly from 1000 to 500 BP. Allen and Barton (1989: 90-91) argue that the rockshelters of Malangangerr and Ngarradj Warde Jobkcng were reoccupied in this period after a period of abandonment of at least 2000 years. Although not dated, the upper levels of Paribari are dominated by freshwater fauna and flora that link the site firmly to the Freshwater Phase (Schrire 1982: 51-55, 58). Silica-polished adze flakes associated with experimental harvesting of Eleocharis dulcis (Akerman 1986), a reed that grows in freshwater swamps, are found in the upper levels of Paribari (Schrire 1982: 72). Polished flakes are also found in the upper levels at Nawamoyn (Schrire 1982: 137). There is a hint that Malakununja II continued to be occupied sporadically or was reoccupied in the Freshwater Phase, as freshwater mussel shell appears in the upper few spits (Kamminga & Allen 1973:45). Recent work on rockshelters in the Wellington Range in western Arnhem Land by Wesley and colleagues also indicates increased use of this environment after 1500 BP (May et al. 2010; Tagon et al. 2010). The resources of the wetlands and billabongs, coupled with a stable coastline, no doubt made these rocksheltcrs desirable habitation sites, particularly during the wet season. The inhabitants of Bomgolo shelter in eastern Arnhem Land continued to exploit a similar range of marine, rocky shore, mudflats and open woodland species, as they did in the lower level of the site (see above). Schrire (1972: 666) notes that fish bones increase in the upper level, while mammal bones decrease, which she concludes represents a change in subsistence patterns with increasing sedentism 10 Holocene settlement of northern Australian coastal plains Figs 9-15. Archaeological sites mentioned in text: 9, Blue Mud Bay. One metre high shell mound dominated by Anadara granosa located at Dilmitjpi on the Dhuruputjpi wetlands (Grindall Bay). Photograph: Pat Faulkner. 10, Hope Inlet (11181) Five metre high Anadara mound, part of large mound complex on laterite ridge overlooking saltflats. Photo Irish Bourke. 11, Blyth River Gulukula shell mound. Photograph: Betty Meehan. 12, Blue Mud Bay. Large Anandara granosa dominated shell mound positioned on the edge of a laterite ridge, Garangarri, Dhuruputjpi wetlands (Grindall Bay). Photograph: Pat Faulkner. 13, Adelaide River. Earth mound MP5. Photograph: Sally Brockwell. 14, Blyth River. Jibena earth mound. Photograph: Betty Meehan. 15, Hope Inlet. Partially buried Anadara granosa midden looking seaward across salt flats. Photograph: Irish Bourke. 11 S. Brockwell et al. over time. Shell fish hooks also appear in the upper level only, about 1000 years ago. Schrire believes that this technology was a result of contact with South-east Asia. However, there is currently no conclusive archaeological evidence for contact between Aboriginal people with South¬ east Asia forthis period. In the Edward Pellew islands, shell middens in rockshelters (Boinmamda, Mushroom Rock, Komandarri-naboya, Turtle Shelter) accumulated steadily indicating more intensive occupation of Vanderlin Island during this period (Sim & Wallis 2008). A large number of dates for shell mounds fall within this period. In the Darwin region, 21 shell mounds, dominated by the open mudflat species Anadara granosa, were located (Hiscock 1997; Bourke 2000; Hiscock & Hughes 2001; Bourke 2004, 2005b; Crassweller 2002, Bourke & Crassweller 2006; Crassweller 2006) (Fig. 10). Excavated shell mounds contain varying frequencies of stone artefacts and other faunal remains in addition to shell, mostly fish and crab with minor quantities of land animals and bird. Turtle is also identified in one mound (MA43) (Crassweller 2006: 20). In western Arnhem Land, two mounds. Field Island 3 and 4, are dominated by mangrove dwellers Terebralia palustris and the sandflat or mudflat species Marcia hiantina (Mowat 1995). One shell mound (V12 Vashon Head) on the Cobourg Peninsula is dominated by Marcia hiantina (Mitchell 1994a: 241). In central Arnhem Land, three shell mounds on the Blyth River, Gulukula (Fig. 11), Muyu- ajirrapa and Ngalijibama, are dominated by the open beach species Dosinia sp. Shell mounds from Milingimbi (Garki, Madanagum, Wallaby Mound (M96), Gadjaw 116) are dominated by Anadara granosa (Roberts 1991). In eastern Arnhem Land, there are seven shell mounds from Blue Mud Bay (BMB 36, 45, 52, 71,92, 101, 116) (Faulkner & Clarke 2004; Faulkner 2006, 2008,2009) (Fig. 12) and one (Barbara Cove Mound) from the Edward Pellew islands (Sim 2005; Sim & Wallis 2008). After 2000 BP, earth mounds proliferated along the margins of newly formed freshwater wetlands (Brockwell 2006a). Further evidence of the trend towards established settlement behaviour on the Reynolds River is demonstrated by the prevalence of earth mounds in the region. Pandyal 2 and Djingurr 2 are today culturally significant to the Werat, especially the Djingurr mounds, which are near the central Dreaming place of the freshwater crocodile (Guse 2005: 114). Other earth mounds dated to this period include HI97 in the Darwin region (Bourke 2000: 225), MP2, MP5 and MP6 on the Adelaide River (Brockwell 2006a) (Fig. 13), Site 40 on the Mary River (Baker 1981: 62-63), and surface mounds Kapalga G and J in western Arnhem Land (WoodrofTe et al. 1988: 97). In central Arnhem Land Ji-bena at the Blyth River contains freshwater turtle in its upper layers, indicating a shift to freshwater conditions of the adjacent swamp Balpilja (Brockwell et al. 2005: 86) (Fig. 14). Numerous dates for shell middens fall within this period, predominantly from 1000 to 500 years BP. Middens in the Darwin region continue to be dominated by Anadara granosa (Bourke 2000) (Fig. 15). Middens in western Arnhem Land include those in the Alligator Rivers region (Kapalga C, L, M) dominated by mangrove species (Woodroffe et al. 1988: 97) and at Black Point, Port Bremer, Vashon Head and Stoneline 3 on the Cobourg Peninsula (Mitchell 1994a: 322). One pre-contact midden (Stoneline 3) contains Lunella cinerea, Nerita chamaeleon (rocky platform) and Gafrarium tumidum (sandy mudflats) (Mitchell 1994a: 321); one post-contact midden (Port Bremer 19) contains Atactodea striata and pottery (Mitchell 1994a: 224-225). In central Arnhem Land, this period is represented by Aningarra, Jilan-gajerra, Larrakun-ajirripa West and Muyu-ajirrapa on the Blyth River (Meehan 1982: 165-168), and Rulku and Balnta83 middens at Milingimbi (Roberts 1994: 178-179). Dates from eastern Arnhem Land, include Gaynada at Cape Arnhem (Bourke 2001), Blue Mud Bay - BMB/3,17,61,67, 84 (Faulkner & Clarke 2004; Faulkner 2006, 2008, 2009), Groote Eylandt - Aburrkbumanja, Dirrangmurumanja and Malmudinga (Clarke 1994); and East Neck Saddle (ENII) on the Edward Pellew islands (Sim 2005; Sim & Wallis 2008). Rock art in this period was prolific and significant (Taqon 1993: 113). Fully developed X-ray art, also called ‘Freshwater’, ‘Later' or ‘Decorative X-ray’, coincides with the arrival of freshwater conditions on the floodplains. It features specific and precise representation of inner organs and is the dominant form in western Arnhem Land and exhibits much more regional variation at this time (Chaloupka 1985; Taqon 1987; Lewis 1988: 55, 99-102; Chaloupka 1993: 114;Taqon 1993) (Fig. 16). Throughout this period, the art becomes increasingly elaborate and decorated and the internal features became more abstract (Chaloupka 1985:277; Taqon 1987; Lewis 1988:99; Taqon 1993). It depicts freshwater fauna and flora, such as magpie geese (Anseranus semipalmata) and lotus lilies ( Nelwnbo nticifera ); and new technology, such as large stone blades ‘leiliras’, ‘goose’ spears and spear throwers (Chaloupka 1985: 277-278; Lewis 1988: 55, 101-102). It should be noted that ‘goose’ spears were not used for spearing geese - but rather were lightweight spears used with a ‘goose’ spearthrower - so-called because the wax hook made it look like a goose head (Lewis 1988: 384; Allen 2011: 78). These spears appear to have been used for dueling. Guse & Woolfe (2006) further demonstrate that the occupation of the coastal plains in western Arnhem Land is reflected by rock art that expanded from gorges and shelters in the escarpment to the outliers with the progression of floodplain evolution. Analysis of the spatial and temporal distribution of these sites indicates a late Holocene reorganisation of land use strategies, social economies and group interactions for the plateau and plains. Guse & Woolfe (2006) conclude that late Holocene changes in the rock art sequence reflect a growing confidence to occupy openly accessible sites in terms of social ecological stability and security, related to ecological abundance and diversity following sea level 12 Holocene settlement of northern Australian coastal plains Fig. 16. Depiction of barramundi (Malarrak, western Arnhem Land). Photograph: Daryl Wesley. stabilisation. The art of this period displays great diversity and regionalisation (Ta^on 1993). Post 500 BP. This period encompasses the later Freshwater Phase and the Contact Period. The Contact Period can be separated roughly into two overlapping phases, the South-east Asian period from at least the 1660s (probably older) through to the early 1900s, and the European contact period beginning in the early 1800s (Macknight 1976; Mitchell 1994a: 56; Clarke 2000; Clarke & Frederick 2008; May et al. 2010; Ta$on et al. 2010; Wesley 2011). The latter part of this period was dominated by environmental changes brought about by European contact, with impacts from feral animal and exotic weed species adversely affecting productivity, especially of the floodplains (Petty et al. 2007; Bradshaw 2008; Walden & Gardener 2008). Dates come from rockshelters, fewer shell mounds and earth mounds, and an increasing number of shell middens. Macassans, Bugis and others from South Sulawesi and nearby islands established a trade in trepang (beche dc mcr), which they obtained and processed on annual visits to northern Australia and sold onto the Chinese who regarded it as a tonic and aphrodisiac. The Southeast Asians supplied the Aborigines with trade goods such as metal, tobacco, beads and cloth in return for access to their lands and other benefits (Macknight 1976; Mitchell 1994: 181). The exact beginnings of this trade are uncertain and debatable (see Discussion below). Two rockshelters from western Arnhem Land, Malakanunja II and Malangangerr, date to this period. Allen & Barton (1989:90-91) interpret this as reoccupation following a long period of abandonment of 2500 years. In eastern Arnhem Land at Borngolo shelter, plant remains in the uppermost levels indicate that the inhabitants continued exploiting the sea shore, monsoon forests, and open woodland, with a range of implements including stone points, scrapers, hammers (Schrire 1972:662-663). Schrire (1972) concludes that the deposit at Borngolo represents pre and post-contact Aboriginal subsistence behaviour. She interprets more intensive occupation in the upper levels as being the desire of the Aborigines to be in close contact with the Macassans for trade benefits. On Groote Eylandt, five rockshelters (Angwurrkbuma, Dadirringka, Lcrrumungumanja, Marngkala Cave, Mungwujirra) belong to this period. There are only four shell mounds recorded. One mound from the lower South Alligator River floodplains ( Kapalga B) includes open beach species (Woodroffe et al. 1988: 97). The others are in central Arnhem Land on the Blyth River (Ngalijibama, Yuluk-adjirrapa), both dominated by open beach species, Mactra sp. and Dosinia sp„ and Blue Mud Bay in eastern Arnhem land (BMB 116) dominated by another open beach species (Anadara antiquata) (Faulkner & Clarke 2004; Faulkner 2006, 2008, 2009). There are dates from seven earth mounds in this period. In the west, Djingurr I is located on the Reynolds River. In the Darwin region on the Adelaide River, a modem date was obtained from NP20, which also contains contact material (Brockwell 2005; 11). On the Maiy River, Site 38 is also dated as modern. In western Arnhem Land, the deposit at Kina adjacent to the freshwater wetlands on the South Alligator River floodplains is dominated by the freshwater mussel Velesunio angasi and polished flakes, similar to those found in the upper levels of Paribari (Meehan et al. 1985: 150). Kina is associated with a large open 13 S. Brockwell et al. Fig. 17. Blyth River. Guna-jenggamidden. Photograph: Betty Meehan. artefact concentration, one of many lining the floodplain margins of the South Alligator River (Meehan et al. 1985; Brockwell 1996). These sites contain hundreds of thousands artefacts, including polished Hakes and trigonal or Teilira’ blades, which are also commonly found in upper levels of rockshelter sites dated to the last 1000 years in outliers of the Arnhem Land escarpment (Jones & Johnson 1985b: 208). Hiscock (1999: 99-100) speculates that there may have been a shift inland from the coast to the freshwater wetlands following the cessation of shell mound building approx. 500 years BR Supporting this theory is intensified occupation at the Adelaide River earth mounds dating to less than 600 years BR (Brockwell 2009). Most of the middens recorded belong to this period. In the Darwin region, two shell concentrations, MA52b and WIN 10, are dominated by mangrove species Telescopium telescopium (Bourkc 2005a) suggesting that the Darwin mudflats were being recolonised by mangroves at this time. In western Arnhem Land, on the floodplains of the lower South Alligator River, seven middens (Kapalga A, B, E, L, M1, M2), contain a mixture of mainly mangrove species (Woodroffe et al. 1988: 97). On Cobourg Peninsula, Site VI at Vashon Head is dominated by Atactodea striata and Marcia hiantina, both sand and mudfiat bivalve species. On the basis of site content, Mitchell defines 17 sites as belonging to pre-Macassan contact (pre 1720 AD) and 12 post-contact midden sites. He concludes that the post-contact midden sites were substantially larger than the pre-contact sites and contain a wider variety of exotic artefacts and remains of large marine fauna (1994a: 377-398). In central Arnhem Land, there are three middens from the Blyth River (Aningarra, Guna-jengga, Jilan-gajerra) (Meehan 1982: 165-168; Brockwell et al. 2009) (Fig. 17). In eastern Arnhem Land, there are four middens at Blue Mud Bay. BMB 16 and 84 are dominated by open beach Fig. 18. Vanderlin Island midden. Photograph: Robin Sim. species, while BMB 15 contains mangrove shellfish and BMB 22 contains rocky coast species (Faulkner & Clarke 2004; Faulkner 2006, 2008). On Groote Eylandt, the 11 middens from this period are dominated by marine shell (Clarke 1994). On the Edward Pellew islands, five surface shell scatters (Investigator Bay, Vanderlin Creek 2, Wabuya Creek, Walala Dunes. Kedgc Point) date from this period and through to the Contact Period (Sim & Wallis 2008) (Fig. 18). Sim & Wallis (2008: 103) report that there is no direct evidence ol Maccassan contact in the Edward Pellew islands archaeology, despite the well-known historical presence of trepanging in the region, as well as the linguistic and rock art record and use of dug out canoes that show contact. Nor is there evidence of intensified occupation of archaeological sites associated with Macassan presence and trade goods, as is argued for the Groote Eylandt, Borngolo Shelter and the Cobourg Peninsula (cfSchrire 1972; Mitchell 1995, 1996; Clarke 2000). However, Sim & Wallis (2008: 103) suggest that the marked decline in the manufacture and use of stone artefacts post 500 years BP may be evidence of replacement with metal tools brought by Macassans. Aboriginal rock art traditions continue through this phase ot occupation. The Contact Period is recorded through images of South-east Asian perahus and items of material culture, and European ships, guns and introduced animals such as horses and buffaloes (Chaloupka 1985:278; Chaloupka 1996; Chippindale & Tatjon 1998; Clarke & Frederick 2006; May et al. 2010; Ta<;on et al. 2010; Clarke 14 Holocene settlement of northern Australian coastal plains 2 Sigma summed probability age ranges by site type Shelter Sites (n = 76) 0 2000 4000 6000 8000 10000 Earth Mounds (n = 31) i ii ■ in □ □ m u 0 2000 4000 6000 8000 10000 Shell Middens (n = 71) ■CEO □ □ □ D 6 T 2000 7 4000 6000 ’ 8000 10000 Shell Mounds (n = 126) HT1W B ~1 □ □□ 6 7 2000 ’ 4000 6000 ’ 8000 10000 Fig. 19. C14 graphs by site type. & Frederick 2011; Wesley in press). This is also the case for the art from Groote Eylandt (Clarke 1994, 2000a, 2000b) and Edward Pellew islands (Sim 2002). In summary, there are four main site types that occur on the coastal plains; rockshelters, shell mounds, earth mounds and shell middens. Rockshelters are the oldest site type and date from the early Holocene period. They arc located on the Reynolds River in the west of the Northern Territory, western Arnhem Land and the Edward Pellew islands in eastern Arnhem Land (Fig. 19). Such sites are located on the edge of the coastal plains and were generally occupied throughout the Holocene. The exceptions arc islands such as Groote Eylandt, where occupation falls largely within the last 500 years, and the Edward Pellew islands, where the evidence indicates an occupational hiatus between about 6700 and 4200 BP, following sea level rise and the cut off of the islands from the mainland. Shell mounds occur right across the Top End. There are a few 4000 4500 year old examples from Reynolds River and western Arnhem Land, but the majority fall between 2500 and 500 BP (Fig. 19). They are usually dominated by shell species from open beach habitats. Earth mounds tend to occur at the junction of a number of resource zones close to the coast, or inland on black soil plains that contain extensive freshwater wetlands systems bordering open woodland. Apart from a few older examples from the Reynolds and Adelaide rivers and Kakadu, the majority of earth mounds are less than 2000 years old. Shell middens occur along the coast and adjacent to palaeochanncls on the floodplains of major rivers across the Top End. They are mostly less than 2000 years old, with the majority being less than 1000 years old (Fig. 19). The older exceptions are in western Arnhem Land, where middens are found buried on the South Alligator River floodplains, and one site in central Arnhem Land, on the Blyth River. DISCUSSION We argue that regional variation in chronology and site distribution is due largely to local topography, and the timing and nature of landscape formation. The evidence for earlier Holocene occupation of these landscapes comes from rockshelters that only occur in some areas where the escarpment is adjacent to the coastal plains. There does appear to be some preservational bias in the record, and it is obvious that it plays a role in site distribution. For example, rockshelters are effective information traps preserving older chronostratigraphic sequences. Early Holocene open sites are rare as unless they are rapidly buried they do not preserve organics. Buried open sites are rarely located and sampled. There is also a geographic bias in our current site sample. Across the broader region, the east coast is the best dated with 134 dates, and the western region, represented by the Reynolds River, the least well-dated with only eight dates. This discrepancy has the potential to skew the data, particularly when combined with a research focus on open sites in a number of regions, making interpretation of settlement patterns through time difficult. That said, however, and given the preliminary nature of the analysis, several interesting patterns begin to emerge. We can see variability in the timing and nature of occupation relative to the geomorphic and climatic parameters outlined. This is particularly evident when considering the differential occurrence of open sites. The pattern of settlement on the coastal plains of the Northern Territory prior to South¬ east Asian contact, more or less tracks the evolution of the landscape and changing opportunities in terms of the creation of new resource zones. With rising seas, evidence of occupation 10,000 years ago was confined mainly to rockshelters on the margins of the coastal plains. Groote Eylandt and the Edward Pellew islands were cut off from the mainland by rising seas approx. 7000 years ago. Shortly after the spread of the mangroves in the Big Swamp Phase, people began foraging along the edges of the vast swamps of the northern floodplains from 7000 BP and the rockshelters adjacent were occupied more intensively. With increasing sedimentation the mangroves declined and after 4000 BP people began to move out onto the floodplains and the rapidly prograding coastline, witnessed by the increasing number of open sites in this period. People also began to 15 S. Brockwell et al. travel offshore, reinhabiting the Edward Pellew islands approx. 4000 years ago. Between 4000 and 2000 BP, there was a diversity of fresh and estuarine environments being exploited on the floodplains, reflected in the faunal assemblages from rockshelters and open sites. Following the Big Swamp Phase, the combination of sea-level recession, continued sedimentation in former shallow embayments built out intertidal mudflats suitable for shellfish biomass. This saw the proliferation of shell mounds in a number of regions across the Territory between approx. 3000 to 500 BP. Figure 2 shows the chronology of shell mounds relative to the environmental data. From 2000 BP, freshwater conditions were established on the sub-coastal plains and earth mounds and artefact concentrations proliferated along the floodplain margins. Between 800 and 500 years BP, environmental change in shoreline characteristics and climatic variability associated with ENSO activity led to a decline of sandy/mudflat shell beds and shell mounding behaviour all but ceased in the Northern Territory (Bourke et al. 2007). Subsequently, mangrove-lined beaches and coastal wetlands were established, economies diversified and middens became more common with molluscs harvested from varied habitats after 500 years BP. There is some suggestion that there may have been some relocation from the coast to the seasonally abundant sub-coastal freshwater wetlands during this period (Hiscock 1999). There are debates about the socio-cultural implications of the archaeological record. Some authors (e.g. Bourke 2005b), quoting the ethnographic literature, have associated mound-building behaviour between approx. 3000 to 500 BP with ceremonial activities and territorial demarcation. Contrary to this, Hiscock and Faulkner argue that there is no historical analogue for shell mounding and the social and economic systems that produced them no longer exist. They hypothesise that the diverse mythology recorded ethnographically that surrounds shell mounds is a recent construction to explain the existence of alien landscape features within modern foraging and social practices (Hiscock & Faulkner 2006: 218). In the sites under discussion, stone unipoints and bipoints appear to be concentrated in a narrow horizon dating from 4000 BP and are rare in the upper levels less than 2000 years old. Stone points are frequently recorded in surface artefact concentrations from the northern coastal plains and although undated may be from this period. The appearance of stone points is part of a widespread technological change across the Top End in the mid Holocene. It is widely agreed to be dated from at least 5000 until 1500 years BP (Allen & Barton 1989: 94-95; Hiscock 1999: 98; Clarkson 2006b: 165-166; Smith & Brockwell 1994: 101). Hiscock (1999: 98) argues that points appear as early as 7000 BP coinciding with the Big Swamp Phase and that mid Holocene changes in stone technology were driven by changes in resource availability created by the evolution of the coastal plains. He goes on to say that unipoints and bipoints are part of a continuum of manufacture, and represent a risk strategy in times of uncertainty (Hiscock 1994a. 1994b, 1999: 98). Clarkson (2006a: 105, 2006b: 165-166) agrees with this analysis and argues that regional technological change was a response to environmental variability created by the onset of ENSO (El Nino Southern Oscillation) from 5000-2000 years ago. Smith & Brockwell (1994: 102) argue that concentrated manufacture of stone unipoints and bipoints at Scotch Creek between 3800 and 2100 cal. BP coincides with the Transition Phase, a period of environmental stress when increasing aridity and the changing nature of the floodplains made resources unpredictable. Smith (1995) associates stone points with spear tips and suggests that tensions over access to resources and inter-group conflict led either directly to warfare or threatening displays at ceremonies. Certainly depictions of inter-group fighting in western Arnhem Land rock art became more common in this period (Ta?on & Chippindalc 1994). However, a phytolith study conducted to investigate environmental change was unable to firmly establish the connection between changes in climate and changes in archaeology (Clarkson & Wallis 2003). It might be noted that there are no ethnographic examples of spears ha fled with small unifacial or bifacial points known from the northern coastal areas, though they do occur in the Kimberley region. It is suggested that South-east Asian contact may have occurred from perhaps as long ago as 800 years BP (Clarke 2000b: 325-328; Clarke & Frederick 2011:151). Macknight (1976, 1986) and Mitchell (1994: 56) argue that it more likely dates from the 1720s when there is documentary evidence for the Macassan trade in trepang with the Chinese. However, recent dating has shown that the image of a Macassan perahu on the walls of a rockshelter in western Arnhem Land has a minimum age of 1664 and may be as old as 1517 AD (Ta?on et al. 2010). Whatever the timing, these visits had a profound impact on Aboriginal social and economic organisation on the northern coastal plains (Clarke 2000b). In eastern Arnhem Land, Schrire (1972) posits that the people of Port Bradshaw replaced traditional stone and bone tools with metal artefacts and became increasingly sedentary as a result of contact with Macassans. Mitchell (1994a, 1994b, 1996) argues persuasively that the economy and settlement patterns of Cobourg Peninsula in western Arnhem Land were completely reorganised and local populations were able to rely much more on exploitation of large marine animals as a result of the adoption of the dugout canoe and access to metal to manufacture harpoons. Metal axes also allowed Aborigines to exploit the hard dupes of Pandanus, which contain an energy-rich kernel (Meehan et al. 1979). These new technologies allowed groups to extend their hunting range and access previously under-utilised resources (Faulkner 2006: 6). It is argued that contact with Macassans stimulated ceremonial exchange cycles and trade networks, involving a variety of objects including stone, shell, metal, spears, cloth, baskets, nets etc, that extended from eastern Arnhem Land into western and southern Arnhem Land and beyond (Allen 1996a, 1997; 16 Holocene settlement of northern Australian coastal plains Bemdt 1951; Davidson 1935; Evans & Jones 1997; Jones & White 1988; Mitchell 1994b, 1996; Paton 1994; Thomson 1949). This network included trade in large stone blades manufactured at Ngilipitji quarry in north-eastern Arnhem Land (Jones & White 1988). Production of similar artefacts, manufactured in the quarries of western Arnhem Land (Jones & Johnson 1985a: 188) and found in the rockshelters and open sites restricted to the last 1000 years, may also have been stimulated by this Arnhem Land wide phenomenon. From 180 years ago, Europeans and later Chinese began penetrating the coastal plains of the Northern Territory. The earliest incursions were British settlements on Bathurst and Melville Islands and on Cobourg Peninsula (Allen 1980; Keen 1980; Mitchell 2005). Apart from the devastating impact of introduced diseases which greatly reduced the Aboriginal population, one effect of these early settlements was the spread of feral buffaloes throughout the coastal plains, especially between Darwin and western Arnhem Land, where they wreaked havoc on the freshwater wetlands. Apart from the damage to wetlands resources, Meehan (1991: 203-205) points out that as Aboriginal people feared buffaloes, this also had the effect of altering their hunting and foraging patterns on the coastal plains in general. In 1858, South Australia annexed the Northern Territory and there was a brief settlement at Cape Hotham near the mouth of the Adelaide River, which subsequently moved to Port Darwin. These initial settlements were followed by pastoralists, buffalo shooters, miners, timber getters, pearlers, fishermen and missionaries. European and Chinese presence had profound effects on Aboriginal social and economic systems and settlement patterns, some of which are reflected in the archaeological record. For example, on Groote Eylandt, as people became more settled on the missions they became more reliant on European food supplies. When they did forage, they targeted easily accessible and favourite bush foods (Clarke 2011). Consequently, midden sites examined on Groote Eylandt relating to this period consist of a limited number of taxa, all of which were available in the immediate environment. As a result of this process, these sites are seen to conform to the criteria outlined by Meehan (1988) for ‘dinner-time’ or temporary camps (Clarke 1994:458). People are presumed to have operated a dual subsistence system - one that had a commodity-based component involving the consumption of European resources, and a hunter-gatherer lifestyle that integrated traditional practice with elements of the new (Clarke 1994: 462). As indicated previously, however, many of these traditional practices arose during the period of contact with Macassans (Faulkner 2006: 8). A similar situation existed on the Adelaide River, where earth mounds from the Contact Period were dominated by estuarine shellfish, as the resources of the freshwater wetlands were impacted negatively by the incursions of feral animals and exotic weeds (Brockwell 2005: 15-16). CONCLUSION The above synthesis is a preliminary attempt to integrate the archaeological evidence within an environmental framework. With only a few exceptions, environmental and archaeological research on the coastal plains of the Northern Territory was not undertaken in tandem. This means that research questions often had different foci and the data sets do not necessarily intersect. We hope that future research is more inter-disciplinary in nature. In this way multiple lines of evidence can be used to reconstruct both the natural and cultural landscape and aid in interpretation of the interactions between the two. Although extensive geomorphological work has been done, there has been little palaeo-climatic research in this region of northern Australia. We envisage that future research programs would explicitly address questions of climatic variability and its impact on human settlement and mobility. Such research would be dedicated to investigating regional and local patterns through isotope analysis and coral proxies. This research could also be integrated with palaeoecological investigations through analysis of pollen, diatoms and phytoliths from both existing archaeological soil samples and coring programs into the many suitable swamps of the Northern Territory. Due to the comparative paucity of work to date in the western region of the Northern Territory, we see this area as a priority area for future research. The archaeological record demonstrates that the last 10,000 years have witnessed recurrent economic and social reorganisation for the occupants of the coastal plains of the Northern Territory. We argue that the evidence clearly suggests this is a result of variable climatic and environmental conditions and landscape evolution combined with contact with peoples from South-east Asia [and later Europeans] within at least the last 400 years. This may seem to be an overly deterministic point of view, but is not unlike the conclusions from other archaeological studies in the coastal areas of northern Australia. They have implied that major environmental changes, rather than strictly social changes, preceded growth in human populations, and that the growing productivity of newly created landscapes combined with higher population levels, were causal factors in cultural change, variations in mobility, and increasing land and sea use during the Holocene (cf. Beaton 1985; Jones 1985b: 291-293; Meehan et al. 1985:153; Sullivan 1996: 7). In explaining the variability of coastal occupation in the north over the I loloccnc, a number of researchers have focused on environmental explanations such as changes in local ecological habitats (e.g. Hiscock 1999; Hiscock & Mowat 1993; Mowat 1995; O’Connor 1999), as well as pointing to the links between the appearance of mounds and evidence for increasing aridity and the northward movement of the northern monsoon on the coast (O’Connor & Sullivan 1994; O’Connor 1999). This particular type of ecological perspective does not view human culture as 17 S. Brockwell et al. being determined by the environment, nor does it assume perfect adaptation of humans to their environment. Rather, it suggests that the archaeological manifestation of a defined set of human behaviours, such as the structure of economic activity, may best be viewed in terms of the formation of a particular environment (Allen 1996b; Cribb 1996:150; Faulkner 2006: 18). ACKNOWLEDGEMENTS The authors would like to thank the Aboriginal communities with which they worked while collecting the data that form the basis of this paper: Werat Traditional Owners (Reynolds River); Larrakia Association (Darwin and Hope Inlet); Wairuk Association (Adelaide River); Gagadju Association and Lamilami family (western Arnhem Land); An-barra community (Blyth River); Yilpara, Rurrangala, Gan-Gan, Djarrakpi and Yirrkala communities (Blue Mud Bay); Anindilyaka Land Council, Angurugu Town Council, Umbakumba Town Council and Milyakburra community (Groote Eylandt and Bickerton Island); and Yanyuwa community (Sir Edward Pellew Group). We would also like to thank Paul Ta^on and Harry Allen, whose constructive comments have greatly strengthened this paper. REFERENCES Akerman. K. 1986. The Western Arnhem Land use of polished flakes: possible functions. Unpublished report. Allan, R., Lindesay, J. & Parker, D. 1996. El Nino, Southern Oscillation and climatic variability. 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It is described from sterile specimens based on distinctive leaf characters as no flowering or fruiting material was available. A key to species of Embelia from the Northern Territory and Timor is provided for comparison. Keywords: Myrsinaceae, Primulaceae, Embelia, new species, Embelia tiwiensis, Tiwi Islands. INTRODUCTION Embelia Burm. f. is a genus of about 130 species (Stahl & Anderberg 2004) occurring in the Old World tropics. It is one of six genera in the family Myrsinaceae occurring in Australia. Recently the Angiosperm Phytogeny Group included the Myrsinaceae in the family Primulaceae where it is treated as a subfamily (Stevens 2011). However, in this paper the Myrsinaceae is still regarded as a separate family. The genus Embelia can be distinguished by habit, being vines or scramblers, and by the flowers, which have five free petals. Vegetativcly Embelia is distinguished from some species of Maesa (now in the family Maesaceae), which also have a scrambling habit, by the secretory glands in the leaves. In Embelia the glands are chiefly globular or shortly lineate, but in Maesa they form long thin cells. Reynolds (1991) recognised four species of Embelia in Australia, of which one, E. curvinervia S.T. Reynolds, occurs in both Queensland and the Northern Territory. Ten species are known from New Guinea and the Solomon Islands (Slcumer 1987) and there are at least three species on the Lesser Sunda Islands. Here I describe a new species from the Tiwi Islands, Embelia tiwiensis Jackcs. This species is known from four collections, one from Melville Island and three from Bathurst Island. Although the material is sterile it can be readily distinguished from other species of Embelia based on leaf features, in particular possession of globular pellucid glands in the leaves. These differences are outlined in the key to species occurring in the Northern Territory and nearby areas. TAXONOMY Embelia tiwiensis sp. nov. (Figs 1,2D,E) Embelia sp. Tiwi Islands (J. Russell-Smith 8240) N.T. Herbarium, in Short, P.S et al. (May 2011), Checklist Vase. PI. Northern Territory, http://www.nl.gov.au/nrcta/wildlife/ plants_herbarium/pdf/200701 nt_checklist.pdf Embelia tiwiensis ab aliis speciebus Embeliae quibus in Australia, Nova Guinea and Insulis Minoribus Sundae habitant, laminis chartaceis cum glandibus numerosis globularibus secrctoriis pellucidis in luce transmissa et pilis ferrugineis in pagina infera differt. Material examined. Type - Australia, Northern Territory, Bathurst Island, Ngaru Swamp, 11 0 43’19”S, 130°20’57”E, 13 June 2011, J.A. Risler 645 & D.J. Dixon (HOLOTYPE: DNAD146206,2 sheets; ISOTYPES: BRI, CNS). Additional specimens examined. NORTHERN TERRITORY: Bathurst Island, Ngaru Jungle, Plot 2839, 28 February 2001, J.A. Risler 583 & R.A. Kerrigan (DNA); Bathurst Island, Ilinga Jungle, 17 June 2001, J.A. Risler 805 & R.A. Kerrigan (DNA); Melville Island, 25 May 1990, J. Russell-Smith 8240 & D. Lucas (DNA, JCT). Description. Vine, semi-prostrate, rarely exceeding 60 cm tall, adventitious roots common, lenticels inconspicuous. Young twigs covered with reddish-brown glandular papillae and/or short hairs. Leaves alternate; petioles 7-21 mm long, papillate; lamina elliptical to broadly elliptical to obovate, (3.5-) 6.5-17.5 cm long, (1.6-) 3.5-7.8 cm wide, chartaceous, discolorous; base 23 B. R. Jackes Fig. 1. Holotype specimen of Embelia tiwiensis sp. nov. (J.A. Risler 645 & D.J. Dixon. DNA D146206 - 2 sheets. For a note on label discrepancies, see text. cuneate, symmetrical to somewhat asymmetrical; apex acuminate; margin entire or coarsely and irregularly toothed in upper half of lamina; glands globular in transmitted light, pellucid, appearing dark in surface view; hairs spherical, red-brown and scale-like, present on both surfaces, more common on lower surface; midrib slightly depressed on upper surface, prominent below; lateral nerves 8-12 on either side of midrib, looping near the margin. Flowers and fruits not seen. Distribution and habitat. Embelia tiwiensis is only known from Bathurst and Melville Islands in the Tiwi Island group. It has been found in seasonal spring-fed swamps and riparian jungles. Here the common dominants are Calophyllum soulattri, Hanguana malayana, Hydriastele wendlandiana . Syzygium angophoroides and S. nervosum. Etymology. The specific epithet reflects the occurrence of the species on the Tiwi group of islands including Bathurst and Melville islands. It is adjectival. Note. Contrary to the labels accompanying the sheets comprising the holotype specimen, an isotype stated to be in JCT is housed in CNS. The specimen label attached to sheet 2 of the holotype specimen was generated in 2011 from the latest Northern Territory Herbarium electronic database. This has resulted in some discrepancies between that label and the one accompanying sheet 1. Importantly, there is a correction in citation of the collection number, from Risler & Dixon 645 to Risler 645 & Dixon. There is also a minor difference, due to computer programming, of the cited latitude of the collection; both sheets are part of the one gathering. Herbarium abbreviations follow Thiers (2011). Key to species of Embelia from the Northern Territory and Timor 1 Glands in lamina pellucid to pale orange when viewed in transmitted light.2 - Glands in lamina chiefly dark red to black when viewed in transmitted light.3 2 Glands pale orange, both globular and lineate, lamina glabrous. E. viridiflora (Timor) - Glands pellucid, globular only, lamina with rusty papillae/hairs on lower surface particularly on the veins. E. tiwiensis (NT) 3 Inflorescence paniculate, much-branched, longer than leaves, 5-merous. E. aff .javanica (Timor) - Inflorescence a cluster of racemes, shorter than leaves, 4-merous.4 24 Embelia tiwiensis (Angiospermae: Myrsinaceae) sp. nov. from the Northern Territory Fig. 2. Embelia curvinervia and E. tiwiensis, from Jackes et al. (in press) Myrsinaceae, Flora of the Darwin Region Vol. 1. A-C, Embelia curvinervia S.T. Reynolds: A, leaf (lower surface) and inflorescence; B, male flower from above; C. fruit (A,B, C.R. Dunlop 5025; C, D.C. Franklin D154622); D, Embelia tiwiensis sp. nov., leaf shapes, both showing lower surface (J.A. Risler 645 & D.J. Dixon, holotype). All illustrations were based on herbarium specimens. 4 Leaves chartaceous, subtending bracts about 1 mm long. E.frangulifolia (Wetar Island) - Leaves firm, subtending bracts 2-3 mm long. . E. curvinervia (NT and QLD) N.B. Embelia frangulifolia Mez has not been recorded from Timor but it does occur on nearby Wetar Island. It may be distinguished from E. curvinervia by the chartaceous versus firm leaves and by subtending bracts about 1 mm long versus bracts 2-3 mm long and the latter is generally a much more robust vine. ACKNOWLEDGEMENTS My sincere thanks to all who have assisted, in particular Emma Short for the Latin diagnosis, Philip Short for general advice and J-F. Vcldkamp from Leiden who answered queries and provided photographs and scans of Embelia specimens from neighbouring areas. Darren Crayn is thanked for his constructive comments. Monika Osterkamp provided the illustration based on the type specimens. REFERENCES Jackes, B.R., Short, PS, Wightman, G.M. & Mangion, C.R 2011. Myrsinaceae. In: Short, RS. (ed.). Flora of the Darwin region, volume 1. In press. Reynolds, S.T. 1991. The genus Embelia N. Burman (Myrsinaceae) in Australia. Austrobaileya 3(3): 361-367. Sleumer, H. 1987. The genera Embelia Bunn. f. and Grenacheria Mez (Myrsinaceae) in New Guinea. Blumea 32: 385-396. Stahl, B. & Anderberg, A.A. 2004. Myrsinaceae. Pp. 266-281. In: Kubitski, K. (ed.). The families and genera of vascular plants , vol. 6. Springer-Verlag: Berlin. Stevens, P.F (2011 onwards). Angiosperm Phylogeny Website, http:// www.mobot.org/MOBOT/research/APweb/ Last accessed July 2011. Thiers, B. (2011, onwards). Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Gardens Virtual Herbarium, http://sweetgum.nybg.org/ih/ Last accessed August 2011. Accepted 16 August 2011 25 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 27: 27-54 New taxa and notes on Helicteres L. (Malvaceae: Helicteroideae) from the Northern Territory, Australia IAN D. COWIE Northern Territory Herbarium, Department of Natural Resources, Environment, The Arts and Sport PO Box 496, Palmerston NT0831, AUSTRALIA ian.cowie@nt.gov.au ABSTRACT Six new species and a new subspecies of Helicteres from the wet - dry tropics of the Northern Territory, Australia are described and illustrated (H. darwinensis, H. kombolgiana, H. macrothrix, H. serpens, H. sphaeivtheca, H. tenuipila and H. cana subsp. latifolia). Methorium integrifolium is reinstated as a species and a new combination is made in Helicteres (H. integrifolia) to accommodate it, while Helicteres dentate is reduced to subspecific level under H. integrifolia. Two varieties of H. dentata are raised to specific level (//. procumbens and H.flagellaris). A key to the 13 species known from the Northern Territory is provided and variation in some widespread taxa is discussed. Keywords: Malvaceae, Sterculiaceae, Helicteroideae, Helicteres, new species, H. cana subsp. latifolia, H. darwinensis, H. fiagellaris, H. integrifolia, H. integrifolia subsp. dentata, H. kombolgiana, H. macrothrix, H. procumbens, H. serpens, H. sphaerotheca, H. tenuipila. INTRODUCTION The family Malvaceae.v. lat. is cosmopolitan, especially tropical, in distribution with nine subfamilies, 113 genera and c. 5000 species (Mabberley 2008). It is widespread in tropical and temperate Australia. While the genus Helicteres L. has traditionally been included in the family Sterculiaceae (e.g. Cronquist 1981), more recent phylogenetic studies support its inclusion in a broadened Malvaceae (APG 1998; APG II 2003; Mabberley 2008). Subfamily Helicteroideae is one of the smallest subfamilies in Australia and is represented only by Helicteres which contains perhaps 60 species. Helicteres has a major centre of diversity in central and South America where some 38 species arc recorded with an apparently lesser one in the Asian-Australian tropics (Christobal 2001). The only Australian treatment for the genus is that of Bcntham (1863) who recognised five taxa, all of which occur in the Northern Territory (N.T.). Additional Australian species have been described since that time by Fitzgerald (1918) and Mueller (1865). Wheeler (1992) provided the only modern Australian flora treatment, for the Kimberley Region of W.A. Recent flora treatments are also available for Thailand with seven species (Phengklai 2001) and for China with ten species (Tang et al. 2007). Four species were recognised for Java by Backer & Van den Brink (1963) and seven for British India by Masters (1874). Many of the east Asian and Malesian taxa appear to be relatively widespread with a few extending to Australia (Phengklai 2001; Tang et al. 2007). In Australia, the incompleteness of available treatments and variability of leaf and indumentum characters especially has resulted in the misapplication of names. It has become evident over the past 25 years that a number of undescribed taxa occur in the northern N.T., Qld and W.A. A number of new taxa are described here in particular for inclusion in the Flora of the Darwin Region. As the type specimens of at least four taxa treated by Bcntham (1863) are from the N.T., the clarification of the taxonomy of the N.T. species is of some value in progressing knowledge of the genus in Australia as a whole. Christobal (2001) recognised seven sections within the genus, with only two of these (section Helicteres and section Orthocarpaea), occurring in Asia and Australia, the others being restricted to South and central America. As circumscribed by her, section Helicteres is monospecific including only H. isora L. while all other Asian-Australian species belonged to section Orthocarpaea. Section Helicteres was distinguished by the geniculate, pseudozygomorphic flowers with the upper petals distinct from the lower; the curved androgynophore which significantly exceeds the calyx; the petals with scattered minute glandular hairs or glabrous, red or orange or white, rarely white-green; stamens 10, usually connate at the base; one of the lower petals longer than the calyx; 2 (-5) llowered axillary cymes with extrafloral nectaries present; the androecium and gynoecium set at an angle on the androgynophore; the capsule spiral and the seeds without fins (Christobal loc. id.). She distinguished section Orthocarpaea as having 10 or fewer stamens; the androgynophore shorter than the calyx or just longer and always straight; the androecium 27 I. D. Cowie and gynoecium set at an angle on the androgynophore; the petals violet, the lower ones clearly differentiated from the upper and with a tuft of long glandular hairs; the capsule not spiral; the seeds without ribs; the axes of inflorescence with nectaries; and the flowers in simple cymes. Herbarium abbreviations follow Thiers (2010). Only duplicates examined in the course of preparation of this treatment have been cited, except for unseen type specimens indicated by the abbreviation n.v. (i.e. non visits -not seen). MORPHOLOGY OF SPECIES IN THE NORTHERN TERRITORY Habit. All species are shrubs or subshrubs with either erect or trailing stems. Two primary resprouting forms are evident. Firstly, there are those with annual above-ground parts which resprout from a perennial rootstock either at or near to ground level. This group contains usually multi-stemmed perennial subshrubs either with erect or prostrate stems and includes H. danvinensis, FI. integrifolia, H. macrothrix and H. sphaerotheca. In at least some members of the group which have been grown in cultivation ( H. integrifolia subsp. dentata and H. integrifolia subsp. integrifolia), the annual dying-back of above-ground parts and the low multi-stemmed, resprouting habit occur even in the absence of fire. The second life form comprises taller, more erect, open, usually few-stemmed shrubs with lower stem regeneration as in for example H. angustifolia L., H. isora and H. kombolgiana. The open, much branched, few-stemmed habit of H. kombolgiana is maintained in cultivation. In some cases H. isora may be multi-stemmed after resprouting from fire but is typically taller and develops an open shrubby habit when fire protected. Helicteres cana and H. hirsuta Lour, are somewhat intermediate being few-stemmed but lower in stature, the former at least resprouting at ground level after annual fire. Indumentum. Most vegetative parts and the calyx have sessile or stipitate stellate hairs with short or long multiangulate aims. The size of these hairs is often diagnostic of a species. The new growth often also has scattered, red, minute, digitiform, glandular hairs. A few species have long simple hairs on the upper surface of the leaf ( H. angustifolia, H. procumbens and sometimes H. sphaerotheca), while most other species have only stellate hairs. Long or short, soft bristles (these bearing stipitate or sessile stellate hairs) are sometimes present on the calyx ( H. kombolgiana) and are often present on the fruit (e.g. H. hirsuta, H. integrifolia, H. macrothrix, H. tenuipila). These bristles appear broadly comparable with those of Triumfetta fruit (Halford 1997). Density of stellate hairs was classified into five classes: very dense, with the underlying surface obscured or nearly so; dense, with the branches touching to overlapping; moderately dense, with the hairs touching to 1 diameter apart; sparse, with the hairs 1-5 diameters apart; scattered, with the hairs more than 5 diameters apart. Leaves. Leaves are simple, almost sessile to pctiolate, the lamina usually has serrate margins or is sometimes entire, and in the case of H. isora (Fig. 1) also with short acute lobes lateral to the apex. Most commonly leaves are lanceolate to broadly ovate, less often linear (H. cana subsp. cana, H. sphaerotheca) or suborbicular (//. cana subsp. latifolia). In //. danvinensis the leaves appear at least partly accrescent, continuing to expand through the wet season until growth is apparently halted by the onset of dry conditions. Consequently leaves of specimens collected late in the wet season may be up to 180 mm long and thus many times larger than those present at flowering. The lower leaf surface in all species has a sparse to dense indumentum of stellate hairs, while the upper surface may be glabrous, or have stellate hairs, simple hairs or a mixture of hair types. Stipules are usually subulate with an indumentum of stellate hairs. Inflorescence. Cristobal (2001) recognised three distinct inflorescence variants, concluding that the partial inflorescence in Helicteres had undergone a progressive reduction from a symmetric or asymmetric double cincinnus to a simple multi-flowered cincinnus (e.g. H. hirsuta) and lastly to a 2-flowered or 1-flowered cincinnus. In turn these types of partial inflorescences may be arranged in a diversity of synflorescences. In the monospecific section Helicteres (//. isora) the cincinni are 2-flowered, rarely 3-flowered, while in section Orthocarpaea cincinni are basically 2-flowered or multi-flowered as in II. hirsuta and the south east Asian H. viscida. The basic inflorescence unit of a I-, 2- or 3-flowered cincinnus is most common and in most taxa these are arranged in cymose inflorescences which may be contracted (e.g. H. macrothrix) to lax (H. serpens, H.flagellaris). In II. hirsuta the inflorescence is indeterminate, appears racemose, and can have up to 19 flowers with one flower, bract and extrafloral nectary per node. Cristobal (2001) interpreted this as a multiflowered cincinnus formed by reduction from a symmetric double cincinnus. There are no species with a symmetric or asymmetric double cincinnus. Extra-floral nectaries. Sessile cxtrafloral nectaries are widespread in the genus and are present at the base of the pedicels on the internal face of the axis of the inflorescence in most species, including those considered here (Cristobal loc. id.). However, in H. hirsuta the gland is lateral to the base of the pedicel and on alternate sides of the rachis on successive flower nodes. These extra-floral nectaries arc present in most taxa, with the notable exception of H. procumbens, and may be green, yellow-green or yellow when fresh but often drying dark to dark red. In all species, glands are sessile to slightly raised, orbicular to oblong or irregular depending on placement, smooth, glossy, glabrous and often depressed and darker in the centre. They appear to be frequented by a number of species of ants, including Green Tree Ants (Oecophylla smaragdina) and meat ants (Iridomyrmex). 28 Helicteres (Malvaceae) from the Northern Territory Flowers. All species have flowers of the ‘geniculate’ type, in this case with the androcciuin and gynoecium set obliquely on the androgynophore rather than with the androgynophore or calyx set at an angle to the pedicel (Cristobal 2001). In these species the calyx is bilabiate and with the flower appearing somewhat zygomorphic (pscudozygomorphic of Cristobal 2001). Local species vary considerably from strongly bilabiate in H. isora and less so in H. cana and H. hirsuta , or weakly bilabiate, to virtually actinomorphic in species such as //. danvinensis and H. procumbens (Figs 2, 6, 9-11, 13, 15, 18, 20). In H. isora , flowers last two days and are functionally male on the first day of opening and functionally female on the second (Winkler 1906, cited in Cristobal 2001). My observations confirm that there is a change in petal colour from pale blue to orange over the two days, and the limb of the petals is strongly reflexed and held against the calyx on the first day becoming perpendicular to it on the second (Fig. 1A.). The anthers are reported to be ripe on the first day and the stigma receptive on the second while nectar is produced by the calyx on the first and by the stigma on the second day. In other N.T. species, no change of flower colour is evident and longevity has not been investigated. Calyx. Helicteres has a 5-lobed, weakly or strongly 2-lipped calyx, with the tube longer than the lobes (Fig. 7L). The calyx is green, subcylindrical, narrowly obconical, ellipsoidal or ovoid to campanulate. The lower three calyx teeth are usually distinct from the upper pair although otherwise similar in appearance. All species have a basal nectary gland but in most species except H. isora this incompletely covers the base of the calyx tube. In these species the gland is usually on the ‘upper’ side adjacent to the adaxial petals or sometimes lateral to the adaxial and abaxial petals and extends from 0.3 to 0.5 mm from the base, or in H. hirsuta 1 mm from the base. In H. isora the nectary extends upwards to c. 3 mm from the base and completely encircles the androecium and gynoecium. The margin of the gland may be irregular or entire. The calyx usually abscises from its base and persists loosely on the androgynophore through into the fruiting stage. Corolla. The corolla of Helicteres is composed of five petals. Characters found to be useful to distinguish taxa included the degree of differentiation between the upper and lower petals, especially in the morphology of the claw and the relative length of the calyx and corolla (Christobal loc. id.). The upper (adaxial) pair and lower (abaxial) three petals may be weakly to strongly dimoiphic but are always clawed with a cuncate limb and one or two appendages at the base of the limb (rarely absent). These structures have been variously referred to as auricles, auriculate appendages, (lateral) teeth, or erase crests (Bentham 1863; Masters 1874; Fitzgerald 1918; Wheeler 1992: Cristobal 2001; Phengklai 2001). The appendages on the lower claws are less robust and less well developed than those on the upper petals. The two upper petals tend to be imperfect mirror images of Fig. 1 . Helicteres isora: A, flowers; B, fruit; C, leaf. (/. D. Cowie 3272). each other, as is also the case for the two outer, lower petals (Figs 3D, E, J, K, 7C,E). The upper petals especially differ slightly in the placement of appendages. When dissected out and laid fiat, the upper pair often curve towards each other, while the outer lower petals curve away from each other (Fig. 7C, E). The central lower petal is usually close to straight and symmetric. In H. isora the limb is strongly geniculate or reflexed, while in H. cana and H. hirsuta the petals are bent at 90 degrees. In other taxa the petals are arcuate so that the limb curves out to about 90 degrees with the claw. The limb of petals is usually mauve to mauve-pink or pink, rarely maroon (H. hirsuta). In H. isora the petals are very pale blue on the first day changing to pale 29 I. D. Cowie orange in the evening and becoming darker orange on the second day. The upper petals in a number of species (H. can a, H. darwinensis, H. integrifolia, H. kombolgiana, H. macrothrix, H. procumbens, H. tenuipild) are white at the base of the limb, corresponding to the glabrous area discussed below. The lower petals are usually uniformly coloured to the base of the limb, although in H. darwinensis frequently longitudinally striate or striped at the base of the limb and in some others darker. Iielicteres hirsuta has a paler area at the base of the limbs of the lower petals. Most species have sparse, patent to appressed, septate, apparently glandular hairs c. 0.1 mm long over the limb and similar but slightly longer hairs on parts of the claw. In three species (//. cana, H. hirsuta and H. isora ), the base of the limb in all petals is smooth or with scattered minute, erect, glandular hairs and lacks an area of matted hairs on the inner surface. However, in other species the lower three (abaxial) petals have a distinct area of matted, simple, septate hairs to 1 mm long at the base of the inner surface of the limb while the upper (adaxial) petals have a smooth, glabrous, thickened and raised area at the base of the limb (Fig. 7). Helicteres isora has dark coloured, capitate, sessile glands of varying size scattered over the claw and base of limb of the upper petals and at the base of the limb on lower petals. In the lower (abaxial) three petals the appendages are ‘lateral’ (on the margin and in a plane with the surface of the petal) whereas in the upper petals one appendage is ‘sublateraT (at the margin but near to perpendicular to the surface of the petal) and the other ‘medial’ (located towards the centre between the margins and near to perpendicular to the surface of the petal). These lateral appendages help to hold the claws in a pseudo-floral tube while the sublateral and medial appendages serve to position the androgynophore and hold it away from the adaxial petals, leaving a clear path over the obliquely set stamens and stigma to the base of the calyx and corolla. This open tube on the upper side of the androgynophore presumably facilitates the access of insect or bird nectivores to the base of the flower and serves to enhance pollen transfer. While flowers, or at least the inflorescences, of most species are frequented by ants it is unclear if these play a role in pollination or harvest nectar from the flowers. Androgynophore. In N.T. species of section Orthocarpaea, the apex of the androgynophore is just exserted from the calyx and floral ‘tube’ although in H. cana it is more strongly exserted. In H. isora (section Helicteres) the androgynophore is curved and greatly exceeds the reflexed corolla lobes. The androecium and gynoecium are mounted obliquely on the androgynophore and orientated adaxially in all species (Fig. 1A). In most species of section Orthocarpaea , the androecium and gynoecium rest on or just past the area of matted hairs on the lower petals. In H. darwinensis and II. macrothrix they may be virtually obscured by these hairs. Androecium. All species have an inner whorl of 5 minute staminodes and an outer whorl of 10 equal stamens, free or fused at the base. The staminodes are shorter than the stamenal filaments and narrowly oblanccolate to spathulate, but acute. Stamens are shorter on the adaxial side of the flower. In most species, anthers are dithecal and transverse but usually inclined slightly inwards at anthesis. In H. hirsuta the anthers are introrsc at anthesis. Nine pollen types have been recognised for the genus (Pire & Cristobal 2001). Gynoecium. The ovary is hairy and the styles are always connate, straight and may be twisted or not, even in species with non-twisted carpels. In H. isora the style and stigma are bent at 90 degrees to the axis of the stamens. In other species the style is parallel or slightly oblique to the stamens. Within the Thai species of section Orthocarpaea , Phengklai (2001) recognised three taxa as having flattened slightly 5-lobed stigmas (H. angustifolia, H. elongata and H. hirsuta), the others having a simple stigma either narrowed to a point or with 5 pin-like teeth. These stigma types can also be recognised in N.T. taxa; they are distinctly flattened in //. hirsuta and straight with 5 cohering pin¬ like lobes in H. cana, H. darwinensis, H. integrifolia and H. isora. In II. cana, H. integrifolia subsp. dentata and II. procumbens at least, the stigmas may diverge slightly at the apex. However, in contrast to Phengklai’s findings, Australian and Malesian material of H. angustifolia sens, hit. that I have seen appears to have a style consisting of 5 pin-like lobes, slightly expanded towards the apex. Capsule. The fruit is a capsule and in Helicteres is composed of five unilocular carpels and may be ovoid to ellipsoid or subcylindrical. With the exception of H. isora (Fig. 1B), all species have straight valves and even in this species valves, although normally twisted spirally, may sometimes be partly straight, incompletely twisted or even with the twist reversing (e.g. Booth 23; Cowie 12406; Scarlett 16). The carpels dehisce along the inner suture, becoming partly separated with age. The indumentum varies considerably in density and hair size and the presence or absence of soft bristles. In descriptions, capsule dimensions include the layer of bristles or other indumentum. In some taxa (e.g. H. angustifolia, //. cana) the styles may at times be accrescent and the apex of the beak becomes apiculate as it develops Seeds. The seeds are arranged in a single row in each carpel, together forming an elongated mass, flattened or angled laterally along the inner angle of the carpel and cut obliquely to transversely between the seeds. Thus the basal and terminal seeds are conoidal or sub-conical and the central ones sub-rhomboidal to sub-cylindrical with the adjoining ends obliquely truncate, or if the capsule has one seed it may be ellipsoidal and flattened. The size and number of seeds varies from numerous and about 2 mm long in //. isora, to few and more than 3.5 mm long in a number of species. 30 Helicteres (Malvaceae) from the Northern Territory RELATIONSHIPS OF NORTHERN TERRITORY SPECIES While H. isora is clearly distinct at the sectional level, within species of Helicteres section Orthocarpaea four loosely-defined groups can be recognised. Helicteres cana group (H. cana ): Plants erect, the indumentum is fine, leaves are virtually entire, with both surfaces densely hairy, the corolla is distinctly 2-lipped, the lower (abaxial) petals lack an area of matted hairs, petals are more strongly bent, relatively long, and the androgynophore is clearly exserted. Helicteres hirsuta group (H. hirsuta ): Similar to H. cana except the indumentum is coarser, leaves are toothed, discolorous, the inflorescence is racemose and indeterminate, the androgynophore is scarcely exserted and the anthers introrse at dehiscence. Helicteres danvinensis group [H. darwinensis, H. macrothrix, Hserpens): Plants are prostrate or erect, the indumentum coarse, leaves serrate, relatively large, weakly discolorous, corolla not or scarcely 2-lipped, relatively large, abaxial petals with matted hairs, androgynophore as long as calyx. Helicteres integrifolia group (//. angustifolia, H. integrifolia, H. kombolgiana, H. procumbens, H. sphaerotheca, H. tenuipila, with H. ftagellaris perhaps transitional to the previous group): Plants arc erect or prostrate, the indumentum often fine to coarse, leaves usually serrate, usually discolorous, corolla weakly 2-lipped, abaxial petals with matted hairs, relatively small. CONSERVATION STATUS OF NORTHERN TERRITORY SPECIES The conservation status of all species has recently been reassessed against IUCN criteria following IUCN guidelines as part of a five-yearly review of the conservation status of the N.T. flora (IUCN 2001, 2003; IUCN Standards and Petitions Subcommittee 2010). Most species have been assessed as IUCN Least Concern. H. tenuipila Fig. 2. Helicteres cana subsp. cana. Flowers and leaves. (/. D. Cowie 11800 ). is regarded as Near Threatened partly because of its restricted distribution. Among the factors contributing to the Endangered status of Helicteres macrothrix are past and projected loss of populations due to land clearing for horticulture, subdivision, railway or road maintenance and quarrying. Both species are threatened to varying degrees by invasion of habitat by the perennial weed Gamba Grass {Andropogon gayanus). The resulting changes in fire regimes, soil hydrology, nitrogen availability and light are serious developing threats (Cowie, Kerrigan & Stuckey in press; J. Westaway & I. Cowie unpubl.). TAXONOMY Helicteres cana (Schott & Endl.) Benth., 1863 (Figs 2, 3A,B, 4) Methorium canum Schott & Endl., Meltemata Botanica 29, t. 5 (1832). Type Citation: “ Habitat in Novae Hollandiae tropicae septentrionalis sinu Carpentaria ” [Australia, N.T., 1803], F. Bauers.n. (Holotype: ?BP, n.v.). Description. Open shrub to 2 m tall, stems 1-several, sparingly branched, often grey or grey-green. Stems and both leaf surfaces hoary, densely and shortly stellate- puberulous with hairs 0.1-0.5(0.7) mm diam.; calyx and capsule densely and shortly stellate-pubescent with stellate hairs and straight or flexuose bristles to 3 mm long. Stipules caducous, linear, 2.2-12 mm long. Petiole 3-13 mm long. Leaf lamina lanceolate to ovate, less often linear, narrowly oblong, suborbicular or very broadly obovate, 27-150 mm long, 7-88 mm wide, 1-11 times longer than wide, concolorous to weakly discolorous, often folded longitudinally, base broadly attenuate or rounded to cordate, 3-5-nerved, closely reticulate and raised on under surface, margins entire, rarely toothed in upper half, apex acute to obtuse or rounded, sometimes apiculate or shortly acuminate. Inflorescence axillary, cymose, contracted, 2- many-flowered, dichasia 2-flowered; extra-floral nectaries present; bracts linear, 1-5 mm long; pedicels 1-3 mm long. Calyx tubular to ellipsoidal, 6-11 mm long, basal nectary incomplete, margin minutely irregular; lobes triangular, 1-3 mm long. Corolla 2-lipped. Petals mauve, slightly dimorphic, 10-20 mm long, 3.5-5 mm wide, long-clawed, cuneate-spathulate, with scattered short simple spreading to patent hairs, apex truncate to obcordate; lower 3 petals narrower than upper pair, on the inner surface with (0) 1 or 2 lateral auricles at base of limb, pad of woolly matted hairs absent; upper pair petals on the inner (adaxial) surface with one lateral and one medial or sublateral auricle at base of limb, with a glabrous glossy, slightly callused area just above auricles. Androgynophore 7.5-14 mm long, straight, medially with scattered, short simple hairs, longer than calyx; stamens 10, scarcely connate at base, 1-1.5 mm long; anthers transverse; staminodes 5, lanceolate to narrowly oblanceolate, 0.5-0.7 mm long. Style 1-1.2 mm long, straight; stigma of 5 pin-like lobes, terete, diverging 31 I. D. Cowie slightly at apex. Capsule ellipsoid to oblong, 7-32 mm long, 7-11 mm diam., apex shortly beaked or not; carpels straight, 2-7-seeded. Seeds variable in shape, sub-rhomboidal to conoidal, laterally compressed, 2-3 mm long, 1.5-2 mm wide, finely rugose, dark brown. Flowers and fruits: most months. Distribution and ecology. The species is endemic to Australia (N.T., Qld). It occurs in the northern N.T. and on nearby islands of the Gulf of Carpentaria, Qld, and grows in open Eucalyptus- dominated vegetation, usually on well drained sandy or stony soils. Notes. The species is characterised by the hoary, concolorous or weakly discolorous leaves, fine indumentum, the lack of a woolly or smooth, glossy callus on the lower (abaxial) petals and the androgynophore which is distinctly longer than the calyx. It differs from II. angustifolia especially in that leaf laminae are concolorous or weakly discolorous and the upper surface of the lamina is densely and shortly stellate-tomentose and hoary; the petals lack a woolly or smooth, shiny callus; the calyx, corolla and androgynophore are all longer than for H. angustifolia. Schott & Endlicher (1832) provide a detailed illustration of H. cana, leaving no doubt as to the application of the name and cited a Bauer specimen from the northern Gulf of Carpentaria as the type specimen. However, it is not immediately clear if this specimen is part of the same gathering as a Brown collection at BM labelled in Brown’s hand “ Orthocarpus incana. North Coast” (duplicate MEL222172) and apparently collected from Sweers Island (island a). Gulf of Carpentaria on 17 Nov 1802 (Steam 1960). The specimen attributed to Bauer could represent either a separate gathering or part of the Brown collection retained by Bauer for illustrative purposes. The taxon is common in eastern Arnhem Land and Bauer would probably have had opportunity to gather material from a number of places from Groote Eylandt north to Arnhem Bay. This area fits better with the ‘northern Gulf of Carpentaria’ locality given by Schott & Endlicher (1832) than does Sweers Island which is in the southern Gulf of Carpentaria. For these reasons I treat the Brown and Bauer collections as distinct gatherings. An additional sheet, MEL (222171), labelled “Helicteres afif. H. cana, North Coast” and collected by Brown is in fact H. integrifolia subsp. dentata. As recognised here, H. cana is variable in the size, shape and degree of folding of leaves, the degree of serration of the margin, the distance the main lateral veins extend up the leaf lamina, and in the length of the fruit and indumentum. Using these characters two geographically distinct entities can usually be distinguished. However, specimens displaying intermediate attributes do exist in the Pine Creek to Nitmiluk N.P. area and this fact, coupled with an absence of critical features on regenerating plants (e.g. Cowie 6068), has led me to recognise them as subspecies, subsp. cana and subsp. latifolia. 1 Leaves mostly strongly channelled (or folded longitudinally when dry), lamina linear to ovate or elliptic, 2-11 times longer than wide, major lateral veins mostly extending less than 50% of lamina length, margin usually entire in upper half; capsule 7-20 mm long.....subsp. cana 1: Leaves not strongly channelled or folded longitudinally, lamina ovate or elliptic to suborbicular or very broadly obovate, 1-3.1 times longer than wide, major lateral veins mostly extending more than 50% of lamina length from base, margin usually at least partly serrate in upper half; capsule 14-32 mm long . .subsp. latifolia Helicteres cana (Schott & Endl.) Benth. subsp. cana Other specimens examined. NORTHERN TERRITORY: McArthur River Station, CSIRO paddocks, 14 June 1985, M. Andrew 1009 (DNA); Cape Arnhem, 7 March 1995, M. Barritt 1741 (DNA, MEL); Nhulunbuy, Whitewood Rd, 3 February 1997, R. Booth 1399 (DNA); Nhulunbuy, Gaynagaru Walk, 5 February 1997, R. Booth 2137 (DNA); Kakadu N.P.. near Coronation Hill gate, Gimbat, 6 March 1991, K. Brennan 1146 (DNA); Kakadu N.P., Gimbat, 6 April 1990, K. Brennan 218 (DNA); 15 km WSW of Cape Arnhem, site B9, 14 February 1994, K. Brennan 2439 (DNA); Kakadu Stage 3 Fauna Survey, site 14, 19 April 1990, K. Brennan 3 & T. On (DNA); English Company Islands, Inglis Is., 7 September 1996, K. Brennan 3321 (DNA); Ml5. MacArthur River mine lease, 7 April 2003, K. Brennan 5889 & K. Metcalfe (DNA); Kakadu N. P., lookout hill near Coronation Hill, 8 August 1990, K. Brennan 694 (DNA); Gove, Latram River. 20 May 2008, K. Brennan 7688 (DNA); Carpentaria, 1802, R. Brown s.n. (MEL, NSW); 3 miles [4.8 km] S of Larrimah, 5 February 1969, AS. Byrnes /354(DNA,NT); Borroloola, McArthur River road, 7 April 1970, N.B. Byrnes 1854 (DNA, NT); 16 km SW of Goyder River crossing, 16 June 1972, N.B. Byrnes 2638 (CANB, DNA, K, L, NSW, NT); Arnhem Land, Maningrida-Oenpelli road, 19 July 1987, M.J. Clark 1171 (DNA); Arnhem Land, 3 km NE of Bulman, 16 September 1987, M.J. Clark 1191 (DNA); NE Arnhem Land, 29 September 1987, M.J. Clark 1521 (DNA, MEL); Nhulunbuy, 200 m S of ovals, 1 March 1985, P. Cleminson 18 (DNA); Gove Peninsula, August 1974, J.F. Cooper s.n. (NSW); Arafura Swamp, old homestead, 19 May 1990, I.D. Cowie 1233 (DNA); west of Old Arafura, 22 May 1990, I.D. Cowie 1311 (DNA, MEL); S of Ramingining, 22 May 1990, I.D. Cowie 1312 (BRI, DNA, MEL, PERTF1); McMinns Bluff, near Pine Creek, 25 January 1991, I.D. Cowie 1472 & C.R. Dunlop (BRI, CANB, DNA, MEL); Groote Eylandt, 5 km N ofAngurugu. 10 September 1991, I.D. Cowie 1997 & PS. Brocklehurst (BRI, DNA); Groote Eylandt, 1.3 km NNE of Angurugu, 11 April 1992, I.D. Cowie 2567 (BRI, DNA, MEL); Arnhem Bay, Raymangirr, 22 May 1992, I.D. Cowie 2788 (DNA); Nth 32 Helicteres (Malvaceae) from the Northern Territory ^ Y 1 V/ 5 mm SNa: 'Yy > /cm V ml if/4 Fig. 3. A, B. Helicteres cana subsp. catui. A, leaf shape variation (Clark 1521, Blackshall 68); B, flowering branchlct (Clark 1521). C. Helicteres cana subsp. latifolici: fruiting branchlct (Cowie 976 & Estbergs). D-G. Helicteres kombolgiana, D, central lower petal; E, upper petals (D-E, Cowic 12530); F, flowering branchlct; G. fruit. (F-G, Egan 4528). 1I-K. Helicteres tenuipila. H, (lowering branchlct (Cowie 3269); I, fruit (Cowie 5347 & Taylor); J, upper petals; K, central lower petal (J-K, Cowic 10361 & Egan, type). Scale bars: A = 3 cm, B = 2 cm, C, F, G, H, I = 1 cm, D, E, J, K = 5 mm. 33 I. D. Covvie H. cana subsp. cana H. cana subsp. Iatifolia H. darwinensis H.flagellaris H. integrifolia subsp. dentata H. integrifolia subsp. integrifolia H. kombolgiana H. macrothrix H. procumbens H. serpens H. sphaerotheca Fig. 4. Distribution of Helicleres species in the N.T. and adjacent parts of W.A. and Qld. H. tenuipila of Oenpelli, Mt Borradaile road, 23 June 1992, I.D. Cowie 3022 <£ R. Booth (BRI, DNA, MEL); Bickerton Is., near airport, 29 April 1993, 1.D. Cowie 3903 & C.R. Dunlop (BRI, CANB, DNA, MEL); Blue Mud Bay, Woodah Is., I May 1993, 1.D. Cowie 4011 & C.R. Dunlop (DNA); Rorruwuy Outstation, Arnhem Bay, 31 August 1995, I.D. Cowie 6068 (DNA); Flinders Peninsula, E side opposite S end of Probable Is., 28 April 1996, l.D. Cowie 6695 & P. Bokarra (BRI, DNA); Inglis Is., north side, 30 April 1996,/.D. Cowie 6828 (DNA); eastern Arnhem Land, near Mirmatja, 16 June 1999, I.D. Cowie 8348 (DNA); central Arnhem Land, near Emu Springs, 18 September 1999, I.D. Cowie 8405 (DNA, MEL); Groote Eylandt, c. 18 km NE of Angurugu, 7 March 2005, I.D. Cowie 10530 & K. Brennan (BRI, DNA, L, MEL); e. 14 km SE of Nhulunbuy, on road to airport, 25 November 2007, I.D. Cowie 11800 (DNA); cultivated at Lucy Ct. Driver (provenance Nhulunbuy area), 7 June 2010, I.D. Cowie 12542 (DNA, SING); McArthur 34 Helicteres (Malvaceae) from the Northern Territory River area, 12 February 1976, L. Craven 3861 (CANB); McArthur River area, 2 June 1976, L. Craven 4042 (CANB, DNA); 10 miles [16 km] SSW of Borroloola, 5 June 1971, C.R. Dunlop 2218 (DNA); Rose River, 3 km W of Policeman Crossing, 1 December 1987, C.R. Dunlop 7405 (DNA); Donydji, Arnhem Land, 20 June 1989, C.R. Dunlop 8495 & N.G. White (BRI, CANB, DNA, MEL, MO, NSW); Kakadu N.P., Coronation Hill, 18 June 1990, C.R. Dunlop 8638 & B. Wilson (DNA); Donydji; Arnhem Land, 24 June 1990, C.R. Dunlop 8667 & N.G. White (DNA, MEL); 55 km from Cape Crawford, 17 April 1993, J. Egan 2083 (DNA, NSW); Roper Hwy, 7 September 1993, J. Egan 2617 & G.J. Leach (DNA); road to Nhulunbuy, 9 September 1993, J. Egan 2648 & G.J. Leach (AD, DNA); Tawallah Ranges, Limmen, 24 May 1996, J. Egan 5363 & C.R. Michell (DNA); 4 km W of Nhulunbuy, 18 November 1989, P. Forster 5958 (BRI, DNA); 67 km NE of Cape Crawford, on road to Borroloola, 23 June 1991, D. Halford569 (BRI, DNA); Batten River crossing, Borroloola to Bing Bong road, 21 May 1984, D. Halford 845110 (DNA); 7 km S of Gove-Ramingining road, Arnhem Land, 20 October 1981, T. Henshall 3881 (DNA); 10 miles NNE McArthur River Station, 6 September 1911, G. Hill562 (MEL); Nhulunbuy, 28 April 1982, R.C. Hinz 106 (DNA, NT); 40 km SSE Nathan River homestead, 18 January 1989, P. Latz 11168 (MEL, PERTH, NT); c. 15 miles SW of El Sharana Mine, 26 February 1973, M. Lazarides 7875 (CANB, DNA, PERTH); track to Koolpin Gorge, 1 May 1994, G.J. Leach 4251 (BRI, DNA); Arafura Swamp area, S of Dog Mountain, 15 September 1998, C.P Martgion 639 <£ l.D. Cowie (DNA); eastern Arnhem Land, Gulf of Carpentaria, outstation, 14 Febaiary 1998, C.P. Mangion 889 (DNA, LD); Cananbirini Park, 26 February 1998, C.R. Michell 1374 & R. Carrow (DNA); Nitmiluk N.P., 4 April 2001, C.R. Michell 3298 & S. Boyce (DNA, PERTH); Nitmiluk N.P., 12 March 2002, C.R. Michell 3617 (DNA, MO, SING); 40m SW of Goydcr River crossing, 16 June 1972, J. Must 1010 (CANB, DNA, NT); 8 km ENE of Urapunga Station homestead, 10 April 1989, T. Orr 268 (DNA); c. 36 km from Borroloola, road to Daly Waters, 18 May 1974, R. Pullen 9304 (CANB, DNA); Djapididjapin, c. 12 km from Nangalala, 15 January 1973,77. Reeves 588 (CANB); SW of Boroloola jump-up, 18 February 1976, B. Rice 2240 (CANB); Nitmiluk N.P., site 451, 5 April 2001, J./L Risler 1300 & M. Waetke (DNA); Nitmiluk N.P., site 453, 5 April 2001, J.A. Risler 1301 tfe M. Waetke (DNA); Mt McMinn Station, Roper Hwy, S of Roper Hwy near Big River Station turn-off, 26 April 2002, J.A. Risler 1850 A A. Fisher (B, BRI, DNA, NT, SING); Big River Station, 18 March 2003, J.A. Risler 2279 (DNA); Groote Eylandt, approx. 24 km NE of Angurugu, 7 March 2005, J.A. Risler 2840 (DNA); Groote Eylandt, 8 km SW of Umbakumba, 15 July 1987, J. Russell-Sntith 2778 & D. Lucas (DNA); 3 km NW of Numbulwar, 1 September 1987, J. Russell-Smith 3089 & D. Lucas (DNA); 5 km W of Policeman Crossing, Rose River, 1 December 1987, J. Russell-Smith 4309 A D. Lucas (BRI, DNA); Doyndji, E Central Arnhem Land, 15 February 1974, N.H. Scarlett 34 (DNA); beside Annie Ck, 12 July 1975, N.H. Scarlett 51 (DNA); North Australia, R.D. Schomburgk s.n. (AD); near Roper River, 9 September 1867, F. Schultz (MEL); Limmen N.P., 50 km SE of Ranger Station along road to Lorella Springs, 19 April 2008, PS. Short 5587 & E.M. Johnston (DNA); Nhulunbuy, 22 October 1986, N.M. Smith 204 (DNA); Groote Eylandt, Hempel Bay, 30 April 1948, R.L. Specht 329 (AD); Gove, 22 August 1948, R.L. Specht 947 (AD); Canopy Rock, Mtn Valley station, 25 February 1963, R.F. Swinbourne 688 (DNA); Groote Eylandt, Eminbinungwa, 17 March 1979, J. Waddy 755 (CANB); Limmen N.P., upper Mountain Ck area, c. 5.8 km SSE of Gibb Bluff, 21 April 2009, J. Westaway 2927 (DNA, LD); Nathan River Station, 5 June 1986, G.M. Wightman 2959 (DNA); 3 km SE of Ramingining, 19June 1987, G.M. Wightman 3871 & N. Smith (DNA); 30 km N ofNgukurr, Arnhem Land, 25 November 1987, G.M. Wightman 4057 (DNA); Nhulunbuy, 21 February 1988, G.M. Wightman 4254 (BRI, CANB, DNA); Nhulunbuy, 22 February 1988, G.M. Wightman 4282 (DNA); Barlalumba Bay road, Groote Eylandt, 12 July 1988, G.M. Wightman 4501 (DNA); Emerald River, Groote Eylandt, 6 October 1988, G.M. Wightman 4618 (DNA); Limmen N.P., St Vidgeon’s block, 20 April 2009, B. Wirf414 (DNA). QUEENSLAND. Mornington Is., 6 November 1979, J. Clarkson 2691 (BRI, DNA, K, MO); Bentinck Is., 14 May 1997, D.J. Liddles.it. (BRI, DNA); Sweers Is., E end of air strip, 16 November 2002, L. Pedley SW/74 (BRI, DNA); SE comer Bentinck Is., 29 May 1963, N.B. Tindale s.n. & P. Aitken (AD - 3 sheets, BRI); Mornington Is., 31 May 1963 ,N.B. Tindale s.n. (AD). Description. Calyx and capsule densely and shortly stellate-puberulous or densely pilose with stellate hairs and straight or flexuose bristles. Stipules caducous, linear, 2.5- 12 mm long. Petiole 3-13 mm long. Leaf lamina lanceolate to ovate, less often linear, narrowly oblong, 37-150 mm long, 7-40 mm wide, 2-11 times longer than wide, more or less discolorous, main lateral veins at base extending 32-50% of leaf length, margin usually entire, rarely toothed in the upper half, apex acute to obtuse. Calyx 6-10 mm long. Petals 10-14 mm long. Androgynophore 7.5- 12 mm long. Capsule 7-20 mm long. Flowers and fruits: most months. Distribution and ecology. Widespread in the eastern part of the Top End, extending from Kakadu N.P. and Gove Peninsula around the Gulf hinterland into the Qld Gulf country. Helicteres cana subsp. latifolia Cowie, subsp. nov. (Figs 3C, 4 ,5) A Helicteres cana Schott & Endl. subsp. cana lamina folii non valde caniculata vel longistrorsum plicata, latiore et ovata vel elliptiea ad suborbiculare vel latissime ovata, 1-1.3-plo longiore quam latiore, venis magnis basis pro parte maxima ultra 50% lamina extensis, margine plus 35 I. D. Cowie minusve serrato dimidio supero ad capsula longioribus (14-32 mm longa) differens. Type: Australia, N.T., Gregory N.P.; near entrance, off Victoria Hwy, 10 February 1992, I.D. Cowie 2209 & PS. Brocklehurst (Holotype: DNA; Isotypes: AD, BRI, CANB, K, MEL, NSW, PERTH). Other specimens examined. NORTHERN TERRITORY. Jarrome Yards, 10 May 1994, M. Barritt 1208 (DNA); Kapalga, SW of mining hut, 21 December 1991, K.G. Brennan 1718 (DNA); Kakadu Hwy, 0.3 km N of turnoff to Goodparla, 22 December 1991, K.G. Brennan 1720{ DNA); Edith Falls, 13 June 1985, J. Brock 87 (DNA); 18 miles [29 km] E Pine Creek, 28 January 1969, N.B. Byrnes 1345 (DNA); Kapalga, 13 December 1988, I.D. Cowie 718 (DNA); near Mining Hut, Kapalga, 15 February 1990, I.D. Cowie 976 & J.A. Estbergs (DNA); Big Nellie Ck, 7 December 1990, I.D. Cowie 1449 & C.R. Dunlop (DNA); near Raft Point, c. 22 km from Mandorah turn¬ off, 1 February 1991, I.D. Cowie 1490 & C.R. Dunlop (DNA); Nitmiluk N.P., near Edith Falls, 23 November 2001, I.D. Cowie 9511 & G.M. Holland (DNA); Kapalga, road to mining hut, 16 August 1991, J. Cusack s.n. (DNA); Edith Falls, 13 April 1995, J. Egan 4724 (DNA); 18 miles [29 km] NE of Pine Creek township, 11 March 1965, M. Lazarides 161 (DNA); Edith River area, 4 February 1999, C.R. Michell 2370 & J.A. Risler (DNA); Nitmiluk N.P., 1 March 2001, C.R. Michell 3297 & S. Boyce. (DNA); Edith Falls area, 7 May 2002, C.R. Michell 3615 (DNA); Frances Ck mine, 23 May 2000, D. Napier 57 (DNA); Nitmiluk N.P., site 472, 6 April 2001, J.A. Risler 1302 & M. Waelke (DNA); 5 km EofWinwuyurrCk crossing, 6 February 1984, J. Russell-Smith 1106 (DNA); Victoria River, Gregory N.P., 25 February 1986, G.M. Wightman 2753 (DNA). Description. Calyx and capsule hoary, densely and shortly stellatc-puberulous with hairs 0.1-0.4 mm diam. Stipules caducous, linear, 2.2-5 mm long. Petiole 3-10 mm long. Leaf lamina ovate to suborbicular, very broadly ovate or very broadly obovate, rarely lanceolate, 42-105 mm long, 17-88 mm wide, 1-3.1 times longer than wide, discolorous, main lateral veins at base extending 46-87% of the length of the lamina, the lamina often irregularly dentate, serrate or sinuate in the upper half, apex acute or acuminate to rounded. Calyx 7-11 mm long; lobes 1-3 mm long. Petals 11-20 mm long. Androgynophore 12-14 mm long. Capsule 14-32 mm long. Flowers: (Aug.) Nov-Feb Fruits: Feb.-June. Distribution and ecology. This subspecies is endemic to the western Top End of the N.T., from the lowlands of Kakadu N.P. and the western margins of Nitmiluk N.P. to Bynoe Harbour and south to Gregory N.P. It grows in woodland dominated by species such as Eucalyptus tectifica, E. miniala , E. patellaris and E. phoenicea, often with perennial grasses on coarse sandy to clay loam soils on granite, sandstone, basalt and Iaterite and in various topographic situations. ql (jMt Bu jj, NSV mTt , MEL , URW Fig. 5. Holotype of Helicteres cana subsp. latifolia. Notes. Collections from near Pine Creek and the western side of Nitmiluk N.P. have narrower leaves and are closer to subsp. cana than those from elsewhere. The isotype specimens were unfortunately dispersed before examination for preparation ot this description and without determinavit slips indicating their type status. Etymology. From the Latin latus, broad and -folius, leaved, referring to the broad leaves of this subspecies. Helicteres danvinensis Cowie, sp. nov. (Figs 4, 6, 7F-I, 8) Ab Helicteres integrifolia habitu prostrato, indumenta grosso, pilis stipitatis stellatis, stipulis persistentibus, longioribus, 7-20 mm longis, inflorescentiis 30-50 mm longis, bracteis, calyce, petal is et androgynophoro omnibus longioribus discrepans. Type: Australia, N.T., Channel Island Rd, SW of Palmerston, 20 November 2008, I.D. Cowie 12221 (Holotype: DNA; Isotypes: AD, B, BRI, CANB, K, L, MEL, MO, NSW, NY, PERTH). Illustrations. Brock, Top End Native Plants 217(1988 and subsequent reprints, inch as Native Plants of Northern Australia ), as Helicteres sp. (lower illustration). Other specimens examined. NORTHERN TERRITORY. E of Fogg Bay, 13 August 1946, S. Blake 16793 (DNA); Mandorah, track off main Darwin road, 21 December 36 Helicteres (Malvaceae) from the Northern Territory 1994, R. Booth 234 & P.F. Mums (DNA); Edith Falls, 10 November 1986, D. Bowman 299 (DNA); Munmalary, 3 May 1986, D. Bowman 367 (DNA); Kakadu N.P., below Mt Brockman massif, 1 April 1990, K.G. Brennan 217 (DNA); Kakadu N.P., Ranger Lease, 10 December 1990, K.G. Brennan 796 (DNA); Ranger Lease, 5 December 1991, K. G. Brennan 1645 (DNA); Murgenella, 3 September 1986, J. Brock 136 (DNA); Darwin River region, Labersheda, 1 October 1983,./. Brock 149 (DNA); Darwin River Dam area, Labersheda, 28 September 1986, J. Brock 154, 155 (DNA); Bcrrimah, Quarantine road, 10 March 1969, N.B. Byrnes 1402 (DNA.); East Ann Rd, near causeway, 16 October 1986, MJ. Clark 705 (DNA); near Marlow Lagoon, Palmerston, 14 March 1990, l.D. Cowie 975 (DNA); near Marlow Lagoon, Palmerston, 12 June 1990, l.D. Cowie 1315 (DNA); E of Pioneer Ck, Mandorah Road, 9 June 1990, l.D. Cowie 1316 (DNA); Old Jim Jim road, 16.4 km from Arnhem Hwy, 8 February 1991, l.D. Cowie 1370 & R. Booth (DNA); Litchfield N.P., 23 November 1990, l.D. Cowie 1425 & C.R. Dunlop (DNA); Jabiru, below lake wall, 30 December 1991, l.D. Cowie 2142 (DNA); c. 2 km N Koongarra Saddle, beside road, I January 1992, l.D. Cowie 2143 (BRI, DNA); N end Koongarra Airstrip, 1 January 1992, l.D. Cowie 2145 (DNA); N of Brogdcn Pt, 10 October 1992, l.D. Cowie 3185 (DNA); Arnhem Hwy, Ocnpclli turnoff, 15 October 1992, l.D. Cowie 3249 (DNA); nearNumbuwah Rock. 13 May 1997, l.D. Cowie 7554 (DNA); Stuart Hwy, c. 35 km N of Katherine, 23 November 2001, l.D. Cowie 9518 & G.M. Holland (BRI, DNA); Daly River road, just S of Survey Ck, 3 December 2002, l.D. Cowie 9658 & C.R Mangion (DNA); 10 km S. of junction Arnhem Hwy-Pine Creek road. 18 May 1980, L. Craven 5576 (CANB, DNA); 35 km SSW Cooinda, Pine Creek road, 19 May 1980, L. Craven 5598 (CANB, DNA); Margin of Jabiru airstrip, 22 March 1984, L. Craven 8238 & G.M. IVightman (DNA); 23 km S of Darwin along Stuart Hwy, 24 May 1987, L. Craven 8589 & J.P. Grace (CANB, DNA); 1 km W of Mt Bundey Ck, Arnhem Hwy, 14 December 1990, C.R. Dunlop 8774 & l.D. Cowie (DNA); Litchfield N.P., along main rd, 24 April 1993, J. Egan 2178 (DNA); Litchfield N.P., 17 December 1993,./. Egan 2911 (DNA); Mount Todd mine site, 17 March 1995, J. Egan 4441 (DNA); road to Umbrawarra Gorge, 5 April 1995, J. Egan 4645 (DNA); Kangaroo Flats-Wangi Falls road, 23 December 2004, J. Egan 5459 (DNA); Edith Falls road, 10.5 km from Stuart Hwy, 3 December 1990, M. Evans 3474 (DNA); Melville Is., 13 April 1986, R. Fensham 86 (DNA); Melville Is., 3 September 1986, R. Fensham 271 (DNA); Mandorah Road, 62 km past Berry Springs, 21 November 1989, P. Forster 6053 (DNA); Takamprimili Ck, Pickertaramoor, Melville Is., 24 November 1989, P. Forster 6085 (DNA); Mt Brockman, 25 October 1974, B. Fox 701 (DNA); c. 12 km N of Daly River Police Stn, 22 May 1983, P. Fryxell 4247 (CANB, DNA); Port Darwin, 26 July 1913, G.F. Hill 97 (MEL); Port Darwin, 1883, M. Iloltze s.n. (MEL); Port Darwin, 1883, M. Holtze 251 (MEL); Port Darwin, 1885, M. Holtze 545 (MEL); Port Darwin, 1888, M. Holtze s.n. (MEL); Port Darwin, s.d., M. Holtze s.n. (NSW); Port Darwin, 5 March 1905, M. Holtze 1134 (MEL); Port Darwin, 5 March 1905, M. Holtze 1235 (MEL); c. 5 km from Dundee Beach turnoff, N along Cox Peninsula Road, 12 October 2001, A.M. Hope 21 & D. Dixon (DNA, SING); Palmerston, near junction of Lambrick Avenue/Stuart Hwy, 23 November 2001, A.M. Hope 32 (DNA); Munmalary Stn, 15 April 1973, P. Latz 3902 (DNA); Melville Is., on Maclcar Ck road, 20 January 1992, G.J. Leach 2942 & l.D. Cowie (DNA); Finniss River road to Litchfield N.P., 21 September 1992, G.J. Leach 3259 (DNA); Emerald Springs, 22 September 1992, G.J. Leach 3264 (DNA); Jindare Station, Stray Ck catchment, Daly Basin, 13 November 2003, D.L. Lewis 595 (DNA); Cox Peninsula, 13 April 1988, K.M. Manning 368 (DNA); Koongarra area,„v.t/., O.H. Marshal s.n. (CANB); 5 miles [8 km] NW of Humpty Doo, 8 November 1971, J.L. McKean 25 (DNA); Darwin Harbour, Wickham Point, 14 January 1997, K. Metcalfe s.n. (DNA); Edith River Area, 12 April 1999, C.R. Michell2340& J.A. /?/.v/cr(DNA); Nitmiluk N.P, 28 March 2002, C.R. Michell3616 (DNA); 12 miles [ 19 km] W.S.W. of Mudginberri H/st, 20 November 1970, J. Must 582 (DNA); Acacia Gap turn-off, 39.6 miles [63.4 km] S of Darwin, 15 June 1964, D.J. Nelson 1195 (DNA); Batchelor area, 17 December 1974, M.O. Parker 614 (DNA); Horns Ck, 8 November 1978, M. Rankin 1554 (DNA); Berrimah Riding School, 9 December 1982, M. Rankin 2680 (DNA); 11 km from Stuart Hwy along Umbrawarra Gorge road, 12 January 1999, J.A. Risler 77 & R.A. Kerrigan (DNA); Charles Darwin N.P., 5 March 1998, PS. Short 4666 & C.R. Dunlop (DNA); Elizabeth River crossing, Channel Is. road, 20 March 1989, A.M Smith 1424 (DNA); near Robin Falls, Mt Bundy station, 3 November 1985, B.W Strong 785 (DNA); Holmes Jungle, 12 km NE Darwin, 18 April 1980, 1.R.H. Telford 7432 & J. Wrigley (CANB); 4 km ESE Jabiru, 18 April 1980, I.R.H. Telford 7547 & J. Wrigley (CANB); Fannie Bay, 1886, J.E. Tenison-Woods s.n. & M. Holtze (MEL); about halfway between Stuart Hwy & Berry Springs, 25 September 1980, J. T. Waterhouse 9834 (NSW); Fig. 6. Helicteres darwinensis. Flowers. (I. D. Cowie 12221). 37 I. D. Cowie Fig. 7. A-E. Helicteres macrothrix. A, (lowering branchlet; B. bristle from fruit (A-B, Dunlop 9805); C. upper petals; D, androgynophore; E, lower petals (C-E, Cowie 9656 & Kerrigan). F I. Helicteres darwinensis. F. flowering branchlet and mature leaf (Dunlop 8774 & Cowie, Rankin 2680); G, fruit (Cowie 1315); H, upper petals; 1, central lower petal (H-I, Cowie 12221, type). J-N. Helicteres serpens. J, flowering branchlet (Evans 3483); K, fruit; (Leach 4379) L, calyx; M, central lower petal; N, upper petals (L-N, Cowie 2139). O-S. Helicteres sphaerotheca. O, flowering branchlet: P. leaf; Q, upper petals; R, central lower petal (O -R, Cowie 2146, type); S, fruit (Dunlop 6120). Scale bars: A, J = 2 cm, F, G, H, I, K, L, O, P, S = 1 cm, C, D, E, M, N, Q, R = 5 mm, B = 2 mm 38 Helicteres (Malvaceae) from the Northern Territory Fig. 8. Holotype of Helicteres darwinensis. 0.5 km W Geringbark scarp, SE comer of Jabiluka outlier, 5 December 1980, J.T. Waterhouse 10938 (DNA, NSW); Mary River Camp, s.ti, M. White 30 (CANB); Berrimah, 7 October 1983, G.M. Wightman 688 (DNA); Lowther Rd, 21 November 1984, G.M. Wightman 1768 (DNA). Description. Multi-stemmed perennial subshrub with prostrate annual above-ground stems to 0.5 m long and a woody perennial root stock. Most vegetative parts and calyx densely stellate-hirsute with sessile and stipitate (0.3 mm long) hairs. 1 lairs sparse to dense on upper surface of leaf lamina, the new growth with scattered digitiform glandular hairs. Stipules persistent, filiform, 7-20 mm long. Petiole 5-10 mm long. Leaf lamina usually ground- hugging, accrescent, lanceolate to ovate or orbicular, 12-180 mm long, 7-110 mm wide, usually discolorous, hairs of the upper surface 1-2.2 mm diam., on the lower surface 1.3-3.5 mm diam., base cordate, asymmetric, 5-nerved, margins irregularly serrate, apex acute to rounded. Inflorescence axillary, congested, a dichasium or panicle of dichasia, 30-50 mm long, 4-22-flowered, extra-floral nectaries present; dichasia 3-flowcred; bracts 7-10 mm long, filiform; pedicels 0-5 mm long. Calyx narrowly obconical, 6-9 mm long, basal nectary incomplete, margin oblique; lobes triangular, 1-4 mm long. Corolla not or scarcely 2-lipped. Petals mauve-pink, weakly dimorphic, 12-16 mm long, claw auriculate, limb cuneate, callused near base, claw and limb with scattered to sparse, simple, septate hairs c. 0.15 mm long; lower 3 petals c. 9.5 mm wide, with 0-2 small appendages on claw, callus with matted simple septate hairs c. 1 mm long; upper pair, c. 5.5 mm wide, with 2 prominent appendages on claw, callus smooth, obscure. Androgynophore 5-7 mm long, straight, sparsely hairy, distinctly shorter than calyx; stamens 10, shortly connate at base, 0.7-1.2 mm long; anthers transverse; staminodes 5, spathulate-acute, c. 0.8 mm long. Style straight, c. 1 mm long; stigma of 5 pin-like lobes, terete. Capsule adpressed to ground, depressed globose to obloid, 5-9 mm long, 6-7 mm diam., sparsely stellate-pilose, brown, styles caducous; carpels straight, 1-3-seeded. Seeds sub-rhomboidal, c. 2.5 mm long, 2 mm wide, irregularly rugose, dark brown. Flowers: Aug.-Jan. Fruits: Dec.-June. Distribution and ecology. The species is endemic to the northern N.T. and very common in the north-western Top End, north of Pine Creek and west of the Goomadeer River, including the Tiwi Islands. It grows in Eucalyptus forest and woodland on lateritic soils. Notes. Helicteres darwinensis differs from II. integrifolia by the prostrate habit, coarse indumentum with stipitate stellate hairs, longer, persistent stipules which are 7-20 mm long, inflorescence 30-50 mm long, and with the bracts, calyx, petals, and androgynophore all longer. While the new shoots are often erect at emergence, they quickly become prostrate. This species has been referred to as Helicteres A78389 Darwin and Helicteres sp. Darwin (S.T. Blake 16793) on herbarium specimen labels and unpublished checklists from DNA. Etymology. A reference to the City of Darwin, Northern Territory, around which the species is a common component of the Eucalyptus woodland. Helicteres flagellaris (Benth.) Cowie, comb, et stat. nov. (Figs 4, 9). Helicteres dentata var. flagellaris Benth., Flora australiensis 1: 233 (1863); H. semiglabra var. flagellaris (Benth.) F.M. Bailey, The Queensland Flora 1: 142 (1899). Type: Australia, N.T., Port Essington, Apr. 1840, J. Armstrong 533 (Holotype: K, n.v., photo DNA). Other specimens examined. NORTHERN TERRITORY. Arafura Swamp, 8 June 1996, R. Booth 1893 (DNA); along track to Black Jungle Spring, 1 January 1992, K. Brennan 1675 (DNA); Kakadu N.P., Arnhem Hwy, 1 km W of Flying Fox Ck, 13 January 1991, K. Brennan 841 (DNA); 1.7 km S of Danger Point. Cobourg Peninsula, 6 May 1987, M.J. Clark 1052 (DNA); 25.1 km E of Kerr Pt, Weipa, 21 April 1991, J. Clarkson 9013 & V.J. Neldner (BRI, DNA, K, L, MBA, PERTH); N of Bertie Ck, 25 February 1992, ./. Clarkson 9202 & V.J. Neldner (BRI, DNA, MBA); Kapalga, 15 December 1976,76 Collins 175 (BRI), CANB, DNA); Kakadu N.P., Kapalga, near Red Lily turn-off, 13 39 I. D. Cowie December 1988, I.D. Cowie 721 (DNA); Kakadu N.P., Arnhem Hwy, Kapalga turn-off, 11 November 1991, l.D. Cowie 2113 (DNA, MEL); Arnhem Hwy, truck stop just W of Kakadu. 10 November 1992, I.D. Cowie 3265 & S. Pilkington (DNA); Port Essington, Wangewanja Cove, 13 April 1993, I.D. Cowie 3371 (BRI, CANB, DNA, MEL); Cobourg Peninsula. Raffles Bay. 20 April 1993. I.D. Cowie 3772 (DNA); Mary River, near Rock Hole, 6 November 1994, I.D. Cowie 5171 (DNA); Central Arnhem Land, Emu Springs area. 17 September 1999, I.D. Cowie 8397 & R.K. Harwood (DNA); Arnhem Land, c. 44 km SSE Maningrida, 16 March 2000, I.D. Cowie 8518 (DNA); near Ranger Station, c. 13 km SSE of Maningrida, Arnhem Land, 6 April 2000, I.D. Cowie 8767 (DNA); Arnhem Land, c. 22 km ESE of Ramingining, 28 June 2001, I.D. Cowie 9462 (BRI, DNA); Arnhem Hwy, 4.4 km E of Point Stuart turnoff, 24 February 2004, I.D. Cowie 9985 (BRI, DNA, MEL, CANB); Mary River, W side near Arnhem Hwy, 4 January 2005, I.D. Cowie 10346 (BRI, CANB, DNA); c. 40 km W of Jabim, 18 March 1981, L. Craven 7853 (CANB, DNA); Kapalga, 23 February 1981,./- Cusack 110 (DNA); Croker Is. (northern), 28 November 2001, D. Dixon 952 & B. Baker (DNA); Koongarra Area, 24 April 1980, C. R. Dunlop 5516 (BRI, CANB, DNA, MEL); Kakadu N.P., Kapalga, 12 March 1982, C.R. Dunlop 6118 (BRI, DNA); Kapalga, 8 December 1992, J. Egan 394 (DNA); Kakadu N.P.; Kapalga, 10 January 1993, J. Egan 859 (CANB, DNA); Kakadu N.P., Kapalga, 7 February 1993, J. Egan 1445 (DNA); Cobourg Peninsula, Record Pt, II April 1993, J. Egan 1996 (CANB, DNA); Cobourg Peninsula, Kangaroo Pt, 11 April 1993,,/. Egan 2008 (CANB, DNA); Kapalga, 16 August 1993, J. Egan 2529 (DNA); Kapalga, Kakadu N.P., 25 January 1994, J. Egan 3052 (DNA); Black Point, Ranger Stn, Cobourg Peninsula Wildlife Sanctuary, 23 May 1983, P. Fiyxell 4256 (CANB, DNA); 16m SSE of Nourlangie Safari Camp, 27 February 1973, M. Lazarides 7884 (CANB, DNA); Arnhem Land, about 2 km NW of Dog Mountain, 12 September 1998, C.P. Mangion 609 & I.D. Cowie (DNA); Mckinlay River Station, 24 August 2001, D. Napier 63 (DNA); Cobourg Peninsula, S of Stewart Point, 15 March 2002, J.A. Risler 1893 & R. Firth (DNA); Arnhem Land, 16 km N ofMirmgatja, 4 November 1987, J. Russell-Smitli 3969, D. Lucas (DNA); Kakadu N.P., Kapalga, 18 January 1995, J. Russell-Smith 9218 (DNA); Murwangi, 10 September 1998, P.S. Short 4807 & C.R. Dunlop (DNA); 15 km from Koongarra Saddle along track to Jabiru, 24 April 1980, I.R.H. Telford 8154 & J. Wrigley (CANB); Kakadu N.P., near Nanguluwurr rock art site, 13 November 2010, J. Westaway 3384 (DNA, MO); Flying Fox Ck, Arnhem Hwy, 9 January 1986, G.M. Wightman 2516 (DNA); Cobourg Peninsula, Gamaragi area, 27 June 1995, G.M. Wightman 6235 & D. M. Jackson (DNA). Description. A subshrub with prostrate annual above¬ ground stems to 0.5 m long and a woody perennial tap root. Most parts with a sparse to very dense indumentum of Fig. 9. Helicteres flagellaris. Flower and leaf bases. (/. D. Cowie 10346). sessile, stellate hairs 0.2-0.4 mm diam. Stipules persistent, subulate, 4-5.5 mm long. Petiole 1.5-5 mm long. Leaf lamina discolorous to strongly discolorous, ovate to suborbicular, 1.8-6.5 cm long, 1.1-5.1 cm wide, 1-1.5 times longer than wide, upper surface usually glabrous or with simple hairs, lower surface often hoary, base cordate, rarely rounded, 5-7-ncrved, margin irregularly dentate or serrate throughout, apex rounded to acute, often shortly apiculate. Inflorescence an interrupted axillary cyme, 30-140 mm long, with 1-6 flowers; bracts subulate, rarely leafy, 3—4 mm long; extrafloral nectaries present, dark red; pedicels 0-4 mm long. Calyx 4.5-9 mm long, tube 4-5 mm long, lobes 1.5-5 mm long. Corolla scarcely 2-lipped. Petals mauve, slightly dimorphic, slightly bilabiate, 10-16 mm long, long clawed; limb broadly cuneate to depressed obovate, callused near base, both surfaces with a sparse indumentum of erect to appressed short septate hairs c. 0.1 mm long; lower 3 petals with lor 2 small appendages on margins of claw; callus woolly hairy; upper 2 with 2 prominent appendages on claw, callus smooth, shiny, glabrous. Androgynophore 4.5-6 mm long; androecium and gynoecium oblique; stamens c. 0.8 mm long. Style 1-1.5 mm long; stigma of 5 pin-like lobes, terete. Capsule ellipsoid to globose, 8-12 mm long. Seeds 1-3 per locule; shape variable, irregularly rugose, dark brown. Flowering & Fruiting: Nov.-June. Distribution and ecology. Common in the northern N.T. from the Mary River, Kakadu N.P., and Cobourg Peninsula to Ramingining, but in the Darwin Region known only from east of the Mary River. Grows in Eucalyptus woodland mostly on sandy or silty soils. Notes. As accepted here, there is some variation in the indumentum of this species. Broadly, specimens fall into those with coarse, stipitate stellate hairs (0.5) 1.0—1.5 mm in diam. Growing on sites with finer textured soils and those with finer, sessile stellate hairs 0.1-0.3 (0.6) mm diam. and growing on sandy soils. Within the large hair group some specimens have calyx lobes longer than the tube (Short 4807 & Dunlop; Cowie 8397 & Harwood), and in Cowie 8767 the lobes are also almost glabrous (rather than densely hairy) 40 Helicteres (Malvaceae) from the Northern Territory on the abaxial surface. Several specimens from the Mary River area {Cowie 5171, 10346; Napier 63) have lanceolate to oblanceolate or very broadly elliptic floral bracts, with long stipitate stellate hairs along their margins. Further examination of material from Queensland is required. A number of specimens e.g. L.J. Brass 19587, ./. Clarkson 10028 & V.J. Neldner appear to be closely related and may be referrable to this taxon. The species differs from H. integrifolia by the prostrate habit, persistent stipules 4-5.5 mm long, leaves typically ovate to suborbicular, the upper leaf surface glabrous or with simple hairs, the cymes 30-140 mm long, petals 10-16 mm long and the longer androgynophore. Helicteres flagellaris has also been referred to as “Helicteres D47082 prostrata” on herbarium specimen labels and unpublished checklists lists from DNA. Helicteres integrifolia (F.Muell.) Cowie, comb. nov. (Fig. 4). Methorium integrifolium F.Muell., Transactions of the Philosophical Institute of Victoria 3: 40 (1859). Type citation: “On rocky declivities of the sandstone tableland of the Upper Victoria River”. Type details: Australia, N. T., upper Victoria River, January 1856, F. Mueller s.n. (Holotype: MEL (MEL222173); Isotype: K, m.v., photo DNA). Helicteres cana auct non. (Schott & Endl.) Benth., Benth., Flora australiensis 1: 232 (1863). Description. Erect to spreading, multi-stemmed subshrub to 0.5 m, with annual above-ground parts and a woody perennial root stock. Most vegetative parts and calyx with moderately dense to very dense sessile stellate hairs 0.1 to 0.6 (-1.2) mm diam., hairs sessile or sometimes shortly stipitate; new growth with scattered red minute glandular hairs. Stipules caducous, subulate, 1.5-6 mm long. Petiole 3-15 mm long. Leaf lamina ovate to elliptic to suborbicular, less often lanceolate, oblong- oblanceolate, or oblanceolate to obovate, 18-115 mm long, 8-50 (-75) mm wide, 1.1-3.2 times longer than wide, scarcely to strongly discolorous, under surface usually hoary or grey-green when young becoming less so with age, upper surface with sparse to dense stellate hairs, green to grey, base cordate to rounded or cuneate, symmetric to sometimes asymmetric, 3-nervcd for 1/3 to 3/4 of length, margins entire near base, usually becoming serrate near apex, apex acute to rounded. Inflorescence axillary, cymose, to 35 mm long, with few-many dichasia, dichasia 2- or 3-flowered; extra-floral nectaries present; bracts 1-4 mm long, subulate; pedicels 0-6 mm long. Calyx tubular to narrowly obconical-campanulate, 3-6 mm long; basal nectary short, incomplete, margin truncate; lobes triangular, 0.5-2 mm long. Corolla weakly 2-lipped or not. Petals mauve-pink, slightly dimorphic, 5-10 mm long, long clawed, limb cuneate, with sparse, minute, erect, glandular hairs, slightly callused near base, apex truncate to obliquely truncate, irregular; lower 3 slightly longer than upper pair, claw with 1-2 small appendages, callus woolly hairy; upper pair with 2 prominent appendages on claw, callus smooth shiny glabrous, claw with sparse longer simple septate hairs. Androgynophore 2.5-4.5 mm long, straight, with scattered minute hairs on adaxial side, shorter than or equal to calyx; stamens 10, shortly connate at base, 0.8-1 mm long; anthers transverse; staminodes 5, oblanceolate to spathulate-acute, c. 0.7-1 mm long. Style 1-1.2 mm long; stigma of 5 pin¬ like lobes, terete, caducous. Capsule ovoid to ellipsoid or globular, 8-19 mm long, 6-14 mm diam., with a mid-dense covering of sessile stellate hairs, stellate-hairy bristles absent or present and then sub-dense or dense, floccose, 1.5-4 mm long; carpels straight, 1-4-seeded. Seeds variously shaped, ellipsoid to conoidal or sub-rhomboidal, often laterally compressed, 1.8-3.8 mm long, 1.7-2.5 mm wide, irregularly rugose, dark brown. Flowers: Sept.-Feb. Fruits: Feb.-June. Notes. Bentham (1863) reduced Methorium integrifolium F.Muell. to synonymy under H. cana. While the leaves of the type of M. integrifolium have almost entire margins and a similar greyish colour and indumentum to H. cana, 1 believe that the species currently known as M integrifolium must be reinstated and transferred, as above, to the genus Helicteres. The type specimen of M. integrifolium has a calyx 4-5 mm long, the petals arc c. 7 mm long, the lower petals have a patch of long, matted, glandular hairs and the androgynophore is 3.5-4 mm long. In contrast, /-/. cana has a calyx 7-11 mm long, the petals 11-20 mm long, the lower ones lack a patch of long matted hairs and the androgynophore is 7.5-14 mm long. These character differences are maintained in populations of H. cana which are sympatric with M. integrifolium Further investigation shows that H. integrifolia and H. dentata form part of a complex varying in leaf width, discolorousness, density of indumentum on the leaf surfaces, length of bristles on the fruit and degree of serration of the leaf margin. Variation appears to be partly related to edaphic factors and partly clinal. I here recognise two subspecies within 11. integrifolia, i.e. subsp. integrifolia and subsp. dentata. However, there are some specimens which do not comfortably fit within either subspecies and for the time being are simply referred by me to H. integrifolia. Thus, a series of collections from southeast Arnhem Land and adjacent Limmen N.R ( Cowie 12368, 12382, Lewis 1008, 1018, Manning 425, Wilson 1285) have relatively small concolorous but bifacial leaves, a coarse open indumentum of hairs 0.5-1.2 mm diam., are almost devoid of short glandular hairs on the new growth and have densely long-bristled fruit. Flowers have not been seen. These may prove to represent a distinct taxon allied to //. integrifolia, H. sphaerotheca and H. tenuipila and the variation exhibited by them is not included in the description above. Similarly, the specimens Brennan 1994, 270 and Cowie 8565 may represent an additional undescribed taxon. They are distinctive in having entire or few-toothed, lanceolate, concolorous leaves, a short, close indumentum and fruit with bristles to 0.5 mm long, although the flower 41 I. D. Cowie size is consistent with H. integrifolia, H. angustifolia, H. sphaerotheca and H. tenuipila. These characters and the small flowers are also consistent with specimens of the southeast Asian H. lanceolata (Tang et al. 2007). A number of collections from the Douglas Springs area, which lack flowers, are also tentatively referred to this species pending further collections from that area. Both subspecies maintain their low, multi-stemmed growth habit in cultivation, regenerating each wet season from a perennial, woody rootstock, even when protected from fire. I Leaf lamina slightly discolorous, upper surface densely hairy; capsule densely covered with floccose, stellate-hairy bristles 1-4 mm long. .subsp. integrifolia 1: Leaf lamina strongly discolorous especially when young, upper surface with sparse to scattered hairs; capsule lacking stellate-hairy bristles or bristles scattered, rarely to 2 mm long .subsp. dentata Helicteres integrifolia (F.Muell.) Cowie subsp. integrifolia Other specimens examined. NORTHERN TERRITORY. Gregory N.R, Newcastle Ranges Sinkhole, 15 April 1996, R. Booth 1624 (DNA, MEL); Timber Creek Lookout Area, 27 January 2011, K. Brennan 90/8 (DNA); Bullita, 24 August 1991, P.S. Brocklehurst 562 (DNA); S of Ramingining, 22 May 1990, l.D. Cowie 1313 (CANB, DNA, MEL, PERTH); 18 Widdup Crcs., Driver, 21 January 1992,/.£>. Cowie 2147 (DNA); Gregory N.P., Bullita station, 13 February 1992, l.D. Cowie 2322 & P.S. Brocklehurst (BR1, DNA, MEL); Gregory N.P., W of Timber Creek, 15 February 1992, l.D. Cowie 2427 & P.S. Brocklehurst (BRI, DNA); just W of Timber Creek, lookout area, 18 May 1997, l.D. Cowie 7574 (CANB, DNA); S of Ramingining, II December 1998, l.D. Cowie 8145 & R.K. Harwood{ BRI, CANB, DNA, MEL, NSW, NY); near SW side of Arafura Swamp, Central Arnhem Land, 11 June 2000, l.D. Cowie 8913 & C.R. Dunlop (DNA); just W of Timber Creek, lookout area, 21 November 2001 ,1.D. Cowie 9395 & G.M. Holland (BRI, DNA); just W of Timber Creek lookout area overlooking town, 21 November 2001, l.D. Cowie 9500 & G.M. Holland (BRI, CANB. DNA, K, MEL, MO); 25 km E of Bulman, 23 June 1990, C.R. Dunlop 8650 & N.G. White (BRI, DNA); Gregory N.P., side gorge, plateau approx. 29 km NW of Bullita outstation. 300 m from head of sandstone, 15 April 1996, M. Duretto 1008 & C. Coles (DNA, MEL); Timber Creek, road to Scarp, V.R.D, 11 April 1990, M. Evans 3120 (DNA, K); Gregory N.P., c. 20 km N of Bullita Ranger Station, 15 April 1995, W. Gaskon 10 & D.B. Foreman (DNA); E of East Baines River, 10 March 1989, G.J. Leach 2402 & C.R. Dunlop (CANB, DNA); Gregory N.P., 10 March 1998, C.R. Michell 1145 (DNA); Gregory N.P., fire plot 22,20 May 1999, C.R. Michell2441 & A.H. Johnson (DNA); VRD, headwaters of Big Horse Ck, 12 March 1989,./. Russell-Smith 7776 (DNA); Gregory N.P., headwater of unnamed tributary of East Baines River, c. 46 km SW of Timber Creek, 15 April 1996, N. Walsh 4359 & G. Jones (DNA, MEL); Milingimbi, Arnhem Land, 16 March 1987, G.M. Wightman 3410& N.M. Smith (DN A). Description. Leaf lamina often grey-green, slightly discolorous, the upper surface densely hairy. Capsule 11-14 mm diam., densely covered with floccose, stellate- hairy bristles 1-4 mm long. Flowers: Nov. Fruits: Feb.-June. Distribution and ecology. This subspecies is endemic to the northern N.T. and grows in Eucalyptus woodland on sandy soils. Notes. Specimens of subsp. integrifolia with almost concolorous leaves can look superficially similar to H. cana subsp. latifolia , but subsp. integrifolia has a shorter androgynophore, calyx and corolla and the lacks long, matted, glandular hairs on the lower petals. This subspecies has been referred to as “ Helicteres D29330 multicaulis" and ‘’'Helicteres sp. many stems {G.M. Wightman 3410)” on herbarium specimen labels and unpublished checklists lists from DNA. Helicteres integrifolia subsp. dentata (F.Muell. ex Benth) Cowie, comb, et stat. nov. (Figs 4, 10) Helicteres dentata F.Muell. ex Benth. Flora australiensis 1: 232 (1863); Helicteres dentata var. dentatata, Benth., Flora australiensis 1: 233; Methorium dentation (F.Muell. ex Benth.) F.Muell., Systematic Census of Australian Plants 15 (1882). Type details: Australia, Northern Territory, Upper Victoria River, October 1855, F. Mueller, (Holotype: K, n.v.. photo DNA; possible Isotype: MEL (MEL222177)). Other specimens examined. NORTHERN TERRITORY. King River, W of Katherine, 15 November 1995, M. Banin 2042 (BRI, DNA, MEL); Daly River Rd, adjacent to Adelaide River crossing, 16 February 1996, R. Booth 1464 & l.D. Cowie (BRI, DNA, MEL); Kakadu N.R, rocky hills just north of Barramundi Gorge turn-off on Kakadu Hwy, 15 May 1991, A'. Brennan 1374 (DNA); Kakadu Hwy, 1.6 km N of Barramundi Ck, 22 November 1991, A. Brennan 1620 (DNA); 1.5 km SE Fisher airstrip, 12 December 1991, A. Brennan 1679 (DNA); Kapalga; E slope of hill, 21 December 1991, K. Brennan 1716 (DNA); Kakadu N.P., on track to Barramundi Gorge, c.200 from Kakadu Hwy turn-off, 13 January 1992, A. Brennan 1777 (DNA); English Company Islands, island just SW of Bosanquet Is., 6 September 1996, A. Brennan 3311 (DNA); Cobourg Peninsula, 9 April 2006, A. Brennan 7116 (DNA); Kakadu N.P., Old Darwin road, 6 November 2008, A. Brennan 7876 (DNA); Melville Bay, 16 February 1803, R. Brown s.n. (BM, CANB); North Coast, R. Brown (DBN, MEL, NSW); Melville Is., Penelli road, 24 June 1987, M.J. Clark 1236 (DNA); Kapalga, 27 April 1977, R. Collins 382 (CANB, DNA, K); Gove Peninsula, Arnhem Land, J.F. Cooper s.n. (NSW); I km past Adelaide River, Daly River Rd, 9 November 1990, l.D. Cowie 1400 & C.R. Dunlop (BRI, CANB, DNA, MEL, PERTH); Melville Is., road to Paru, 42 Helicteres (Malvaceae) from the Northern Territory Fig. 10. Helicteres integrifolia subsp. dentata. Flower and leaves. (/. D. Cowie 11801). 1.5 km below jump-up, 21 January 1992, 1.D. Cowie 2179 A G.J. Leach (BRI, CANB, DNA, MEL); Melville Is., Pickertaramoor, 21 January 1992, 1.D. Cowie 2186 A G.J. Leach (DNA); Victoria Hwy, W of Katherine, 10 February 1992, I.D. Cowie 2207 A P.S. Brocklehurst (CANB, DNA, MEL); Gregory N.P., near Matt Wilson Lookout, 10 February 1992, l.D. Cowie 2232 A P.S. Brocklehurst (BRI, DNA, MEL); Arnhem Bay, mouth of Cato River, 25 May 1992, I.D. Cowie 2890 (DNA, MEL); 34 km N of Katherine, Stuart Hwy, 31 December 1992, l.D. Cowie 3273 (CANB, DNA, MEL); Melville Is., Pickertaramoor, 31 March 1994, l.D. Cowie 4837 (CANB, DNA); Garalja Is., 30 April 1996, l.D. Cowie 6808 (DNA); Macassan pictures, c. 10 km SE ofNhulunbuy, 22 May 1996, l.D. Cowie 7018 (DNA);NitmilukN.P., Douglas Springs area, 22 May 2001, l.D. Cowie 9233 A D. Dixon (DNA); Victoria Hwy, c. 2 km SW of Scott Ck, 20 November 2001, I.D. Cowie 9390 A G.M. Holland{ BRI, CANB, DNA); Victoria Hwy, Kuwang Lookout, c. 30 km E of Timber Creek, 21 November 2001, l.D. Cowie 9502 A G.M. Holland (BRI, CANB, DNA, MO); Menngen (Innesvale) Station, 22 November 200!,/.£). Cowie 9505 A G.M. Holland (DNA, MEL, NSW); Gregoiy N.P., near Matt Wilson Lookout, 22 November 2001, I.D. Cowie 9507 & G.M. Holland (B, BRI, DNA, MEL, MO); Yinberric Hills, N of Edith River, 23 November 2001, I.D. Cowie 9514 A G.M. Holland (B, BRI, CANB DNA, MEL, MO, NSW); Yinberric Hills, N of Edith River, 23 November 2001, ID. Cowie 9516 A G.M. Holland (DNA, MEL); Gunn Point area: 6.2 km SE of Gunn Point, 5 December 2003, l.D. Cowie 9975 A R.K. Harwood (BRI, CANB, DNA, MO); Bradshaw Field Training Area, c. 70 km NE of Timber Ck, near fire plot 2, 30 March 2007, I.D. Cowie 11458 A B. Stuckey (B, DNA); c. 14 km SE ofNhulunbuy, on Central Arnhem road, 25 November 2007, l.D. Cowie 11801 (BRI, DNA, MO); road to Mata Mata Outstation, c. 7.5 km N of Central Arnhem Road, W ofNhulunbuy, 4 December 2007, l.D. Cowie 11840 (BRI, DNA, MEL, B); Kapalga, 2 May 1990, J. Cusack 572 (DNA); Kakadu N.P., Kapalga, top of transect G. 8 November 1991, J. Cusack s.n. (DNA); Mitchell Range, Arnhem Land, 1 April 1999, C.R. Dunlop 10252 A N.G. White (DNA); Melville Is., Poonali Beach, 18 December 1975, C.R. Dunlop 3996 (DNA, NT); Kambolgie Ck, 5 km E of, 19 April 1993, J. Egan 2128 (DNA); Kambolgie Ck, 5 km E of. 19 April 1993, J. Egan 2132 (DNA); Kakadu N.P., Kapalga G plot, 15 March 1994, J. Egan 3286 (BRI, CANB, DNA); Hayes Ck, 5 April 1995, J. Egan 4641 (BRI, DNA, MEL); Victoria Hwy, near Kuwary lookout. 12 May 1995,,/. Egan 4958 (DNA); Melville Is., 10 km SE of Milikapiti, 8 December 1995, J. Egan 5212 (DNA, MEL); Stuart Hwy/Edith Ck crossing, 17 December 1990, M. Evans 3515 (DNA, K); Melville Is. on road to Poonali, 1 February 1992, R. Fensham 1206 (DNA); Melville Is., 18 September 1986, R. Fensham 308 (DNA); Melville Is., 25 October 1986, R. Fensham 357 (DNA); road to Matt Wilson lookout, Gregory N.P., 12 January 1998, R.K. Harwood 284 (DNA); 20 km WSW of Dorisvale homestead. Plot 776,25 September 1998, R.K. Harwood 509 (DNA); Gove, 1 June 1981, D. Hinz 81168 (DNA); Nhulunbuy, 9 June 1982, R.C. Hinz 127 (DNA); Glencoe Stn, 1892, N. Holtze 1362 (MEL); 41 km N of Pine Creek turn-off on Stuart Hwy, 5 October 2001, A.M. Hope 6 A D. Dixon (DNA); Nitmiluk N.P., near Douglas Springs, Marrawal Plateau, 5 May 2002, R.A. Kerrigan 587 (DNA); 3 miles [4.8 km] N of Katherine, 15 August 1963, M. Lazarides 6994 (DNA); c. 29 miles [46 km] SW of Nourlangie Safari Camp, 25 February 1973, M. Lazarides 7846 (CANB, DNA, PERTH); Fossil Head, 23 February 1994, G.J. Leach. 4225 (BRI, DNA); 10 miles [16 km] NW of El Sharana, Pine Creek road, 22 January 1973, P. Martensz 465 A R. Schodde (BRI, CANB, DNA, NT); Bathurst Is., Point Fawcett, 1 May 1998, C.R. Michell 1415 A R.K. Harwood (DNA); Edith River area, railway transect 11.1, 13 March 1999, C.R. Michell 2323 A J.A. Risler (DNA); Edith River area, 16 March 1999, C.R. Michell 2399 A J.A. Risler (DNA); Gregory N.P.. Matt Wilson sector, fire plot 2, 17 May 1999, C.R. Michell 2401 A A.H. Johnson (DNA); Nitmiluk N.P., Marrawal Plateau Nth, 8 April 2002, C.R. Michell 3741 (DNA); Nitmiluk N.P., Marrawal Plateau Nth, 9 April 2002, C.R. Michell 3742 (DNA); Nitmiluk N.P., NE comer, 11 April 2002, C.R. Michell3743 (DNA); Cape Wessel, W ofNhulunbuy, 16 December 1998, A.A. Mitchell 5602c (DNA); Daly River road, near Adelaide River, 5 October 1962, J.S. Muspratt 70 (DNA); Fenton Airstrip, 23 October 1974, J. Must 1284 (CANB, DNA, MO, NT); Melville Is., E side of Brenton Bay, Plot 2797, 14 April 2000, J.A. Risler 396 A C.P. Mangion (DNA); Tipperary, Mt Pleasant, 17 February 1968, C.S. Robinson 42 (DNA); Buckingham R catchment, 5 December 1987, ./. Russell-Smitli 4413 (BRI, DNA); 1 km S of Adelaide River Crossing on Daly River road, 19 October 1988, J. Russell-Smith 6323 (DNA); Mount Wilson area, Gregoiy N. P., 27 December 1990,./. Russell-Smith 8452 A D. Lucas (DNA, PERTH); Eastern Arnhem Land. Baralminan, 2 April 43 I. D. Cowie 1992, J. Russell-Smith 8590 (BRI, DNA); Manbulloo Stn, Limestone Ck area, 27 November 1996, N.M. Smith 3924 (DNA); Darwin, W.B. Spencer s.n. (NSW); Kintore Caves Reserve, 5 March 1996, S. Taylor 362 & D.J. Liddle( DNA); Gregory N.R, 6 km NE of Bullita Outstation, 8 February 1986, B.G. Thomson 1124 (DNA); Gregory N.R, 15 km E of Victoria River Inn, 27 February 1986, B.G. Thomson 1238 (DNA); Victoria River, Gregory N.P., 25 February 1986, G.M. Wightman 2771 (DNA); Cape Fewcett, Bathurst Is., 8 April 1987, G.M. Wightman 3586 & N.M. Smith (DNA); Cato River mouth area, Djurlpungbuy, NE Arnhem Land, 16 February 1988, G.M. Wightman 4214 (DNA, MEL). Description. Leaf slightly to strongly discolorous, undersurface usually hoary or grey-green when young becoming less so with age, upper surface with sparse to scattered hairs. Capsule 6-12 mm diarn., with a moderately dense covering of sessile stellate hairs, stellate-hairy bristles usually absent or scattered, rarely to 2 mm long. Flowers: Sept.-Feb. Fruits: Feb.-June. Distribution and ecology. The subspecies is endemic to the northern N.T., from Melville Island to Victoria River and east to Gove Peninsula. It grows in Eucalyptus woodland on sandy, clayey or lateritic soils. It flowers profusely on the new growth early in the wet season. Notes. This subspecies has also been referred to as Helicteres D20342 Gove (G.M. Wightman 3410) on herbarium specimen labels and unpublished checklists lists from DNA. The holotype bears Mueller’s field label with the name “Me thorium dentation n. sp.” and Bentham (1863) clearly ascribes the epithet to Mueller. There is an additional collection of Mueller’s at MEL labelled “Sea Range [Yambarran Range], Oct [ 18]55” (MEL222177). While the corner of the label of this collection has been annotated with a 'B' to indicate it was apparently seen by Bentham and presumably formed part of the original material used to describe H. dentata, he does not cite it specifically. Specimens on both the K and MEL sheets are at a similar stage of growth and it is plausible that they are in fact part of the same gathering. Despite the Upper Victoria River locality on the label of the K sheet, the October collection date indicates that it was probably collected a little further inland than the vicinity of the depot established by Gregory on the upper Victoria River estuary c. 12 km west of the modem township of Timber Creek (Gregory & Gregory 1884). This locality is only 30 km or so from Yambarran Range, the location on the MEL sheet. Because of this uncertainty, the Sea Range specimen at MEL is included here as a possible isotype. Helicteres kombolgiana Cowie, sp. nov. (Figs 3D-G, 4, 11, 12) Helicteres integrifoliae affinis, sed habitu erecto, caulibus paucis, partibus supra terrain perennibus, ramulis et calyce setis pilos stellatos ferens, foliis et ramulis pilos duarum classium amplitudinum ferens, foliis parvioribus acutis, carpellis rostris longis, inflorescentiis brevibus, floribus paucis distinguit. Type: Australia, N.T., cultivated at 18 Widdup Cres., Driver, 13 May 2007,/.D. Cowie 11740 (Holotype: DNA; Isotypes: BRI, CANB, K, L, MEL, MO, NSW). Other specimens examined. NORTHERN TERRITORY. Mt Brockman Range, 13 km S of Jabiru, 23 February 1973, L.G. Adams 3089 (CANB, DNA); Nourlangie Rock car park, 16 May 1995, R. Booth 960 (DNA); Kakadu N.P., below Brockman massif, 1 April 1990, K.G. Brennan 216 (DNA); Nourlangie Rock near main galleries, 28 December 1991, K.G. Brennan 1741 (DNA); Kakadu fire plot 132, Leichhardt Spring, 23 March 1999, K.G. Brennan 3809 (DNA); Upper Deaf Adder Ck, Kakadu fire plot 63, 21 March 2009, K.G. Brennan 7930 (DNA); Upper Liverpool River, on road between Kulnguki and Mirbik, 22 April 2009, K.G. Brennan 8011 (DNA); Kakadu N.P., Birdie Ck, 18 April 1990, I D. Cowie 1091 & G.J. Leach (DNA); Koongarra Saddle, 1 January 1992, I.D. Cowie 2144 (DNA); Arnhem Land, Magela Ck upper catchment, 11 April 1995, I.D. Cowie 5604 & K.G. Brennan (DNA); Kakadu N.P., near East Alligator River, W of Rock Holes, 20 April 1999, I.D. Cowie 8299 (DNA); c. 24 km E Myra Falls, Western Arnhem Land, 17 March 2000, I.D. Cowie 8578 (DNA); Western Arnhem Land, Magela Falls Gorge, 23 August 2004, I.D. Cowie 10271 & Crase, B. (DNA); cultivated at Palmerston from seed collected at Magela Falls, 6 June 2006, I.D. Cowie 10644 (DNA); Palmerston, Widdup Cres,cultivated, 5 January 2010, I.D. Cowie 12530 (CANB, DNA, SING); Little Nourlangie Rock, 19 March 1979, C.R. £>»/;/op 4767 (DNA); Little Nourlangie Rock, 12 May 1978, C.R. Dunlop 4830 (DNA); Jim Jim Falls, 26 March 1982, C.R. Dunlop 6227 (BRI, DNA); Mount Brockman, Kakadu N.P., 27 March 1995, J. Egan 4528 (DNA); Kakadu N.R, 10 km NNE of Jabiru, 28 April 1995, J. Egan 4855 & Knox. S.F. (DNA); Kakadu N.P., Baroalba Springs, 25 May 1983, P. Ftyxell 4272 (CANB, DNA); Kakadu N.P., Mt Brockman, 22 May 1980, M. Lazarides 8908 (AD, CANB, DNA, MEL); Kakadu Nat. Park, 14 km E of Sleisbcck, 18 April 1990, G.J. Leach 2755 & I.D. Cowie (BRI, DNA, MEL); 2 km W of Nabarlek Airstrip, 24 April 1979, M. Rankin 2170 (DNA); Koongarra Jump-up, 12 June 1978, B. Rice 2925 (CANB, DNA); 5 km from Oenpelli, 21 July 1983, J. Russell-Smith 777 (DNA); Goomadeer, escarpment country, 28 October 1987, J. Russell-Smith 3830 & D. Lucas (DNA); Upper East Alligator River, Arnhem Land, 19 April 1988, J. Russell-Smith 5243 c£ D. Lucas (DNA); Kakadu, 16 June 1988, J. Russell-Smith 5681 (DNA); Mt Brockman outlier. Kakadu N.P., 20 April 1989,./. Russell-Smith 8039 (DNA); 12 km E. ofMudginbcrri Homestead, Kakadu N.P., 7 January 1991,/. Russell-Smith 8411 & J. Brock (DNA); Upper Magela Ck valley, Arnhem Land, 3 May 1991, J. Russell-Smith 8461 &J. Brock (DNA); Upper Magela Ck valley, Arnhem Land, 3 May 1991, J. Russell-Smith 8467 &J. Brock (DNA); Kakadu N.P., Koongarra Area, 16 April 44 Helicteres (Malvaceae) from the Northern Territory 1992, J. Russell-Smith 8638 & D. Lucas (DNA); Magela Ck catchment, Kakadu N.P., plots 7B2,6B1, 13 April 1995, J. Russell-Smith 10348 & D. Lucas (DNA, SING); Deaf Adder Ck, Gorge, 22 April 1980, I.R.H. Telford 7997 & J. Wrigley (CANB); 6.5 km SSW Mt Brockman, 23 April 1980, I.R.H. Telford 8040 & J. Wrigley (CANB); 0.5 km NEKoongarraSaddle, 16August 1980. I.R.H. Telford8421 & J. Wrigley (CANB). Description. Open, perennial shrub to 3 m tall, stems few, much branched. Branchlets, young growth, stipules and leaf surfaces covered with a mid-dense to very dense, white indumentum (hyaline on leaf upper surface) of sessile stellate hairs of 0.1-0.3 mm diam., grading into an overlying layer of stipitate stellate hairs 0.5-1.2 mm diam. Calyx also with scattered bristles to 0.7 mm long, bearing stellate hairs. Stipules caducous, subulate, 2.5-5 mm long. Petiole 3-10 (19) mm long. Leaf lamina antrorse, lanceolate to ovate or elliptic, 25-70 (105) mm long, 10-35(60) mm wide, 1.6-3 times longer than wide, slightly to strongly discolorous, hoary or grey-green below when young, base cordate to rounded, symmetric to slightly asymmetric, 3-5-nerved, margins serrate, apex acute to sometimes obtuse. Inflorescence axillary, to 10 mm long, few-flowered (often a single dichasium), extra-floral nectaries present; dichasia 1-2-flowered; bracts 2.5-5 mm long, subulate; pedicels 0-1 mm long. Calyx tubular to obovoid-urceolate, 4-6.5 mm long, basal nectary incomplete, truncate; lobes triangular or narrowly triangular, 1-4 mm long. Corolla weakly 2-lipped. Petals mauve, the upper pair paler than lower 3, slightly dimorphic, 5.5-9 mm long, long clawed, limb cuneate, with scattered minute erect hairs, with a callus near base, apex truncate to rounded; lower 3 slightly longer than upper pair, claw with 2 small appendages, callus woolly hairy; upper pair broader, w'ith 2 prominent appendages on claw (one adaxial, one lateral), callus glabrous, smooth. Androgynophore 3-5 mm long, straight, shorter than calyx; stamens 10, shortly connate at base, c. 0.7 mm long; anthers transverse; staminodcs 5, spathulate-acute. Style straight, 1-1.5 mm long; stigma of 5 pin-like lobes, terete. Capsule ovoid to subglobular, 10-22 mm long, c. 12 mm diam., densely covered with floccose bristles to 4 mm long, bristles bearing stellate hairs 0.6-0.8 mm diam., styles persisting and forming a beaked apex; carpels straight, 2-5-seeded. Seeds variously shaped, ellipsoid to obconoidal or subrhomboidal, often laterally compressed, c. 3 mm long, c. 2.5 mm wide, irregularly rugose, dark brown. Flow'ers: Jan.-May. Fruits: Jan.-July. Distribution and ecology. Endemic to the northern N.T., in Kakadu N.P. and adjacent areas of western Arnhem Land, almost to Maningrida. Grows in Allosyncarpia forest or Eucalyptus woodland, on sandy soils on scree slopes or closely associated with other outcropping sandstone. Notes. This species is allied to Helicteres integrifolia but distinguished by the erect habit, few-stemmed perennial above-ground parts, branchlets and calyx with bristles Fig. 11 . Helicteres kombolgiana. Flower and leaf. (I. D. Come 10644). Fig. 12. Holotype of Helicteres kombolgiana. bearing stellate hairs, hairs of two size classes on the leaves and branchlets; the smaller and acute leaves; long, beaked apices of the carpels and short, few-flowered inflorescences. The open, much branched, few-stemmed growth habit of this taxon is maintained in cultivation. The type collection was grown from seed gathered from l.D. Cowie 10271 , collected at Magela Gorge Falls, N.T. This species has been referred to as Helicteres A63558 Kakadu and Helicteres sp. Kakadu (L.G. Adams 3089) on 45 I. D. Covvie herbarium specimen labels and unpublished checklists from DNA. Etymology. A reference to the Kombolgie sandstone of the western Arnhem Land Plateau, on which geological unit the plant occurs. Helicteres macrothrix Cowie, sp. nov. (Figs 4, 7A-E, 13, 14) Ab Helicteres integrifolia et specierum cognatis inflorescentiis congestis, indumento densissimo, pilis grandioribus, daimetro 1.5-4.5 mm, androgynophoro breviore, capsulis dense hirsutis diagnoscenda. Affinis Helicteres darwinensis Cowie, sed ea habitu erecto, multicaule, inflorescentiis congestis, indumento densissimo, pilis grandioribus distincta. Type: Australia, N.T., Stuart Hwy, near Glenluckie Creek, 9 November 1990, I D. Cowie 1410 & C.R.Dunlop (Holotype: DNA; Isotypes: BR1, CANB, K, NSW, MEL, PERTH). Other specimens examined. NORTHERN TERRITORY. 54 miles [86 km] S of Darwin, Stuart Hwy, 10 January 1969, N.B. Byrnes 1280 (DNA); near Glenluckie Ck, Stuart Hwy, 25 January 1991, /.D. Cowie 1481 & C.R. Dunlop (DNA); near Glenluckie Ck, Stuart Hwy, 15March 1991,/.D. Cowie 1565 & RF. Munns (DNA); Marrakai Road, W of turnoff to Lake Bennett, 29 November 2002, 1.D. Cowie 9654 (DNA); near Lake Bennett, E side, 29 November 2002, 1.D. Cowie 9656 & R.A. Kerrigan (DNA); Lake Bennett, c. 70 km S Darwin, 26 January 1994, C.R. Dunlop 9805 (BRI, DNA); 8.4 km W ofAnnaburroo, 12 September 1992, J. Egan 402 (DNA); 8 km W of Annaburroo, 12 September 1992, J. Egan 403 (DNA); 8 km W of Annaburroo, 1 October 1993, J. Egan 902 (DNA); Annaburroo, 8 km E of dam, 10 January 1 993, J. Egan 903 (DNA); Arnhem Hwy, near Annaburroo, 27 March 1995, J. Egan 4472 (DNA); Mt Bundey area, 8 km W of Annaburroo, near Arnhem Hwy, 14 November 2001, A.M. Hope 23 & I.D. Cowie (DNA, SING): Mt Bundey area, 8 km W of Annaburroo, near Amhcm Hwy, 14 November 2001 ,A.M. Hope 24 & I.D. Cowie (DNA); Mt Bundey area, 6.5 km W of Bark Hut/Annaburroo, 14 November 2001, A.M. Hope 27 & I.D. Cowie (DNA). Description. Erect, multi-stemmed subshrub to 0.6 m, with annual above-ground parts and a woody perennial rootstock. Vegetative parts, calyx and bracts usually with a dense to very dense indumentum of multiangulate stellate trichomes 1.5^1.5 mm diam., sessile or on stipes to 1.5 mm long. Stipules linear, 6-12 mm long. Petiole 0-5 mm long. Leaf lamina ovate, occasionally elliptic or broadly ovate, 50-170 mm long, 40-110 mm wide, 1.3-1.7 times longer than wide, discolorous, base shallowly cordate or rounded, margins senate to dentate, apex acute to obtuse. Inflorescence axillary, congested, cymose, to 40 (70) mm long, with up to 12 flowers, dichasia 2-flowered; extra-floral nectaries present; bracts linear, 8-13 mm long; pedicels 0-1 mm long. Calyx ellipsoid or ovoid, 6-11 mm long, basal nectary incomplete, margin entire; lobes 2-6 mm Fig. 13. Helicteres macrothrix. Flowers and leaf base. (I. D. Cowie 1481). long, triangular. Corolla not or scarcely 2-lipped. Petals mauve-pink, weakly dimorphic, 9-14 mm long, glabrous on exterior (abaxial) surfaces, limb cuneate to obcordatc, calluscd near base; lower 3 petals with 1 or 2 small appendages on margin of claw, callus woolly hairy; upper pair with 2 prominent appendages on claw, callus smooth, shiny, glabrous. Androgynophore 3-4 mm long, straight; stamens 10. shortly connate at base, c. 1 mm long; anthers transverse; staminodes 5, spathulate-acute, c. 0.7 mm long. Style c. 1.5 mm long; stigma of 5 pin-like lobes, terete. Capsule ellipsoid, 13-17 mm long, indumentum very dense, of long-stalked stellate hairs or bristles to 6 mm long each bearing 2-5 stellate hairs, apex attenuate; carpels 3-5-seeded. Seeds rhomboidal, truncate, 3-3.5 mm long, c. 2 mm wide, finely rugose, brown. Flowers: Nov.-Mar. Fruits: Jan.-Mar. Distribution and ecology. The species is endemic to the northern N.T. and known from three subpopulations between Batchelor and Mt Bundey. It grows in Eucalyptus tectifica or E. miniata woodland on clayey soils derived from siltstone or sandy soil derived from granite. Notes. This species may be distinguished from II. integrifolia and related species by the erect multi- stemmed habit, congested inflorescences, very dense indumentum of long stellate hairs and comparatively small flowers. It is allied to II. darwinensis , but distinct from it by the erect multi-stemmed habit, more compact inflorescences, and the indumentum of long stellate hairs, denser and coarser on most parts. The isotype specimens were unfortunately dispersed before examination for preparation of this description and are without determinavit slips indicating their type status. This species has been referred to as Helicteres D2164 Glenluckie Ck and Helicteres sp. Glenluckie Ck (N.B. Byrnes 1280 ) on herbarium specimen labels, unpublished checklists lists, other literature (c.g. Kerrigan & Cowie 2007) and listings under the N.T. Government Territory Parks and Wildlife Conservation Act and Commonwealth 46 Helicteres (Malvaceae) from the Northern Territory Fig. 14 . Holotype of Helicteres macrothrix. Government Environment Protection and Biodiversity Conservation Act. Etymology. From the Greek macro - large and thrix, a hair, a reference to the unusually large hairs present in this species. Helicteresprocumbens (F. Muell ex Benth.) Cowie, comb, et stat. nov. (Figs 4, 15). Helicteres dentata var. procumbens F. Muell. ex Benth., Flora australiensis 1: 233 (1863); II. semiglabra var. procumbens (F. Muell. ex Benth.) F.M. Bailey, The Queensland Flora 1: 142(1899). Type citation: Macadam Range. Type details: Australia, N. T., M’Adam Range (K), between Macadam Range and Providence Hill (MEL), Oct. 1855, F. Mueller (Holotype: K, n.v., photo DNA; Isotype: MEL(MEL222174)). Other specimens examined. WESTERN AUSTRALIA. 58 miles [93 km] N of Kununurra, 2 June 1969, F.C.J. Lullfitz & N. Mackenzie (PERTH); Vicinity of Kimberley Research Station, D.H. Mackenzie 691102-19 (CANB). NORTHERN TERRITORY. Litchfield Station, 2 miles [3.2 km] S W of homestead, 3 May 1968, N.B. Byrnes 665 (DNA, NT); headwaters of Lalngang Ck, 16 May 1994, l.D. Cowie 5075 & N. Walsh (CANB, DNA, MEL); Daly River road, c. O. 5 km N of Silver Mine Ck, 3 December 2002, l.D. Cowie 9660 & C.P. Mangion (B, BRI, CANB, DNA, MEL, MO, NSW); Daly River road, c. 10.7 km N of Daly River Police Stn, 3 December 2002, l.D. Cowie 9662 & C.P. Mangion (B, BRI, CANB, DNA, MEL, MO, NSW); Daly River road. c. 12.7 km N of Daly River Police Stn, 3 December 2002, l.D. Cowie 9663 & C.P. Mangion (BRI, DNA, SING); Daly River road, 12.7 km N of Daly River Police Stn, 17 December 2004, l.D. Cowie 10343 (B. DNA, L, MEL, MO); c. 12 km N of Daly River Police Stn, 22 May 1983, P. Ftyxell4246 (CANB. DNA); Peppimenarti, Moyle River, 31 October 2000, A.A. Mitchell 6406 (CANB); Kurrowa Ck, Daly River Reserve, 16 October 1988,./. Russell-Smith 6295 (DNA); Litchfield N.P., 27 April 1995, S. Taylor 280 (DNA); Daly River Mission area, 26 February 1992, G.M. Wiglitman 5642 (DNA); Daly River Mission area, 24 March 1993, G.M. Wightman 6026 ( DNA). Description. Multi-stemmed subshrub, with prostrate annual above-ground stems to 0.5 m long and a woody perennial root stock. Upper leaf surface with sparse long simple or stellate hairs, leaf undersurface and branchlets with sparse sessile stellate hairs, moderately dense on calyx, stipitate stellate hairs usually absent. Calyx with bristles present towards apex. Stipules caducous or persistent, subulate, 2-6 mm long. Petiole 1.5-5 mm long. Leaf lamina patent, ovate to oblong-ovate or broadly ovate, rarely elliptic, 2.5-8.5 cm long, 1.7-4.8 cm wide, 1.4-2.2 times longer than wide, slightly discolorous, base cordate to rounded, asymmetric, 3-nerved, margins entire to serrulate, apex acute to rounded. Inflorescence axillary, cymose, to 40 mm long, few-many flowered, glands absent; bracts 1-4 mm long, subulate; pedicels 1-6 mm long. Calyx tubular, 3.5-5.5 mm long; lobes triangular, 1.5-2 mm long. Corolla not 2-lipped. Petals mauve-pink, slightly dimorphic, 6-9.5 mm long, long clawed, limb cuneate, with scattered minute erect hairs, with a callus near base, apex truncate to obliquely truncate, irregular; lower 3 slightly longer than upper pair, claw with 2 small appendages, callus woolly hairy; upper pair with 1 or 2 prominent appendages on claw, callus glabrous, smooth. Androgynophore 4-4.5 mm long, straight; stamens 10, c. 0.5 mm long; staminodes 5. Style c. 1 mm long; stigma of 5 pin-like lobes, terete. Capsule ellipsoid, 6-14 mm long, 6-8 mm diam., with a moderately dense covering of sessile stellate hairs, and stellate-hairy bristles to 1 mm long; carpels straight, 2-3-seeded, styles caducous. Seeds not seen. Flowering: Oct.-Dee. Fruiting: Mar.-May. Distribution and ecology. In the northern N.T. and adjacent W.A., from near Kununurra to Litchfield N.P. In D.R., known from near Daly River settlement and along the Daly River road to the western side of Litchfield N.P. Grows in Eucalyptus woodland on sandy soils. Notes. The MEL sheet bears three labels, the middle clearly in Bentham’s hand writing reading "Both I believe forms of H. ( Methorium ) dentata of which we have many” 47 I. D. Cowie Fig. 15 . Helicteres procumbens. Flowers and leaves. (/. D. Cowie 9660 ). and the K sheet with Bentham’s determination “H. dentata var. procumbens ” written on the sheet, indicating both were seen by him. While the type details apparently vary slightly between the K and MEL sheets, one is regarded here as an abbreviated form of the other, perhaps written in haste during the preparation of duplicates. The specimens are a good match and it appears that they form part of the same gathering. The holotype bears Mueller’s label with the name “Methoriwn procumbens n. sp.”. Although Bentham (1863) does not ascribe the epithet to Mueller, he has clearly used Mueller’s name and the authorship can thus be cited as “Helicteres procumbens F.Muell. ex Benth.”. Helicteres procumbens differs from H. integrifolia by the prostrate habit, lack of inflorescence glands, leaves with a sparse indumentum below (not hoary or grey-green) and presence of long simple or sessile stellate hairs on the upper surface. It is also similar to H.flagellaris, but H. procumbens has a short inflorescence which lacks glands, shorter petals, a shorter androgynophore and caducous stipules. Helicteres serpens Cowie, sp. nov. (Figs 4, 7J-N, 16) Helicteres darwinensi similis, sed stipulis deciduis, petiolis saepc longioribus, inflorescentiis apertis, paniculatis, 30-140 mm longis, bractcis 4-7 mm longis, petalis 8-12 mm longis, androgynophoro breviore, capsula setis ad 2.5 mm longis dense tecta distans. Type: Australia, N.T., Cutta Cutta Caves Reserve, 17 December 1992,./. Egan 55b (Holotype: DNA; Isotypes: BRI, n.v., CANB, n.v., MEL, n.v., MO, n.v., K, n.v.) Other specimens examined. NORTHERN TERRITORY. Gunlom-Pine Creek road. 1.5 km toward S Alligator River from creek marked Plum Tree Ck, 12 December 1991, K.G. Brennan I6S1 (DNA); Kakadu N.P., in hills W of Barramundi Gorge, 12 January 1992, K.G. Brennan 1770 (DNA); Kakadu N.P., fire plot 33, 29 February 2000, K.G. Brennan 4874 (DNA); Nitmiluk N.P., veg site 130, 13 February 2001, K.G. Brennan 5254 (DNA); Katherine Gorge N.P., 24 April 1968, N.B. Byrnes 627 (DNA, NT); Plum Tree Ck, 20 January 1969, N.B. Byrnes 1370 (BRI, DNA); Waterfall Ck, S Alligator, 5 January 1972, N.B. Byrnes 2449 (DNA); road to El Sharana, 20 March 1987, M.J. Clark 915 (DNA); Kakadu N.P., near Kurundie Ck, 20 April 1990, l.D. Cowie 1174 & G.J. Leach (DNA); near Mary River Ranger Station, 7 December 1990, l.D. Cowie 1456 & C.R. Dunlop (DNA); c. 10 km S of Cutta Caves T/ofT, Stuart Hwy, 10 December 1991, l.D. Cowie 2139 (DNA, SING); Arnhem Land, headwaters of Cadell River, c. 99 km S Maningrida, 22 March 2000, l.D. Cowie 8708 (DNA); Headwaters of Cadell River, c. 82 km SSE Maningrida, Arnhem Land, 14 April 2000, l.D. Cowie 8843 (DNA); Katherine Gorge N.P., 7 April 1981, L. Craven 6721 (DNA, MEL); King River Valley road, 3 January 1993, J. Egan 821 (DNA); Cutta Cutta Caves Reserve, 18 January 1993,./. Egan 1130 (DNA); King River Valley road, 14 February 1993, J. Egan 1460 (DNA); Cutta Cutta Reserve, 9 March 1993, J. Egan 1816 (DNA); King River Valley road, 9 March 1993, J. Egan 1817 (DNA); 50 km N of Mataranka, 15 April 1993, J. Egan 2059 (DNA); Edith Falls N.P., 15 April 1995, J. Egan 4759 (DNA); Nitmiluk, near visitor centre, 14 December 1990, M. Evans 3483 (BRI, DNA); c. 12 km N Korlobidahda, plot 2243, 23 March 2000, R.K. Harwood 881 (DNA); El Sharana Rd, 43 miles [69 km] from Pine Creek, 12 January 1973, D. Hearne 390 (DNA); Nitmiluk N.P., Marrawal Plateau, 3 May 2002, R.A. Kerrigan 586 (DNA); Nitmiluk N.P., Eva Valley Boundary, 8 March 2002, R.A. Kerrigan 588 (DNA); Fiorina site, 19 miles [30 km] SE of Katherine, 9 December 1963, M. Lazarides 6969 (CANB, DNA, PERTH); 24.5 miles [39 km] NNW El Sharana Mine, 25 February 1973, M. Lazarides 7852 (CANB); 13 miles [21 km] E of El Sharana Mine, 2 March 1973, M. Lazarides 7976 (BRI. CANB, DNA, MEL); Headwaters Koolpin Ck, 20 April 1995, G.J. Leach 4379 (DNA); Kakadu N.P., S of Plum Tree Ck, 6 January 1995, D. Lucas 9166& J. Russell-Smith (DNA); 4m [6 km] NNW of El Sharana, 18 January 1973, P. Martensz 406 & R. Schodde (BRI, DNA); UDP Falls, near El Sharana, 13 January 1973, J.L. McKean 868 (DNA); NitmilukN.P., 28 February 2001, C.R. Michell3299 (DNA); Nitmiluk N.P., Nit 2000, 9 February 2001, C.R. Michell 3300 (DNA); Nitmiluk N.P., SE comer of park, 14 March 2002, C.R. Michell 4172 (DNA); Kakadu Stage 3, SE of Sleisleck, 18 April 1990, T. Orr370 (DNA); Kakadu Stage 3, E of Coronation Hill, 18 April 1990, T. Orr 400 (DNA); Mataranka, 31 July 1980, M. Rankin 2543 (DNA); Nitmiluk N. P., Site 432, 3 April 2001, J.A. Risler 1303 & M. Waetke (DNA); Nitmiluk N.P., site 491, 17 April 2001, J.A. Risler 1304 c£ M. Waetke (DNA); Kakadu, Graveside Gorge, 3 May 1987, J. Russell-Smith 2270 (DNA); Kakadu N.P., Round Jungle, 5 February 1995, J. Russell-Smith 9444 & D. Lucas (DNA); Kakadu N.P.. near Kombolgie Ck crossing on UDP Falls road, 1 May 1990, A. V. Slee 3071 L. Craven (AD, DNA, MEL); 3 km W Barunga Community, 2 February 1987, N.M. Smith 342 (DNA); White stone area near Barunga Community, 12 December 1996 ,N.M. Smith 3947 (DNA); Eva Valley Station, 16 January 1989, B.G. Thomson 48 Helicteres (Malvaceae) from the Northern Territory HOLOTYPE Fig. 16 . Holotype of Helicteres serpens. 2824 (DNA); Cutta Cutta Caves Reserve, 6 February 1989, B.G. Thomson 3268 (DNA); Gorge between Twin Falls & Jim Jim Falls, 24 March 1984, G.M. Wightman 1328 (DNA); Nitmiluk N.P. Ranger H.Q, 13 December 1990, G.M. Wightman 5204 & T. Presnell { DNA). Description. Multi-stemmed subshrub, with prostrate annual above-ground stems to 0.7 m long and a woody perennial root stock. Most vegetative parts and calyx hirsute with sessile or stipitate stellate hairs, sparse to dense sessile stellate hairs on upper surface of leaf lamina. Stipules deciduous, subulate, 4-10 mm long. Petiole 3-40 mm long. Leaf lamina sometimes ground-hugging, lanceolate to ovate, 40-140 mm long, 23-90 mm wide, discolorous when young, hairs of the upper surface 0.4-1.2 mm diam., on the lower surface 0.4-1.7 mm diam., base cordate, asymmetric, 5-nerved, margins irregularly serrate, apex acute to rarely rounded. Inflorescence axillary, open, paniculate, 30-140 mm long, 6-21-flowered, dichasia 1- 2-flowered; extra-floral nectaries present; bracts 4-7 mm long, filiform; pedicels 0-3.5 mm long. Calyx funnel-shaped or ellipsoidal-cylindrical, 5-9 mm long, hairs 0.3-1.1 mm diam., basal nectary incomplete; lobes narrowly triangular, 2- 4 mm long. Petals mauve-pink, dimorphic, 8-12 mm long, clawed to long clawed, limb cuneate, minutely hairy, calluscd near base; lower 3 petals slightly longer, with 0-2 small appendages on claw; callus woolly hairy; upper pair with 2 prominent appendages on claw, callus smooth. Androgynophore 3.5-5.5 mm long, straight, distinctly shorter than calyx; stamens 10, shortly connate at base, c. 1.5 mm long; anthers transverse; staminodes 5, lanceolate or elliptic, 0.5-0.7 mm long. Style c. 1.3 mm long; stigma of 5 pin-like lobes, terete. Capsule ovoid to ellipsoid, 8-20 mm long, 11-14 mm diam., densely covered with stellate-hairy bristles to 5 mm long, hoary, hairs 0.8-1.2 mm diam., styles caducous; carpels straight, 3-5-seeded. Seeds sub-rhomboidal, c. 2.7 mm long, 2 mm wide, irregularly rugose, dark brown. Flowers: Dec.-Apr. Fruits: Feb.-June. Distribution and ecology. The species is endemic to the northern N.T., where it is common from Kakadu N.P. to near Mataranka and east to near Maningrida. It grows in Eucalyptus woodland mostly on deep sandy soils. Notes. Similar to IT. darwinensis but separated by the deciduous stipules, petioles often longer than in H. darwinensis (to 40 mm), the open, paniculate inflorescences 30-140 mm long, the bracts 4-7 mm long, the petals 8-12 mm long, the androgynophore shorter, and the capsule densely covered with bristles to 2.5 mm long. As with H. darwinensis , this species flowers early in the wet season on the shoots of the new growth arising from the woody rootstock. The new growth is very densely hairy and at first acquaintance strikingly different from the sparser indumentum of the mature leaves present later in the wet season. The species shows clitial variation in density of the indumentum with it becoming sparser along a gradient from south of Katherine to Kakadu and Arnhem Land. Specimens from southern Kakadu also have smaller hairs than those from elsewhere. This species has been referred to as Helicteres D4247 elongate and Helicteres sp. elongate ( J. Must 887 ) on herbarium specimen labels and unpublished checklists from DNA. Etymology. From the Latin serpens - creeping, a reference to the prostrate habit. Helicteres sphaerotheca Cowie, sp. nov. (Figs 4, 70-S, 17) Ab Helicteres integrifolia et specibus cognatis stipulis persistentibus, lamina folii angustc lanceolata ad anguste oblanceolate, 2-7 mm lata, 5-8.5-plo longiore quam latiore, concolora, pagina supera pilis sparsis simplicibus, capsula globosa, setis densis, usque ad 2 mm Iongis vestita discrepans. Type: Australia, N.T., Arnhem Hwy, 300 m E of West Branch of West Alligator R, 4 January 1992, l.D. Cowie 2146 (Holotype: DNA; Isotypes: AD, B, BR1, CANB, CNS, K, L, MEL, MO, NSW, NT, NY, PERTH. SING). Other specimens examined. NORTHERN TERRITORY. Arnhem Hwy, 300 m E West Branch of W Alligator River, 3 December 1990, K.G. Brennan 1642 (DNA); Arnhem Hwy, 300 m E of West Branch of W Alligator R, 11 49 i. D. Cowie December 1991, K.G. Brennan 1676 (AD. BRI, CANB, DNA, K, L, MEL. MO, NSW, PERTH); Arnhem Hwy,c. 0.5 km W ofW branch of Flying Fox Ck, 29 March 1991, l.D. Cowie 1594 (BRI, DNA, CANB, PERTH); Kakadu N.P., c. 500 m E of W branch of West Alligator River, 9 April 2003, l.D. Cowie 9750 & D. Dixon; (DNA); 60 km W of Jabiru, 19 March 1981, L. Craven 7854 & G. Whitbread (CANB, DNA, MEL); Kapalga, 12 March 1982, C.R. Dunlop 6120 (BRI, DNA); Wildman River Reserve, 29 January 1997, C.R. Michell 348 (DNA). Description. Erect, multi-stemmed subshrub to 0.6 m, with annual above-ground parts and woody perennial root stock. Most vegetative parts and calyx with sparse sessile stellate hairs, sparse simple hairs on upper surface of leaf lamina. Stipules persistent, setose, 1-2 mm long. Petiole 0.5-1 mm long. Leaf lamina patent to recurved, narrowly lanceolate to narrowly oblanceolate, 15-35 mm long, 2-7 mm wide, 5-8.5 times longer than wide, concolorous, base rounded, symmetric, 3-nerved, margins entire, or irregularly serrate near apex, apex acute to rounded. Inflorescence axillary or terminal on short lateral shoots, cymose, 10-15 mm long, few-flowered, dichasia 3-flowered; extra-floral nectaries present; bracts c. I mm long, setose; pedicels !^L5 mm long. Calyx tubular to funnel-shaped. 3.5-4.5 mm long, basal nectary short, incomplete, truncate; lobes triangular, c. 0.5 mm long. Petals mauve-pink, scarcely dimorphic, 7-9.5 mm long, long clawed, limb cuneate. minutely hairy, callused near base, apex obliquely truncate; lower 3 petals slightly longer than upper pair, claw with 1-2 small lateral appendages, callus woolly hairy; upper pair with 2 prominent appendages on claw, callus glabrous, smooth. Androgynophore 4.5-5 mm long, straight, equal in length to calyx; stamens 10, shortly connate at base, c. 0.8 mm long; anthers transverse; staminodes 5, spathulate-acute, c. 0.7 mm long. Style c. 1.3 mm long; stigma of 5 pin-like lobes, terete. Capsule globular, 8-11 mm diam.. densely covered with stellate- hairy bristles to 2 mm long, hoary, styles caducous; carpels straight, 1-3-seeded. Seeds sub-rhomboidal, c. 2.5 mm long, 2 mm wide, irregularly rugose, dark brown. Flowers: Dec.-Jan. Fruits: Mar. Distribution and ecology. The species is endemic to the northern N.T., between the Mary River and the South Alligator River. It grows in Eucalyptus ininiata woodland on sand. Notes. Differing from H. integrifolia and related species by the persistent stipules; the lamina narrowly lanceolate to narrowly oblanceolate. 2-7 mm wide, 5-8.5 times as longer than wide, concolorous, upper surface with sparse simple hairs and the globular capsule with bristles to 2 mm long. This species has been referred to as “ Heiicteres D21039 linifolia” and “ Heiicteres sp. linear leaves (L.A. Craven 7854)" on herbarium specimen labels and unpublished checklists lists from DNA. Etymolygy: from the Greek, sphaera - a ball or globe and thece - a case or container, in reference to the globular capsule. Heiicteres tenuipila Cowie, sp. nov. (Figs 3H-K, 4, 18, 19) Ab H. integrifolia et speciebus cognatis indumento sparsiore et grossiore, foliis oblanceolatis grandis, pilis slcllatis sparsioribus in paginis ambabus, parum discoloribus; calyce pilis stipitatis stellatis frequentibus secedus. Type: Australia, N.T., Litchfield N.P., opposite turnoff to Green Ant Creek, l.D. Cowie 10361 & J.L. Egan (Holotype: DNA (3 sheets); Isotypes: AD, B, BRI, CANB, K, L, MEL, MO, NSW, PERTH). Other specimens examined. NORTHERNTERRITORY. Stapleton Park, 20 May 1985, D. Bowman 185 B. Wilson (DNA); Litchfield N.P, track to Lost City, 23 November 1990, l.D. Cowie 1436 & C.R. Dunlop (BRI, CANB, DNA, K, PERTH); Litchfield Park, near Blythe homestead ruins, 26 November 1992, l.D. Cowie 3269 (BRI, DNA, MEL); Litchfield N.P, Tabletop Range, 23 March 1995 J.D. Cowie 5347 & S. Taylor (DNA); Litchfield N.P., turnoff to Lost City, 20 January 2005, l.D. Cowie 10362 & J. Egan (CNS, DNA, SING); Tolmer Plateau, 5 April 1991, C.R. Dunlop 50 Helicteres (Malvaceae) from the Northern Territory Fig. 18. Helicteres tenuipila. Flowers and leaves. Inset. Flower (I. D. Cowie 10361). 8809 & I.D. Cowie (I3RI, DNA, MEL); Litchfield N.P., Lost City road, 10 December 1992, J. Egan 410 (DNA); Litchfield N.P., road to Lost City, 26 December 1992, J. Egan 646 (DNA); Litchfield N.P., Lost City, 24 January 1993, J. Egan 1161 (DNA); Litchfield N.P., Lost City road, 17 March 1993, J. Egan 1762 (DNA); Litchfield N.P., Lost City road, 24 April 1993, J. Egan 2182 (DNA); Litchfield N. P., 8 March 2006, J. Egan 5726 & D. Lucas (DNA); Litchfield N.P., 2 November 1995, S. Taylor 328 (DNA). Description. Erect, multi-stemmed subshrub to 0.5 m, with annual above-ground parts and a woody perennial root stock. Most vegetative parts with sparse to moderately dense sessile stellate hairs, denser on calyx, stipitate stellate hairs occasional on stems, common on calyx. Stipules caducous, setose, 5-7 mm long. Petiole 4-7 mm long. Leaf lamina elliptic to oblanccolate or obovate, rarely ovate or lanceolate, 45-95 mm long, 15-40 mm wide, 2.4-3.3 times longer than wide, slightly discolorous, hairs of the upper surface 0.6-1.5 mm diam, on the lower surface O. 4-1.5 mm diam., base cuneate to rounded, symmetric or asymmetric, 3-nerved, margins entire near base, becoming serrate near apex, apex acute to rounded. Inflorescence axillary, umbellate, c. 15 mm long, few flowered, extra-floral Fig. 19. Holotype of Helicteres tenuipila. nectaries present; bracts c. 4 mm long, linear; pedicels c. 1 mm long; dichasia 2-flowered. Calyx cupular to urceolate, ellipsoid or tubular, c. 6 mm long, basal nectary incomplete, margin oblique; lobes triangular, c. 1 mm long. Corolla weakly 2-lipped. Petals mauve-pink, the upper pair paler than lower, slightly dimorphic, 7-9 mm long, long clawed, limb cuneate, minutely hairy, calluscd near base, apex truncate to obliquely truncate; lower 3 slightly longer than upper pair, claw with 0-2 small appendages, callus woolly hairy; upper pair with 2 prominent appendages on claw, callus glabrous, smooth. Androgynophore c. 3.5 mm long, straight, shorter than calyx; stamens 10, shortly connate at base, c. 0.8 mm long; anthers transverse; staminodes 5, spathulate-acute, c. 0.7 mm long. Style 0.8-1.5 mm long; stigma of 5 pin-like lobes, terete, sometimes slightly divergent. Capsule globular to ellipsoid, 12-20 mm long, 10-13 mm diam., densely covered with stellate-hairy bristles to 5 mm long, styles caducous or persistent; carpels straight, 1-3-seeded. Seeds variously shaped, conoidal to depressed cylindric or fusiform, often laterally compressed, 1.5-3.5 mm long, c. 2.5 mm wide, irregularly rugose, dark brown. Flowers; Nov.-Dee. Fruits: Jan.-May. 51 I. D. Cowie Distribution and ecology. The species is endemic to Litchfield N.P. in the northern N.T., where it is found on the surface and side slopes of the Mt Tolmer Plateau. It grows in Eucalyptus ininiata / E. tetrodonta open forest on sandy soils. Notes. It is separated from H. integrifolia and allied species by the sparser and coarser indumentum, larger leaves which are oblanceolate, more coarsely and irregularly serrate with sparser stellate hairs on both surfaces, only slightly discolorous, and with stipitate stellate hairs common on the calyx. Several collections from the Wingate Mountains - Fish River Conservation Reserve are tentatively placed here 0 Cowie 12604, 12606; Walsh 3671). They differ in having narrower, linear leaves 50-100 mm long, 5-11 mm wide, 6.5-19 times longer than wide and a coarser indumentum of stellate hairs 0.7-2 mm diam. with the branches touching to distinctly overlapping, but the underlying surface is highly visible. No flowers have been seen, although the length of both the remains of the persistent calyx and the androgynophore are consistent with H. tenuipila. This species has been referred to as “ Helicteres D27003 Litchfield” and “. Helicteres sp. Litchfield (D.M.J.S. Bowman 185)" on herbarium specimen labels and unpublished checklists from DNA. Etymology. From the Latin tenuis , thin or fine and pilus, a hair, referring to the sparse indumentum of the leaves ot this species. Notes on other species Helicteres angustifolia L. (not illustrated). The name Helicteres angustifolia L. is applied here in a broad sense and may include a number of both described and undescribed taxa (Masters 1874; Backer & van den Brink 1963; Cristobal 2001; Phengklai 2001; Tang et at. 2007). Overseas and most Australian material share the following common features of a short, close indumentum on the stems and lower surface of leaves; stipules caducous, 3.5-6 mm long, leaves strongly discolorous, lanceolate to narrowly elliptic (sometimes oblong, ovate or oblanceolate), 2-6.3 times longer than wide, usually glabrous on the upper surface and hoary below, margins entire, inflorescences short, axillary, 2-11-flowered; cincinni 2-flowered, extra¬ floral nectaries present; calyx 4-8 mm long, weakly 2-lipped, sub-cylindrical; petals 5-11 mm long, weakly dimorphic, mauve or purplish, the lower three with a woolly hairy callus at base of limb; the upper pair with a smooth glabrous area at base of limb, androgynophore 3-7.5 mm long, style apparently with 5 pin-like lobes; capsules ovate to oblong, 7-23 mm long, densely stellate-pilose with bristles absent or up to 4 mm long; and carpels straight, 2-7-seeded. Stellate hairs of the stem and calyx are 0.2-0.8 mm diam. The growth habit is an erect open shrub. In Australia and overseas, there is variation in indumentum colour, the size and apex of the leaf, the indumentum of the upper leaf surface and in size, apex and indumentum of the fruit. A number of entities can be recognised. Some entities have leaves with the upper surface densely hairy with stellate, or stellate and simple hairs, or may be united by a similar capsule apex and indumentum, but are otherwise not readily distinguished by other characters. Much, but not all, of the variation in individual characters seen in Australian material falls within the range observed in overseas material. Some Australian entities have a localised geographic distribution while others are geographically more scattered, morphologically more variable and scarcely distinguishable from some Malesian material. The name Corchorus allenii F.Muell., described by Mueller (1892) from a specimen collected in the Prince Regent River area by Bradshaw and Allen, represents a species of Helicteres and is likely to apply to one of the above entities. Further examination of these entities is needed and especially in relation to Malesian material from the Java to New Guinea area where relatively few specimens have been seen. Helicteres hirsuta L. (Fig. 20). Australian (i.e. N.T.) material of Helicteres hirsuta may be worthy of formal recognition as a distinct taxon at some level but further examination of material from India and Lesser Sunda Islands is required. Compared to material from China, Vietnam, Laos, Thailand, Peninsular Malaysia, Borneo, Sumatra, Philippines and Java (largely at SING), N.T. material mostly has a hyaline to stramineous indumentum rather than a hyaline to stramineous or apparently ferruginous one, the leaves arc often less densely hairy below, the calyx and corolla are mostly shorter, the androgynophore is almost always shorter (5-12 mm long as opposed to 12-19 mm long) and leaf shape and petiole length are not as variable (although included within the range of variation). However, the apparently darker indumentum may well be an artefact of storage conditions as considerable darkening of specimens within a few years of collection has been observed by the author where specimens are subject to un-airconditioned storage under tropical conditions. Most of the Malesian material seen was collected prc-1940. Also, specimens of Fig. 20. Helicteres hirsuta : A, flowers; B, leaf. (/. D. Cowie 10654). 52 Helicteres (Malvaceae) from the Northern Territory similar age from China, Vietnam and Philippines at both AD and LUND had only white to stramineous indumentum. Some Philippine material (e.g. Borromeo s.n. Aug 1915 (SING), Acebicio 35463 (SING); Merrill 506, 547 (SING); Ramos 1437 (SING), s.n. Nov. 1909 (LD)) has an indumentum density on leaves and stems close to N.T. specimens, with a shorter calyx down to 10 mm long (7-11 mm long in Australian material) and petals down to 15 mm long (11-18 mm long in Australian material). Also, Chow 78282 (at AD) from China has leaf size and indumentum very similar to Australian material, but with a longer calyx and corolla. No material from the islands east of Java has been seen and it is not clear if the taxon occurs in the Lesser Sunda Islands. Further study of any material from these areas is needed to determine if Australian material is morphologically distinct. KEY TO SPECIES OF HELICTERES IN THE NORTHERN TERRITORY 1 Leaves linear, less than 8 mm wide (Fig 7P-S). . H. sphaerotheca 1: Leaves lanceolate, ovate, elliptic, orbicular or obovate, more than 8 mm wide .2 2 Androgynophore 8 mm or longer; all petals lacking an area of matted hairs at base of limb.3 2: Androgynophore 7 mm or shorter; lower 3 (abaxial) petals with a distinct area of matted hairs at base of limb.5 3 Leaf margins entire, rarely toothed in upper half (Figs 2, 3A-C). H. cana 3: Leaf margins toothed throughout.4 4 Calyx more than 15 mm long; carpels twisted spirally; inflorescence cymose (Fig. 1). H. isora 4: Calyx less than 15 mm long; carpels straight; inflorescence racemose (Fig. 20). H. hirsuta 5 Upper surface of leaves glabrous or nearly so or simple hairs present, with or without stellate hairs.6 5: Upper surface of leaves stellate hairy, sometimes sparsely so, simple hairs absent.8 6 Leaf margins entire; erect shrub; leaves white below . H. angustifolia sens. lat. 6: Leaf margins serrate; plant prostrate; stems long and trailing in mature plants.7 7 Petals 10-16mmlong;inflorescence30-140mmlong; extra-floral nectaries present (Fig. 9)... H. flagellar is 7: Petals 6-9.5 mm long; inflorescence to 40 mm long; extra-floral nectaries absent (Fig. 15). . H. procumbens 8 Plants prostrate, stems long and trailing in mature plants.9 8: Plants ascending to erect when well developed ... 10 9 Inflorescence elongate, interrupted; usually more than 30 mm long (Fig. 7J-N). H. serpens 9: Inflorescence contracted, often congested; usually less than 30 mm long (Figs 6, 7F-I). H. danvinensis 10 Leaf indumentum coarse, hairs mostly more than 1.3 mm diam.; petals 11 mm or more long; stipules more than 6 mm long (Figs 7A-E, 13). . H. macrothrix 10: Leaf indumentum fine to medium, hairs mostly less than 1.3 mm diam.; petals mostly 10 mm or less long; stipules 6 mm or less long (rarely longer in H. tenuipila ).11 11 Leaves with sparse hairs on both surfaces, the lower surface green (Figs 3H-K, 18). H. tenuipila 11: Leaf with the lower or both surfaces densely hairy and obscured, the lower surface often white or grey-green below.12 12 Leaf hairs of two size classes; calyx with bristles and stellate hairs; few-stemmed perennial shrubs to 2 m tall (Figs 3D-G, 11) . H. kombolgiana 12: Leaf hairs of one size class; calyx with stellate hairs only; multi-stemmed subshrubs to 0.5 m tall resprouting each year from a woody rootstock (Fig. 10). IL. integrifolia ACKNOWLEDGEMENTS Greg Leach generously obtained photos of many east Asian and Australian type specimens and literature from K, L, and BM during his time as Australian Botanical Liaison Officer at K. Gordon Guymer and Philip Short reviewed the paper and their constructive comments resulted in considerable improvements. Richard Willan saw it through the editorial process. David Halford kindly drew the name Corchus allenii F.Muell. to my attention. Raelee Kerrigan, Chris Mangion and Andrea Hope accompanied me on field trips specifically to collect and assess the conservation status of Helicteres. The curators of AD, CANB, MEL, NSW and PERTH provided loans of specimens. At BRI and SING, the Curator and Keeper, respectively, provided access to collections, allowed use of facilities and made me welcome. Russell Barrett provided input on W.A. entities and records. Emma Short checked the Latin diagnoses. Clyde Dunlop provided guidance in the early stages of the project and passed on much knowledge both on various field trips and in the herbarium. The help of all these people is greatly appreciated. 53 I. D. Cowie REFERENCES Angiosperm Phylogeny Group. 1998. 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Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium, http://sweetgum.nybg.org/ih/ W.A. Herbarium 2011. FloraBase. http://florabase.calm.wa.gov.au/ [Accessed 29 June 2011], Wheeler, J.R. 1992. Sterculiaceae. Pp. 182-196. In: Wheeler, J.R. (ed.) Flora of the Kimberley region. Department of Conservation & Land Management: Como. Winkler, H. 1906. Beitrage zur morphologic und biologic tropischer bluten und friichtc. Botanische Jahrbiicher fur Systematick, Pflanzengeschichte undPflanzengeographie 38: 233-271. Accepted 8 November 2011 54 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 21: 55-84 Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of northern Australia. Family Halichondriidae BELINDA ALVAREZ' and JOHN N. A. HOOPER 2 ‘Museum and Art Gallery of the Northern Territory. GPO Box 4646, Darwin NT 0801, AUSTRALIA belinda.glasby@nt.gov.au 2 Queensland Museum. PO Box 3300, South Brisbane QLD 4101, AUSTRALIA john.hooper@qm.cild.gov. an ABSTRACT Fifteen species in six genera of the family Halichondriidae, including two new species, Halichondria ( Halichondria) carotenoiclea sp. nov. and Halichondria ( Halichondria) microbiana sp. nov., are recorded for northern Australia as part of a revision of the order Halichondrida (Porifera: Demospongiae) in this region. Descriptions and discussion of those species are presented here. Eight new combinations within the family Halichondriidae arc here established, i.e. Amorphinopsis fenestrata (Ridley, 1884, as Leucophloeus), Amorphinopsis maculosa (Pulitzer-Finali, 1996, as Topsentia ), Axinyssa bergquistae (Hooper et ai. 1997, as Halichondria ), Axinyssa mertoni (Hcntschel, 1912, as Ciocalypta ), Axinyssa gracilis (Hentschel, 1912, as Ciocalypta rutila gracilis ), Axinyssa terpnis (De Laubenfcls, 1954, as Phycopsis ), Ciocalypta vansoesti (Hooper et ai, 1997, as Halichondria) and Topsentia ridleyi (Hooper et ai. 1997 as Halichondria) and one species is relocated into the family Dictyonellidae, i.e, Stylissa vernonensis (Hooper et ai, 1997, as Hymeniacidon). A lectotypc is designated for Ciocalypta stalagmites Hentschel, 1912. Keywords: sponge, Porifera, Halichondrida, Halichondriidae, northern Australia, new species, taxonomy. INTRODUCTION The family Halichondriidae was revised by Erpenbeck & Van Soest (2002). Its definition is based entirely on a few skeletal characters, i.e. presence of an ectosome with specialised skeleton, a disorganised choanosomal skeleton, dimensions of oxeas and styles, absence of microscleres. Other morphological characters (c.g. general shape of megascleres, presence or size categories among the megascleres, spicule density and its relation to consistency, orientation of spicules at the cctosomal level) arc used in combination with the diagnostic characters to separate species and genera within this family. All these characteristics are very simplistic, and often displayed as a gradient of variation which makes separation of species very subjective as reflected by the large number of synonyms (see Van Soest et al. 2008). The family includes 14 genera and at least 296 valid species (Van Soest et al. 2008); however, the status and generic allocation of many of the species listed on this database on the Internet needs to be revised and validated against the current concept of the genera. The Australian Faunal Directory (Hooper 2005) currently lists a total of 42 species of Halichondriidae. Twenty species were reported and described for the Beagle Gulf (northern Australia) by Hooper et al. (1997). No other revision of species of Halichondriidae in Australia or adjacent areas is presently available. The present paper represents the third part of a revision of the order Halichondrida in the northern Australian region and includes the family Halichondriidae. Alvarez & Hooper (2009.2010) provided details and presented an introduction to the revision of the order and the families Axinellidae and Dictyonellidae. MATERIALS AND METHODS This revision includes material of the family Halichondriidae recorded for the tropical northern Australian waters of the Northern Territory and Queensland coast (from Admiralty Gulf in the west to Torres Strait in the east, approx, between the 125° E and 142° E). Complete locality and collection data for non-type voucher material deposited at the Queensland Museum and the Museum and Art Gallery of the Northern Territory are available in Appendix 1. The distribution of species is given according to the marine provinces defined by Spalding et al. (2007). Spicule measurements arc in micrometres and are based on 25 spicules (unless indicated in square brackets) and denoted as range (and mean ± 1 S.E.) of spicule length and width. All other methods as discussed in Alvarez & Hooper (2009.2010) . 55 B. Alvarez and J. N. A. Hooper Fig. 1. A, Amorphinopsis excavans , specimen in situ at South Shell I., Darwin Harbour, N.T; B, Amorphinopsisfenestrata, specimen in situ at Weed Reef, Darwin Harbour, N.T. C, Amorphinopsis maculosa: Specimen (QM G313577) dredged at Groote Eylandt, Gull of Carpentaria, Queensland; D, Axinyssa bergquistae, specimen (NTM Z. 5976) in situ at Stevens Rock, Darwin Harbour, N.T.; E. Axinyssa mertoni , specimen in situ at South Shell I.. Darwin Harbour, N.T.; F, Ciocalypta heterostyla , specimen in situ at Channel Rock, Darwin Harbour, N.T.; G.H, C. stalagmites, specimens with different colouration in situ at Channel Rock and Stevens Rock, respectively; I. C. vansoesti, specimen in situ at Channel Rock, Darwin, N.T. Photographs: A,B - M. Browne; D,G-1 - H. Nuguyen, H,E,F - B. Alvarez. 56 Halichondriidae from northern Australia ABBREVIATIONS Abbreviations used in the manuscript are: AIMS, Australian Institute of Marine Sciences, Townsville; BMNH, Natural History Museum, London (formerly British Museum (Natural History)); CRRF, Coral Reef Research Foundation, Palau; GBR, Great Barrier Reef, MAGNT, Museum and Art Gallery Northern Territory; MNHN, Musee National d’Histoire Naturelle, Paris, France; MSNG, Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy; MAGNT/NTM, Museum and Art Gallery Northern Territory (formerly Northern Territory Museum), Darwin, Northern Territory, Australia; NHMB - Naturhistorisches Museum, Basel, Switzerland; SMF, Senckenberg Research Institute and Natural History Museum, Frankfurt; QLD, Queensland, Australia; QM, Queensland Museum, Brisbane; WA, Western Australia; ZMA, Zoologisch Museum, University of Amsterdam, ZMB, Museum fur Naturkunde and der Universitat Humboldt zu Berlin, Berlin, Germany. Numbers prefixed with Q666C, OCDN, 0M9H are the cross-reference sample number collected for the United States National Cancer Institute, under the ‘Collection of shallow-water organisms’ programme, by the Australian Institute of Marine Sciences, CRRF and MAGNT (subcontracted through CRRF) respectively. TAXONOMY Family Halichondriidae Gray, 1867 Genus Amorphinopsis Carter, 1887 Gender feminine. Type species, by monotypy, Amorphinopsis excavans Carter, 1887. Recent, Indian Ocean. Amorphinopsis excavans Carter, 1887 (Figs 1A, 2) Amorphinopsis excavans Carter, 1887: 77; Hooper & Wiedenmayer 1994:205; Hooper etal. 1997: 25; Erpenbeck & Van Soest 2002: 791; Lim, de Voogd & Tan 2008: 115. Amorphinopsis sacciformis- Hooper et al. 1997: 27. Material examined. Darwin Harbour: NTM Z.2215, Z.4093, Z.4125 (OCDN -8016-W), Z.5213 (0M9H2184-Q); Z.5222 (0M9H2251-O), Z.5736. Vernon Is: G303658. Description Shape (Fig. 1A). Thinly to thickly encrusting (up to 50 mm thick), massive to lobate, or developing short projections and small lumps, generally growing in patches and following substrate, semi-buried in substrate. Colour. Olive green, yellow inside. Oscula. Round to ovate, inconspicuous, 10 mm diameter. Surface. Hispid, bumpy. Skeleton. Ectosomal skeleton (Fig. 2A) thin, detachable tangential layer, composed by a disorganised criss-cross reticulation of paucispicular-multispicular tracts of oxeas, up to 100 pm thick, with small styles tangentially to paratangentially oriented, sometimes in disorganised tufts. Choanosomal skeleton (Fig. 2B) halichondroid, with large oxeas oriented in all directions, sometimes grouped in directionless multispicular tracts; slightly cavernous at subectosomal area with short multispicular tracts supporting the ectosomal skeleton. Spicules (Fig 2C). Oxeas, hastate, in a large range of sizes, 213.4-945.3 pm (598±221) x 5.9-25.1 pm (16.3±5.5). Smaller ectosomal styles 140.8-264.9 pm (193.8±41.3) x 4.1-7.5 pm (5.5±0.9). Remarks. The material examined here agrees with the description of the syntype by Erpenbeck & Van Soest (2002: 790). The specimen described under Amorphinopsis saccifonnis by Hooper et al. ( 1997) is better allocated to A. excavans. It does not agree with the syntype of Ciocalypta sacciformis Thiele which, as mentioned below (sec under remarks of A. carpentariensis), should be interpreted as Halichondria. The material described by Flooper et al. (1997) is a thin crust covering a bivalve shell with skeleton similar to the rest of the material assigned here to A. excavans. Species considered synonyms of Amorphinopsis excavans by Hooper & Wiedenmayer 1994, following Burton (1959), were excluded by Flooper et al. (1997) and Erpenbeck & Van Soest (2002). Distribution. The species was recorded originally from the Mergui Archipelago (Andaman province). Records from Singapore and northern Australia extend the distribution of the species to the Sunda and Sahul Shelf provinces. The northern Australian and Singaporean populations seem to be common at the intertidal region associated with piers and wharfs. The species is found also subtidally between 9 and 16 m. Amorphinopsis fenestrata (Ridley, 1884) comb. nov. (Figs IB, 3) Leucophloeus fenestratus Ridley, 1884: 464; Dendy 1922: 124; Burton 1928: 127. Leucophloeus fenestratus unnamed variety. - Ridley 1884:464. Suberites oculatus Kieschnick, 1896: 534. Hymeniacidon fenestratus. - Lindgren 1897: 483; Lindgren 1898: 312 [?]. Ciocalypta oculata. - Thiele 1900: 75. Ciocalypta oculata maxima Hentschel, 1912: 428. Axinyssafenestratus. -Van Soest, Diaz& Pomponi 1990: 27; Hooper & Bergquist 1992: 102. Ciocalypta confossa Hooper et al., 1997: 23. Ciocalypta fenestrata .- Hooper et al. 1997: 17. Ciocalypta oscitans Hooper et al., 1997: 20. Amorphinopsis foetida. - Hooper et al. 1997: 28. Material examined. Type material - Leucophloeus fenestratus, Holotype, BMNH 1882.2.23.255, Darwin Harbour, N.T., 16-24 m depth, October 1881, HMS Alert. Leucophloeus fenestratus unnamed variety, BMNH 57 B. Alvarez and J. N. A. Hooper Fig. 2. Amorphinopsis excavans (NTM Z.5213): A, light microphotograph of tangential section of ectosomal skeleton, showing oxeas organised in bundles and small styles, paratangentially oriented in disorganised brushes; B, light microphotograph of section perpendicular to surface, showing choanosomal skeleton; C, diagram of spicules. Scale bars: A,C, 100 pm; B, 500 pm. 1882.2.23.195, Arafura Sea, 64-72 m depth, 18 October 1881, coll. HMS Alert. Suberites oculatus, syntype SMF 680, Ternate, Maluku Sea, coll. Kiikenthal, W, 1894. Ciocalypta oculata maxima, SMF 971, Aru Is, between Meriri and Leer, 6-10 m depth, 31 March 1908, coll. Merton, FI. Ciocalypta oscitans, Holotype, QM G303560, Bynoe Harbour, E Fish Reef, N.T., 12°24.1334’ S, 130°28.16’ E, 17 m depth, 6 October 1993, coll. CCNT Ocean Rescue 2000 Program, dredge. Ciocalypta confossa, NTM Z.3106, Parry Shoals, Arafura Sea, N.T., 11° 12.5167’S, 129°42.07’ E., 20 m depth, 15 August 1987, coll. Mussig, AM and NCI team. Additional specimens - Melville Is: QM G313543. Shoal Bay, Vernon Is, Cape Hotham: QM G303558, G303677, G303541. 3ynoe Harbour: NTM Z.5207 (0M9FI2319-N), Z.5208 (0M9H2543-H). Darwin Harbour: QM G303287, G310170; NTM, Z.2018, Z.4085, Z.4122, Z.4123, Z.5215 (0M9H2137-P), Z.5206 (0M9H2192-Y). Cobourg Peninsula, NTM Z. 1391. Gulf of Carpentaria: QM G314246, G314247, G315205. Description. Shape (Fig. 1B). Massive to subspherical, with tapering, hollow, rudimentary, subconical or volcano-shaped fistules, up to 18 mm long and 30 mm diameter; basal portion buried beneath sediment and fistules protruding through substrate. 58 Halichondriidae from northern Australia Fig. 3. Amorphinops is fen eslrata: A, light microphotograph of tangential section of ectosomal skeleton (NTM Z.5206), showing spicule tracts criss-crossing and forming a reticulation of polygonal meshes; B, light microphotograph of section perpendicular to surface (NTM Z.5215) showing choanosomal skeleton; C. diagram of spicules. Scale bars: A-B, 500 pm; C, 100 pm. Colour. Yellow, brown or pale mauve, alive. Oscula. Large, up to 10 mm in diameter or, grouped on a terminal sieve-plate, on apex of fistules. Texture and consistency. Compressible, harsh, easily torn. Surface. Irregular, rugose, translucid, hispid, marked in some specimens with longitudinal channels. Skeleton. Ectosomal skeleton (Fig. 3A) detachable, supported by subcctosomal multispicular tracts. Formed by multispicular tracts of larger choanosomal styles, up to 3 spicules abreast, lying tangential to surface, directionless or criss-crossing, forming a nearly regular reticulation of polygonal meshes; and, irregular bundles of smaller ectosomal styles arranged mostly paratangential to surface as plumose brushes or tufts. Choanosomal skeleton disorganised, halichondroid criss-cross of both unispicular and multispicular tracts, containing 5-20 spicules abreast, with larger choanosomal styles mainly confined to central region; becoming more wide-meshed, paratangentially oriented and cavernous at subectosomal region. In fistules choanosomal tracts more compressed in central region (Fig. 3B), cavernous towards periphery. Single spicules, including smaller styles scattered through choanosome. Spicules (Fig. 3C, Table I). Choanosomal styles and styloids (thicker in apical third and with basal ends narrower than the centre), slender, straight or slightly curved at centre, fusiform, in a great size range (163-895 x 3.3-17.4 pm). Smaller styles can be transitional to subtylostyles. Relative proportions of styles, styloids and subtylostyles vary among populations. Remarks. The species was originally described by Ridley (1884) under Leucophloeus, a junior synonym of Ciocalypta (Van Soest el al. 1990; Erpenbeck & Van Soest 2002) and related to species of Ciocalypta by Hooper et al. (1997) based on the characteristics of the ectosomal skeleton, and the presence of styles. As described above, the growth form of A morphinopsis fenestrata is characterised by the presence offistule-like projections of several shapes 59 B. Alvarez and J. N. A. Hooper Table 1 . Comparison of spicule dimensions among specimens of Amorphinopsis fenestrata. Specimen Locality Styles BMNH 1882.2.23.255 (Holotype) Darwin Harbour 193.6-611.8pm (390.4±136.4) x 5.3-12.7pm (8.6±2) BMNH 1882.2.23.195 (Ridley’s unnamed variety specimen) Arafura Sea 252.9-895.3pm (617.7±217.3) x 5.9-15.4pm (9.6±2.6) SMF 680 (Syntype of Suberites oculatus) Temate, Indonesia 210.2-761.6pm (43l.lil60) x 4.7-17.4pm (8.1±2.8) SMF 971 (Syntype of Ciocalypta oculata maxima) Aru Is, Indonesia 306.4-892.8pm (563.3±176) x 3.3-11.6pm (6.6±1.8) NTM Z.3106 (Holotype of Ciocalypta confossa) Parry Shoals 217.8-649pm (448.9±149.7) x 6.4-12.6pm (9.2±1.9) QM G303560 (Holotype of Ciocalypta oscitans) Bynoe Harbour 175.l-690.4pm (502.7±155.8) x 5.2—13.3pm (8.7±2.2) NTM Z.5215 Darwin Harbour 178.5-891.1pm (427.1±199.8) x 4-12.3pm (7.7±2.3) NTM Z.5207 Bynoe Harbour 163.5-833.1pm (432.4±214.6) x 3.7-13.7pm (7.3±2.3) QM G314247 Gulf of Carpentaria 218.8-599.3pm (412.2±120.3) x 4.3-11.1pm (7.2±2.2) (e.g. pointy, hollow, pyramidal or volcano-shaped), some with a system of exhalant channels. They project from a semi-buried massive base and protrude through the substrate/sand. The skeletal architecture in these fistulose projections, although similar, is considered here not to be homologous to the one observed in Ciocalypta species, which is characterised by a central spicular axis and extra- axial secondary tracts supporting the ectosomal skeleton (see below under Ciocalypta). Instead, the skeleton of the fistulose projections of this species is formed by multiple axes, oriented longitudinally and radiating towards the surface. Based on these arguments, we suggest that the species is better allocated to the genus Amorphinopsis. It needs to be noted however, that the skeleton of the present species is composed totally by styles with a great size range and lacks of ‘true’ oxeas, a diagnostic characteristic of this genus. In the majority of specimens examined, including the holotype, a great proportion of the styles arc ‘subacerate’ (styloids) similar to those observed in Aaptos (Hadromerida: Suberitidae) where the thickest part of the spicule occurs in the apical third and the basal end is substantially narrow. The relative proportion of styloids varies among the populations examined. Subtylostyle modifications, especially in the smaller styles arc also common. We are unable to confirm whether this type of styles might be considered a derived form of oxea (as stated in the current definition of the genus) and therefore we propose to expand slightly the definition of Amorphinopsis to accept species with spicules differentiated into oxeas and/or styles in a large size range, with smaller ones concentrated at the surface. The examination of additional specimens allowed us to understand the concept of this species better and to conclude that the material from Beagle Gulf described by Hooper et al. (1997) under Ciocalypta oscitans, C. confossa and Amorphinopsisfoetida, and the Indonesian species Suberites oculatus Kieschnick and Ciocalypta oculata maxima Hentschel are all conspecific with A. fenestrata. Amorphinopsissuhacerata (Ridley & Dendy, 1886) from the Philippine Islands is very similar to A. fenestrata and as stated by those authors they share many characteristics including the type of styles but differ in external form and in having larger and thicker styles. Study of local populations and examination of the type material is required to establish whether the two species are conspecific. Distribution. Amorphinopsis fenestrata is common throughout the Northern Territory coast (Sahul Province), from Parry Shoals (NW of Darwin Harbour) to the Gulf of Carpentaria. The records of Lindgren (1897, 1898) from Vietnam and China, of Dendy (1922) from the Indian Ocean and of Burton (1928) from the Malay Archipelago need to be verified. Amorphinopsis maculosa (Pulitzer-Finali, 1996) comb. nov. (Figs 1C, 4) Topsentia maculosa Pulitzer-Finali, 1996: 114. Axinyssa aplysinoides. - Hooper et al. 1997: 4. Not Axinyssa aplysinoides Dendy, 1922: 39. Material examined. Holotype - MSNG 48701, Laing Is., 4° 09’ S, 144° 52’ E, Papua New Guinea, 6 m depth, 23 August 1986. Additional specimens - Gulf of Carpentaria: QM G300854, G301034, G313577, G314255, G314267, G315207, G320819, G320904. Shoal Bay: QM G303561. Description Shape. Thickly encrusting, following substrate with convoluted ridges and short projections. Colour. Light grey or yellow alive; yellow inside beige in ethanol. Oscula. Inconspicuous, of different diameter, flushed and irregularly distributed or aggregated in top of the short projections (Fig. 1C). Surface. Smooth, lumpy, with dermal skin of reticulated appearance. Consistency and texture. Finn but crumbly. 60 i Halichondriidae from northern Australia hig. 4. Amorphinopsis maculosa (QM G313577): A, light microphotograph ot tangential section of ectosomal skeleton showing brushes of small styles with ends projecting through surface; B, light microphotograph ot section perpendicular to surface showing choanosomal skeleton; C, diagram of spicules. Scale bars: A, 100 pm; B, 500 pm; C, 100 pm. Skeleton. Ectosomal skeleton (Fig. 4A) consisting of tangential to paratangential crust, approx. 200-300 pm thick, supported by choanosomal tracts of spicules and with disorganised brushes of small styles with ends projecting through surface, forming a discontinuous palisade spaced roughly at regular distances. Choanosomal skeleton (Fig. 4B) halichondroid, forming oval to round lacunae, 300-1000 pm in diameter, becoming compact towards surface, with very little collagen and abundant spicule content. Spicule tracts long and multispieular, running either towards surface or parallel to surface at the subectosomal area. Spicules (Fig. 4C, Table 2). Oxeas and much less frequent styles in a large size variation, 207-994 x 5-38 pm; small ectosomal styles (and transitional to oxeas), 139-274x 3-8 pm. Remarks. The holotypc of Topsentia maculosa Pulilzer-Finali was examined and is comparable in external morphology and skeletal characteristics to material collected in the Gulf of Carpentaria and Shoal Bay. The species is redescribed using this material and assigned to the genus Amorphinopsis. Table 2. Comparison of spicule dimensions among specimens of Amorphinopsis maculosa. Specimen Locality Oxeas Styles MSNCi 48701 (Holotypc of Topsentia maculosa) Papua New Guinea 240.2-994.6pm (633.7±255.1) x 7.4-37.6pm (21.7±9.9) l38.5-259.2pm (211±35.5) [16] x 3-9.7pm (6.6±1.8) [16] QM G313577 Gulf of Carpentaria 369.l-863.5prn (675.8±124.6) x 7.9-20.3pm (15.2±3.2) 143.7-273.8pm (187.7±35.2) x 3.2-7pm (4.8±1) QM G314255 Gulf of Carpentaria 207.9-819.5pm (560.1 ±166.3) x 4.5-22.1 pm (12.6±4) 147.5-240.4pm (174.2±20.9) x 3-7.8pm (5.6±l) 61 B. Alvarez and J. N. A. Hooper Amorphinopsis maculosa is very similar in habit and skeletal organisation to A. fenestrata (Ridley, 1884) but that species has only styles as megasclcres and lacks the oxeas of variable sizes present in this species (see above). Fistulous projections, as observed in A. fenestrata, are not present in the examined material of A. maculosa. The specimen assigned to Axinyssa aplysinoides (Dendy) by Hooper et. al. (1997) was re-examined and has characteristics (i.e. habit, ectosomal skeleton, oxeas and smaller styloids) that agree with Amorphinopsis maculosa. Amorphinopsis maculosa was also compared to other species of Amorphinopsis recorded for northern Australia and adjacent biogeographical regions - Amorphinopsis excavans (Dendy, 1889) from the Indian Ocean, A. foetida (Dendy, 1889) from the Gulf of Manaar, A. maza (De Laubenfels, 1954), A. oculata (Kieschnick, 1896), and A. sacciformis from Indonesia. Ambrphinopsis excavans has similar habit, skeletal structure and spicule composition, but differs in the organisation of the ectosomal skeleton; in A. excavans it consists of a tangential layer of thick intercrossing tracts of large oxeas, and loose oxeas of all sizes with small styles filling up the spaces (Erpenbeck & Van Soest 2002). The material of A. excavans from northern Australia described above also has a clearly tangential and detachable ectosomal skeleton of large oxeas grouped in bundles and small styles tangentially to paratangentially oriented, sometimes in disorganised tufts. Other characteristics that differentiate /). maculosa from A. excavans are the predominance ot small styles in the ectosome (instead oxeas and styles as reported in most populations of.4. excavans) and the lacunar appearance of the choanosomal skeleton. Amorphinopsis foetida (type specimen BMNFI 1889.1.21.55, examined) is also a massive and slightly lobose species but with a skeletal architecture and spicule geometry different to A. maculosa. The ectosomal skeleton of A. foetida is formed by a halichondroid dermal crust, tangential-paratangetially oriented, 500-900 pm thick, with a mixture of oxeas and small styles projecting through the surface, and with vague tracts and small rounded meshes approx. 100-300 pm. The choanosomal skeleton is halichondroid, cavernous, with large rounded lacunae, 500-700 pm approx., and ill-formed multispicular tracts and fibres irregularly oriented through the skeleton; less compact than the ectosome and bounded by little collagen. The spicules are mixture of oxeas of different thickness and sizes, characteristically curved or sinuous (165.4-720.6 pm x 3.3-16.6 pm). The smaller styles arc curved, and some are transitional to oxeas (154-281.7pm x 2.7-7.9pm). Amorphinopsis maza was re-examined by Erpenbeck & Van Soest (2002) and differs from A. maculosa in habit, skeletal organisation and spicule composition. Suberites oculatus Kieschnick, 1896, accepted as Amorphinopsis oculata (Van Soest et al. 2008) is a synonym of Amorphinopsis fenestrata (see above). Ciocalypta sacciformis Thiele, 1900 (syntypes, SMF 685, 1818, Ternate Maluku Sea, 1894, coll. Kiikenthal W„ examined) was transferred from its original genus to Ciocalapata by De Laubenfels (1936) and related it to species-like Ciocalypta including oxeas and styles. The species was interpreted as Amorphinopsis by Hooper et al. (1997). The revision of the type material indicates that the specimens described by Thiele agree better with the current concept of Halichondria but not with Ciocalypta, Ciocalapata or Amorphinopsis as suggested by previous authors. The syntypes examined are small fragments with a pouch-like shape as described by Thiele. The ectosomal skeleton is a thick tangential to paratangential layer up to 500-800 pm, with single spicules and short tracts criss¬ crossing in a disorganised manner. The choanosomal skeleton is disorganised with bundles of spicules irregularly spaced and running towards the surface where they merge with the ectosomal skeleton. The skeleton is formed entirely by oxeas in a great size range 195-706 pm. The skeleton of SMF 1818 includes styles in low frequency 463-525 pm (491.2±17.4) x 6-16 pm. As admitted by Thiele, the styles might be modifications of oxeas. Distribution. Papua New Guinea (Eastern Coral Triangle Province), Gulf of Carpentaria and outer region of Shoal Bay (Sahul Shelf province) between 6-28 m depth. It is also found in Torres Strait (Northeastern Australian Province) (Alvarez & Hooper unpublished data). Remarks on Amorphinopsis. Amorphinopsis includes approx. 13 valid species (Van Soest et al. 2008) distributed mainly throughout the Indian Ocean but with species also recorded from the Atlantic and Mediterranean oceans. The genus is represented in northern Australia (Sahul Shelf Province) by three species - Amorphinopsis excavans, A. fenestrata comb. nov. and A. maculosa comb. nov. Amorphinopsis foetida was recorded by Hooper et al. (1997) from the Beagle Gulf but it is concluded here that the specimen is better allocated to A. fenestrata. The species originally described as Leucophloeus fenestratus Ridley, 1884 is common in the study area. Its habitat and skeletal characteristics have been interpreted as typical from the genus Ciocalypta but as discussed above, these similarities are considered not homologous and we propose to include the species in Amorphinopsis. The choanosomal skeleton of A. fenestrata is formed entirely by styloids and lacks ‘true’ oxeas as seen in other species of Amorphinopsis and other genera of Halichondriidac. We propose to expand the definition of Amorphinopsis slightly to include species with spicules differentiated into oxeas and/or styles. Genus Axinyssa Lendenfeld, 1897 Gender feminine. Type species, by original designation, Axinyssa topsenti Lendenfeld, 1897: 116. Recent, western Indian Ocean. Axinyssa bergquistae (Hooper et ah, 1997) com. nov. (Figs 1D, 5) Halichondria bergquistae Hooper et al., 1997: 45. 62 Halichondriidae from northern Australia Material examined. Holotype - QM G303351, East Point Bommies, Darwin Harbour, Northern Territory, Australia, 12°24.083’ S, 130°48.14’ E, 10 m depth, 23 September 1993, coll. Hooper, JNA & Hobbs, LJ. Additional specimens - Cartier Island: QM G301059. Bynoe Harbour: Z.5224 (0M9H2375-X); Z. 5901. Darwin Harbour: Z.5976. Gulf of Carpentaria: QM 313572. Description Shape (Fig ID). Massive-lobate, bulbous-digitate; with erect columns or irregular coalescent plates. Individuals approx. 70-100 mm high, 45-200 mm thick. Colour. Purple-mauve, grey-brown; some individuals with lighter tinges. Oscula. Variable in size (3-7mm diameter), conspicuous, discrete, with raised white and opaque membranous lips (Fig. 1D), irregularly distributed. Surface. Smooth to microconulose at base with shallow and meandering channels covered by a translucent membrane; becoming distinctively conulose at erect columns or digits. Conules up to 2 mm long organised in longitudinal rows along erect columns and digits, with brushes of larger choanosomal spicules protruding through surface. Texture and consistency. Hispid due to projection of spicules through surface, firm and incompressible. Skeleton (Fig. 5A,B). Ectosome without specialised skeleton, with lightly coloured collagenous skin. Choanosomal skeleton halichondroid, with high spicule density in deeper regions; becoming more organised at subcctosomal region, with multispicular spicule tracts, 50-200 pm running longitudinally and ascending towards surface, becoming more radial and plumose near periphery; ending in disorganised brushes that project through ectosome. Spicules (Table 3, Fig. 5C). Mixture of oxeas of variable thickness and length (354-948 x 4—27.8 pm), slightly bent Fig. 5. Axinyssa bei-gquislae: A, light microphotograph of perpendicular section (NTM Z. 5976) showing choanosomal spicule tracts projecting through surface; B, light microphotograph of perpendicular section through surface (NTM Z.5224), showing choanosomal skeleton with longitudinal plumose tracts; C, diagram of spicules. Scale bars: A-B, 500 pm; C, 100 pm. 63 B. Alvarez and J. N. A. Hooper and sometimes slightly sinuous. Styloid modifications common. Remarks. The species was initially described under Halichondrici. Examination of additional material considered conspecific with the specimen described by Hooper et al. (1997) allowed us to conclude that the species is better allocated in Axinyssa. The plumose conules observed in some areas of the surface of A. beigquistae resembles Axinyssa mertoni (described below), but that species has a more lax skeleton with less spicular density and more collagen. It differs also in general shape, colour and the oscula morphology. Axinyssa bergquistae is comparable tod. valida (Thiele, 1899: 12) [holotype NHMB 13, examined] in external morphology, skeletal organisation and size of spicules [(290.3-859.3 pm (599±201.8) x 9.7-34.8 pm (20.6±8.3)] and could possibly be conspecific. But both the skeleton and the size of spicules of most Axinyssa species are very similar and the separation of species is difficult and subjective (see below under remarks on the genus). The examination of more material from Indonesia (Alvarez & De Voogd in progress) will help to determine whether the northern Australian populations belong to the same species. Distribution. Darwin Harbour and Bynoe Harbour (Sahul Shelf Province). Probably present also in the Northeast Australian Shelf Province (Alvarez & Hooper unpublished data). Table 3. Comparison of spicule dimensions among specimens of Axinyssa bergquistae. Specimen Locality Oxeas QM G303351 (Holotype of Halichondria bergquistae) Darwin Harbour 407.3-793.8pm (619.7±82.8) x 10.3-27.8pm (15.5±4.3) NTM Z.5224 Bynoe Harbour 366.3-948.1pm (749.2±135.7) x4.5-33.2pm(21.3±7) QM G313572 Gulf of Carpentaria 353.6-747.3pm (618.3±77.5) x 4-20.9pm (14.7±4.2) Axinyssa mertoni (Hentschel, 1912) com. nov. (Figs IE, 6) Ciocalypta mertoni Hentschel, 1912:424; Burton 1934: 564. Halichondria tyleri. - Hooper & Wiedenmayer 1994: 209. Halichondria mertoni. — Hooper et al. 1997: 52. Pseudaxinyssapity’s De Laubentels, 1954:178; Bergquist 1965 : 175 [?]. Axinyssa pitys. — Hooper & Bergquist 1992; 102. Material examined. Type material - Ciocalypta mertoni, holotype, SMF 1608, Am Is, North of Penambulai; station 10, 8 m depth, 2 April 1908, coll. Merton exp. 1908. Pseudaxinyssa pity’s, holotype, USNM 23103, Caroline Islands, Palau Is, Koror 1., Iwayama Bay, 2 m depth, 1 September 1949, coll. De Laubenfels, M.W. Additional specimens -Darwin Harbour: NTM Z.5221 (0M9H2189-V). Description Shape (Fig. IE, 6A). Massive with conspicuous fistules up to 50 mm long, 1-2 mm thick, projecting from semi- buried basal portion up to 100 mm diameter. Colour. Grey alive. Oscula. On top of fistules, flushed, less than 5 mm diameter. Surface. Regularly conulose with transluscent membrane stretching over conules; marked with choanosomal axes. Skeleton (Fig. 6B). Ectosomal skeleton absent. Choanosomal skeleton plumose to halichondroid with multispicular axes of spicules running longitudinally, nearly parallel and close to each other, anastomosing and diverging towards surface and becoming dendritic, bounded with collagen and ending in disorganised plumose brushes that project through surface. Spongin fibres ill-defined and short, direction-less, embedding spicule tracts. Spicules (Fig. 6C, Table 4). Oxeas, hastate, slightly bent, straight or crooked, 476-1470.2 x 11.2-29.7 pm. Style and strongylote modifications also present. Remarks. Ciocalypta mertoni is conspecific with Pseudaxinyssa pity’s and is better allocated to the genus Fig. 6. Axinyssa mertoni: A, preserved specimen (NTM Z.5221); B, light microphotograph of perpendicular section through surface showing choanosomal skeleton with longitudinal tracts projecting through surface (Holotype, SMF 1608); C, diagram of spicules. Scale bars: A,B. 500 pm; C, 100 pm. 64 Halichondriidae from northern Australia Axinyssa. The type material of both species were examined here, and they are identical in all their characteristics. Ciocalypta mertoni was considered a junior synonym of Halichondria lyleri by Hooper & Wiedenmayer (1994) (following Burton 1959), but this synonym was later rejected by Hooper et al. (1997) who considered it a valid species of Halichondria , but admitted that the lack of an ectosomal skeleton was atypical of that genus. The species agrees well with the concept of Axinyssa , as it lacks an ectosomal skeleton and has a halichondroid to vaguely plumose choanosomal skeleton bounded by relatively high amounts of collagen and formed by oxcas of variable size and common styles and strongyles. The growth form of this species however, is not reported for other species of Axinyssa, thus a slight expansion in the diagnosis of the genus is necessary to accommodate species with fistulose projections. A single specimen found in Darwin Harbour is assigned to this species. It differs from the type material examined only on spicule dimensions. The oxeas of the Darwin specimen are in average longer and thicker, but this might correspond to intraspecific variation within the species. Distribution. Axinyssa mertoni is rare within the study area with only one specimen recorded here. The distribution of the species is extended to the Sahul Shelf and the Tropical Northwestern Pacific provinces. The record of Bergquist (1965) from Palau needs to be verified as there are several sympatric species of Axinyssa occurring in the area (Lori Bell Colin pers. comm.). Remarks on Axynissa. Axinyssa is represented in the area of the Indo-Pacific by several species: A. aculeata Wilson, 1925 (Philippines); A. aplysinoides (Dendy, 1922) (Seychelles); A. oinops (De Laubenfels, 1954) (central West Pacific); A. topsenti Lendenfeld, 1897 (Tanzania); A. variabilis Lindgren, 1897 (Malaysia); and A. valida (Thiele, 1899) (Indonesia). The new combinations established in this revision extend the list by two additional species -A. mertoni (Sahul Shelf and Tropical Northwest Pacific provinces) and A. bergquistae (Sahul Shelf and probably Northeast Australian Shelf). Two additional species from the central Indo-Pacific region are also referred to Axinyssa after examination of type material - Ciocalypta nitila gracilis Hentschel, 1912 (SMF 1566, Aru Is, examined; see under remarks of Ciocalypta), and Phycopsis terpnis De Laubenfels, 1954 (Caroline Is, Central Pacific, USNM 23061, examined). We were not able to revise all the type material of the Indo-Pacific species of Axinyssa thoroughly, therefore it remains inconclusive whether A. mertoni and A. bergquistae might be conspecific with other species recorded from the region. As mentioned above, the skeletal organisation and the size of spicules among Axinyssa species is very similar and separation of species is subjective. The external morphological characteristics seem to be more discrete, but study of individual populations is necessary to determine the actual range of variability present within these species. Table 4. Comparison of spicule dimensions among specimens of Axinyssa mertoni. Specimen Locality Oxcas SMF 1608 Aru Island, Indonesia 515.6-779.6pm (691.4±61) x 11.2-21.7pm (18±2.5) USNM Caroline Is, 514.3-871 pm (788.2±74.9) 23103 (holotype of A. pitys) Central West Pacific x 8.6-18.5pm (I5±2.5) NTM Z.522I Darwin Harbour, NT 476-1470.2pm (979.5±285.9) x 14.7-29.7pm (21.6±4.8) Study of different populations ofAv/nyssa through Indonesia (Alvarez & De Voogd unpublished data) is currently in progress and will help to re-define the limits of Axinyssa species. The diagnosis of Axinyssa is here expanded to include species Vk& Axinyssa mertoni with fistulose projections. We note however, that the skeletal organisation of the fistules observed in A. mertoni is considered not to be homologous with the organisation observed in species of Ciocalypta. The fistulose projections of Ciocalypta spp. are transparent with the skeleton formed by a central axis of spicules and extra- axial tracts diverging towards the surface. The fistulose projections of A. mertoni are opaque, tough-cartilagineous, arising from a massive base, and without a central axis of spicules which is diagnostic for Ciocalypta. Axinyssa mertoni shares with the Indian Ocean Ciocalypta digitata (Dendy, 1905) the lack of an ectosomal skeleton, and the presence of fistulose projections; however, the fistules of C. digitata are transparent with a spicular axis from which thick bundles of spicules diverge towards the surface ending in conulcs (Erpcnbeck & Van Socst, 2002). In our view the expansion of the definition of Axinyssa to include species with fistular projections such as A. mertoni does not affect the current position of C. digitata or the definition of the Ciocalypta. The current position of the genus Axinyssa within the family Halichondriidae is debatable as molecular data (Erpcnbeck et al. 2005) indicate that some species currently allocated to this genus arc related to other dictyoncllid genera such as Acanthella and Dictyonella. From a morphological point of view, the lack of an ectosomal skeleton, the presence of abundant collagen in the skeleton and the common occurrence of styles, strongyles and transitional forms, also points to possible relationships with members of Dictyonellidae. These relationships should be further explored to confirm the placement of this genus within the family Halichondriidae. However, taxonomic verification of the species of Axinyssa used in the molecular analyses should also be taken in consideration, particularly given the paucity of morphometric characters in this group and our still rudimentary understanding of character homology. Genus Ciocalypta Bowerbank, 1862. Gender feminine. Type species, by monotypy, Ciocalypta penicillus Bowerbank, 1862. Recent, eastern Atlantic Ocean. 65 B. Alvarez and J. N. A. Hooper Ciocalypta heterostyla Hentschel, 1912 (Figs IF, 7) Ciocalypta heterostyla Hentschel, 1912: 424; Hooper etal. 1997:36. Material examined. Holotype - SMF 1569, Aru Is, N Penanbuli, 8 m depth, 2 April 1908, coll. H. Merton. Additional specimens - Darwin Harbour, NTM Z.5902. Description Shape (Fig. 1F). Fistulose, with semi-buried and massive base. Fistules pointed, projecting perpendicularly from base, 20-30 mm long, less than 10 mm diameter wide, slightly translucent especially at tips. Colour. Light yellow. Oscula. Apical on fistules. Surface. Smooth, microconulose. Ectosomal skeleton (Fig. 7A). Thin layer formed by tangentially oriented pauci- to multispicular tracts of spicules, crossing over and forming a reticulation of triangular meshes; supported by choanosomal tracts. Choanosomal skeleton. Differentiated at fistules, with central column formed by thick multispicular tracts oriented longitudinally and expanding into thick brush at tip of fistule (Fig. 7B). Secondary tracts, 20-100 pm thick, slightly plumose, diverging nearly perpendicularly from central column toward surface, regularly spaced and connected irregularly by unispicular-paucispicular tracts of spicules; becoming thicker and ending in fan-like brushes at the subectosomal area, generally with smaller spicules oriented with their ends towards surface. Skeleton at base Fig. 7. Ciocalypta heterostyla (Holotype SMF 1569): A, light microphotograph of tangential section of ectosomal skeleton, showing tracts of spicules, forming a reticulation of triangular meshes; B, light microphotograph showing choanosomal skeleton at a fistule. C, diagram of spicules. Scale bars: A-B, 500 pm; C, 100 pm. 66 Halichondriidae from northern Australia halichondroid, with directionless multispicular tracts and single spicules in confused reticulation. Spicules (Fig. 7C, Table 5). Mixture of styles in a large range of sizes, straight to slightly curved, some sinuous, some with rounded ends, 184-809 x 4—19 pm. Remarks. Ciocalypta heterostyla was originally described from the Aru Is (Indonesia) and it has not been re- dcscribcd or recorded until this present study. The material from Darwin Harbour agrees closely with the type. Only the spicule dimensions are on average slightly larger in the specimen from Darwin. The species is not very conspicuous due to its cryptic habit. It is apparently rare in the study area with only one specimen collected so far. Hooper &Wiedenmayer (1994) followed Burton (1959) and considered this species a synonym of Ciocalypta tyleri Bowerbank. The two species are certainly similar in external morphology and skeletal organisation, however the skeleton of C. heterostyla is formed exclusively by styles instead of oxeas, and the skeleton within the fistules is much more organised with a regular reticulation. Because of these characters we considered that C. heterostyla is not only a valid species but also it can be easily differentiated from its South African relative. Distribution. Known only from the type locality (Aru Is, Indonesia) and from Darwin Harbour (Sahul Shelf province). It is found between 8-12 m depth. Tabic 5. Comparison of spicule dimensions between specimens of Ciocalypta heterostyla. Specimen Locality Styles SMF Aru Is, Indonesia 199.9-584.1 pm (387.3±137) 1569 x 3.5-14.8pm (8.3±3.2) NTM Darwin Harbour 184-809.2pm (466.5±229.1) Z.5902 x 3.9-18.6pm (9.4±5.5) Ciocalypta stalagmites Hentschel, 1912 (Figs 1G-H, 8) Ciocalypta stalagmites Hentschel, 1912: 426. Halichondria tyleri- Hooper et al. 1997. Material examined. Lectotype - SMF 1567 (here designated, a representative specimen of the species, chosen from six examined specimens deposited at SMF and registered as syntypes with a single accession number), Aru Is, Mimien I., Indonesia, 15 m depth, 9 April 1908, coll. Meron, H. Paralectotypes - SMF 1537, Aru Is, Sungi Manumbai (Kapala Sungi), Indonesia, station 17, Merton Expedition Aru and Kei Is 1907-1908, 20 m depth, 5 May 1908, coll. Merton, H. SMF 1574, Am I., SW Lola I, Indonesia, station 9, Merton Expedition Aru and Kei Is 1907-1908, 8-10 m depth, 1 April 1908, coll. Merton, H. SMF 1595, Am Is, Bambu I., Indonesia, station 11, Merton Expedition Am and Kei Is 1907-1908, 10 m depth, 3 April 1908, coll. Merton, H. SMF 1623, Am Is, SW Mariri and Leer Is, Indonesia, Merton Expedition Am and Kei Is 1907-1908, 6-10 m depth, 31 March 1908, coll. Merton, H. SMF 1627, Am Is, N Penambulai I., Indonesia, Merton Expedition Aru and Kei Is 1907-1908,8 m depth, 2 April 1908, coll. Merton, H. Additional specimens - Parry Shoals, NT, NTM Z.3133. Charles Point, NT, NTM Z.5230 (0M9H2574-P). Bynoc Harbour: NTM Z.5226 (0M9H2511-V). Darwin Harbour: NTM Z.941, Z.5210 (0M9H2571 -M), Z.5229 (0M9H2299-Q), Z.5977. Cobourg Peninsula: NTM Z.592, Z. 1358, Z. 1395, Z.3286. Wessel Is: Z.3920, Z.5218 (0M9H2666-P). Description Shape (Figs IG-H, 8A). Flat cushion-shaped or massive base, buried or semi-buried, strongly attached to substrate, up to 35 mm thick, 4-100 mm diameter, with fistules projecting perpendicularly above surface. Fistules tig. 8. Ciocalypta stalagmites (Lectotype, SMF 1567): A, Preserved lectotype; B, light microphotograph showing choanosonial skeleton at fistule; C, light microphotograph showing choanosomal skeleton at base; D, diagram of spicules. Scale bars: A, 2 cm; B-C, 500 pm; D, 100 pm. 67 B. Alvarez and J. N. A. Hooper sharply pointed, globular or flattened, mammiform, tubular, rounded, 5-125 mm long, laterally fused sometimes or branching at tips. Colour. Base generally pale-beige. Fistules, transparent, mauve-brown-purple, greenish-yellow. Internally, beige. Oscula. Generally at apex of fistules or tubes but also observed irregularly distributed along fistules, with membranous rims. Consistency and Texture. Compressible-spongy, easily tom. Surface. At fistules, smooth to slightly conulose, marked with longitudinal rows of minute conules. Choanosomal tracts of spicules, visible through the ectosome of translucent specimens. At base smooth, opaque, rough, spiculous. Skeleton. Ectosomal skeleton, tangential layer of variable thickness (5-300 pm) easily peeled in some specimens, formed by a dense mass of smaller oxeas and supported by choanosomal skeleton. Choanosomal skeleton (Fig. 8C) at base, halichondroid, densely spiculous, with mixture of large and smaller oxeas and no distinct tracts of spicules. In fistules (Fig. 8B), becoming distinctive and organised, with central column of large spicules oriented longitudinally and smaller spicules criss-crossing. Multispicular tracts generally formed by medium-size spicules radiating from central column towards surface, connected by shorter tracts and single spicules, forming a regular reticulation. Large subectosomal spaces (up to 5 mm diameter) in some areas. Spicules (Fig. 8D, Table 6). Two size classes of oxeas: I, smaller, thinner, fusiform, straight or slightly flexuous (147-321 x 4-10 pm); II, large, thick and slightly curved, fusiform, occasionally crooked-sinuous (378-886 xl 1-40 pm), and styles in equivalent size categories are common. Remarks. The growth form of Ciocalypta stalagmites is remarkably variable. Fortunately this variation is also well represented in the type material. The external colouration is also variable and is possibly related to the presence of different cyanobacterial associations. The skeletal organisation, composition and size of spicules, however, are consistent and very similar among specimens with different habits and colouration. A very consistent characteristic through all the populations we examined is that the oxeas are differentiated in size categories not only by their length but also by their thickness. Proper statistic and morphometric analyses could be employed to demonstrate whether this is diagnostic to separate C. stalagmites from other Ciocalypta species. Ciocalypta stalagmites is very similar to C. tyleri Bowerbank, 1873 from South Africa, and it was interpreted as such by Hooper et al. (1997) but allocated to the genus Halichondria (following Van Soest et al. 1990; Van Soest 1991). Both species are now referred to Ciocalypta under the revised concept of Halichondriidae (Erpenbeck & Van Soest 2002). The two species are similar in external morphology and skeletal organisation, however, the skeleton of C. stalagmites seems to be less organised. Despite similarities between these two species, it is unlikely that the Indonesian and northern Australian populations are conspccific with their South African relatives, thus we propose to reserve C. stalagmites lor those populations inhabiting the Sahul Province and adjacent areas. Future independent evidence might demonstrate whether C. tyleri and C. stalagmites belong to a complex of cryptic species that cannot be easily separated using traditional morphological characters, or are instead conspccific and genuinely a widely distributed species. Ciocalypta stalagmites is also very similar to C. vansoesti (Hooper et al., 1997). comb. nov. The two species have similar growth form and skeletal characteristics, but they can be differentiated from each other by some distinctive characteristics (see below). Distribution. Ciocalypta stalagmites is very common throughout the northern Australian localities ot the Sahul Shelf province and its distribution extends to adjacent provinces including the Northeast Australian Shelf, Papua New Guinea (Alvarez & Hooper unpublished), and Indonesia (Alvarez & De Voogd unpublished). It is found from the intertidal zone to 40 m. Ciocalypta vansoesti (Hooper et al., 1997) comb. nov. (Figs II, 9) Halichondria vansoesti Flooper et al., 1997: 37. Material examined. As listed by Hooper et al. (1997). Additional specimens - Darwin Harbour: Z.5904, Z.5217 (0M9H2568-J), Z.5903, Z.5978. Bynoe Harbour: Z.5212 (0M9H2498-I).Gulf of Carpentaria: QM G303524. Remarks. This species was well described by Hooper et al. (1997), but allocated to the genus Halichondria (following Van Soest et al. 1990; Van Soest 1991). It is Table 6. Comparison of spicule dimensions between specimens of Ciocalypta stalagmites. Specimen Locality Oxea type I Oxea type II SMF 1567 (Lectotype) Ant Is, Indonesia 167.8-240.2pm (200.8±17.4) x 4.4-10pm (7.9±1.3) 377.7-837pm (626.2±102.8) x 12.3-37.1pm (18.7±4.7) NTM Z.5212 Bynoe Harbour 165.3-320.6pm (238.2±34) x 5.5-10.3pm (7.6±1.2) 411.4-659.5pm (524.9±56.8) x 11.1-26.3 pm (18.9±4.1) NTM Z.5210 Darwin Harbour 188.3-258.7pm (214± 19.1) x 5.6-9.2pm (7.4±0.9) 440.1-662.8pm (601.8±53.9) x 11.3-38.5pm (21.9±7.7) NTM Z.5218 Wessel Is 147.2-202.7pm (169.1± 12.8) x 5.3-9.4pm (6.7±0.9) 437.7-886.5pm (588.1±135.1) x 8.7-39.5pm (21,7±7.9) 68 Halichondriidae from northern Australia Fig. 9. Ciocalypta vansoesti: A, preserved paratype specimen (QM G303450); B, light microphotograph showing choanosomal skeleton at fistule. Scale bars: A, 20 mm; B, 500 pm. now referred to Ciocalypta under the revised concept of Halichondriidae (EqDenbeck & Van Soest 2002). As mentioned by Hooper et al. (1997) and above, Ciocalypta vansoesti is closely related to C. stalagmites, both with similar growth form, skeletal organisation and type of spicules. Both species have fistules projecting from a buried-semiburied mass, but in the case of C. vansoesti the fistules are transluscent-white with the surface regularly conulose and subectosomai tracts and central column visible beneath (Figs 11,9A). However, the skeleton in the fistules, when compared to C. stalagmites, is denser and slightly disorganised; the central column is not as condensed and the extra axial reticulation is vague (Fig. 9B, Floopcr <?/«/. 1997, figs. 22a, 23a). The dimensions of the oxeas arc similar and overlap with those of C. stalagmites, but in C. vansoesti there is not a clear difference in size categories, with the two classes of spicules overlapping both in length and in width (Table 7). Ciocalypta vansoesti is also similar in some of the field characteristics to Axinyssa mertoni, indicating once again that fistule-like growth forms are common among halichondrids and so are not useful to differentiate species unless they are used in combination with other skeletal characters. Distribution. Ciocalypta vansoesti is common in Darwin Harbour and Bynoe Harbour. It is also recorded from Cobourg Peninsula and the Gulf of Carpentaria. It is found intertidally and subtidally to 40 m depth. Remarks on Ciocalypta. Burton (1959) considered many species of halichondriid genera with a fistulose habit conspecific with Ciocalypta penicillus (type species of Ciocalypta ). These synonyms were not properly substantiated and some of them have been rejected by Hooper et al. (1997) and the present revision, i.e. Ciocalypta heterostyla, C. stalagmites, C. tyleri, C. oeulata maxima (referred here to Amorphinopsisfenestrata), and C. mertoni (referred here to Axinyssa mertoni). Additional species of Ciocalypta recorded from the Sahul Shelf Province and adjacent areas besides the three revised here (i.e. C. heterostyla. C. stalagmites and C. vansoesti) are C. rutila gracilis Hentschel, 1912 (see below), C. digitata (Dendy, 1905, as Collocalypta), C. melichlora Sollas, 1902, C. rutila Sollas, 1902, and C. simplex Thiele, 1900: 76. One of the syntypes of Ciocalypta rutila gracilis Hentschel, 1912 (SMF 1566, examined) belongs in Axinyssa. Both the external morphology (based on Hentschel’s description) and the arrangement of skeleton agree with the concept of that genus. The skeleton of the material examined is formed by two classes of oxeas: straight 480.8-706.4 pm (601.6±76.7)x 9-24 pm (17±3.1) and vermicular, crooked sinuous, relatively thinner, and often bent up to a 90 degree angle, 193-600.7 pm (406.7±110.6) x 5.3-16 pm (10.6±3) [18]; less often styles are also present. Specimens recorded for Northeast Australian Shelf and Indonesia correspond with the type examined (Alvarez & Hooper unpublished data; Alvarez & Dc Voogd unpublished data) and will be redcscribed under the name of Axinyssa gracilis in forthcoming publications. Ciocalypta digitata (Dendy, 1905) resembles Ciocalypta stalagmites and C. vansoesti in habit but differs in skeletal Tabic 7. Comparison of spicule dimensions between specimens of Ciocalypta vansoesti. Specimen Locality Oxea type 1 Oxea type 11 NTM Z.2648 Darwin Harbour, East Point 185.3—459.7pim (320±60.9) x 6.4-9.5pm (8±0.9) 475.2-616.4pm (546.2±38.3) x 14-21.6pm (I7.9±2.3) QM G303450 Bynoe Harbour 194.1-361.7pm (275.3±45.6) x 2.1-9.8pm (6.8±1.8) 382-662.1pm (536.1±78.2) x 6.8-30.8ptn (16.8±6) QM G303524 Gulf of Carpentaria 237.5-429.2pm (3I4.9±56.5) x 6.4-11.9pm (8.4±1.3) 432.8-677.3pm (541,2±58.7) x 8.2-31.9pm (17.5±4.8) 69 B. Alvarez and J. N. A. Hooper organisation and composition and size of spicules. This species lacks an ectosomal skeleton, but the fistular processes have spicular axis and extra-axial tracts as in other species of Ciocalypta. Genus Halichondria Fleming, 1828 Gender feminine. Type species, by subsequent designation (of Bowerbank 1862), Spongiapanicea Pallas, 1766. Recent, East England. Subgenus Halichondria (Halichondria ) Fleming, 1828 Halichondria (Halichondria) carotenoidea sp. nov. (Figs 10A,B, 11) Halichondria stalagmites. - Hooper et al. 1997: 43 [in part, not Z. 131, Z.1097, Z.1991 = Halichondria (Halichondria) microbiana sp. nov.] Phakellia sp. 614. - Hooper et al. 1992. Axinella sp. 244. - Hooper et al. 1992. Material examined. Holotype - NTM Z.5909 , off Dudley Point, Fannie Bay, Darwin Harbour, N.T., 12°24.96’ S, 130°48.83’ E, 4-7 m depth, 4 June 2002, coll. Alvarez, B and party. Paratypes- NTM Z.4451 (0M9H2036-G), Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, N.T., 12°29.103’S, 130°47.1irE, 8-14 m depth, 7 May 2002, coll. Alvarez, B and party. NTM Z.5225 (0M9H2462-S), Spencer Point, Indian 1., Bynoe Harbour, N.T., 12°35.351 ’ S, 130°31.454’ E, 6-8 m depth, 11 June 2003, coll. Alvarez, B and party. Additional specimens - N Adele I., Collier Bay, NW Shelf: Z.712. Parry Shoals, NT: NTM Z.3147. Darwin Harbour NT: NTM Z.194, Z.919, Z.934, Z.986, Z.1979, Z.2086, Z.2245, Z.2651, Z.2697. Description Shape (Figs 10 A,B). Fan shaped, digito-palmate, multiplanar, arborescent, erect, stalked, bifurcated at base, with complex branching; branches or digits flattened generally with pointed tips but round tips also common; specimens reaching up to 350 mm high, 300 mm wide. Colour. Orange. Blue-grey, or beige in alcohol Consistency and texture. Soft, easily tom, rubbery. Oscula. Regularly distributed along lateral side of branches (Fig 10B) less than 5 mm in diameter, generally in rows, with membranous rims slightly elevated. Some specimens with subectosomal thin canals ending in oscula. Surface. Smooth, wrinkled, evenly covered with micro- conules when exposed to air. Marked with subectosomal drainage canals in some specimens. Skeleton (Fig 11 A, B). Ectosomal skeleton formed by 100-200 pm thick continuous layer, supported by choanosomal skeleton and packed mainly with smaller category of oxeas, in short, ill-defined and criss-crossing bundles, oriented tangentially- paratangentially, protruding occasionally through surface in disorganised manner (Fig. 11 A). Choanosomal skeleton, halichondroid to plumose, formed by ill-defined bundles of larger spicules, criss-crossing in all directions in the inner region of choanosome. Skeleton becomes more organised at subectosomal region, with a vague reticulation of ascending tracts connected by short bundles and single spicules. Near surface, spicule tracts become more defined and even slightly plumose, supporting ectosomal skeleton and occasionally protruding through surface (Fig. 11B). Skeleton of some specimens is obscured by granular pigments and cyanobacteria when examined under light microscope. Spicules (Fig. 11C, Table 8). Mixture of oxeas not separable into size categories 112-389 x 2-12 pm, hastate, pointed, straight or occasionally slightly bent in middle. Remarks. Halichondria carotenoidea is diagnosed by a unique combination of features (i.e. growth form, organisation of the skeleton, and the size and composition ol spicules) not found in other species of Halichondria revised here or reported for adjacent areas (see Tabic 9). The skeleton and spicule composition of all the material examined is almost identical to Halichondria (H.) microbiana sp. nov. (see below). Both species are sympatric and include a high density of cyanobacteria in the choanosome. The two species clearly differ in shape, which is branching to arborescent in //.(//.) carotenoidea and massive to cushion shape in H. (//.) microbiana. Both species can be further distinguished by the size ot the oxeas, which are separated into class categories in H.(H.) microbiana but not in //.(//.) carotenoidea. The material described by Hooper et al. (1997) as Halichondria stalagmites (Hentschel) is a composite of //■ (H.) carotenoidea (specimens NTM Z.2651 and Z.3147) and H(H.) microbiana (specimens NTM Z. 131, Z. 1097, Z. 1991), and it does not agree with the concept of the species described by Hentschel (1912) as Ciocalypta stalagmites (see above). Distribution. Halichondria carotenoidea is common in Darwin and Bynoe Harbours, and it was also observed at Wessel Is (Raragala I.). Etymology. Named after carotenoid, a pigment which might give the red-orange colour to the species. It is intended as a noun in apposition. Table 8. Comparison of spicule dimensions between specimens of Halichondria (Halichondria) carotenoidea. Specimen Locality Oxeas NTM Darwin Harbour, 113.2-388.9pm (263.2±87.i) Z.5909 Dudley Point x 2.3-12.3pm (7.3±3) NTM Bynoe Harbour 111.7-379.5pm (245.3±80.2) Z.5225 x 2.3-10.7pm (6.2±2.4) Halichondria ( Halichondria ) danvinensis Hooper et al., 1997 (Fig. 10C) Halichondria danvinensis Hooper et al., 1997: 49. Material examined. Specimens as listed in Hooper et al. (1997). Additional specimens - Darwin Harbour, NTM Z.5211. Remarks. Halichondria danvinensis was described by Hooper et al. (1997). Additional material, an in situ 70 Halichondriidae from northern Australia Fig. It). Halichondria ( Halichondria ) carotenenoidea sp. nov.: A, specimen in situ at Dudley Point, Darwin Harbour, N.T.: B, specimen in situ at Spencer Point, Bynoe Harbour, N.T., showing oscula distributed along lateral side of branches. C, Halichondria (H.) danvinensis, specimen in situ at Weed Reef, Darwin Harbour, NT. Halichondria (//.) phakellioides: D, specimen in situ exposed at the reef flat of East Arm. Darwin Harbour during the low tide of 18 October 2001; E, specimen in situ at Raragala I. Wessel Is, 20 m depth. F, Halichondria (//.) microhiana sp. nov, specimen in situ at Lee Point, Darwin Harbour, N.T. G, Topsentia dura , specimen in situ at Nightcliff bommies, Darwin Harbour, N.T. Photographs: A,D,G - B. Alvarez; B - M. Browne; C,F - H. Nguyen; E P. Colin. 71 B. Alvarez and J. N. A. Hooper Table 9. Halichondria (Halichondria ) species from the Central Indo-Pacific (after Van Soest et al. 2008), with remarks on current taxonomic allocation, type material and description. Species__Remarks_ Halichondria (Halichondria) urmata Unrecognisable from the description; no type material available. Lindgren. 1897 Halichondria (Halichondria) bergquistae Referred here to Axinyssa (see Remarks of Axinyssa this revision). Hooper, Cook, Hobbs & Kennedy, 1997 Halichondria (Halichondria) cartilaginea Valid, common species through Indo-Pacific. Always associated to symbiotic green algae (Esper, 1794) (Lim etal. 2008). Halichondria (Halichondria) darwinensis Valid (see under genus Halichondria , this revision). Hooper, Cook, Hobbs & Kennedy, 1997 Halichondria (Halichondria) fragilis Unrecognisable from the description; no type material available. Kieschnick, 1896 Halichondria (Halichondria) incrustans Kieschnick, 1896 Halichondria (Halichondria) pelliculata Ridley & Dendy, 1886 Halichondria (Halichondria) ridleyi Hooper, Cook, Hobbs & Kennedy, 1997 Halichondria (Halichondria) stalagmites (Hentschel, 1912) Halichondria (Halichondria) syringea Pulitzer-Finali, 1996 Halichondria (Halichondria) vansoesti Hooper, Cook, Hobbs & Kennedy, 1997 Unrecognisable from the description; no type material available. Valid, recognisable from the description. Type material not examined. Distinctive shape and surface characteristics, not comparable to the new species recorded here. Referred in this revision to Topsentia (see Remarks of Topsentia this revision) Referred here to Ciocalypta (see remarks of Cioclaypta this revision) Valid. Type material examined (MSNG 48700). Coalcscent fistules on a buried base [?]. Detachable ectosome differentiated into a thick layer, with spicules oriented paratan- gentially, disorganised. Choanosome halichondroid with some ill defined, multispieular, ascending to surface and supporting ectosome. Oxcas several sizes, 256.4-706.7pm (539.3±137) x 5.2-19|itn (13±3.7); few styles. Refered here to Ciocalypta (see under genus Ciocalypta, this revision) photograph (Fig 10C), and spicule measurements (Table 10) are provided here to complement the original description. Halichondria darwinensis is very inconspicuous and represented so far by only three individuals (including the holotype and the paratype). Its encrusting habit with small and insubstantial digits makes it inconspicuous and hard to find. The re-examination of the available material and additional measurements of spicules, indicate that the differences in the thickness of the spicules reported by Hooper et al. (1997) as a distinctive character for the species, no longer appear to be significant (see Table 10). Both the length and the thickness of the oxeas are variable, but not divisible into size classes. The species seems to have affinities with the genus Axinyssa. The skeleton is poorly developed, with very little Table 10. Comparison of spicule dimensions among specimens of Halichondria ( Halichondria ) darwinensis. Specimen Locality Oxcas NTM Z.3205 Darwin 272.3-659.6pm (Holotype) Harbour, East (467.2±112) Point x 3.7-13.2pm (8.5±2.5) QM G303252 Darwin 307-627.5pm (538.8±90.6) (Paratype) Harbour, East Ann x 4.3-16pm (10.2±3.2) NTM Z.5211 Darwin 386.2-663.7pm (524±68) Harbour, Weed Reef x 7-14.9ptn (11.8±1.9) spongin and relatively less spicule density when compared to other Halichondria species; the ectosomal skeleton is quite undifferentiated and the spicules are similar in shape and dimensions to other Axinyssa species. Therefore, the assignment of this species to the genus Halichondria is inconclusive. Distribution. Halichondria darwinensis is presently known only from Darwin Harbour. It occurs intertidally and subtidally to 10 m. Halichondria (Halichondria ) phakellioides Dendy & Frederick, 1924 (Figs 10 D, E, 12) Halichondria phakellioides Dendy & Frederick, 1924: 498; Burton 1934: 600; Hooper et al. 1997. Material examined. Specimens as listed in Hooper et al. (1997). Additional specimens - Bynoc Harbour, NT: Z.241, Z.5223 (0M9H2321-P). Darwin Harbour, NT: Z.2026, Z.4100, (0CDN-8022-F), Z.5219 (0M9H2057-C), Z.5905, Z.5906, Z.5915, Z.5923, Z.5925, Z.5928, Z.5948, Z.5949. Z.5950, Z.5965, Z.5970. Parry Shoals, N.T., QM G310137. Boucat Bay, E of Maningrida, Arnhem Land. N.T.: Z.5623. Wesscl Is: Z.5228 (0M9H2655-C). Remarks. Halichondria phakellioides was described by Hooper et al. (1997). Additional material, illustrations (Figs 10D, E, 12), and spicule measurements (Table 11) are provided here to complement the original description. 72 Halichondriidae from northern Australia Fig. II. Halichondria ( Halichondria ) carotenenoidea sp. nov. (NTM Z.5909): A, light microphotograph of tangential section of ectosomal skeleton showing ill-defined and criss-crossing bundles of spicules; B, light microphotograph of perpendicular section through surface showing organisation of choanosomal skeleton and part of ectosomal skeleton layer (left upper corner of section); C, diagram of spicules Scale bars: A, 100 pm; B, 500 pm; C, 50 pm. Distribution. Halichondria phakellioides is widely distributed through the Northwest Australian Shelf and Sahul Shelf provinces. It is found from the intertidal to 20 m depth. Halichondria (.Halichondria ) microbiana sp. nov. (Figs 10F, 13) Halichondria stalagmites- Hooper el al. 1997: 43 [in part; not Z.2651 and Z.3147 = H. (H.) carotenoidea]. Material examined. Holotype. -NTM Z.5907, Lee Point, Darwin, N.T., 12°20.538’S, 130°52.184' E, 9-12 m depth, 7 August 2003, coll. Nguyen, H. Paratypes- NTM Z.1097, Dudley Point Reef, East Point, Darwin, N.T., 12°25.000r S, 130°48.0r E, 6-7 m depth, 22 December 1982, coll. Hooper, JNA. NTM Z.5908, Moira Reef, Bynoe Harbour, N.T., 12°30.799’ S, 130°30.527’ E, 5-8 m depth, 25 June 2003, coll. Nguyen, H. Additional specimens- Darwin Harbour NT: NTM, Z.1991. Cobourg Peninsula, N.T.: NTM Z.131. Description Shape (Fig. 1 OF). Cushion-shaped with globular, subspherical or massive base, sometimes semi-buried Table 11. Comparison of spicule dimensions among specimens of Halichondria (Halichondria) phakellioides. Specimen Locality Oxea type I Oxea type II NTM Z.5223 Bynoe Harbour 170.1-308.4pm (232±30.7) x 4.1-10.3pm (8±1.5) 363.2-524.6pm (440.6±40.8) x 11.5-24.4pm (17.9±3.7) NTM Z.5219 Darwin Harbour 154.2-293.2pm (228.6±34.5) x 5-10.3pm (7.3±1.4) 356.7-499.6pm (425.5±38.7) x 10.5-20.2pm (16.3±3) NTM Z.5228 Wessel Is 162.1-313.9pm (242.4±48.6) x 3.7- 7.5pm (5.6±0.9) 326.5-519.3pm (440.5±45.5) x 10.6-21.1 pm (15.6±3) 73 B. Alvarez and J. N. A. Hooper Fig. 12. Halichondria ( Halichondria ) phakellioides (NTM Z.5228): A, light microphotograph of tangential section showing organisation of ectosomal skeleton; B, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton. Scale bars: A-B, 500 pm. in substrate, and small tapering erect projections (from 10 mm long and less than 5 mm diameter) at apex. Erect projections vary in shape, i.e. fistulose-like, convoluted, bifurcating and lobate; generally small 30-70 mm diameter, 10-20 mm thick. Colour. Orange. Blue-grey, in alcohol. Consistency and texture. Soft, easily tom, rubbery. Oscula. Apical and conspicuous on small digits, with membranous rims. Surface. Semitransparent at apex of digits in some specimens. Skeleton (Fig. I3A,B). Ectosomal skeleton, halichondroid, formed by a thin (less than 50 pm), wavy, loose membranous layer, with oxeas tangentially oriented and supported by choanosomal tracts (Fig. 13A). Choanosomal skeleton, halichondroid, formed by ill-defined bundles of larger spicules, criss-crossing and without any particular orientation connected by short bundles and single spicules. In subectosomal region, skeleton becoming cavernous with large lacunae, 100-600 pm diameter (Fig. 13B) and vague reticulation of pauci-multispicular, ill- defined tracts, diverging towards surface or condensed and running nearly parallel to it, supporting ectosomal skeleton. High densities of filamentous cyanobacteria present in both ectosome and choanosome, especially near surface. Spongin or collagen scarce. Spicules (Fig. 13C; Table 12). Oxeas, hastate, straight, possibly in two size categories, larger and thicker (249-587 x 5-14 pm) and smaller and thinner (88-217 x 3-7 pm). Style modifications slightly common within larger category. Remarks. We compared Halichondria inicrobiana with the valid species recorded for the Central West Pacific (Table 7) and those reported in this revision. It is diagnosed by a unique combination of features (i.e growth form, organisation of the skeleton, size and composition of spicules and presence of cyanobacteria in the ectosome and the choanosome) not found in those species. All the specimens examined here have a high density of filamentous cyanobacteria in the choanosome, generally concentrated at the subectosomal region, a character shared with //. (//.) carotenoidea. As mentioned above, this new species is also very similar in skeletal characteristics to H.(H.) carotenoidea sp. nov., but it differs in habit and size of spicules, which are larger and separated in size categories in H. (//.) inicrobiana. Hooper et al. (1997) interpreted this species as Halichondria stalagmites. However, a thorough re-examination of the type material of that species indicates that it is not conspecific with H. (H .) inicrobiana (see description of C. stalagmites above). Distribution. Halichondria inicrobiana is relatively common at Darwin and Bynoe Harbours, and it was also observed at Wessel Is (Raragala I.) and recorded from the Northeastern Australian Shelf (Alvarez and Hooper, unpublished data). It occurs between 6-12 m depth. Etymology. Named after the symbiotic microorganisms hosted by the species. The specific name is intended as a noun in apposition. Remarks on Halichondria ( Halichondria ). The subgenus Halichondria is large with approx. 95 valid species (Van Soest et al. 2008). Table 7 lists species distributed within the Central Indo-Pacific Realm (following Spalding et al. 2007) which includes the study area and adjacent areas. Some of those species, in particular the Table 12. Comparison of spicule dimensions between specimens of Halichondria ( Halichondria ) inicrobiana. Specimen Locality Oxea type I Oxea type II NTM Z.5907 Darwin Harbour, NT 87.9-203pm(126.5±24.8) 249.2-587pm (482.9±75.1) x 3.3-7.1pm (5±0.9) x 6.3-14.2pm (8.9±1.8) NTM Z.5908 Bynoe Harbour, NT 91.3-217gm (137.8±32) 282.5-572.1pm (446.6±93.5) x2.9-7.2pm(5.1±l) x 5.2-13.2pm (8.8±2) 74 Halichondriidae from northern Australia Fig. 13. Halichondria ( Halichondria ) microbiana sp. nov. (NTM Z.5908): A, light niicrophotograph of tangential section of ectosomal skeleton, showing oxeas on membranous thin layer; B, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton and large lacuna at subectosomal level (left upper comer of section); C, diagram of spicules. Scale bars: A, 200 pm; B. 500 pm; C, 50 pm. ones described by Kiesehnick (1896) and Lindgren (1897), are unrecognisable and their type material has never been relocated. Other species reported for the area have been allocated to other genera as result of this revision, thus Halichondria (Halichondria) is represented within this realm by nine species including the two new species described above. The subgenus Eumastia Schmidt, 1870 is not represented in the study area at all and it is reserved for Halichondria- like species from high latitudes (Erpenbeck & Van Soest 2002 ). Genus Hymeniacidon Bowerbank, 1858 Gender: feminine. Type species, by subsequent designation of Bowerbank (1864), Hymeniacidon canmcula Bowerbank, 1958. Recent, Tenby, Wales. Hymeniacidon gracilis (Henstschel, 1912) (Fig. 14) Stylotella digitata gracilis Henschel, 1912: 356. Hymeniacidon gracilis. - Hooper et al. 1997. Material examined. As listed by Hooper et al. (1997). Remarks. The species was well described by Hooper et al. (1997) and elevated to full species rank in the genus Hymeniacidon. We provide a new illustration of the type material (Fig. 14) and additional spicule measurements (Table 13) to complement the description given by Hooper et al. (1997). Only three specimens arc recorded for the study area; no new material was located in recent collections. Distribution. As recorded by Hooper et al. (1997). Remarks on the genus Hymeniacidon. Hymeniacidon gracilis is the only valid species of the genus from the study area. Other species of Hymeniacidon recorded for the 75 B. Alvarez and J. N. A. Hooper Fig. 14. Hymeniaddon gracilis (Synype SMF 970). A. light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton; B, diagram of spicules. Scale bars; A, 500 pm; B, 50 pm. Sahul Shelf Province and adjacent areas are H. vernonensis Hooper et al., 1997 and H. flaccida Pulitzer-Finali, 1996. The material described under H. vernonensis by Hooper et al.( 1997) was revised and it does not agree with the current concept of the genus. The species is formally transferred here to the dictyonellid genus Stylissa. As admitted by Hooper et al. (1997), is very similar to Sty 7 issa jlabelliforniis but S. vernonensis includes distinctive styles, curv ed at the centre, sinuous or rhabdose and frequent anisoxeas with one telescoped point. It should be noted however, that these spicule modifications are common among species of Stylissa and therefore they are not reliable for the delimitation of species. Both species are similar in growth form, surface characteristics and skeletal organisation. The choanosomal skeleton of S. vernonensis however, is nearly halichondroid with only vague tracts of spicules and with a much higher spicule density and no spongin fibres (whereas in S. flabelliformis is vaguely plumo-reticulated and with well developed spongin fibres). The type specimen of Hymeniaddon flaccida (MSNG 48703) from Laing Is., Papua New Guinea, was re-examined and it does not correspond to the genus Hymeniaddon. The species belongs also to the dictyonellid genus Stylissa and is likely to be conspecific with Stylissa massa (Carter, 1887). The specimen examined however, includes distinctive subtylostyles transitional to strongyles with tylote modifications. As is the case with 5. vernonensis, Table 13. Comparison of spicule dimensions between specimens of Hymeniaddon gracilis. Specimen Locality Styles SMF 970 Indonesia 220.5-261.1 pm (238.4±9.6) x 3.3-8.9pm (6.1±1.5) [25] NTM Z.883 Darwin 243.1-279.1 pm (265.6±11.1) Harbour x 3-7.4pm (5.1±1.1) these style modifications are common among the genus. Population genetics and additional morphometric analyses might reveal whether these style modifications of Stylissa are reliable characters for the separation of species. Genus Topscntia Gender feminine. Type species, by original designation, Anisoxya glabra Topsent, 1898. Recent, Azores Is. Topsentia dura (Lindgren, 1897) (Figs 10G, 15) Halichondria dura Lindgren, 1897 : 480. Topsentia dura - Hooper et al. 1997: 14. Material examined. As listed by Hooper et al. (1997). Additional material - Darwin Harbour, N.T.: Z.5209, Z.5233. Wessel Is: Z.5234. Remarks. Hooper et al. (1997) assigned material from the Beagle Gulf to this species under the genus Topsentia. Additional specimens from recent collections agree also with this material and are assigned to this species. Further illustrations (Figs 10G, 15) and spicule measurements (Table 14) are provided here to complement that description. The description agrees with the current concept of Topsentia, however it remains inconclusive whether the material from the Beagle Gulf is conspecific with Lindgren’s species from Indonesia as the type was not examined. Examination of additional specimens from Indonesia (Alvarez & De Voogd unpublished data) and a re-description of the type might provide additional evidence to confirm if these populations belong in the same species. Distribution. Indonesia [?], Darwin Harbour and Wessel Is. Topsentia dura is occurs the intertidal zone to 25 m depth. Remarks on Topsentia. Hooper et al. (1997) described material under the Red Sea species Topsentia halichondrioides (Dendy, 1905) that seems very similar 76 Halichondriidae from northern Australia Fig. 15. Topsentia dura (NTM Z.5209): A, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton and palisade of erect oxcas at surface; B, diagram of spicules. Topsentia halichondrioides (QM G303442); C, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton and oxcas oriented perpendicularly at surface level; I), diagram of spicules. Topsentia ridleyi (QM G303309, Holotype); E, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton; F, diagram of spicules. Scale bars: A,C,E, 500 pm; B,D,F, 100 pm. to T. dura (Fig. 15B,C). These species are massive, of hard consistency with skeletons made of a confused mass of oxcas of similar dimensions, not clearly differentiated into size classes (see Table 14) and differing only in colouration and shape of osculcs (i.e. volcano-shaped in T. halichondrioides, and sunken and small in T. dura). Halichondria ridleyi Hooper et al., 1997 is referred here to the genus Topsentia (comb, nov.) and it is also very similar to T. dura (Fig. 15E,F). It differs from T. dura in having some surface fistules and processes. 77 B. Alvarez and J. N. A. Hooper Table 14. Comparison of spicule dimensions among specimens assigned to Central Pacific species of Topsenlia. Species Specimen Locality Oxea type 1 Oxea type 11 Topsentia ridleyi QM G303309 Darwin Harbour 142.3-273.5pm (207.7±39.4) x 3.3-7.6pm (5.4±1.2) 335.6-576.7pm (443.9±52.7) x 6.7-16.6ptn (12±2.6) Topsentia ridleyi NTM Z.3262 Cobourg Peninsula 503-848.1 pm (653.8±83.5) x 13.4-31.5pm (20.9±4.6) Topsentia halicltondrioides G303442 Bynoe Harbour 160.6-331.2pm (257.4±50.3) x 4.2-12.3pm (8.2±2.3) 421.6-716.5pm (555.3±62.7) x 14-24.4pm (19.3±2.6) Topsentia lialichondrioides NTM Z.5233 East Point 111.4—165.9pm (142.3±13.8) x 5.9-8.9pm (7.2±0.9) 487.6-914.9pm (711.2±99.1) x 13-30pm (18.9±4.4) Topsentia dura NTM Z.5234 Wessel Is 142.4-333.5pm (220.1±47.2) x 3.7-10.3pm (7.3±1.7) 348.6-568.1pm (448.9±57.8) x ll-23.7pm(16.8±3.2) Topsentia dura NTM Z.5209 Darwin Harbour 151.2-399.7pm (248.3±61.9) x 3-11.7pm (7.6±2.3) 373.9-656.5pm (477.9A79.2) x 10-23.9pm(15±3.2) Topsentia dura NTM Z.3178 Darwin Harbour 189.4-357.5pm (269.9±41.9) x 4.1-9.7pm (7.1±1.4) 352.5-525.1pm (441.4±46.2) x 9.9-18.5pm (13.7±2.4) Topsentia dura NTM Z.1442 Gunn Point 189.6-387.1 pm (254.5±52.5) x 5.5-12.5pm (7.8±2) 341.2-611.2pm (482.6±69.2) x 8.3-22.7pm (15.6±3.4) Topsentia indica SMF 997 Aru Is, Indonesia 493.7-1078.1 pm (734.4±125) x 11.1-39.4pm (21.7±5.5) An additional species of Topsenlia recorded from Aru Is, Indonesia, is T. indica Hentschel, 1912 (syntype SMF 995 and 997, examined). The differentiation of these species using traditional morphological characters is extremely subjective. Molecular and morphometric studies of local populations might contribute to a better understanding of the concept of this species. Other species of Topsenlia recorded for the Sahul Province and adjacent areas that are better placed elsewhere include Topsentia maculosa Pulitzer-Finali, 1996 from Papua New Guinea (it belongs in Amorphinopsis, see above) and Topsentia plurisclera Pulitzer-Finali, 1996 (holotype, MSNG 48702, examined) is a species of Petrosia. DISCUSSION This revision of species of Halichondriidae from northern Australia recognises a total of 15 species belonging to the genera Amorphinopsis, Axinyssa, Ciocalypta, Halichondria ( Halichondria ), Hymeniacidon and Topsentia. Other genera of the family (i.e. Epipolasis, Laminospongia, Vosmaeria , Ciocalapata and Spongosorites are not represented in the Sahul Shelf Province. Epipolasis and Spongosorites are however represented in the Northeast Australian Shelf (Alvarez and Hooper, unpublished data). Of the species reported in this revision, Axinyssa bergquistae, Ciocalypta vansoesti and the two new species Halichondria ( Halichondria ) carotenoidea and H (H.) microbiana arc so far known only from northern Australian waters. The rest of the species have extralimital distributions through the Central Indo-Pacific realm. Axinyssa mertoni (Hentschel, 1912) in its new generic combination is recorded for northern Australia and it represents a new record for the study area. As in other members of the order Flalichondrida, particularly in Axinellidae and Dictyonellidae, species within and across all genera of Halichondriidae arc extremely di fficult to delimit. This problem is demonstrated by the large number of misidentifications in previous studies due to the lack of adequate generic definitions and also to the poor understanding of the importance, or indeed relevance, of some of the alleged pivotal characters that currently differentiate both species-groups and genera within the Halichondriidae. The additional information obtained from the collection of new material plus the revised generic definitions of the family (Erpenbeck & Van Soest 2002) allowed us to clarify the concept of halichondriid species of northern Australia and to allocate them to more appropriate genera. Nevertheless, differentiating species within Halichondriidae continues to be ambiguous based solely on the present limited suite of accepted morphological characters, with a number of them shared among species and even genera. For example, Ciocalypta heterostyla, C.vansoesti. and Axinyssa mertoni are species with fistulc- like growth form, nearly indistinguishable in the field (Fig 1). However, they all are easily diagnosed based on skeletal characteristics. This suggests that the growth form of these species in particular, might be an adaptation to the habit where they occur (i.e. soft and muddy sediments). Amorphinopsisfoetida and A maculosa are also very similar in habit and in their skeletal characteristics but they can be distinguished by the predominance of oxeas and styles. Separation of species based on the dominance of styles or oxeas however might be debatable, as this could possibly be related to intraspecific variation as seen in some species of Axinellidae (Alvarez et al. 1998; Alvarez & Hooper 2009). Spicule morphometries (i.e. size variation of spicules) within Halichondriidae seems to be a useful tool for the 78 Halichondriidae from northern Australia differentiation of species. Ciocalypta stalagmites, for example, is distinguished from the other species by the distinctive and nearly constant size of two categories of oxeas. Similarly, Halichondria (II.) microbiana can be distinguished from H. (//.) carotenoidea by the size of the larger category of oxeas. On the other hand, all the Axinyssa species we have studied have always a mixture of oxeas in a large size range, thus spicule sizes is not a useful character for the distinction of species within this genus. These examples indicate that the characters currently used to separate genera and species within the family are extremely homoplastic and suggest that genera and species within Halichondriidae might be non-monophyletic. The study of local populations using both morphological and genetic methods will help to clarify whether these taxa are monophylctic. This study represents the final contribution to the present taxonomic revision of the order Halichondrida of northern Australia, restricted to the marine province identified as the Sahul Shelf in the classification of Spalding et al. (2007). The result of this and previous studies (Alvarez & Hooper 2009, 2010) indicates that the group is represented in the study area mainly by the families Axinellidae (Alvarez & Hooper 2009), Dictyonellidae (Alvarez & Hooper 2010) and Halichondriidae (this revision). One additional family, the Heteroxyidae (formerly Desmoxyidae see Van Soest & Hooper 2005), is represented in the area by two very common species: Myrmekioderma granulatum (Esper, 1830), which is documented and illustrated by Hooper et al. (1997); and Higginsia mixta (Hentschel, 1912). The type of H. mixta (SMF 968) was examined and it agrees with material deposited at the collections of QM and NTM and recorded for the area of study. The cctosomal skeleton of the specimens studied have a relatively thick tangential crust formed by a dense mass of spined microxeas and interrupted by disorganised brushes of thin raphidiform oxeas and extra long styles- strongylcs (mostly broken in the preparations) projecting through surface. The similarities of this type of skeletal organisation with raspailiid genera such as Ceratopsion are remarkable and worth further investigation. Phylogenetic affinities based on molecular data (Erpenbcck et al. 2005) indicated that members of Heteroxyidae (i.e. Didiscus and Myrmekioderma) are closely related to the axinellid genera Reniochalimi and Ptilocaulis. The position of these genera within Axinellidae (Halichondrida) and its relationships based on molecular data with other raspailiid genera such as Axechina has already been discussed (Alvarez 2009 and references within). Sequencing data of additional genera currently allocated to Heteroxyidae, and in particular of the species represented in the Sahul Shelf province will help to clarify these relationships and classification. The family Bubaridae is the only family of Halichondrida not represented in the area of study. ACKNOWLEDGEMENTS This work was funded by an Australian Biological Research Studies Research grant (No 205-10) and by the ‘Collection and Taxonomy of Shallow Water Marine Organisms’ program for the U.S. National Cancer Institute (Contract N02-CM-27003 and Contract N02CO-2009-00012) subcontracted to the M AGNT through CRRF. We thank the following people: Michael Browne and Huy Nguyen, for their invaluable assistance during 2002-2004 MAGNT field collections; Dr Pat Colin, CRRF, and Don DeMaria, for their assistance and photographic work during the MAGNT field collections in the 2004 Wessel Is; Terry Yumbuluy, Wessel Is, for allowing collections of sponges in his sea area; Merrick Ekins, QM, for his assistance interrogating QM database and making specimens available for study and Dr Kathryn Hall for substantial re-checking voucher specimens housed in the collections of the QM to confirm or refute previous species diagnoses. Drs Rob W.M. Van Soest (ZMA) and Nicole De Voogd are thanked for their input and valuable discussions on Indo-Pacific halichondrid sponges. Mr Swee-Cheng Lim is thanked for his assistance locating type material at MSNG. Drs Richard Willan and Chris Glasby (MAGNT) are thanked for their continuous advice and suggestions during the preparation of this manuscript. 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Cook, SD. on CSIRO RV Southern Surveyor G301034 SW Vrilya Point, SW, Gulf of Carpentaria, QLD. 11°29.0167’ S, 142°55.09' E, 18 m , 1 Dec 1991, coll. Cook, SD. on CSIRO RV Southern Surveyor G303252 South Shell I., reef N of boatramp. East Ann, Darwin Harbour, NT, 12°29.1334’ S. 130°53.09’ E, 0 m , 19 Sep 1993, coll. Hooper, JNA & Hobbs, L.J. G303287 South Shell I., reefN ofboatramp. East Ann. Darwin Harbour, NT, 12°29.1334" S, 130°53.09' E, 0 m, 19 Sep 1993, coll. Hooper, JNA & Hobbs, LJ G303309 Dudley Point Reef, East Point, Darwin, NT, 12°25.05' S, 130°49.0r E, 0 in , 20 Sep 1993, coll. Hooper, JNA & Hobbs, L.J. G303351 East Point Bommies, Darwin Harbour, NT, 12°24.0834' S, 130°48.14' E, 10 in , 23 Sep 1993, coll. Hooper, JNA & Hobbs, LJ G303442 Fish Reef, west side, Bynoe Harbour, NT, 12°26.0167’ S, 130°26.09’ E, 11 nr, 26 Sep 1993, coll. Hooper, JNA & Hobbs, LJ G303450 Fish Reef, west side, Bynoe Harbour, NT, 12°26.0167’ S, 130°26.09’ E, 11 in , 26 Sep 1993, coll. Hooper, JNA & Hobbs, LJ G303524 Duyfken Point. W Gulf of Carpentaria, QLD, 12°41.050rS, 141 °3.01 ’ E, 42 m, 11 Nov 1993,coll. Cook, SD. & Kennedy,J. on CSIRO RV. Southern Surveyor G303541 Vernon Is, W ofSouth West Vernon I„ NT, I2°6.15’ S, I31°4.I4’ E, 13 m, 10 Oct 1993. G303558 Cape Hotham, NW of cape, NT, 12° 1.05’ S, 131 °13.16’ E, 34 m. 9 Oct 1993. G303560 Bynoe Harbour, 2 nmls E Fish Reef, NT, 12°24.1334’ S, 130°28.I6' E, 17 m, 6 Oct 1993, coll. CCNT Ocean Rescue 2000 Program G303561 Shoal Bay, outer region of bay, NT, I2°6.15’ S, 130°49.I6’ E, 18 in, 12 Oct 1993, coll. CCNT Ocean Rescue 2000 Program G303595 Vernon Is, WofKnight Reef, NT, 12°1.0334'S, 131°3.16’ E, 22 in, 11 Oct 1993. G303658 Vernon Is, N marsh Shoal, NT, 12°07.0001'S, 130°56.rE, 16 m, 11 Oct 1993, coll. CCNT stn. 138. Dredge G303677 Shoal Bay, middle of bay, NT, 12° 13.0167’ S, 130°56’ E, 17 m, 12 Oct 1993, coll. CCNT Ocean Rescue 2000 Program G310137 Parry Shoals 35nm W Bathurst I., NT, 11°7.0321' S, 129°25.9’ E, 16 m, 12 Aug 1987, coll, mussig, AM and NCI team G310I70 Darwin Harbour, NT, 12° 15.1834’ S., 130°29.11’E., 9 m depth, 17 August 1987, coll, mussig, AM and NCI team G313543 N Bathurst I., Timor Sea, NT, 11°13.98’ S, 130°34.21’ E, 41.2 m, 5 Oct 1997, coll. Cook, SD. on RV Southern Surveyor G313572 SW Groote Eylandt, NT, 14°25.0801’ S, 135°58.51’ E, 20.3 m, 13 Oct 1997, coll. Cook. SD. on RV Southern Surveyor G313577 SW Groote Eylandt, NT, 14°20.22’ S, 136°34.98’ E, 19.6 m, 14 Oct 1997, coll. Cook, SD. on RV Southern Surveyor G314246 N Groote Eylandt, Gulf of Carpentaria, NT, 13°32.2801’ S, 136° 18.13’ E, 20 m, 27 Sep 1998, coll. Leys, SP. on RV Southern Surveyor G314247 N Groote Eylandt, Gulf of Carpentaria, NT, 13°32.2801’ S, 136° 18.13’ E, 21.7 m, 27 Sep 1998, coll. Leys, SP. on RV Southern Surveyor G314255 W of Groote Eylandt, Gulf of Carpentaria, NT, 14°8’S, 136°8’E, 13 m, 6 Oct 1998, coll. Leys, SP. on RV Southern Surveyor G314267 SW ofGroote Eylandt, Gulf of Carpentaria, QLD, I4°20’ S, 136°2’ E, 22.1 m, 6 Oct 1998, coll. Leys, SP. on RV Southern Surveyor G315205 W Groote Eylandt, Gulf of Carpentaria, NT, 14°8.5667’S, 136° 16.51' E, 21.4 m, 130ct 1998, coll. Wassenberg T G315207 SW ofGroote Eylandt, Gulf of Carpentaria, NT, 14°22.5’S, 136°9.12’ E, 22.2 m, 130ct 1998, coll. Wassenberg T G320819 Gulf of Carpentaria, QLD, 15°20.037’ S. 140° 19.84’ E, 28 m, 24 may 2003, coll. Bartlett C, Cook S on RV Southern Surveyor 2380403 CSIRO “Effects of Trawling” G320904 Gulf of Carpentaria, QLD, 15°20.037’ S, 140°19.84’ E, 28 m, 11 mar 2003, coll. Bartlett C, Cook S on RV Southern Surveyor 2380403 CSIRO “Effects of Trawling” 81 B. Alvarez and J. N. A. Hooper APPENDIX 1 (continued) Collection and locality data of material examined in the collections of QM and NTM NTM material Z.84 Coral Bay, Port Essington, Cobourg Peninsula, NT, ll°11.500rS, 132°2’E, 180ct 1981,coll. Hooper, JNA& Alderslade, PN Z.131 Sandy I. No.2. Cobourg Peninsula. NT, 11°5.5001'S, 132°17’E, 10 m, 21 Oct 1981, coll. Hooper, JNA & Alderslade, PN Z.194 Dudley Point Reef. East Point. Darwin, NT, 12°25.0001’S, 130°48.01’E, 0-0.5 m, 13 Sep 1981, coll. Hooper, JNA and party Z.241 Indian I„ Bynoc Harbour, NT, 12°35’S, 130°33.0rE, 3 m, 18 Nov 1981, coll. Byers.P.,F.V. Skeleton Z. 592 Table Head, Port Essington, Cobourg Peninsula, NT. 11°13.5’S., 132 o 10.51’E., 3 m depth, 4 May 1982, coll. Hooper, JNA Z 712 N Adelc I.,Collier Bay, NW Shelf, WA 15°58.0167’S„ 122°39.07’E„ 59 nr depth, 21 April 1982, coll. R.V.SPRIGHTLY, dredge. Z.919 Dudley Point Reef. East Point, Darwin, NT, 12°25.0001’S, 130°48.0I ’E, 10 m, 31 Aug 1982, coll. Hooper, JNA Z.934 East Point Reef, East Point, Darwin, NT, 12°24.05'S, 130°48.01 'E, 12 m, 13 Sep 1982, coll. Hooper, JNA Z.941 East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01'E, 12 nt. 13 Sep 1982, coll. Hooper, JNA Z.945 East Point Reef, East Point, Darwin, NT, 12°24.05‘S, 130°48.01'E, 12 m. 13 Sep 1982, coll. Hooper, JNA Z.986 Dudley Point Reef, East Point, Darwin, NT, 12°24.5'S, 130°48.01 'E, tn, 26 Oct 1982, coll. Hooper, JNA Z.987 Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, I30 0 48.01’E, tn. 26 Oct 1982, coll. Hooper, JNA Z.1097 Dudley Point Reef, East Point, Darwin, NT, I2°25.000l ’S, I30°48.01'E, m. 22 Dec 1982, coll. Hooper, JNA Z.1358 Coral Bay, Port Essington, Cobourg Peninsula. NT, 11° 11,3’S, 132°3.71’E, .5-6 m, 16May 1983,coll. Hooper, JNA Z.1391 Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3’S, 132°3.71'E, 6 m. 17 May 1983, coll. Hooper, JNA Z. 1395 Coral Bay, Port Essington, Cobourg Peninsula, NT. 11°10.4’S, 132°2.8’E, 2 m, 19 May 1983, coll. 1 Iooper, JNA Z.1442 Blue Hole, Gunn Point, NT, 12°9.000rS, 131°0’E,25 in, l9Aug 1983, coll. Alderslade, PN Z.1979 West side of Weed Reef, Darwin, NT, 12°29,2001 ’S, 130 o 47.1’E, m, 1 1 May 1984, coll. Hooper, JNA and party Z.1991 West side of Weed Reef, Darwin, NT, 12°29,200rS, 130°47.1’E, m, 1 1 May 1984, coll. Hooper, JNA and party Z.2018 West side of Weed Reef, Darwin, NT, 12°29.2001 ’S, 130°47.1'E, m, 11 May 1984, coll. Hooper, JNA and party Z.2026 West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.I’E. 11 May 1984, coll. Hooper, JNA and party Z.2086 Dudley Point Reef. East Point, Darwin, NT, !2°24.5'S, 130°48.01'E, tn, 20 Jul 1984, coll. Hooper, JNA Z.2215 Vestey’s Beach. Bullocky Point, Darwin, NT, 12°26.2’S, I30 C ’49.89’E 21 Jan 1985, coll. Hooper, JNA Z.2245 Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01'E, 10 m, 12 Apr 1985, coll. Hood, C and party Z.2648 Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01'E, m. 3 Apr 1986, coll. Hooper, JNA and party Z.2651 Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 3 Apr 1986, coll. Hooper, JNA and party Z.2697 Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 3 Apr 1986, coll. Hooper, JNA and party Z.3106 Parry Shoals, Arafura Sea, NT, 11° 12.5167’S, I29°42.07'E. 20 m, 15 Aug 1987, coll. Mussig, AM and NCI team Z.3133 Parry Shoals. Arafura Sea, NT, 11 0 11.4’S., 129°43.01’E., 18 m depth, 13 August 1987, coll. Mussig, AM and NCI team Z.3147 Parry'Shoals, Arafura Sea, NT, 11 0 12.5167’S, I29°42.07’E, 16m, 15Aug 1987, coll. Mussig, AM and NCI team Z.3178 East Point Reef, East Point, Darwin, NT, 12°29.5’S, 130°48.0I ’E, 0.5 m, 10 Sep 1987, coll. Smit, N Z.3195 Dudley Point Reef, East Point, Darwin, NT, I2°24.5’S, 130°48.01’E, 9 m, 16 Sep 1987, coll. Smit, N Z.3205 Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.0rE, 25 Sep 1987, coll. Smit, N Z.3262 Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5’S, 132 o 10.51’E, 11 Sep 1986, coll. Hooper, JNA & Johnson.C Z.3920 Cumberland Strait. NE bay, Wessel Is, Gove Peninsula, NT, 11°26.8’S, I36°30.2’E, 13 m, 14 Nov 1990, coll. Hooper, JNA Z.4085 Near Boat Ramp, East Arm Port, Darwin, NT, Australia, 12°29.8’S, I30°53.5’E, intertidal, 20 September 2001, coll. B. Glasby & party, by hand Z.4093 Near Boat Ramp, East Arm Port. Darwin. NT, 12°29.8’S, 130°53.5’E, intertidal, 20 September 2001, coll. B. Glasby & party, by hand Z.4100 Near Boat Ramp, East Arm Port, Darwin, NT, 12°29.8’S, I30°53.5’E, intertidal, 18 October 2001, coll. B. Glasby & party, by hand Z.4122 Near Boat Ramp, East Arm Port, Darwin, NT, 12°29.8’S, 130°53.5’E, intertidal, 18 October 2001, coll. B. Glasby & party, by hand Z.4123 Near Boat Ramp, East Arm Port, Darwin, NT, I2°29.8’S, I30 1 ’53.5’E, intertidal, 19 October 2001, coll. B. Glasby & party, by hand Z.4125 Near Boat Ramp, East Arm Port, Darwin, NT, 12°29.8’S, 130°53.5’E, intertidal. 19 October 2001, coll. B. Glasby & party, by hand Z.4451 Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.103’S, 130°47.11 l’E, 8-14 nt, 7 May 2002, coll. Alvarez, B and party Halichondriidae from northern Australia Z.5206 South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.14rE, 10-12 m, 21 Aug 2002, coll. Alvarez, B and party Z.5207 Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.238’S, 130°35.557’E, 5-10 m, 23 May 2003, coll. Alvarez, B and party Z.5208 Dawson Rock. 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.207’S, 130°35.459’E, 7-12 m, 24 Jul 2003, coll. Alvarez, B and party Z.5209 Nightcliff bommies, off Nightcl iff jetty, Darwin Harbour. NT, 12°22.751’S, 130°50.116’E, 5-8 m, 8 Aug 2003, coll. Alvarez, B and party Z.5210 Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8-14 m, 22 Aug 2003, coll. Alvarez, B and party Z.5211 Weed Reef, entrance to West Arm, Darwin Harbour, NT, 12°29.25’S, I30°47.54’E, 9-12 nt, 6 Sep 2003, coll. Nguyen, H Z.5212 off Herbert Point, Indian I., Bynoe Harbour. NT, 12°34.586’S, 130°31.419’E, 0-5 m, 24 Jun 2003, coll. Alvarez, B and party Z.5213 South Shell I„ East Ann, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 7-11. m, 19 Aug 2002, coll. Alvarez, B and party Z.5215 West Ann, 2.5 km N of Stokes Point, Darwin Harbour. NT, I2°31.300’S, 130°48.500’E, 4-5 m, 3 Aug 2002, coll. Alvarez, B and party Z.5216 Wickham Point, 2.5 km SW of East Ann Wharf, East Arm, Darwin Harbour, NT, 12°30.12’S, 130°52.39’E, 4-7 nt, 15 Sep 2002, coll. Alvarez, B and party Z.5217 Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8-14 m, 22 Aug 2003, coll. Alvarez, B Z.5218 Raragala E, bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17-20 m. 30 Mar 2004, coll. Alvarez, B and party Z.5219 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2001 ’S, 130°47.1'E, 5-19 nt, 8 May 2002, coll. Alvarez, B and party Z.5221 South Shell 1., East Arm. Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 7-14 m, 20 Aug 2002, coll. Alvarez, B and parly Z.5222 East Arm Wharf, East Arm, Darwin Harbour, NT, 12°29.19’S, 130°53.35’E, 0.6 m, 1 Mar 2002, coll. Alvarez, B and party Z.5223 Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.238’S, 130°35.557’E, 5-10 m, 23 May 2003, coll. Alvarez, B and party Z.5224 Dawson Rock. 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.207’S, 130°35.459’E, 3-7 m, 25 May 2003, coll. Alvarez. B and party Z.5225 Spencer Point, Indian 1., Bynoe Harbour, NT, 12°35.35 i ’S, 130°31,454’E, 6-8 m, 11 Jun 2003, coll. Alvarez, B and party Z.5226 Moira Reef, Bynoe 1 larbour, NT, 12°30.799’ S, 130°30.527’E, 5-8 nt, 25 Jun 2003, coll. Browne, M Z.5228 Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839'E, 17-20 m, 30 Mar 2004, coll. Alvarez, B and party Z.5229 Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04'E, 12-18 m, 16 Sep 2002, coll. Alvarez, B and party Z.5230 Approx. 3 km NE Charles Point, Cox Peninsula, NT, 12°22.782’S, I30 0 38.371’E, 9-12 m, 23 Aug 2003, coll. Browne M Z.5233 off Dudley Point. Fannie Bay, Darwin Harbour, NT, 12°24.96’S, 130°48.83’E, 4-7 m, 4 Jun 2002, coll. Alvarez, B and party Z.5234 Raragala L, bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, I36°17.839’E, 17-20 m, 30 Mar 2004, coll. Alvarez, B and party Z.5736 Mandorah jetty, NW Cox Peninsula, Darwin Harbour, NT, I2°26.55’S, 130°46.05’E, 9-12 m, 5 Sep 2003, coll. Alvarez, B and party Z.5901 Moira Reef, Bynoe Harbour, NT, 12°30.799’S, 130°30.527’E, 5-8 m, 25 Jun 2003, coll. Alvarez, B and party Z.5902 Channel Rock, 4 km NE West Pi on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 12-24 m, 3 Sep 2002, coll. Alvarez, B and party Z.5903 Approx. 3 km NE Charles Point, Cox Peninsula, NT, 12°22.782’S, 130°38.371 ’E, 9-12 m, 23 Aug 2003, coll. Nguyen, H Z.5904 West Arm, 2.5 km N of Stokes Point, Darwin 1 larbour, NT, 12°31.300'S, 130°48.500’E, 4-5 nt, 3 Aug 2002, coll. Alvarez. B and party Z.5905 Channel Island, 100-400 nt N of bridge, Middle Ann, Darwin Harbour, NT, 12°33.09’S, 130°52.43’E, intertidal 0.02 m, 7 Nov 2006, coll. Alvarez, B Z.5906 Channel Island, 100-400 m N ofbridge. Middle Ann, Darwin Harbour, NT, 12°33.09’S, 130°52.43’E, intertidal 0.02 m, 7 Nov 2006, coll. Alvarez, B Z.5907 Lee Point. Darwin, NT, 12°20.538'S, 130°52.184’E, 9-12 m, 7 Aug 2003, coll. Nguyen, LI Z.5908 Moira Reef, Bynoe Harbour, NT, 12°30.799'S, I30°30.527’E, 5-8 m, 25 Jun 2003, coll. Nguyen, H 83 B. Alvarez and J. N. A. Hooper Z.5909 off Dudley Point, Fannie Bay, Darwin Harbour, NT, 12°24.96’S, 130°48.83’E, 4—7 nr, 4 Jun 2002, coll. Alvarez, B and party Z.5915 Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12°28.04’S., 130“49.77'E., 20 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Z.5923 Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12°28.04’S., 130°49.77’E., 19.5 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Z. 5925 Larrakeyah sewerage outfall. Darwin Harbour, NT, Australia, 12°28.04’S., 130°49.77’E., 19.5 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Z.5928 Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12°28.04’S., 130°49.77’E., 19.5 nr depth, 22 April 2009, coll. Sultana, S. SCUBA. Z.5948 Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12°29.17'S„ 130°47.19'E., 10-16 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA Z.5949 Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12°29.17’S., 130“47.I9’E., 10-16 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA Z.5950 Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12°29.17'S., 130°47.19’E„ 10-16 nr depth, 21 May 2009. coll. Alvarez, B and Sultana, S, SCUBA. Z.5965 East Point, Fannie Bay, Darwin Harbour, NT, Australia, 12°25.01'S., 130°48.88’E., 6 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA Z.5970 East Point, Fannie Bay, Darwin Harbour, NT, Australia, 12°25.01'S., 130°48.88’E., 6 m depth, 21 May 2009, coll. Alvarez. B and Sultana, S, SCUBA Z.5976 Stevens Rock. 1.25 km SETalc Head, off Cox Peninsula, Darwin Harbour, NT. 12°29.I88'S, 130°47.110’E, 8-14 m, 22 Aug 2003, coll. Alvarez, B and party Z.5977 Channel Rock, 4 km NE West Pt on Cox Peninsula. Darwin Harbour, NT, 12°24.94\S, 130°47.04’E, 13-16 m, 5 Sep 2003, coll. Alvarez, B and party Z.5978 Larrakeyah sewerage outfall, Darwin Harbour, NT, 12°28.04'S, 130°49.77’E, 5.8 m depth, 22 July 2010, coll. Sultana, S, SCUBA 84 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 21 : 85-105 New larval food plant associations for some butterflies and diurnal moths (Lepidoptera) from the Northern Territory and eastern Kimberley, Australia MICHAEL F. BRABY Museum and Art Gallery of the Northern Territory’, GPO Box 4646, Darwin NT 0801, and Research School of Biology’, Australian National University’, Canberra ACT0200, AUSTRALIA Present address: Biodiversity Conservation, Natural Resources, Environment, The Arts and Sport, PO Box 496, Palmerston NT0831, AUSTRALIA michael.braby@nt.gov.au ABSTRACT This paper documents 122 insect-plant associations for 10 families of butterflies/diurnal moths (Castniidae, Sessidae, Immidae, Geometridae, Hesperiidae, Papilionidae, Pieridae, Nymphalidae, Lycaenidae and Noctuidae (Agaristinae)) from the Northern Territory and eastern Kimberley, of which 76 associations are newly recorded for Australia and 46 are newly recorded for the Northern Territory and/or Western Australia. Of particular note are the first recordings of the families Lecythidaceae for Antltene (Lycaenidae), Santalaceae for Theclinesthes (Lycaenidae), Vitaceae and Rubiaceae for Periopta (Agaristinae), and Vitaceae for Radinocera (Agaristinae). For Cephrenes augiades and Borbo impar (Hesperiidae), the native larval food plants and natural breeding habitats in the Northern Territory are documented for the first time. For Candalidesdelospila, C. erinus and C. geminus (Lycaenidae), errors in previously reported larval food plants are corrected, and for two of these species (C. delospila, C. erinus) facultative ant associations are recorded for larvae for the first time. New attendant ant associations are also recorded for several lycaenids, notably Ogyris zosine and Theclinesthes miskini, in which the larvae have obligate relationships with ants. Keywords: butterfly, day-flying moth, insect-plant associations, larval food plant, Lepidoptera. INTRODUCTION The following catalogue of larval food plants for butterflies and day-flying moths is based on field observations and rearing of the early stages from the central arid zone and the ‘Top End’ of the Northern Territory (NT) and the eastern Kimberley of Western Australia (WA) by the author during the five year period, October 2005 to December 2010. The eastern Kimberley region extends from Pinkerton Range-Spirit Hills, NT, to north-western WA. The Lepidoptera covered here include the Castniidae, Sessidae, Immidae, Geometridae, Hesperiidae, Papilionidae, Pieridae, Nymphalidae, Lycaenidae and Noctuidae (Agaristinae). The new records are in addition to those reported in the review by Meyer (1996) for part of the region, and contribute to the growing body of knowledge of insect-plant associations for the Australian Lepidoptera as a whole (see Common 1990; Braby 2000 for review). The catalogue is presented in annotated form for each association: the (scientific and common) name of the lepidopteran appears first as a subheading in bold, followed by the larval food plant and voucher number, and then the relevant field observations. The field observations summarise details of locality, date, early stages and, in some cases, rearing data and other biological notes (e.g. attendant ants for Lycaenidae) where these are poorly documented. The new Lepidoptera-larvai food plant associations are arranged into two groups; firstly, those that comprise new records for Australia as a whole, and secondly, those that are new for the NT and/or WA. In many cases, samples of the early stages of Lepidoptera were preserved, photographed and/or reared to adulthood in captivity to confirm species level identification, and vouchers of these specimens are lodged in the Musuem and Art Gallery of the Northern Territory, Darwin (NTM) or the Australian National Insect Collection, Canberra (ANIC). In some cases, however, the early stages were not reared, and identifications of eggs, larvae or pupae were determined on the author’s field experience and knowledge of the relevant species. Nomenclature for butterflies follows Braby (2010), while that for moths follows Nielsen et at. (1996). The ants were identified by Alan Anderson. Voucher specimens of the ants have been lodged in the CSIRO Tropical Ecosystem Research Collection, Darwin. Botanical nomenclature follows a recent updated checklist of vascular plants published by the Northern Territory Herbarium (Short et at. 2011). Voucher numbers refer to plant specimens lodged in the Northern Territory Herbarium, Palmerston (DNA). Introduced, naturalised and ornamental plants are designated by an asterisk (*). For consistency, geocoordinates are given 85 M. F. Braby in, or have been converted to, decimal degrees, followed by datum (e.g. AGD66, WGS84 or GDA94). The level of precision of coordinates is not given, though in general the spatial area of sites sampled varied from 10-500 m radius, though was usually about 50-100 m. NEW LEPIDOPTERA LARVAL FOOD PLANT ASSOCIATIONS FOR AUSTRALIA The following catalogue of species comprises plants that, to the author’s knowledge, have not previously been documented as larval food plants for Australian Lepidoptera. CASTNIIDAE Synemonphaeoptilu Turner, 1906. No common name. Clirysopogon latifolins S.T.Blake (Poaceae). (vouchers M.F. Braby 19, 31,DNA). Caiman Creek, Cobourg Peninsula, NT (11.22504°S, 132.20236°E; WGS84), 4 February 2007, M.F. Braby & V. Ressner. Several females were observed at 1215-1245 hr ovipositing on Chtysopogon latifolius growing in savannah woodland that was regenerating after fire; numerous eggs were laid in soil at the base of thick stems/rhizomes of C. latifolius. Victoria Hwy, Pinkerton Range, NT (15.98698°S, 129.51213°E; WGS84), 1 February 2008, M.F. Braby & V. Kessner. Two empty pupal exuviae were collected protruding from soil at the base of separate tussocks of C. latifolius growing in savannah woodland on flat low-lying terrain adjacent to a sandstone outcrop; four pairs of adults were also collected mating around midday as they sat in copula on these and other nearby tussocks of C. latifolius. Two of the females had almost certainly emerged from the pupal exuviae. Synemon sp. ‘Roper River’. No common name. Chrysopogonfallax S.T.Blake. (Poaceae). (voucher M.F. Braby 51, DNA). Nathan River Ranger Station, Limrnen National Park, NT (15.57681°S, 135.42783°E; WGS84), 14 May 2009, M.F. Braby & N. Collier. Four pupal exuviae were collected protuding from soil close to clumps of Chrysopogon fallax growing in grassy open woodland on flat terrain in a valley between sandstone ridges/outcrops; one freshly emerged female was also collected at rest drying its wings within a clump of C. fallax. The habitat had been extensively burnt three days earlier. SESIIDAE Pseudosesia oberthuri (Le Cerf, 1916). No common name. Ampelocissus acetosa (F.Muell.) Planch. (Vitaceae). (not vouchered). Lee Point, Casuarina Coastal Reserve, NT (12.33166°S, 130.89444°E; WGS84), 28 March 2009, M.F. Braby & A. Rallies. Numerous larvae and pupae were collected from within the main stem of several vines of Ampelocissus acetosa, which grew in eucalypt open forest, and reared to adult in captivity. East Point, Darwin, NT (12.41140°S, 130.82500°E; WGS84), 31 January 2010, M.F. Braby. One pupal exuvia was collected protruding from the stem of A. acetosa growing about 1.5 m distant from where a freshly emerged male was perched, at 1055 hr, on a large leaf of a vine growing along the edge of coastal semi-deciduous monsoon vine-thicket. Ampelocissus frutescens Jackes. (Vitaceae). (not vouchered). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 11 April 2009, M.F. Braby & L.J. Aitchison. Numerous larvae were collected from within the stem of a single shrub of Ampelocissus frutescens which had not yet shed its leaves and which was growing at the base of a hill; several larvae were reared to adult in captivity. Most of the other short-lived aerial stems of this perennial vine were already senescent following onset of the dry season, suggesting that A. frutescens may not be a widely used food plant. IMMIDAE Birthana cleis (R. Felder & Rogenhofer, 1875). No common name. Dendrophthoe odontocalyx (F.Muell. ex Benth.) Ticgh. (Loranthaceae). (voucher M.F. Braby 29, DNA). Gurrumuru outstation, Gove Peninsula, NT (12.59302°S, 136.23194°E; WGS84), 21 June 2007, M.F. Braby. A cohort of 15 early instar larvae was recorded on a leaf of Dendrophthoe odontocalyx, a mistletoe parasitising the host tree Grevilleapteridifolia growing in savannah woodland; three larvae were reared to adult in captivity. The remains of an old cohort of Delias argenthona were evident on the same mistletoe clump (see below). HESPERDDAE Borbo impar lavinia (Waterhouse, 1932). Yellow Swift. Hymenachne acutigluma (Steud.) Gilliland. (Poaceae). (voucher M.F. Braby 41, DNA). Yellow Water, Jim Jim Creek, Rakadu National Park, NT (12.89689°S, 132.51749°E; WGS84), 20 April 2008, M.F. Braby, R. Nishida & L.J. Aitchison. Numerous larval shelters and adults were recorded on Hymenachne acutigluma growing in standing water of floodplain wetland. The site was revisited on 8 June 2008 and one mid-instar larva was collected from H. acutigluma. Taractrocera ina Waterhouse, 1932. No-brand Grass- dart. Cymhopogon procerus (R.Br.) Domin. (Poaceae). (voucher M.F. Braby 3, DNA). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 6 May 2006, M.F. Braby & D.A. Young. One late instar larva was collected 86 New Lepidoptera food plant associations from within its shelter on Cymbopogon procerus growing in low woodland at the summit of a rocky hill; the larva was subsequently reared to adult in captivity. The site was revisited on 6 March 2010 and two larvae were collected from C. procerus ; one developed directly and was reared to adult in captivity within a few weeks, the other remained dormant as a final instar and did not feed for six months until it was removed by hand from its shelter. Sorghum sp. (Poaceae). (not vouchered). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 6 March 2010, M.F. Braby & S. Kcates. Two larvae were collected from within their shelters on Sorghum sp., an annual grass growing in savannah woodland on the lower slope; one larva developed directly and was reared to adult in captivity within a few weeks, the other remained dormant as a final instar and did not feed for many months. Sorghum macrospermum E.D.Garber. (Poaccac). (not vouchered). Stuart Hwy, 10 km NW of Katherine, NT (14.40684°S, 132.19710°E; WGS84), 26 January 2009, M.F. Braby & V. Kessner. Six larvae were collected from within loose shelters on new growth of Sorghum macrospermum growing in degraded mixed vine-thicket-woodland on limestone; four larvae were reared to adult in captivity. Sorghum macrospermum is a localised annual grass endemic to the Katherine district (Lazarides et al. 1991). Suniana lascivia larrakia L.E. Couchman, 1951. Dark Grass-dart. Ischaemum australe R.Br. (Poaceae). (voucher M.F. Braby 60, DNA). El Questro Gorge, El Qucstro Wilderness Park, WA (16.02135°S, 128.02356°E; WGS84), 14 July 2009, M.F. Braby. A female was observed at 1050 hr depositing a single egg on a broad fresh blade of Ischaemum australe, which grew as a dense understorey grass along the edge of a creek. Adults were locally common at the site where /. australe grew in sunlit areas. Telicota colon argea (Plot/, 1883). Pale-orange Darter. *Paspalum scrobiculatum L. (Poaceae). (voucher M.F. Braby 77, DNA). Fogg Dam Conservation Reserve, NT (12.55769°S, 131.29701°E; WGS84), 29 May 2010. M.F. Braby, S. Keates & D. Bisa. One mid-instar larva was collected from within its shelter on Paspalum scrobiculatum growing in paperbark swampland and reared to the pupal stage. Cephrenes augiades (C. Felder, 1860) ssp. ‘Top End’ (M.F. Braby 354, ANIC). Orange Palm-dart. Carpentaria acuminata (H.Wendl. & Drude) S.T.Blake (Arecaceae). (not vouchered). Greenant Creek, Litchfield National Park, NT (13.20015°S, 130.69971°E; WGS84), 6 November 2010, M.F. Braby. A pupa was collected from within its shelter on Carpentaria acuminata growing in riparian wet monsoon forest, and a male emerged the following morning; males were commonly recorded between 1500 hr and 1600 hr as they perched on foliage of understorey trees 7-8 m above the ground in sunlit patches along the edge of the habitat where they appeared to establish mating territories. Wangi Falls, Litchfield National Park, NT (13.16305°S, 130.68166°E; WGS84), 7 November 2010, M.F. Braby & L.J. Aitchison. One late instar larva was collected from within its shelter on C. acuminata growing abundantly in wet monsoon forest; the larva was reared to adult in captivity. A female of this hesperiid was previously recorded at this location on 17 April 2006, but the larval food plant was not determined. Robin Falls, creek upstream of falls, NT (13.34877°S, 131.12622°E; WGS84), 29-30 April 2011, M.F. Braby & J.J. Armstrong. Three larvae were collected from within shelters on C. acuminata growing in riparian evergreen monsoon forest below a sandstone escarpment, one of which was reared to adult in captivity. Males were commonly recorded during mid-afternoon (1530-1600 hr) establishing territories in the mid-canopy along the western edge of the breeding habitat that received direct sunlight. Cephrenes trichopepla (Lower, 1908). Yellow Palm- dart. Livistona humilis R.Br. (Arecaceae). (not vouchered). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 21 February 2009, M.F. Braby & D.A. Young. A female was observed ovipositing on Livistona humilis growing in low woodland at the summit of a rocky hill. The site was revisited on 21 November 2009 and a late instar larva close to the prepupal stage was recorded within its shelter on L. humilis growing on the slope of a hill. On 6 March 2010 another larva in early instar was recorded on L. humilis growing at the summit of the hill. Livistona mariae F.Muell. (Arecaceae). (not vouchered). Palm Valley, Finke Gorge National Park, NT (24.03333°S, 132.69528°E; WGS84), 13 October 2005, M.F. Braby & L.J. Aitchison. Several early instar larvae were recorded from within their shelters on small individuals of Livistona mariae growing along Palm Creek; one female was reared to adult in captivity. Livistona nasmophila Dowe & D.L.Jones. (Arecaceae). (not vouchered). El Questro Gorge, El Questro Wilderness Park, WA (16.02135°S, 128.02356°E; WGS84), 14 July 2009, M.F. Braby & L.J. Aitchison. Several larvae were collected from within their shelters on Livistona nasmophila growing in riparian monsoon forest and reared to adult in captivity. Pentecost River, 4 km NW of El Questro Wilderness Park, WA (15.97504°S, 127.95461°E; WGS84), 2 April 2010, M.F. Braby & L.J. Aitchison. One final instar larva was recorded within its shelter on L. nasmophila growing in riparian woodland. Livistona victoriae Rodd. (Aracaceae). (not vouchered). 87 M. F. Braby Echidna Chasm, Purnululu National Park, WA (17.32272°S, 128.41183°E; WGS84), 2 July 2009, M.F. Braby & L.J. Aitchison. One empty egg shell, two larvae and numerous empty shelters were recorded on Livistona victoriae growing in depauperate monsoon forest along a gorge. Mini Palm Gorge, Purnululu National Park, WA (17.33083°S, 128.40916°E; WGS84), 2 July 2009, M.F. Braby & L.J. Aitchison. Two early instar larvae and one empty pupal shell were collected from within their shelters on L. victoriae growing in mixed monsoon forest-riparian woodland; one larva was reared to adult in captivity. PAPILIONIDAE Graphium eutypylus nyctimus (Waterhouse & Lyell, 1914). Pale Triangle. *Polyalthia longifolia (Sonn.) Thwaites. (Annonaceae). (not vouchered). Casuarina, Darwin, NT (12.366°S, 130.867°E; AGD66), 29 September 2010, M.F. Braby. Four larvae comprising one early instar and three late instars were recorded on new leaf growth of young ornamental trees of Polyalthia longifolia, which grew adjacent to Casuarina Shopping Centre. The site was revisited on 3 October 2010 and a further three mid¬ instar larvae were collected and reared to adult in captivity. PIERIDAE Eurema hecabe (Linnaeus, 1758). Large Grass- Yellow. * Senna alata (L.) Roxb. (Fabaceae). (not vouchered). Dhamiyaka outstation, 17.5 km SSE of Gapuwiyak, Gove Peninsula, NT (12.65666°S, 135.85833°E; WGS84), 16 June 2006, M.F. Braby & I. Morris. A female was observed ovipositing on a shrub (< 1 m high) o {'Senna alata growing near the edge of monsoon vine-forest; eggs were laid singly on the upperside of the leaves. *Senna obtusifolia (L.) H.S.lrwin & Bameby. (Fabaceae). (voucher M.F. Braby 35, DNA). Berrimah (CSIRO complex), Darwin, NT(12.41333°S, 130.92194‘ E; WGS84), 2 April 2008, M.F. Braby. Two females were observed at 1215 hr ovipositing on a low' shrub of Senna obtusifolia, which grew in degraded savannah woodland; several eggs and larvae were also recorded on the foliage of S. obtusifolia. Elodina padusa (Hewitson, 1853). Narrow-winged Pearl-white. Capparis lasiantha R.Br. ex DC. (Capparaceae). (voucher M.F. Braby 79, DNA). 12 km S of Dunmarra, NT (16.78726°S, 133.43216°E; WGS84),4 August 2010, M.F. Braby. One egg was collected from the underside of a mature leaf of a large scrambling vine of Capparis lasiantha, which grew into the canopy of mixed lancewood-vine-thicket; adults w ; ere also present in the habitat, but were not as numerous as during a previous visit on 16 October 2005. Elodina walkeri, the other species of Elodina that occurs in the NT, was not recorded at this site on either occasions, and its geographic range may not extend this far south in the Top End. Appias paulina ega (Boisduval, 1836). Yellow Albatross. Capparis sepiaria L. (Capparaceae). (voucher M.F. Braby 15, DNA). Black Point, Cobourg Peninsula, NT (11.15515°S, 132.14391°E; WGS84), 3 February 2007, M.F. Braby. A female was observed ovipositing on Capparis sepiaria growing in coastal monsoon vine-thicket; several eggs were laid singly on new shoots. Ceporaperimale (Donovan, 1805). Caper Gull. Capparis umbonata Lindl. (Capparaceae). (vouchers M.F. Braby 4, 64, DNA). Education campground, Litchfield National Park, NT (13.11361°S, 130.79805 U E; WGS84), 17 April 2006, M.F. Braby & L.J. Aitchison. A female was observed ovipositing on Capparis umbonata growing in savannah woodland on a gentle rocky slope; five mid-instar larvae were also recorded on the larval food plant. 1 km ENE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84375°S, 132.82558°E; WGS84), 20 December 2009, M.F. Braby. Four late instar larvae were recorded feeding on an almost leafless, spiny shrub of C. umbonata, which grew on sandy-loam at the base of an escarpment. The larvae were feeding together with those of Belenois java (see below). The site was revisited on 24 January 2010 and one final instar larva was recorded on C. umbonata. Deliasargenthonafragalactea (Butler, 1869). Scarlett Jezebel. Dendrophthoe odontocalyx (F.Muell. ex Benth.) Tiegh. (Loranthaceae). (voucher M.F. Braby 29, DNA). Gurrumuru outstation, Gove Peninsula, NT (12.59302°S, 136.23194°E; WGS84), 21 June 2007, M.F. Braby. One remnant pupal exuvia was found attached to a leaf of Dendrophthoe odontocalyx, a mistletoe parasitising the host tree Grevilleapteridifolia, growing in savannah woodland; several cast larval skins and larval damage to leaves were also evident. No other mistletoe species grew on the host tree or on adjacent eucalypt trees. A cohort of Birthana cleis was also present on the same mistletoe clump (see above). NYMPHALIDAE Danauspetilia (.Stoll, 1790). Lesser Wanderer. Cynanchum pedunculatum R.Br. (Apocynaceae). (voucher M.F. Braby 84, DNA). Nanguluwur Art site, Nourlangie Rock, Kakadu National Park, NT (12.84262°S, 132.81895°E; WGS84), 13 November 2010, M.F. Braby & J. Westaway. A female was observed at 1008 hr for several minutes searching and inspecting a vine of Cynanchum pedunculatum, which grew New Lepidoptera food plant associations amongst footslope sandstone boulders; a single egg was eventually laid on the underside of a leaf of the food plant. Euploea Sylvesterpelor Doubleday, 1847. Two-brand Crow. Parsonsia alboflavescens (Dennst.) Mabb. (Apocynaceae). (not vouchercd). Rocky Bay, 5 km SSE of Yirrkala, Gove Peninsula, NT (12.29583°S, 136.90305°E; WGS84), 3 July 2006, M.F. Braby. A female was observed at 1215 hr depositing a single egg on a vine of Parsonsia alboflavescens growing in mixed tall paperbark swampland with rainforest elements in the understorey; the larva was reared to mid-instar. Euploea corinna (W.S. Macleay, 1826). Common Crow. Marsdenia geininata (R.Br.) P.I.Forst. (Apocynaceae). (voucher M.F. Braby 1, DNA). 5.5 km NE of Adelaide River, NT (13.21222°S, 131.14750°E; AGD66), 4 June 2006, M.F. Braby & V. Ressner. A female was observed depositing a single egg on a vine of Marsdenia geminata growing in regenerating riparian forest along the bank of the Adelaide River. Parsonsia alboflavescens (Dennst.) Mabb. (Apocynaceae). (not vouchercd). Rocky Bay, 5 km SSE of Yirrkala, Gove Peninsula, NT (12.29583°S, 136.90305°E; WGS84), 30 August 2007, M.F. Braby, P. Wise & B. Marika. Five eggs and two early instar larvae were collected from Parsonsia alboflavescens growing in mixed tall paperbark swampland with rainforest elements in the undcrstorcy; the early stages were reared to the pupal or adult stage in captivity. Melanitis leda bankia (Fabricius, 1775). Evening Brown. *Cynodon radiatus Roth ex Roem. & Schult. (Poaceae). (voucher M.F. Braby 49, DNA). Nathan River Ranger Station, Liminen National Park, NT (15.57681°S, 135.42783°E; WGS84), 12 May 2009, M.F. Braby & N. Collier. Numerous larvae and several pupae were recorded on Cynodon radiatus, as well as on Imperata cylindrica (Poaceae) (see below), growing along the bank of a river; one pupa was reared to adult in captivity. Ischaemum australe R.Br. (Poaceae). (not vouchered). Buffalo Creek, Leanyer swamp, NT (12.35353°S, 130.90407°E; WGS84), 6 February 2010, M.F. Braby. Five mid-instar larvae were recorded feeding on Ischaemum australe growing in paperbark swampland adjacent to mangroves; the larvae were clustered into two groups on the underside of blades of /. australe. Hypocysta adiante antirius Butler, 1868. Orange Ringlet. *Sporobolus sp. (Poaceae). (voucher M.F. Braby 9, DNA). Berrimah (CSIRO complex), Darwin, NT (12.41333°S, 130.92194“E; WGS84), 23 February 2006, M.F. Braby. A female was observed at 1240 hr depositing a single egg on the underside of a blade of an undetermined introduced species of grass growing in savannah woodland; one late instar larva was also noted at rest on the same blade of Sporobolus sp., which showed signs of recent larval damage. Chloris sp. (Poaceae). (not vouchered). Lee Point, Casuarina Coastal Reserve, NT (12.33166°S, 130.89444°E; WGS84), 11 March 2006, M.F. Braby. A female was observed during the early afternoon depositing a single egg on the underside of a blade of Chloris sp. growing along the edge of coastal semi-deciduous monsoon vine-thicket. Arundinella nepalensis Trin. (Poaceae). (voucher M.F. Braby 61, DNA). King Edward River, WA (14.88834°S, 126.20372°E; WGS84), 11 July 2009, M.F. Braby & L.J. Aitchison. A female was observed at 1315 hr depositing a single egg on Arundinella nepalensis growing along the bank of a river in riparian paperbark woodland; the egg was laid on new fresh growth at the base of a large tussock of this grass. Ischaemum australe R.Br. (Poaceae). (voucher M.F. Braby 60, DNA). El Questro Gorge, El Questro Wilderness Park, WA (16.02135°S, 128.02356°E; WGS84), 14 July 2009, M.F. Braby & L.J. Aitchison. One late instar larva was recorded feeding on a blade of Ischaemum australe growing in a dry seasonal gully. LYCAENIDAE Hypochrysops apelles (Fabricius, 1775). Copper Jewel. Ceriops australis (C.T.White) Bailment, T.J.Sm. & J.A.Stoddart. (Rhizophoraceae). (voucher M.F. Braby 65, DNA). Buffalo Creek, Leanyer swamp, NT (12.35353°S, 130.90407°E; WGS84), 27 April 2008, M.F. Braby. Four larvae were collected from foliage of Ceriops australis growing in mangroves some distance from the landward edge and reared to adult in captivity; larvae were attended by black ants, Crematogaster sp. (species group C). Hypochrysops igitita erythrina (Waterhouse & Lyell, 1909). Fiery Jewel. Gardenia megasperma F.Muell. (Rubiaceae). (not vouchered). Litchfield National Park, NT (13.12555°S, 130.80305°E; WGS84), 15 April 2006, M.F. Braby & L.J. Aitchison. Two larvae were collected feeding on large, broad leaves of shrubs of Gardenia megasperma growing on a rocky sandstone plateau; the larvae were attended by numerous coconut ants, Papyrius sp. ( nitidus species group), and were reared to adult in captivity. Arhopala eupolis asopus Waterhouse & Lyell, 1914. Purple Oak-blue. Corymbia bella K.D.Hill & L.A.S.Johnson. (Myrtaceae). (voucher M.F. Braby 46, DNA). 89 M. F. Braby Gunlom, Kakadu National Park, NT (13.43555°S, 132.41565°E; WGS84), 28 August 2008, M.F. Braby & L. J. Aitchison. Two recent empty pupal exuviae and one parasitised pupa were recorded within curled leaves of a sapling (approx. 1.5 m high) of Corymbia bellci growing in open flood plain of South Alligator River; a pair of freshly emerged adults was also observed at 1700 hr mating on the same plant; nests of the attendant green-tree ant, Oecophylla smaragdina, were present. Adults oiArhopala micale were not detected on C. bella, or elsewhere at the site. Termincilia fenlinandiana Excell. (Combretaceae). (not vouchered). Nanguluwur Art site, Nourlangie Rock, Kakadu Nationar Park, NT (12.84262°S, 132.81895°E; WGS84), 7 December 2009, M.F. Braby & J.J. Armstrong. Several larvae were recorded on a shrub (1.5 m high) of Terminalia ferdinandiana growing in savannah woodland; larvae were attended by green-tree ants, Oecophylla smaragdina, and were noticed to have caused considerable damage to the foliage; adults alighted frequently on the foliage of T. ferdinandiana. Arhopala micale Blanchard, 118481. Shining Oak- blue. Brachychiton diversifolius R.Br. (Malvaceae), (not vouchered). Berrimah (CSIRO complex), Darwin, NT (12.41333°S, 130.92194°E; WGS84), 10 January 2007, M.F. Braby. Three eggs were collected from mature leaves of Brachychiton diversifolius growing in savannah woodland; one egg hatched and was reared to final instar on new fresh foliage of B. diversifolius. Sterculia quadrifida R.Br. (Malvaceae), (voucher M.F. Braby 90, DNA). Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 18 December 2010, M.F. Braby & J. Westaway. Five late instar larvae were recorded feeding on soft new leaf growth of medium sized trees of Sterculia quadrifida, which grew in monsoon vine-thicket on a low lateritic cliff above the beach; larvae were attended by Oecophylla smaragdina ants. The site was revisited on 20 December 2010 and four additional larvae, one pupa and a freshly emerged adult were recorded on S. quadrifida ; the pupa was located inside a silken shelter comprising a rolled mature leaf and attended by numerous ants. Ogyris amaryllis meridionalis (Bethune-Baker, 1905). Satin Azure. Diplatia grandibractea (F.Mucll. & Tate) Tiegh. (Loranthaccae). (voucher M.F. Braby 78, DNA). Near Lake Mary Ann, 5 km NNW of Tennent Creek, NT (19.60678°S, 134.20479°E; WGS84), 18 August 2010, M. F. Braby. Numerous eggs were recorded on haustoria of a mistletoe Diplatia grandibractea parasitising smooth white barked cucalypts growing in woodland with a ground layer of Triodia sp.; fresh larval feeding scars were also evident on the foliage and bracts of the mistletoe. Hypolycaena pltorbas phorbas (Fabricius, 1793). Black-spotted Flash. Decaisnina signata (F.Muell. ex Benth.) Tiegh. (Loranthaceae). (not vouchered). Black Point, Cobourg Peninsula, NT (11.15515°S, 132.14391 °E; WGS84), 23 April 2008, M.F. Braby & K. Nishida. One final instar larva was recorded feeding on foliage of the mistletoe Decaisnina signata parasitising a host tree Alstonia actinophylla (A.Cunn.) K.Schum. (Apocynaceae) growing in degraded monsoon vine-thicket; the larva was attended by three green-tree ants, Oecophylla smaragdina', one pupal exuvia was also collected from within a curled leaf shelter on the mistletoe. Anthene lycaenoidesgodeffroyi (Semper, 118791). Pale Ciliate-blue. Barringtonia acutangula (L.) Gaertn. (Lecythidaceae). (not vouchered). Oolloo Crossing, Daly River, NT (14.06946°S, 131.25052°E; WGS84), 12 April 2009, M.F. Braby & L.J. Aitchison. Several females were observed at 1415-1430 hr ovipositing on flower buds of Barringtonia acutangula growing in riparian monsoon forest on a steep sandy bank; one final instar larva was also collected feeding on soft new leaves of B. acutangula and reared to adult in captivity; the pupal duration was six days. Candalides margarita gilberti Waterhouse, 1903. Trident Pencil-blue. Decaisnina triflora (Span.) Tiegh. (Loranthaceae). (voucher M.F. Braby 12, DNA). Nawurlandja (Little Nourlangie Rock), Kakadu National Park, NT (12.85701°S, 132.79053°E; WGS84), 19 April 2008, M.F. Braby & K. Nishida. Two hatched eggs were recorded on new leaves of Decaisnina triflora', feeding damage from first instar larvae was also evident, but no larvae were detected. Candalides gemiitus E.D. Edwards & Kerr, 1978. Twin Dusky-blue. Cassytha filiformis L. (Lauraceae). (vouchers M.F. Braby 27, 082, DNA). Ovipositing females, eggs and larvae were frequently recorded (and subsequently reared to adult in captivity) on Cassytha filiformis growing in heathland or woodland on sandstone from many locations in western Arnhem Land, including Gubara Track, Nourlangie Rock and Gunlom of Kakadu National Park, and Katherine Gorge of Nitmiluk National Park, NT, during January, May, July, August, November and December 2007-2009. Cassytha capillaris Meisn. (Lauraceae). (not vouchered). Nourlangie Rock, Kakadu National Park, NT (12.85463°Sr 132.81677°E; VVGS84), 4 May 2008, M.F. Braby & L.J. Aitchison. Females were observed at 1410- 1412 hr ovipositing on Cassytha capillaris parasitising Triodia sp. growing in heathy woodland on sandstone; eggs were laid singly on flowers or new soft tendrils; two 90 New Lepidoptera food plant associations mid-instar larvae were also collected on C. capillaris and reared to adult in captivity. Candalides heathi heathi (Cox, 1873). Rayed Blue Prostanthera striatiflora F.Muell. (Lamiaceae). (voucher M.F. Braby 8, DNA). Larapinta Drive, approx. 95 km W of Hermannsburg, NT (23.88722°S, 131,90805°E; WGS84), 12 October 2005, M.F. Braby & L.J. Aitchison. Nine larvae of various instars were collected from Prostanthera striatiflora and reared to adult in captivity; eggs were also recorded singly on leaves, flowers and developing fruits; the pupal duration varied from six months to three years. Nesolycaena nnimclia (Tindale, 1922). Spotted Opal. Boronia laxa Duretto. (Rutaceae). (vouchers M.F. Braby 25, 026, DNA). Nourlangie Rock, Kakadu National Park, NT (12.85930°S, 132.81754°E; WGS84), 27 January 2007, M.F. Braby & L.J. Aitchison. A female was observed at 1140 hr ovipositing on a herb of Boronia laxa, which grew in heathy woodland on sandstone; eggs were laid singly on the underside of broad new leaves of separate plants; a second female was observed at 1315 hr depositing a single egg on the underside of a leaf of B. laxa. The site was revisited on 4 May 2008 and a female was observed at 1350 hr ovipositing on petals of B. laxa. I km ENE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84375°S, 132.82558°E; WGS84), 31 January 2009, M.F. Braby & L.J. Aitchison. A female was observed at 1330 hr depositing a single egg on a flower bud of B. laxa ; numerous eggs were also recorded on the underside of leaves and one mid-instar larva was noted feeding on the foliage of B. laxa. The site was revisited on 24 January 2010 and one pupal exuvia was recorded on the underside of a leaf of B. laxa. Prosotas dubiosa dubiosa (Semper, |1879|). Purple Line-blue. Millettia pinnata (L.) Panigrahi. (Fabaceae). (not vouchered). Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 15 September 2009, M.F. Braby. A female was observed ovipositing on flower buds of a small tree (approx. 5 m high) of Millettia pinnata, which grew in coastal parkland; numerous freshly emerged females were also observed settled on M. pinnata. Catopyrops florinda estrella (Waterhouse & Lyell, 1914). Speckled Line-blue. Mallotus nesophilus Muell.Arg. (Euphorbiaceae). (voucher M.F. Braby 22, DNA). East Point, Darwin, NT (12.41140°S, 130.82500°E; WGS84), 30 September 2006, M.F. Braby. Females were observed ovipositing on fresh leaves of Mallotus nesophilus growing along the edge of coastal semi-deciduous monsoon vine-thicket; additional eggs and early instar larvae were also recorded on the underside of leaves of M. nesophilus. The site was revisited on 8 October 2006 and an egg and first instar larva were collected from M. nesophilus and reared to adult in captivity. Dodonaea hispidula Endl. (Sapindaceae). (vouchers M.F. Braby 37, 054, DNA). Mt Cahill, Kakadu National Park, NT (12.86488°S, 132.70468°E; WGS84), 3 May 2008, M.F. Braby & L.J. Aitchison. Numerous eggs were recorded on the underside of leaves of Dodonaea hispidula growing in eucalypt woodland with a heath understorey on a rocky quartzite- sandstone slope; a female was also observed displaying oviposition behaviour (i.e. trailing and probing her abdomen on M. nesophilus), but she did not lay any eggs. Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131,03572°E; WGS84), 14 March 2009, M.F. Braby. Three larvae were collected from the underside of leaves of D. hispidula growing in low woodland at the summit of a rocky hill; the larvae were subsequently reared to adult in captivity. The site was revisited on 17 April 2010 and a female was observed at 1125 hr depositing a single egg on the underside of a new leaf of D. hispidula-, one mid-instar larva was also recorded on the underside of a leaf of D. hispidula. Amalia Gorge, El Questro Wilderness Park, WA (15.98124°S, 128.03708°E; WGS84), 2 April 2010, M.F. Braby & L.J. Aitchison. Empty egg shells and one late instar larva were recorded on foliage of D. hispidula growing near the entrance of the sandstone gorge. Theclinesthes sulpitius (Miskin, 1890). Samphire Blue. Tecticornia halocnemoides (Nees) K.A. Sheph. & Paul G. Wilson. (Amaranthaceac). (voucher M.F. Braby 2, DNA). 12.5 km SW of Palmerston, NT (12.56246°S, 130.90619°E; WGS84), 25 April 2007, M.F. Braby. Several egg shells and larval feeding damage were recorded on Tecticornia halocnemoides growing in saltmarsh. The site was revisited on 10 June 2006 and a female was observed at 1325 hr depositing a single egg on T. halocnemoides-, several additional eggs were also recorded on T. halocnemoides. Theclinesthes miskini miskini (T.P. Lucas, 1889). Wattle Blue. Acacia difficilis Maiden. (Fabaceae). (voucher M.F. Braby 5, DNA). Litchfield National Park,NT(13.12555°S, 130.80305°; WGS84), 15 April 2006, M.F. Braby & L.J. Aitchison. Six prepupae and pupae were collected from foliage of Acacia difficilis growing on a rocky sandstone plateau; the early stages were attended by meat ants, Iridomyrmex reburrus, and were subsequently reared to adult in captivity. 1 km ENE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84375°S, 132.82558°E; WGS84), 24 January 2010, M.F. Braby & L.J. Aitchison. All early stages were recorded in abundance on A. difficilis growing in a sandstone gully; larvae and pupae were attended by small brown ants, Iridomyrmex pallidus. 91 M. F. Braby Acacia platycaipa F.Muell. (Fabaceae). (not vouchered). Gurrandalng, Keep River National Park, NT(15.87478°S, 129.05144°E; WGS84), 2 February 2008, M.F. Braby & V. Kessner. Numerous eggs, three larvae and one pupa were recorded on several shrubs of Acacia platycaipa ; larvae were attended by swarms of meat ants, Iridomyrmex sp. Corymbia bella K.D. Flill & L.A.S. Johnson. (Myrtaceae). (not vouchered). Flora River Nature Park,NT(14.75744°S, 131.59694°E; WGS84), 31 January 2008, M.F. Braby & V. Kessner. Several larvae were recorded on a sapling of Coiyitibia bella growing in open vine-thicket on limestone; larvae were attended by numerous meat ants, Iridomyrmex sp. Corymbia ferruginea (Schauer) K.D. Hill & L.A.S. Johnson. (Myrtaceae). (not vouchered). Cabbage Gum, Charles Darwin University, Katherine, NT (14.39932°S, 132.14186°E; WGS84), 25 January 2009, M.F. Braby & V. Kessner. All stages were recorded on a sapling (approx. 1 m high) of Corymbia ferruginea growing in monsoon vine-thicket on limestone karst; larvae were attended by numerous sugar ants, Camponotus crozieri, at night as well as during daytime. Santalum lanceolatum R.Br. (Santalaceae). (voucher M.F. Braby 36, DNA). Jasper Gorge, approx. 58 km SW of Victoria River Roadhouse, NT (16.03119°S, 130.80275°E; WGS84), 21 March 2008, M.F. Braby & V. Kessner. A female was observed at 1450 hr ovipositing on a tall shrub (approx. 3 m high) of Santalum lanceolatum, which grew in woodland along a dry rocky sandstone seasonal gully; numerous eggs were also present on new leaf growth; one pupa was found at the base of the stem where it was well protected beneath debris; a pupal exuvia was also recorded on the upperside of a leaf of S. lanceolatum. Jamides phased (Mathew, 1889). Purple Cerulean. Sesbania simpliciuscula F.Muell. ex Benth. (Fabaceae). (voucher M.F. Braby 44, DNA). Escarpment Walk, Gregory (Jutpurra) National Park, NT (15.61325°S, 131.11597°E; WGS84), 16 March 2008, M.F. Braby & V. Kessner. Females were observed ovipositing on Sesbania simpliciuscula growing in savannah woodland at the base of an escarpment; numerous eggs were also recorded on flower buds of S. semipliciuscula. Catochrysops panormus platissa (Herrich-Schiiffer, 1869). Pale Pea-blue. Sesbania simpliciuscula F.Muell. ex Benth. (Fabaceae). (voucher M.F. Braby 45, DNA). Duncan Road, Hicks Creek crossing, NT (16.19774°S, 129.05383°E; WGS84), 19 March 2008, M.F. Braby & V. Kessner. A female was observed ovipositing on Sesbania simpliciuscula, which grew in abundance along a roadside. Duncan Road, Rosewood Station, NT (16.46031°S, 129.03348°E; WGS84), 19 March 2008, M.F. Braby & V. Kessner. Several females were observed ovipositing on S. simpliciuscula, which grew in abundance along a roadside. Zizeeria karsandra (Moore, 1865). Spotted Grass- blue. Tribulopis bicolor F.Muell. (Zygophyllaceae). (voucher M.F. Braby 75, DNA). Intersection of Buchanan Hwy and Victoria Hwy, 20 km SE of Timber Creek, NT (15.74320°S, 130.64830°E; WGS84), 1 April 2010, M.F. Braby & L.J. Aitchison. Several eggs were recorded on the underside of leaves of Tribulopis bicolor growing in a disturbed roadside verge in savannah woodland; numerous adults were observed flying in close proximity or settling on T. bicolor. Famegana alsulus alsulus (Herrich-Schiiffer, 1869). Black-spotted Grass-blue. Vigna vexillata (L.) A. Rich. (Fabaceae). (voucher M.F. Braby 6, DNA). Magnetic Termite Mounds, Litchfield National Park. NT (13U0500°S, 130.84250°E; WGS84), 15 March 2006, M.F. Braby & D.A. Young. A female was observed during mid-afternoon ovipositing on a twining creeper of Cigna vexillata, which grew in savannah woodland; numerous additional eggs were also present on tendrils of V. vexillata. NOCTU1DAE: AGARISTINAE Agarista agricola biformis Butler, 1884 (Fig. 1). Painted Day-moth. Leea rubra Blume ex Sprcng. (Vitaceae). (voucher M.F. Braby 83, DNA). Ml Bundy, Mary River National Park, NT (12.85990°S, 131.60281 °E; WGS84), 30 October 2010, M.F. Braby. A female was observed at 1415 hr ovipositing on Leea rubra growing along the edge or ecotone of decidous monsoon vine-thicket during overcast conditions with light rain; a second female was subsequently observed at 1600 hr actively searching L. rubra on which to lay eggs during bright sunny conditions; eggs were bright green and laid on stems and dry leaf bracts, and hatched five days later; four early instar larvae were also collected from the foliage of three separate shrubs and reared to adult in captivity on L. rubra, as well as on L. indica (Burm.f.) Merr. The site was revisited on 14 November 2010 and a further four early instar larvae were collected from L. rubra growing in the ecotone or in savannah woodland adjacent to decidous monsoon vine-thicket; one larva was reared to adult in captivity. Comocrus behri (Angas, 1847) (Fig. 2). Mistletoe Day-moth. Amyema maidenii (Blakely) Barlow. (Loranthaceae). (not vouchered). Alice Springs Desert Park, Alice Springs, NT (23.700°S. 133.833°E; WGS84), 13 Febaiary 2010, M.F. Braby & L.J. Aitchison. Three larvae were collected from clumps of the mistletoe Amyema maidenii that were parasitising Acacia aneura and reared to adult in captivity; a final instar larva was also collected crawling rapidly along the ground 92 New Lepidoptera food plant associations beneath the mulga host tree in search of a pupation site. Several adults were present at the breeding site. Amyema sanguined (F.Muell.) Danser. (Loranthaceae). (not vouchered). Limbunya Station, Victoria River District, NT (17.52339°S, 130.05701 °E; WGS84), 25 July 2010, M.F. Braby. A cohort of nine late instar larvae was recorded feeding openly, but not gregariously, on a single clump of the mistletoe Amyema sanguinea that was parasitising the host tree Corymbia sp. growing in eucalypt open woodland with a ground layer of Triodia sp. on red sandy soil; six larvae were collected, two of which were reared to adult in captivity. Cruria tropica (T.P. Lucas, 1891) (Fig. 3) Typhonium flagelliforme (Lodd.) Blume. (Araceae). (voucher M.F. Braby 40, DNA). Howard Springs Nature Reserve, NT (12.45240°S, 131.05347°E; WGS84), 16 and 24 February 2008, 8 and 24 March 2008, M.F. Braby. Several females were observed during mid-afternoon (1430-1530 hr) during the wet season ovipositing on Typhonium flagelliforme, a seasonal geophytic herb, which grew in paperbark-pandanus swamp adjacent to wet evergreen monsoon vine-forest; eggs were laid singly on the underside of leaves; numerous larvae were also collected from T. flagelliforme and reared to adult in captivity. Idalima aethrias (Turner, 1908). No common name. (Fig. 4) Hibbertia cistifolia R.Br. ex DC. (Dilleniaceae). (voucher M.F. Braby 23, DNA). Caiman Creek, Cobourg Peninsula, NT (11,22504°S, 132.20236°E; WGS84), 22 February 2007, M.F. Braby & D.A. Lane. One late instar larva was recorded on the foliage of Hibbertia cistifolia growing in savannah woodland. Hibbertia brevipedunculata Toelken. (Dilleniaceae). (voucher M.F. Braby 21, DNA). 3 km N of Berry Springs, NT (12.67554°S, 131,00994°E; WGS84), 14 January 2007, M.F. Braby. Two females were observed at 1440 hr and 1535 hr ovipositing on the foliage of Hibbertia brevipedunculata or on nearby objects (i.e. ground, leaves of other plants) in savannah woodland. The site was revisited on 2 November 2008 and 11 larvae in various instars were recorded on the larval food plant, several of which were reared to adult in captivity. This food plant was only recently described, and is listed as Hibbertia sp. Mt Finniss in Kerrigan & Albrecht (2007) and was vouchered in DNA under that name. Idalima leonora (Doubleday, 1846). No common name. (Fig. 5) Hibbertia brownii Benth. (Dilleniaceae). (voucher M.F. Braby 24, DNA). Caiman Creek, Cobourg Peninsula, NT (11.22364°S, 132.21402°E; WGS84), 21 February 2007, M.F. Braby & D.A. Lane. Females were observed at 1520 hr and 1745 hr ovipositing on Hibbertia brownii growing in savannah woodland; eggs were laid singly on flower buds or dead twigs; one mid-instar larva was collected from the underside of a leaf and reared to the pupal stage. The site was revisited on 15 March 2007 and females were observed at 1543 hr, 1550 hr and 1655 hr ovipositing on H. brownii; empty egg shells were also located on H. brownii; all eggs were laid singly, mostly on flower buds or sometimes on leaf bracts of developing flower spikclcts and on new leaf shoots at the apex of a branch of H. brownii. Hibbertia candicans Benth. (Dilleniaceae). (voucher M.F. Braby 67, DNA). 1 km ENE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84375°S, 132.82558°E; WGS84), 20 December 2009, M.F. Braby. Two larvae were collected from Hibbertia candicans growing in sandy-loam at the base of an escaipment; one larva was reared to adult in captivity. 0.4 km NE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84535°S, 132.82072°E; WGS84), 24 January 2010, M.F. Braby & L.J. Aitchison. Two early instar larvae and one late instar larvae were collected from separate shrubs of //. candicans growing in open woodland on a sandstone plateau; one larva was reared to adult in captivity. 0.6 km W of Gubara Track, Nourlangie Rock, Kakadu National Park, NT (12.83721°S, 132.84970°E; WGS84), 27 November 2010, M.F. Braby. One final instar larva was collected from II. candicans and reared to adult in captivity. Periopta ardescens (Butler, 1884). No common name. (Fig. 6) Ampelocissus frutescens Jackes. (Vitaceae). (vouchers M.F. Braby 57, 058, DNA). BullitaRd, 11 km SSEofTimber Creek, NT (15.75712°S, 130.51135°E; WGS84), 10 December 2008, M.F. Braby & V. Kessner. Numerous larvae were recorded feeding on new regenerating leaves of Ampelocissus frutescens growing in savannah woodland; several larvae were reared to adult in captivity; most larvae developed directly and emerged a few weeks later, but others emerged one or two years later. The larvae occurred together with those of Radinocera vagata (see below). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 12 December 2008, M.F. Braby. One late instar larva was collected from A. frutescens growing in woodland at the base of a hill and reared to adult in captivity. Ampelocissus acetosa (F.Muell.) Planch. (Vitaceae). (not vouchered). Buffalo Creek, Lee Point, NT (12.33639°S, 130.90389°E; WGS84), 28 November 2009, M.F. Braby. Seven larvae in various instars were collected from Ampelocissus acetosa growing in savannah woodland regenerating after dry season bums and pre-monsoon storms; a few larvae were reared to adult in captivity. The larvae occurred together with those of Radinocera vagata (see below). 93 M. F. Braby Figs 1-8. Larvae ofNottuidae: Agaristinae from the Northern Territory discussed in this paper: 1, Agarista agricola on Leea rubra, Mt Bundy. Mary River National Park; 2, Comocrus behri on Amyema sanguined, Limbunya Station, Victoria River District; 3, Cruria tropica on Typhonium flagelliforme, Howard Springs Nature Reserve; 4, Idalima aethrias on llibbertia brevipedunculata. Berry Springs; 5, Idalima leonora on Hibbertia brownii. Caiman Creek, Cobourg Peninsula; 6, Periopta ardescens on Ampelocissus acetosa, Buffalo Creek, Lee Point; 7, Periopta diverse on Spermococephalloides, Leanyer, Darwin; 8, Radinocera vagata on Ampelocissus acetosa, Buffalo Creek, Lee Point. Photos by M.F. Braby. 94 New Lepidoptera food plant associations Nanguluwur Art site. Kakadu National Park, NT (12.84262°S, 132.81895°E; WGS84), 7 and 19 December 2009, M.F. Braby & J.J. Armstrong. Females were observed at 1730 hr and 1845-1850 hr (just before sunset) ovipositing on A. acetosa growing in woodland near the base of an escarpment; numerous larvae in various instars were also recorded on A. acetosa. Periopta diversa (Walker, 11865]). No common name. (Fig. 7) Spermococe phalloides Harwood. (Rubiaceae). (not vouchercd). Leanyer Recreation Park, Darwin, NT (12.39055°S, 130.91167°E; WGS84), 21 December 2005, 5, 20 and 21 January 2006, M.F. Braby. Numerous larvae were recorded on Spermococe phalloides growing in disturbed grassland, several of which were reared to adult in captivity. Oldenlandia sp. (Rubiaceae). (vouchers M.F. Braby 10, 11, DNA). Leanyer Recreation Park, Darwin, NT (12.39055°S, 130.91167°E; WGS84), 4, 20 and 21 January 2006, M.F. Braby. Numerous females were observed at 1710-1855 hr (within a few hours of dusk) ovipositing on Oldenlandia sp., as well as on other small herbs, including grass blades, growing in disturbed grassland; eggs were laid singly; several larvae were also collected from Oldenlandia sp. and reared to adult in captivity. Holmes Jungle National Park,NT(12.400°S, 130.916°E; AGD66), 6 February 2010, M.F. Braby. Several females were observed at 1720-1740 hr ovipositing on Oldenlandia sp. growing in a disturbed area along a track; eggs were laid singly and directly onto leaves of Oldenlandia sp.; two larvae were also recorded on Oldenlandia sp. Radinocera vagata (Walker, 1865). No common name. (Fig. 8) Ampelocissus acetosa (F.Muell.) Planch. (Vitaceae). (voucher M.F. Braby 17, DNA). Caiman Creek, Cobourg Peninsula, NT (11.22504°S, 132.20236°E; WGS84), 7 and 22 February 2007, M.F. Braby & V. Kessner. Three larvae were collected from vines of Ampelocissus acetosa growing in savannah woodland; two larvae were reared to adult in captivity. Lee Point, Casuarina Coastal Reserve, NT (12.33166°S, 130.89444°E; WGS84), 16 February 2008, M.F. Braby. Two late instar larvae were recorded on a vine of A. acetosa growing around Corymbia bella in parkland adjacent to coastal semi-deciduous monsoon vine-thicket. Buffalo Creek, Lee Point, NT (12.33639°S. 130.90389°E; WGS84), 28 November 2009, M.F. Braby. Ten larvae in various instars were collected from A. acetosa growing in savannah woodland that was regenerating after a dry season burn and pre-monsoon storms; all larvae were feeding singly, with a maximum of two per vine; several larvae were reared to adult in captivity. The early stages occurred together with those of Periopta ardescens (see above). Ampelocissus frutescens Jackes. (Vitaceae). (voucher M.F. Braby 58, DNA). BullitaRd, 11 km SSE of Timber Creek, NT (15.75712°S, 130.51135°E; WGS84), 10 December 2008, M.F. Braby & V. Kessner. Six larvae were collected feeding openly on Ampelocissus frutescens growing in savannah woodland; most larvae were on separate plants and several were reared to adult in captivity. The larvae occurred together with those of Periopta ardescens (see above). 9 km SE ofTimber Creek, NT (15.57325°S, 130.51804°E; WGS84), 10 December 2008, M.F. Braby & V. Kessner. Several larvae were recorded on A. frutescens growing in savannah woodland. Cayratia maritime Jackes. (Vitaceae). (voucher M.F. Braby 14, DNA). 2 km ENE of Black Point, Cobourg Peninsula, NT (11.15015°S, 132.16125°E; WGS84), 13 March 2007, M.F. Braby. One late instar larva was collected from Cayratia maritima growing in paperbark woodland adjacent to a swamp and reared to adult in captivity; the pupal duration was approx, seven months. Buffalo Creek, Lee Point, NT (12.34497°S, 130.90578°E; WGS84), 8 March 2008, M.F. Braby. Two late instar larv ae were recorded on a vine of C. maritima, which grew around the trunk of a dead Acacia sp. along the edge of a paperbark swamp. The site was revisited on 7 November 2010 and two early instar larvae were collected from new leaf growth of C. maritima growing on the trunk of a Melaleuca sp. tree and reared to adult in captivity. Cayratia trifolia (L.) Domin. (Vitaceae). (voucher M.F. Braby 89, DNA). Mt Bundy, Mary River National Park, NT (12.85990°S, 131.60281°E; WGS84), 30 October 2010, M.F. Braby. Four early instar larvae were collected from new growth of Cayratia trifolia growing on the trunk of a cucalypt in savannah woodland and reared to adult in captivity. NEW LEPIDOPTERA LARVAL FOOD PLANT ASSOCIATIONS FOR THE NORTHERN TERRITORY AND/OR NORTHERN WESTERN AUSTRALIA The following catalogue of species comprises plants that have previously been recorded as larval food plants from eastern Australia (e.g. Common 1990; Braby 2000) but, to the author’s knowledge, have not been previously documented for the NT and/or the Kimberley region of western NT and north-western WA. GEOMETRIDAE Dyspltaitia numana (Cramer, 1779). Six O’clock Moth. Carallia brachiata (Lour.) Merr. (Rhizophoraceae). (not vouchered). 95 M. F. Braby Ovipositing females and/or the early stages were frequently recorded on Carallia brachiata growing in riparian monsoon forest and semi-deciduous monsoon vine- thicket from many locations, including Darwin. Litchfield National Park, Beatrice Hill, Kakadu National Park and Gove Peninsula. Females were observ ed ovipositing during the late afternoon (1540-1745 h); eggs were laid on the underside of mature leaves and stems of C. brachiata ; larvae fed solitarily on young soft foliage and pupated within a shelter made from two or more mature leaves of C. brachiata. HESPERIIDAE Chaetocneme clenitza (Hewitson, 1867). Ornate Dusk-flat. Planchonia careya (F.Muell.) Kunth. (Lecythidaceae). (not vouchered). Black Point, Cobourg Peninsula, NT (11.15515°S, 132.14391°E; WGS84), 19 February 2007, M.F. Braby & D.A. Lane. One first instar larva was collected from a small regenerating sapling of Planchonia careya growing in disturbed savannah woodland and reared to adult by DAL. Berrimah (CS1RO complex), Darwin, NT (12.41333°S, 130.92194°E; WGS84), 11 March 2008, M.F. Braby. A freshly emerged male and a pupal shelter were collected from a small tree of Planchonia careya growing in degraded savannah woodland; the adult perched upside-down on the underside of a leaf at midday. Taractrocera sp. ( ina Waterhouse, 1932, or anisomorpha Lower, 1911). Grass-dart. Eulalia aurea (Bory) Kunth. (Poaceae). (voucher M.F. Braby 42, DNA). Standley Chasm, West Macdonnell Ranges, NT (23.72150°S, 133.46982°E; WGS84), 4 March 2007, M.F. Braby & C.M. Palmer. Numerous larvae were recorded within their tubular shelters on Eulalia aurea growing in water or damp areas in riparian eucalypt open forest. Adults of both species were present in the habitat and either may have been using this native grass. Taractrocera sp. (ina or anisomorpha). Grass-dart. *Cenchrus ciliaris (Kuntze) L. (Poaceae). (not vouchered). Standley Chasm, West Macdonnell Ranges, NT (23.72150°S, 133.46982°E; WGS84), 2 March 2007, M.F. Braby & C.M. Palmer. Several eggs and larvae were recorded on Cenchrus ciliaris growing in riparian eucalypt open forest. Adults of both species were present in the habitat and either may have been using this invasive, introduced, perennial tussock grass. Taractrocera sp. (ina or anisomorpha). Grass-dart. *Paspalum conjugation Bcrgius. (Poaceae). (voucher M.F. Braby 43, DNA). Standley Chasm, West Macdonnell Ranges, NT (23.72150°S, 133.46982°E; WGS84), 4 March 2007, M.F. Braby & C.M. Palmer. Numerous larvae were recorded within their tubular shelters on the broad-leaved tufted perennial grass Paspalum conjugation, which grew in water or damp areas in riparian eucalypt open forest. Adults of both species were present in the habitat and either may have been using this introduced grass. Telicota colon argea (Plotz, 1883). Pale-orange Darter. Ischaemum australe R.Br. (Poaceae). (not vouchered). Amalia Gorge, El Questro Wilderness Park, WA (15.98124°S, 128.03708°E; WGS84), 15 July 2009, M.F. Braby. Three females were observed during mid- afternoon ovipositing on Ischaemum australe growing along a creek; one larva was also collected from within its shelter on I. australe and reared to adult in captivity. Cephrenes trichopepla (Lower, 1908). Yellow Palm- dart. Livistona benthamii F.M.Bailey. (Arecaceae). (not vouchered). Cahills Crossing, East Alligator River, Kakadu National Park, NT (12.42542°S, 132.96458°E; WGS84), 6 November 2009, M.F. Braby & L.J. Aitchison. Fourteen larvae were recorded from within their shelters on Livistona benthamii growing along the edge of riparian monsoon forest; several larvae were reared to adult in captivity. PAPILIONIDAE Graphium eurypylus nyctimus (Waterhouse & Lyell, 1914). Pale Triangle. Miliusa traceyi Jessup. (Annonaceae). (voucher M.F. Braby 85, DNA). Mt Bundy, Mary River National Park, NT (12.85990°S, 131.60281 °E; WGS84), 28 November 2010, M.F. Braby. Two females were observed at 1250 hr for several minutes ovipositing on shrubs of Miliusa traceyi growing along the edge of semi-deciduous monsoon vine-thicket; numerous eggs were laid on the underside of leaves, including new growth as well as older mature leaves; two early instar larvae were also noted feeding on the new soft leaves of M. tracei. Papilio aegeus aegeus (Donovan, 1805). Orchard Swallowtail. *Citrus sp. (Rutaceac). (not vouchered). Nhulunbuy, Gove Peninsula, NT (12.17805°S, 136.78361°E; WGS84), 6 September 2007, M.F. Braby & P. Wise. Two larvae were collected on Citrus sp. growing in a residential garden and reared to adult in captivity; the larvae pupated on 13 September 2007 and emerged as adults 11 days later. The site was revisited on 24 June 2008 and a further three larvae were collected from Citrus sp. and reared to adult in captivity; the larvae pupated the following month and emerged 13 days later. Papilio fuscus canopus Westwood, 1842. Fuscous Swallowtail. *Citrus sp. (Rutaceac). (not vouchered). Yirrkala, Gove Peninsula, NT (12.25611°S, 136.89778°E; WGS84), 26 January 2008, M.F. Braby & P. Wise. Numerous New Lepidoptera food plant associations eggs and several first instar larvae were collected from new leaf growth of Citrus sp. growing in an orchard adjacent to monsoon vine-thicket; 15 larvae were collected and reared in captivity, with adults emerging up to 10 months later. Wanguri, Darwin, NT (12.37308°S, 130.88657°E; WGS84), 29 June 2008, M.F. Braby. One pupal exuvia was recorded on a stem of Citrus sp. growing on a nature strip adjacent to a residential garden. Papilio demoleus stiieneliis W.S. Macleay, 1826. Chequered Swallowtail. Cullen cinereum (Lindl.) J.W. Grimes. (Fabaceae). (not vouchered). Victoria River crossing, approx. 2 km S of Victoria Hwy, NT (15.63072°S, 131.13261°E; WGS84), 4 April 2010, M.F. Braby & L.J. Aitchison. Several females were observed ovipositing on seedlings of Cullen cinereum growing in open dry floodplain near the Victoria River. PIER1DAE Catopsiliapomona (Fabricius, 1775). Yellow Migrant. * Cassia fistula L. (Fabaceae). (not vouchered). Eggs and larvae were frequently recorded on Cassia fistula, an ornamental tree in suburban areas throughout Darwin, during October to December when the plant produces buds, flowers and new leaf growth during the pre-monsoon period. Alice Springs, NT(23.700°S, 133.866°E; GDA94), 14 February 2010, M.F. Braby. Six final instar larvae were recorded feeding on new foliage of C. fistula growing in the Central Buisness District. Catopsilia scylla etesia (Hewitson, 1867). Orange Migrant. Senna leptoclada (Benth.) Randell. (Fabaceae). (voucher M.F. Braby 69, DNA). Near Twin Falls, Kakadu National Park, NT (13.26680°S, 132.81320°E; WGS84), 7 November 2009, M.F. Braby & L.J. Aitchison. A female was observed at 1245 hr ovipositing on Senna leptoclada (approx. 1.5 m high) growing in woodland on sandy soil; three eggs were laid singly on new soft terminal leaf growth, two of which were collected and reared to adult in captivity. Eurema Iterla (W.S. Macleay, 1826). Pink Grass- yellow. Chamaecrista mimosoides (L.) Greene. (Fabaceae). (voucher M.F. Braby 76, DNA). 1.6 km NNW of Pine Creek, NT (13.80997°S, 131,82852°E; WGS84), 1 May 2010, M.F. Braby & L.J. Aitchison. A female was observed at 1445 hr ovipositing on a herb (150-200 mm high) of Chamaecrista mimosoides, which grew in the ground layer of savannah woodland; two eggs were laid on the upperside of pinnae of separate plants. Eurema alitha novaguineensis Shriozu & Yata, 1982. Scalloped Grass-yellow. Galactia tenuiflora (Willd.) Wight & Am. (Fabaceae). (vouchers M.F. Braby 32, 33, 55, DNA). Town Lagoon, Nhulunbuy, Gove Peninsula, NT (12.17805°S, 136.78361 °E; WGS84), 27 January 2008, M.F. Braby & L.J. Aitchison. A female was observed at 1140 hr ovipositing on leaves of a trailing creeper of Galactia tenuiflora, which grew along the edge of paperbark swamp adjacent to monsoon vine-thicket; numerous additional eggs were also present on leaves of G. tenuiflora. Coolibah Station, Victoria River District, NT (15.58340°S, 130.97647°E; WGS84), 18 March 2008, M.F. Braby & V. Kessner. Females were observed at 1000 hr, 1109 hr and 1110 hr ovipositing on a creeper of G. tenuiflora, which grew over rock scree on a sheltered slope in monsoon vine-thicket; all the eggs were laid singly on new soft leaf growth; additional eggs were also present on leaves of the larval food plant. Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131,03572°E; WGS84), 21 February 2009, M.F. Braby & D.A. Young. A female was observed at 1100 hr ovipositing on a leaf of a twiner of G. tenuiflora, which grew in eucalypt woodland on a rocky slope of a hill; additional eggs were also present on new leaves; several early instar larvae were also collected on the new leaf growth and reared to adult in captivity; the larvae pupated the following month and emerged as adults five or six days later. Eurema hecabe (Linnaeus, 1758). Large Grass- Yellow. Sesbania cannabina (Retz.) Poir. (Fabaceae). (voucher M.F. Braby 72, DNA). Rapid Creek, Darwin, NT (12.38083°S, 130.86462°E; WGS84), 20 March 2010, M.F. Braby. One late instar larva was recorded on Sesbania cannabina, which grew in profusion in a disturbed area along the edge of mangroves; numerous females were also observed alighting on the larval food plant but they did not appear to oviposit. The early stages occurred together with those of Theclinesthes miskini (see below). Breynia cerntta (Poir.) Muell.Arg. (Phyllanthaceae). (voucher M.F. Braby 7, DNA). Eggs and larvae were frequently recorded (and subsequently reared to adult in captivity) on new soft leaf growth of Breynia cernua growing in monsoon vine-forest or riparian monsoon forest from many locations, including Darwin, Mary River Park and Fogg Dam Conservation Reserve, usually from November to June (i.e. wet season- early dry season). Belenoisjava teutonia (Fabricius, 1775). Caper White. Capparis umbonata Lindl. (Capparaceae). (voucher M.F. Braby 64, DNA). Leanyer, Darwin, NT (12.37500°S, 130.89667°E; WGS84), 14 December 2005, M.F. Braby. Several larvae and numerous pupae (approx. 200) were recorded on a small 97 M. F. Braby tree of Capparis umbonata growing in parkland, and a series was reared to adult in captivity. The site was revisited on 17 October 2007 and more than 100 late instar larvae and approx. 20 pupae were recorded on the same tree. The tree was subsequently destroyed by Tropical Cyclone Helen on 5 January 2008. 1 km ENE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84375°S, 132.82558°E; WGS84), 20 December 2009, M.F. Braby. Two late instar larvae were recorded on an almost leafless, spiny shrub of C. umbonata, which grew on sandy-loam at the base of an escarpment. The larvae were feeding together with those of Cepora perimale (see above). NYMPHALIDAE Danaus petilia (Stoll, 1790). Lesser Wanderer. Cynanchum pedunculatum R.Br. (Apocynaceae). (voucher M.F. Braby 62, DNA). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 21 February 2009, M.F. Braby & D.A. Young. A female was observed around midday depositing a single egg on the underside of a leaf of Cynanchum pedunculatum growing in savannah woodland. Tylophoraflexuosa R.Br. (Apocynaceae). (voucher M.F. Braby 63, DNA). Emma Gorge Resort, El Questro Wilderness Park, WA (15.90753°S, 128.12909°E; WGS84), 17 July 2009, M.F. Braby & L.J. Aitchison. A female was observed at 1130 hr ovipositing on Tylophara flexuosa growing on sandstone rock scree in open woodland above Emma Creek; numerous additional eggs and two larvae were also present on the larval food plant. Euploea corinna (W.S. Macleay, 1826). Common Crow. Marsdenia viridiflora R.Br. (Apocynaceae). (voucher M.F. Braby 68, DNA). 1 km ENE of Nanguluwur, Nourlangie Rock, Kakadu National Park, NT (12.84375°S, 132.82558°E; WGS84), 20 December 2009, M.F. Braby. A female was observed at 1020 hr depositing a single egg on the underside of a leaf of a vine of Marsdenia viridiflora, which grew in woodland at the base of an escarpment; two early instar larvae were also recorded feeding on new soft leaves of M. viridiflora. Secamone elliptica R.Br. (Apocynaceae). (voucher M.F. Braby 74, DNA). Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 23 February 2010, M.F. Braby. One mid-instar larva was recorded feeding on a vine of Secamone elliptica, which grew around Capparis sepiaria in degraded monsoon vine-thicket. Acraea andromacha andromacha (Fabricius, 1775). Glasswing. Hybanthus enneaspermus (L.) F.Muell. (Violaceae). (not vouchered). ‘Ryala’, Noonamah, NT (12.62694°S, 131.09527°E; AGD66), 6 March 2006, M.F. Braby & 1. Morris. Numerous final instar larvae (>20) were recorded feeding on Hybanthus enneaspermus growing in woodland; larvae were noted to feed singly or in small groups. *PassiJlora foetida L. (Passifloraceae). (voucher M.F. Braby 88, DNA). Nanguluwur Art site, Nourlangie Rock, Kakadu National Park, NT (12.84262°S, 132.81895°E; WGS84), 13 November 2010, M.F. Braby & J. Wcstaway. A female was observed at 1145 hr ovipositing on Passiflora foetida growing in profusion over footslope sandstone boulders; a cluster of four eggs was laid on the underside of a new leaf. Junonia orithya albicincta Butler, 1875. Blue Argus. Buchnera linearis R.Br. (Orobanchaceae). (voucher M.F. Braby 39, DNA). Education campground, Litchfield National Park, NT (13.11361°S, 130.79805°E; WGS84), 17 April 2006, M.F. Braby & L.J. Aitchison. Two final instar larvae were recorded on Buchnera linearis , a herb growing in savannah woodland. Nawurlandja (Little Nourlangie Rock), Kakadu National Park, NT (12.85701°S, 132.79053°E; WGS84), 19 April 2008, M.F. Braby & K. Nishida. Two larvae were recorded on B. linearis growing close to a creek near the car park. Maguk Plunge Pool (Barramundie Gorge), Kakadu National Park, NT (13.31841 °S, 132.43828°E; WGS84), 30 August 2008, M.F. Braby & L.J. Aitchison. A female was observed at 1245 hr ovipositing on B. linearis growing in moist sand near the edge of a large pool of water. Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 28 February 2009, M.F. Braby. A single larva was recorded on B. linearis growing in savannah woodland. Pseuderanthemum variabile (R.Br.) Radik. (Acanthaceae). (voucher M.F. Braby 87, DNA). Nanguluwur Art site, Nourlangie Rock, Kakadu National^ Park, NT (12.84262°S, I32.81895°E; WGS84), 27 November 2010, M.F. Braby. Eggs and three final instar larvae were recorded on Pseuderanthemum variabile growing in the ground layer in savannah woodland; two larvae were collected and reared to adult in captivity. Junonia liedonia zelinta (Fabricius, 1775). Chocolate Argus. Hygrophila angustifolia R.Br. (Acanthaceae). (voucher M.F. Braby 38, DNA). Howard Springs Nature Reserve, NT (12.45240°S, 131,05347°E; WGS84), 8 and 24 March 2008, M.F. Braby. A female was observed during early afternoon ovipositing on Hygrophila angustifolia (and on adjacent objects), a herb growing in paperbark-pandanus swamp adjacent to wet evergreen monsoon vine-forest; one early instar larva and one final instar larva were also recorded feeding on H. angustifolia. 98 New Lepidoptera food plant associations Fogg Dam Conservation Reserve, NT (12.55769°S, 13 1,29701°E; WGS84), 29 May 2010, M.F. Braby, S. Keates & D. Bisa. A female was observed at 1145 hr ovipositing on H. angustifolici, as well as on adjacent objects, including a dead twig of an Acacia sp. sapling and dead dry leaves of grass, in mixed paperbark swampland adjacent to monsoon vine-forest. Hypolimnas bolina nerina (Fabricius, 1775). Varied Eggfly. *Synedrella nodiflora (L.) Gaertn. (Astcraceae). (voucher M.F. Braby 71, DNA). Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 23 March 2010, M.F. Braby. Two females were observed at 1245-1255 hr ovipositing on Synedrella nodiflora , a herb growing in shade beneath a large African Mahogany tree; all eggs were laid singly on the underside of leaves. The site was revisited on 24 March 2010 by M.F. Braby & S. Keates and a female was observed at 1240-1250 hr ovipositing on introduced Tridaxprocumbens (Astcraceae), as well as a species of grass on which a cluster of five eggs were laid on the underside of a blade. Since both plants were growing amongst numerous herbs of S. nodiflora, the suitability of these species as larval food plants requires confirmation. On 7 April 2010 one late instar larva was collected from the ground underneath S. nodiflora and reared to adult in captivity on this food plant. Melanitis leda bankia (Fabricius, 1775). Evening Brown. Imperata cylindrica (L.) Reausch. (Poaceae). (not vouchered). Nathan River Ranger Station, Limmcn National Park, NT (15.57681°S, 135.42783°E; WGS84), 12 May 2009, M.F. Braby & N. Collier. Numerous larvae and several pupae (mostly diseased) were recorded on Imperata cylindrica, as well as on introduced Cynodon radiatus (Poaceae) (see above), growing along the bank of a river. LYCAENIDAE Ogyris oroetes oroetes (Hewitson, 1862). Silky Azure. Amyema bifurcata (Benth.) Tiegh. (Loranthaceae). (not vouchered). Cockatoo Lagoon, Keep River National Park, NT (15.96950°S, 129.04099°E; WGS84), 1 February 2008, M.F. Braby. One pupa was collected from under the bark of a eucalypt host tree supporting several clumps of the mistletoe Amyema bifurcata growing in the canopy; a male was also netted perched in the canopy of the host tree. Victoria Flwy, approx. 35 km E of Kununurra, WA (15.96229°S, 128.96069°E; WGS84), 6 July 2009, M.F. Braby. One pupa was collected (and reared to adult in captivity) from under loose bark at the base of the host tree Corymbia grandifolia (R.Br. ex Benth.) K.D.Hill & L.A.S.Johnson, which supported numerous clumps of A. bifurcata, growing in savannah woodland on a sandplain. The pupa was found together with a cohort of O. zosine larvae, which were attended by sugar ants, Camponotus sp. (humilior species group). Ogyris oroetes apiculata (Quick, 1972). Silky Azure. Amyema bifurcata (Benth.) Tiegh. (Loranthaceae). (not vouchered). Larapinta Drive, NNE of Kings Canyon, NT (23.99472°S, 131.48583°E; WGS84), 12 October 2005, M.F. Braby & L. J. Aitchison. One mid-instar larva was collected (and reared to adult in captivity) from a mallee eucalypt host tree supporting clumps of the mistletoe A myema bifurcata. The larva was found together with a cohort of O. amaiyllis meridionalis larvae. Amyema miquelii (Lam. ex Miq.) Tiegh. (Loranthaceae). (not vouchered). 6 km S of Hermannsburg, NT (23.99111°S, 132.77555°E; WGS84), 14 October 2005, M.F. Braby & L.J. Aitchison. One late instar larva and one prepupa were collected (and reared to adult in captivity) from loose bark at the base of the host tree Eucalyptus camaldulensis Dehnh., which supported clumps of the mistletoe Amyema miquelii, growing along the Finke River; the larva was attended by three ants, Crematogaster sp. ( longiceps species group). Trephina Gorge National Park, East Macdonnell Ranges, NT (23.52805°S, 134.38000°E; WGS84), 30 April 2006, M. F. Braby. Ten larvae in various instars were collected (and reared to adult in captivity) from a mallee eucalypt supporing clumps of the A. miquelii ; a few larvae were attended by ants, Crematogaster sp. ( longiceps species group); larvae were polymorphic in colour, varying from green, through pale green-brown and pale brown, to dark brown. Luritja Road, 30 km N of Lasseter Hwy, NT (24.93221 °S, 132.27826°E; WGS84), 16 August 2010, M.F. Braby. One mid-instar larva was collected from under loose bark of a mallee eucalypt host tree supporting numerous clumps of A. miquelii-, the larva was not attended by ants. Several adult males were collected during the afternoon as they settled on dead branches in the canopy of the breeding trees where they appeared to be defending encounter sites to detect receptive females. Ogyris amaryllis meridionalis (Bethune-Baker, 1905). Satin Azure. Amyema miquelii (Lam. ex Miq.) Tiegh. (Loranthaceae). (not vouchered). Trephina Gorge National Park, East Macdonnell Ranges, NT (23.52805°S, 134.38000°E; WGS84), 30 April 2006, M.F. Braby. One final instar larva was collected (and reared to adult in captivity) from a mallee eucalypt host tree supporting the mistletoe Amyema miquelii growing adjacent to a breeding colony of O. oroetes (see above). 99 M. F. Braby Ogyris zosiite (Hewitson, [1853|). Northern Purple Azure. Amyema bifurcata (Benth.) Tiegh. (Loranthaceae). (not vouchered). Mary River Billabong, Mary River National Park, NT (12.89245°S, 131.63890°E; WGS84), 19 July 2008, M. F. Braby & S. Keates. A female was observed at 1500-1515 hr ovipositing on the host tree Corymbia bella supporting a small clump of the mistletoe Amyema bifurcata growing in the canopy (approx. 8 m from ground level) in savannah woodland; eggs were laid on patches of rough bark of the trunk below the mistletoe clump. The site was revisited the following week and six recently hatched egg shells were collected; no attendant ants were present on the host tree. Ebony Creek, Limmen National Park, NT (16.35714°S, 135.69812°E; WGS84), 11 May 2009, M.F. Braby & N. Collier. A cohort of nine larvae in various instars and one pupa were collected (and reared to adult in captivity) from soil at the base of the trank of the host tree Corymbia grandifolia supporting two clumps of A. bifurcata in savannah woodland; the early stages were attended by numerous sugar ants, Camponotus sp. ( humilior species group). Amyema sanguinea (F.Muell.) Danser. (Loranthaceae). (voucher M.F. Braby 81, DNA). Limbunya Station, Victoria River District, NT (17.30592°S, 129.77728°E; WGS84), 24 July 2010, M.F. Braby, V. Ressner & T. Parkin. A cohort comprising 13 early to mid-instar larvae and one final instar larva was collected (and reared to adult in captivity) from under a small rock at the base of a Corymbia sp. host tree, which supported a single clump of the mistletoe Amyema sanguinea growing near the canopy (approx. 7 m from ground level) in woodland on a steep mixed limestone outcrop; the larvae were attended by numerous pale brown sugar ants Camponotus sp. ( humilior species group). Of eight females reared to adult in captivity, four had the basal areas on the upperside of the wings bright purple, while the remaining four were bright blue. Limbunya Station, Victoria River District, NT (17.55211 °S, 130.07086°E; WGS84), 25 July 2010, M.F. Braby. One pupa was collected from the base of a small Corymbia sp. host tree, which supported three clumps of A. sanguinea, growing in eucalypt open woodland with a ground layer of Triodia sp.; the pupa was attended by five sugar ants, Camponotus sp. ( crozieri species group), a darker brown species compared with Camponotus sp. (humilior species group). Buntine Hwy, 31 km SW of Top Springs, NT (16.75994°S, 131.61429°E; WGS84), 28 July 2010, M.F. Braby. A cohort of 10 larvae was found at the base of a eucalypt host tree supporting a large clump of A. sanguinea growing 1.5 m above ground level; most larvae were parasitised by braconid wasps and only two pre-pupae, which were attended by several sugar ants, Camponotus sp. ( novaelwllandiae species group), were not parasitised and reared to adult in captivity. Antheneseltuttus affinis (Waterhouse & R.E. Turner, 1905). Dark Ciliate-blue. Cupaniopsis anarcardioides (A.Rich.) Radik. (Sapindaceae). (not vouchered). Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 18 December 2010, M.F. Braby & J. Westaway. Five larvae in various instars were recorded feeding on soft new leaf growth of Cupaniopsis anarcardioides ; larvae were attended by Oecophylla smaragdina ants. Candalides erinus erittus (Fabricius, 1775). Small Dusky-blue. Cassytha capillaris Meisn. (Lauraceae). (voucher M.F. Braby 34, DNA). Jinumum Gorge, Keep River National Park, NT (15.83376°S, 129.11018°E; WGS84), 2 and 4 February 2008, M.F. Braby & V. Kessner. A female was observed at 1005 hr depositing a single egg on a flower bud of Cassytha capillaris parasitising grass growing in savannah woodland on sandy loam on a gentle slope adjacent to a gorge; three larvae were also collected from new growth of the larval food plant and reared to adult in captivity; one of these larvae was attended by a small black ant, Iridomyrmex sp. (mattiroloi species group), and everted its eversible organs in presence of the ant. The site was revisited on 20 March 2008 and a female was observed at 1048 hr ovipositing two eggs on flower buds of the food plant; two larvae and one pupa, neither of which were attended by ants, were also collected from C. capillaris and reared to adult in captivity. Overall, the larvae were polymorphic in colour, varying from green, through pale green-brown and pale brown, to dark brown. The early stages were found together with those of Candalides delospila (see below). Joe Creek, Gregory (Jutpurra) National Park, NT (15.59754°S, 131,07042°E; WGS84), 16 March 2008, M.F. Braby. One mid-instar larva, which was not attended by ants, was recorded on C. capillaris. Scenic Lookout, Timber Creek, NT (15.64580°S, 130.45827°E; WGS84), 18 March 2008, M.F. Braby. A female was observed at 1455 hr depositing a single egg on a young tendril of C. capillaris parasitising Triodia sp. (Poaceae) growing in eucalypt woodland on sandstone; one late instar larva, which was not attended by ants, was also collected feeding openly on C. capillaris and it was reared to adult in captivity. Amalia Gorge, El Questro Wilderness Park, WA (15.98124°S, 128.03708°E; WGS84), 2 April 2010, M.F. Braby & L.J. Aitchison. One final instar larva, which was not attended by ants, was collected from C. capillaris parasitising Triodia sp. growing on a lower slope of the gorge with a north facing aspect and it was reared to adult in captivity. 100 New Lepidoptera food plant associations Candalides delospila (Waterhouse, 1903). Spotted Dusky-blue. Cassytha capillaris Meisn. (Lauraceae). (vouchers M.F. Braby 28, 034, DNA). The early stages were frequently recorded (and subsequently reared to adult in captivity) on Cassytha capillaris parasitising Triodia spp. and other grasses growing in open woodland on rocky sandstone plateaux or on sandy loam from many sites in Gregory (Jutpurra) and Keep River National Parks, NT, during visits in February, March and July 2007 and 2008. Eggs were collected from, and females were observed ovipositing on, new growth of C. capillaris ; all eggs were laid singly; larvae were found at rest or feeding openly on C. capillaris during the day (i.e. they were not concealed amongst clumps of the host grass); most larvae were not attended by ants, but one was attended by a single black ant, Iridomyrmex sp. ( gracilis species group); pupae were collected from stems of the host plant Triodia sp. around which the larval food plant grew. The larvae were polymorphic in colour, varying from green, through pale green-brown and pale brown, to dark brown. At Keep River, the early stages were found together with those of Candalides erinus (see above). At all other sites where Candalides delospila was recorded (e.g. Limbunya Station, Riveren Station, Timber Creek, Jasper Gorge, near Elliott, NT; Picaninny Creek Purnululu National Park, WA) adults flew in close proximity of C. capillaris. Prosotas dubiosa dubiosa (Semper, [1879|). Purple Line-blue. Acacia auriculiformis A.Cunn. ex Benth. (Fabaceae). (not vouchered). East Point, Darwin, NT (12.41140°S, 130.82500°E; WGS84), 1 April 2006, M.F. Braby. A female was observed at 1500 hr ovipositing on flower buds of Acacia auriculiformis growing in coastal semi-deciduous monsoon vine-thicket. Tliecliitesthes miskini miskini (T.P. Lucas, 1889). Wattle Blue. Acacia holosericea A.Cunn. ex G.Don. (Fabaceae). (not vouchered). The early stages were frequently recorded on Acacia holosericea growing in a variety of woodland habitats from many locations, including Channel Island, Beatrice Hill, near Black Point Cobourg Peninsula, Muirella Park Kakadu National Park, NT, and the Pentecost River El Questro Wilderness Park, WA. The early stages were usually attended by meat ants, Iridomyrmex reburrus, or sometimes small black ants, Iridomyrmex sp. ( mattiroloi species group), or medium-sized black ants, Iridomyrmex sp. ( anceps species group). Atalaya variifolia F.Muell. ex Benth. (Sapindaceae). (voucher M.F. Braby 30, DNA). Victoria Hwy, Pinkerton Range, NT (15.98698°S, 129.51213°E; WGS84), 1 February 2008, M.F. Braby & V. Kessncr. A female was observed ovipositing on a sapling of Atalaya variifolia (approx. 1.5 m high) growing in savannah woodland on a sandstone outcrop; about 40 additional eggs were also present on new soft terminal growth of A. variifolia ; numerous meat ants, Iridomyrmex sp., were present on the food plant. BullitaRd, 11 km SSEofTimber Creek, NT (15.75712°S, 130.51135°E; WGS84), 10 December 2008, M.F. Braby & V. Kessner. Numerous larvae and pupae were recorded on A. variifolia growing in savannah woodland; the early stages were attended by small black ants, Iridomyrmex sp. ( minor species group). Sesbania cannabina (Retz.) Poir. (Fabaceae). (voucher M.F. Braby 72, DNA). Rapid Creek, Darwin, NT (12.38083°S, 130.86462°E; WGS84), 20 March 2010, M.F. Braby. Numerous females were observed ovipositing on new leaf growth of Sesbania cannabina growing in a disturbed area along the edge of mangroves adjacent to Rapid Creek. The early stages were found together with those of Eurema hecabe (see above). Jamidesphaseli (Mathew, 1889). Purple Cerulean. Canavalia rosea (Sw.) DC. (Fabaceae). (not vouchered). Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 28 May 2009, M.F. Braby. A female was observed at 1340 hr ovipositing on flower buds of Canavalia rosea , which grew amongst Ipomeapes-caprae (Convolvulaceae) in coastal sand dunes adjacent to mangroves; three larvae were also recorded feeding inside flower buds and flowers of C. rosea. Zizeeria karsandra (Moore, 1865). Spotted Grass- blue. Tribulus cistoides L. (Zygophyllaceae). (voucher M.F. Braby 20, DNA). Gunn Point, NT (12.22833°S, 131.03000°E; WGS84), 10 December 2006, M.F. Braby. All stages were recorded on the underside of leaflets of Tribulus cistoides growing in coastal open beach shrubland above high tide mark; the larvae were not attended by ants; two pupae were reared to adult in captivity. 5.1 km ENE of Black Point, Cobourg Peninsula, NT (11.14695°S, 132.19014°E; WGS84), 23 April 2008, M.F. Braby & K. Nishida. A female was observed at 1230 hr ovipositing on the underside of leaflets of T. cistoides growing in beach sand along the edge of coastal semi- deciduous monsoon vine-thicket; several larvae were also recorded on the underside of foliage of T. cistoides. Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 25 May 2010, M.F. Braby. Eggs and one early instar larva were recorded on the underside of leaflets of T. cistoides growing in profusion on beach sand. The site was revisited on 11 June 2010 and a female was observed at 1230 hr ovipositing on T. cistoides. On 1 September 2010, a mid-instar larva was collected from the underside of leaves of T. cistoides and reared to adult in captivity. 101 M. F. Braby Euchrysopscttejuscnidus Waterhouse& Lyell, 1914. Spotted Pea-blue. Vigna vexillata (L.) A.Rich. (Fabaceae). (voucher M.F. Braby 52, DNA). Mt Burrell, Tipperary Station, Daly River district, NT (13.49623°S, 131.03572°E; WGS84), 11 April 2009, M.F. Braby & L.J. Aitchison. A female was observed at 1045 hr ovipositing on Vigna vexillata growing in a dry gully at the base of a hill; adults were common around V vexillata. *Macroptilium atropurpureum (DC.) Urb. (Fabaceae). Buffalo Creek, Lee Point, NT (12.33639°S, 130.90389°E; WGS84), 16 August 2008, M.F. Braby. A female was observed ovipositing on a low-spreading vine oiMacroptilium atropurpureum , which grew in a paperbark swamp; additional eggs were also present on flower buds of this introduced plant. Bullocky Point, Darwin, NT (12.43777°S, 130.83377°E; WGS84), 23 June 2010, M.F. Braby. Females were observed at 1230 hr ovipositing on M. atropurpureum growing on a beach cliff; numerous additional eggs were collected from leaves, new soft stems and flower buds; two early instar larvae were also collected from M. atropurpureum , feeding on flowers or developing pods, and were reared to adult in captivity. DISCUSSION In total, 122 insect-plant associations are documented, of which 76 are new for Australia and 46 are new for the NT and/or WA. Of particular note are the first recordings of two plant families for two lycaenid species: the Lecythidaceae is recorded for the genus Anthene , and the Santalaceae is recorded for the genus Theclinesthes. New ant associations are also recorded for the larvae and/or pupae of several lycaenids, notably of Ogyris zosine and Theclinesthes miskini. In addition, new larval food plant associations are reported for six species of agaristines ( Cruria tropica, Idalima aethrias, /. leonora, Periopta ardescens, P. diversa, Radinocera vaga) for which the life histories have not previously been documented. The final instar larvae of these six species are illustrated for the first time in Figs 3-8. Several food plant records are based only on oviposition behaviour of females, and further observations may be required to determine their suitability. In my experience, however, females rarely make mistakes when ovipositing, and usually when they oviposit on ‘new’ food plants those species belong to plant families that arc well-known to be exploited and the larvae are able to feed and develop to adulthood on these plants in captivity. An association of the aposematic diurnal moth Birthana cleis with the Loranthaceae has only recently been established, and in the Darwin area it breeds commonly on Decaisnina signata (Anderson & Braby 2009; M.F. Braby unpublished data). Decaisnina is distantly related to Dendrophthoe (Vidal-Russel & Nickrent 2008), the new food plant recorded from north-eastern Arnhem Land, suggesting that the host range of this immid may be considerably wider than available data indicates. A broad host range within the Loranthaceae may also occur in Delias argenthona, which in the Top End was previously known to be associated only with Decaisnina signata in the Darwin area (Wade 1978; Meyer 1996; Anderson & Braby 2009). In the NT, Borbo impar has been recorded only from a limited number of locations at and near Darwin, but the native larval food plant and natural breeding habitat have up until now remained unknown. All previous food plant associations (Meyer 1997b) comprise naturalised grasses from Africa that grow commonly in suburban areas of Darwin. The occurrence of B. impar in the floodplains of the South Alligator River suggests that the hesperiid is a specialist of lowland floodplains and was possibly formerly restricted to that habitat, but following European settlement has colonised non-natural areas through its ability to utlise introduced grasses. The native larval food plants of Taractrocera ina have also not previously been recorded; all known food plants comprise introduced or naturalised grasses (Braby 2000). The introduced Paspalum conjugation has been previously recorded as a larval food plant for the species from central Qld (De Baar 1979), but not from the NT. Atkins (1975) observed a female Taractrocera sp. lay an egg on Buffcl Grass Cenchrus ciliaris in the Gogango Ranges, Queensland, and considered the species to be T. anisomorpha based on the presence of males in the immediate vicinity. Atkins (1991) subsequently recorded the native Eulalia aurea as the main food plant for this hesperiid in the Alice Springs area. Telicota colon is the only grass-feeding member of the genus represented in the Kimberley and Top End, the other species, T. augias (Linnaeus, 1763), specialises on Flagellaria indica (Flagellariaceae) (Meyer 1996). The two other species of Telicota reported from the Northern Territory, T. mesoptis Lower, 1911 and T. ancilla (Herrich- Schiiffer, 1869), appear to be erroneous (Braby 2008a; M.F. Braby unpublished data). The native larval food plant and natural breeding habitat Cephrenes augiades ssp. ‘Top End ’, which is endemic to the NT, have not previously been recorded, and there are few records outside Darwin. The natural habitat of this species comprises riparian evergreen monsoon vine-forest, but the species often breeds on cultivated palms in suburban areas of Darwin, an artificial habitat which it has no doubt invaded in recent decades following the extensive cultivation of palms in gardens and parklands since Tropical Cyclone Tracy. Dunn (1996) listed Carpentaria acuminata , which is endemic to the Top End, as a food plant for Cephrenes augiades , but this was based on the presence of early stages of the eastern form utilising cultivars in the Mount Coot-tha Botanic Gardens, Brisbane, Qld. Dunn (1996) listed Livistona marine as a food plant for Cephrenes trichopepla based on observations of a single larva in the Mount Coot-tha Botanic Gardens, Brisbane; 102 New Lepidoptera food plant associations however, the natural utilisation of this palm, which is restricted to the West McDonnell Ranges, has not been confirmed until now. Dunn (2009) also observed two mature larvae of this hesperiid in shelters on a species of Livistona at Cahills Crossing, and this record almost certainly refers to L. benthamii. Papilio aegeus has become established on the mainland in north-eastern Arnhem Land only in the last decade since about 2000 (L. Wilson pers. comm.). Previously, P. aegeus was restricted to Groote Eylandt and Marchinbar Island in the Wessel Islands where it breeds on Micromelum minutum (G.Forst.) Wight & Am. (Rutaceae) (Fenner 1991). The usual larval food plants for Papilio fuscus in the Top End are M. minutum and Glycosmis trifoliata (Blume) Spreng. (Rutaceae) (Hall 1976; Meyer 1996). Cultivated Citrus sp. has previously been recorded for this species from the eastern Kimberley at Kununurra, WA (Common & Waterhouse 1981). Meyer (1996) listed Senna leptoclada as a larval food plant of Catopsilia scylla in the Kununurra district of the eastern Kimberley, WA, but it has not previously been recorded as a food plant from the NT. Moss (2008) recently reviewed and updated the larval food plants for Eurema alitha in Queensland and noted that Galactia tenuiflora and G. muelleri are the preferred legumes compared with Glycine tabacina, the previously known food plant. The larval food plant in the Top End, however, has not previously been reported. In the NT, Elodina padusa appears to be parapatrie with E. walkeri Butler, 1898, which occurs predominantly in the higher rainfall areas of the monsoon tropics where it specialises on Capparis sepiaria, whereas E. padusa occurs mainly in the arid zone. The two species have a narrow range of overlap in the drier semi-arid inland areas of the monsoon tropics, but it is not known if they share the same larval food plants in this region. Available data suggests they may exploit different species of Capparis. The usual food plant of Appias paulina in the Top End is Diypetes deplanchei (Brongn. & Griseb.) Merr. (Putranjivaceae) (Meyer 1996; Braby et al. 2010). The butterfly is known to utilise Capparis spp. in eastern Australia (Braby 2000) and elsewhere within its geographical range. The extent to which A. paulina uses Capparis in the NT is not known, but is possibly infrequent. The usual larval food plant of Euploea alcathoe in north¬ eastern Arnhem Land is Parsonsia alboflavescens (Braby 2009), but it appears that occasionally larvae of E. corinna and E. Sylvester also feed on this vine. The usual larval food plant of E. Sylvester is Marsdenia geminata (Meyer 1997a), which is widespread across the Top End, but the extent to which E. corinna uses this vine is not known. Grand and Hunt (2001) recorded the introduced weed Passi/lora foetida as a larval food plant for Acraea andromacha in the northern Kimberley, WA, where females were observed to lay large batches of eggs on it in the Kalumburu area, but it has not previously been reported as a food plant from the Top End. Hypoclnysops apelles in the Top End is restricted to mangrove communities where the usual larval food plant near Darwin is Lumnitzera racemosa Willd. (Combretaceae) (Meyer 1996; Eastwood et al. 2008), which typically grows along the landward edge of the habitat. Despite the presence of many other species of mangroves, which are utilised as food plants elsewhere in the geographical range of //. apelles in eastern Queensland, these plants do not seem to be utilised in the Northern Territory. However, at Buffalo Creek in the early dry season when the species is particularly abundant, larvae of H. apelles were found to be associated mainly with L. racemosa and less frequently with Ceriops australis, whereas later in the dry season (May-September) larvae were found only on L. racemosa. These observations suggest that species of mangroves other than L. racemosa are exploited but only during periods of high population densities. Reared adults of Ogyris oroetes from the eastern Kimberley and other material collected elsewhere in the Top End arc similar to specimens from northern Queensland in that the males have narrow black margins and the upperside ground colour iridescent pale lilac, and are better placed with the nominate subspecies than with the ‘arid form’ proposed by Braby (2000). However, reared adults and material collected from central Australia arc similar to specimens from southern Australia in that the males have a broad black apex of the fore wing. In many respects they resemble O. oroetes apiculata (Quick, 1972) described from Western Australia, except that the upperside ground colour is iridescent dark lilac or purple, rather than blue as in the type specimen. However, the holotype appears to be abberant or not representative of populations elsewhere in southern Australia in which the males are typically dark lilac. Specimens from central Australia are therefore provisionally placed with O. oroetes apiculata rather than with the ‘arid form’ proposed by Braby (2000). The usual food plant for Ogyris amaryllis in the central arid zone appears to be Amvema maidenii, which often parasitises Acacia aneura F.Muell. ex Benth. (see also Grand 2005), but several other species have now been recorded, viz: Amvema miquelii, Amyema bifurcata and Diplatiagrandibractea. In addition, Amyema sanguinea has been recorded as a larval food plant in the Schwerin Mural Crescent, WA (M.F. Braby & J.J. Armstrong unpublished data). The extent to which these four other mistletoes are utilised in central Australia is not well understood. In the Northern Territory, the purple morph of female Og\ris zosine has only rarely been recorded, the blue phenotype being the predominate morph. Le Souef (1971) listed the species from Elliott, and examination of his material (reared to adult in October 1969) in the ANIC revealed that two of the three female specimens collected are bright purple above. Elliott is located at similar latitude to the sites further west at Limbunya Station, both locations being situated in semi-arid areas at 103 M. F. Braby the southern boundary of the monsoon tropics biome. More recently, Weir et al. (2011) recorded the purple morph from a population in suburban coastal Darwin in which five females (18%) from a sample of 28 pupae reared during the wet season (December) were of the purple phenotype. Weir et al. (2011) suggested that female polymorphism in this species may be influenced by environmental factors, such as humidity (with high humidity promoting the purple morph); however, the purple morphs from Limbunya were reared to adult during the late dry season (August to September) when conditions were comparatively cool and dry. These observations suggest that phenotypic variation in O. zosine may be under genetic control but the allele for the femal purple morph occurs in low frequency. The usual food plant of Hypolycaena phorbas in the Top End is Clemdendmmfloribundam R.Br. (Lamiaceae), but larvae have also been recorded on Smilax australis R.Br. (Smilaceae) (Meyer 1996). Elsewhere, the butterfly is associated with a wide range of plant families (Braby 2000), but rarely has it been recorded from mistletoes (Valentine & Johnson 1988). The only larval food plant previously recorded for Candalides margarita in northern Australia is Decaisnina signata (Loranthaceae) (Samson & Wilson 1995; Braby 2008b); however, given that the geographical distribution of the butterfly is wider than that of this mistletoe, other food plants arc expected to be utilised. Therefore, the new record for D. triflora as a host plant is not unexpected. In an earlier publication (Braby 1995). I listed Cassytha pubescens R.Br. as the larval food plant for Candalides geminus in sandstone habitats on the Burra Range, Qld, but this record is incorrect and refers to Cassythafiliformis. in that same publication (Braby 1995) I listed Cassytha filiformis as the larval food plant for Candalides delospila from the Burra Range, Qld; however, this record is also in error and refers to Cassytha capillaris. In another publication (Braby 1997), 1 listed Cassytha pubescens as a larval food plant for Candalides erinus based on records from several localities in northern Queensland; however, these records also refer to Cassytha filiformis. All of these determinations of Cassytha species were made prior to the monograph of Weber (2007), which explains the misidentifications. The usual food plant of Candalides erinus in the Northern Territory is Cassytha filiformis (M.F. Braby, unpublished data), but in the drier semi-arid areas of the Top End the butterfly was occasionally found breeding on Cassytha capillaris , which grows as a finer, less vigorous hemiparasitic vine on Triodia spp. and other grasses on sandstone or sandy soils. Candalides delospila is endemic to the monsoon tropics (Braby 2008a) where it appears to be ecologically specialised on Cassytha capillaris, which likewise is restricted to northern Australia. Grund & Hunt (2001) recorded Cassytha capillaris as a larval food plant for both Candalides erinus and Candalides delospila in the Tanami Desert at the Northern Territory-Western Australian border. Facultative ant associations are here recorded for the larvae of Candalides erinus and C. delospila , two species not previously known to associate with ants, and these records add to the growing list of ant-butterfly associations among theAustralianLycaenidae (Eastwood & Fraser 1999; Eastwood et al. 2008). The usual food plants of Nesolycaena urumelia in the Top End are Boronia lanceolata F.Muell. in western Arnhem Land and areas closer to Darwin (Edwards 1980; Meyer 1996) and B. lanuginosa Endl. in subcoastal heathy woodland on sand in Gove Peninsula and Elcho Island (Braby 2000; M.F. Braby, unpublished data), but in the drier rocky areas of western Arnhem Land, the butterfly was found breeding on both B. laxa and B. lanceolata. Meyer & Wilson (1995) listed Tecticornia indica (Willd) K.A.Sheph. & Paul G.Wilson as one of two larval food plants for Theclinesthes sulpitius in the Top End based on observations near Darwin. Although this saltbush grew together with Tecticornia halocnemoides at the site near Palmerston, neither eggs nor larvae were present on Tecticornia indica. The Mistletoe Day-moth Comocrus behri has not previously been recorded from the Northern Territory, although there is a small series of adults in the Museum and Art Gallery of the Northern Territory from Alice Springs collected by I. Archibald. This species is widespread and abundant in the temperate areas of south-eastern Australia, but it appears to be rare and seasonal in the Top End. ACKNOWLEDGEMENTS I am indebted to K. Brennan, I.D. Cowie, P.S. Short, G.M. Wightman, J. Westaway and R. Flarwood for their botanical advice and competent determinations of many of the larval food plants listed in this paper, and to A. Andersen for identification of the attendant ants. J. Westaway kindly read a draft of the manuscript and greatly assisted with plant nomenclature and higher classification; A.F. Atkins and A.A.E. Williams critically reviewed the text and provided constructive comments, and R.C. Willan provided substantial editorial advice. 1 thank L.J. Aitchison, J.J. Armstrong, D. Bisa, N. Collier, A. Kallies, S. Keates, V. Kessner, D.A. Lane, B. Marika, I. Morris, K. Nishida, C.M. Palmer, J. Westaway, P. Wise and D.A. Young for their assistance in the field. 1 am also grateful to the Northern Territory Parks and Wildlife Service and the Dhimurru Aboriginal Land Management Corporation for their co-operation and access to lands under their management and control. Field work in Kakadu National Park, Northern Territory, was undertaken under Scientific Research Permit numbers RK 704 and RK 729 under the auspices of Commonwealth EPBC Act and for Purnululu National Park, WA, under a scientific purposes licence administered by the Department of Environment and Conservation under the provisions of the Wildlife Consen’ation Act. 104 New Lepidoptera food plant associations REFERENCES Anderson, S.J. & Braby, M.F. 2009. Invertebrate diversity associated with tropical mistletoe in a suburban landscape from northern Australia. Northern Territory Naturalist 21: 2-23. Atkins, A.F. 1975. Larval food plants of some Queensland butterflies. News Bulletin of the Entomological Society of Queensland 3: 117-119. Atkins, A.F. 1991. Observations on the biology of Taractrocera anisomorpha (Lower) (FIcspcriidae: Hesperiinae). Australian Entomological Magazine 18: 121 123. Braby, M.F. 1995. The life history of Zetona delospila (Waterhouse) (Lepidoptera: Lycaenidae). The Australian Entomologist 22: 37-42. Braby, M.F. 1997. New larval food plants for some butterflies (Lepidoptera) from northern and central Queensland, Australia. The Australian Entomologist 24: 97-108. Braby, M.F. 2000. Butterflies of Australia. Their identification, biology and distribution. CSIRO Publishing: Melbourne. Braby, M.F. 2008a. Biogeography of butterflies in the Australian monsoon tropics. Australian Journal of Zoology* 56: 41-56. Braby, M.F. 2008b. Taxonomic review of Candalides absimilis (C. Felder, 1862) and C. matgarita (Semper, 1879) (Lepidoptera: Lycaenidae), with descriptions of two new subspecies. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 24: 33-54. Braby, M.F. 2009. The life history and biology of Euploea alcathoe enastri Fenner, 1991 (Lepidoptera: Nymphalidae) from northeastern Arnhem Land, Northern Territory, Australia. The Australian Entomologist 36: 51-62. Braby, M.F. 2010. The merging of taxonomy and conservation biology: a synthesis of Australian butterfly systematics (Lepidoptera: Hcsperioidea and Papilionoidea) for the 21st century. Zootaxa 2707: 1-76. Braby, M.F., Lane, D.A. & Weir. R.P. 2010. The occurrence of Appiasalbinaalbino (Boisduval, 1836) (Lepidoptera: Pieridae: Picrinae) in northern Australia: phenotypic variation, life history and biology, with remarks on its taxonomic status. Entomological Science 13: 258-268. Common, I.F.B. 1990. Moths of Australia. Melbourne University Press: Melbourne. Common, I.F.B. & Waterhouse, D.F. 1981. Butterflies of Australia. Revised. Angus & Robertson: Sydney. De Baar, M. 1979. Some new food plants for Australian Lepidoptera with life history notes. Australian Entomological Magazine 5: 87-89. Dunn, K.L. 1996. Notes on the biology and new larval hosts of Cephrenes (Lepidoptera: Hesperiidae) - Part III. Victorian Entomologist 26: 47-55. Dunn, K.L. 2009. Notes on Cephrenes augiades (C. Felder) in the Northern Territory, Australia (Lepidoptera: Hesperiidae). Victorian Entomologist 39: 53-56. Eastwood, R.G., Braby, M.F., Lohman, D.J. & King, A. 2008. New ant-lycacnid associations and biological data for some Australian butterflies (Lepidoptera: Lycaenidae). The Australian Entomologist 35: 47-56. Eastwood, R.G. & Fraser, A.M. 1999. Associations between lycaenid butterflies and ants in Australia. Australian Journal of Ecology’ 24: 503-507. Edwards, E.D. 1980. The early stages of Adaluma urumelia Tindale and Candalides geminus Edwards and Ken' (Lepidoptera: Lycaenidae). Australian Entomological Magazine 1: 17-20. Fenner, T.L. 1991. A new subspecies of Euploea alcathoe (Godart) (Lepidoptera: Nymphalidae) from the Northern Territory, Australia. Australian Entomological Magazine 18: 149-155. Grund, R. 2005. Some new butterfly observations for central Australia. Victorian Entomologist 35: 111—114. Grund, R. & Hunt, L. 2001. Some butterfly observations for the Kimberley and Tanami regions. Western Australia. Victorian Entomologist 31:19-23. Hall, M.C. 1976. Native food plant records for some Northern Territory butterflies. Australian Entomological Magazine 3:41. Kerrigan, R.A. & Albrecht, D.E. 2007. Checklist of NT vascular plant species. Northern Territory Herbarium: Palmerston. Lazarides, M., Hacker, J.B. & Andrew, M.H. 1991. Taxonomy, cytology and ecology of indigenous Australian Sorghums (Sorghum Moench: Andropogoneae: Poaceac). Australian Systematic Botany 4: 591-635. Le Souef, J.C. 1971. Winter insect collecting in the Northern Territory. The Victorian Naturalist 88: 350-356. Meyer, C.E. 1996. Butterfly larval food plant list for the Northern Territory and the Kununurra District in Western Australia. Victorian Entomologist 26: 66-72. Meyer, C.E. 1997a. Notes on the life history and variations in adult forms of Euploea Sylvester pelor Doubleday (Lepidoptera: Nymphalidae: Danainae). The Australian Entomologist 24: 73-77. Meyer, C.E. 1997b. Notes on the life history of Borbo itnpar lavinia (Waterhouse) (Lepidoptera: Hesperiidae). The Australian Entomologist 24: 78-80. Meyer, C.E. and Wilson, D.N. 1995. A new distribution record for Theclinesthes sulpitius (Miskin) (Lepidoptera: Lycaenidae) in the Northern Territory and notes on the life history. The Australian Entomologist 22: 63. Moss, J.T. 2008. Further notes on the Scalloped Grass-yellow (Eurema alitha) and its hostplants. Metamorphosis Australia. Magazine of the Butterfly and Other Invertebrates Club 51: 13—14. Nielsen, E.S., Edwards, E.D. & Rangsi, T.V. (eds) 1996. Checklist of the Lepidoptera of Australia. Monographs on Australian Lepidoptera, Volume 4. CSIRO Publishing: Melbourne. Samson, PR. & Wilson, D.N. 1995. The life history of Candalides gilberti Waterhouse (Lepidoptera: Lycaenidae). The Australian Entomologist 22: 71-73. Short, P.S., Albrecht, D.E., Cowie, I.D., Lewis, D.L. & Stuckey, B.M. 2011. Checklist of the vascular plants of the Northern Territory. Northern Territory Herbarium, Department of Natural Resources, Environment, The Arts and Sport: Palmerston. Valentine, P.S. & Johnson, S.J. 1988. Some new larval food plants for north Queensland Lycaenidae (Lepidoptera). Australian Entomological Magazine 14: 89-91. Vidal-Russel, R. & Nickrent, D.L. 2008. Evolutionary relationships in the showy mistletoe family (Loranthaceae). American Journal of Botany 95: 1015-1029. Wade, A. 1978. Notes on the habits of the Northern Jezabel butterfly. Northern Territory Naturalist 1: 13-14. Weber, J.Z. 2007. Cassytha , Lauraceae. Pp 117-136. In: Wilson, A.J.G. (cd.) Flora of Australia. Volume 2, iVinteraceae to Platanaceac. Australian Biological Resources Study and CSIRO Publishing: Canberra and Collingwood. Weir, R.P., Meyer, C.E. & Brown, S.S. 2011. Notes on the biology of Ogyris zosine (I lewitson, 1853) (Lepidoptera: Lycaenidae: Theclinae), including the first record of the purple female form from the Northern Territory, Australia. The Australian Entomologist 38: 97-100. Accepted 1 September 2011 105 ' The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 21: 107-112 Dimorphothynnus (Hymenoptera: Thynnidae: Rhagigasterini) newly recorded from the Northern Territory, Australia GRAHAM R. BROWN Research Associate, Museum and Art Gallery of the Northern Territory GPO Box 4646, Darwin NT0801, AUSTRALIA graham.brown@nt.gov.au (thynnini@hotmail.com) ABSTRACT The predominantly southern Australian genus Dimorphothynnus Turner is newly recorded from the Northern Territory, and two new species, D.fulvicms and D. nigricrus, are described. Both species arc relatively small in size, and only known from male individuals. A key to males of 15 species of Dimorphothynnus is given; that is, all but two of the known species. A key to females of eight species of Dimorphothynnus is given under the assumption of species fidelity of mates. Two new combinatins are made - Rhagigaster haemorrhoidalis Guerin is transferred to Rhytidothynnus, and R. obtusus Smith is transferred to Dimorphothynnus. Keywords: Northern Australia, Thynnidae, Rhagigasterinini, Dimorphothynnus. INTRODUCTION The genus Enteles was erected by Westwood (1844). However, this name was preoccupied by the weevil genus Enteles Schoenherr (1833), and so in 1910 Turner proposed the replacement name Dimorphothynnus. This genus has never been revised and the most recent key to species is over a century old (Turner 1907). Males are very similar to those of Rhagigaster Guerin, and the related genera Rhytidothynnus Brown and Umbothynnus Brown. The males of these genera are distinguished by a long, cylindrical metasoma with strong constrictions between the metasomal segments. However, those of Dimorphothynnus have the clypeus with an inverted V-shaped carina which is not sharply defined (Fig. 1), the propodcum has a transverse carina separating the dorsal and posterior surfaces, the epipygium is flat with the apical margin straight or almost so (Fig. 2), and the genitalia have the cuspides long with the outer surface denticulate apically. Females however, arc unique within the tribe in that metasomal tergite 2 is transversely multicarinate. The latter character is more typical of the tribe Thynnini, where it occurs in most genera (Turner 1910). As in all Thynninae the sexes of Dimorphothynnus are completely dimorphic; males are typical winged wasps, while females are wingless and strongly modified for bun-owing in soil. This makes the association of sexes on morphological grounds impossible. Matching the sexes is further complicated by the prevalence of miscoupling (Brown 1993, 2001) in the subfamily. Miscoupling occurs when two different species couple, and thus there is no guarantee that pairs collected in copula are actually the same species. This makes it difficult to compare species if the types are of different gender, as discussed for the genus by Given (1959). Nothing is known of the biology of Dimorphothynnus. As with most species of Australian Thynninae, they are presumed to be parasites of scarab larvae (Bun-ell 1935; Given 1953; Ridsdill Smith 1970), but this is based on fewer than 20 records. Adults have a preference for feeding on the nectar of myrtaceous flowers but they are attracted to other flowers as well as insect exudates. This biological information was summarised by Brown (2005). There are now 15 described species of Dimorphothynnus including the two new species described here. Most are restricted to coastal regions of southern Australia except D. barnardi (Turner) and D. integer (Fabricius) from tropical Queensland as well as the two Northern Territory species described here. Fig. 1. Dimorphothynnus nigricrus, male: head. Scale line = 0.1 mm. 107 G. R. Brown Fig. 2. Dimorphothynnus fulvicnts , male: epipygium. Scale line = 0.05 mm. Terminology follows Snodgrass (1941), Brown (1997a, b) and Naumann (1991). Relative terms relating to microsculpture are interpreted as follows: sparsely punctate = punctures greater than two puncture-diameters apart; punctate = punctures at most two puncture-diameters apart, but never confluent; closely punctate = punctures almost confluent; rugosely punctate = punctures partially confluent; finely punctate = punctures small and shallow; coarsely punctate = punctures large and deep; obscurely punctate = punctures small, sparse, shallow and only visible at certain angles. Abbreviations. Morphological characters: Tl-7, metasomal tergites 1-7; Sl-8, metasomal stemites 1-8. Specimen repositories; ANIC, Australian National Insect Collection, CS1RO, Canberra; MNG, Museo Civicodi Storia Naturale di Genova, Genoa, Italy; NTM, Museum and Art Gallery of the Northern Temtory (formerly Northern Territory Museum), Darwin; OUM, Oxford University Museum, Oxford, United Kingdom. SYSTEMATICS Dimorphothynnus Turner, 1910 Enteles Westwood, 1844. Gender masculine. Type species, by original designation, Enteles bicolor Westwood, 1844. Recent, King George’s Sound, Western Australia. Preoccupied by Enteles Schoenherr, 1833 (Insecta: Coleoptera). Dimorphothynnus Turner, 1910. Replacement name for Enteles Westwood (1844). Gender masculine. As explained above, Dimorphothynnus is a replacement name for the preoccupied Enteles Westwood (1844). Dalla Torre (1897: 100) used Enteles as a subgenus of Thynnus and Ashmead (1899: 50,52, 1903: 103) inadvertently misspelt Enteles as Entelus. I have examined the holotype of Enteles bicolor Westwood, 1844, which is deposited in the OUM. Dimorphothynnus fulvicnts sp. nov. (Figs 2, 3, 5, 7) Material examined. Holotype - NTM 1.6201, cf, Northern Territory, Adelaide River, 13°14'S, 131°06'E, Adelaide River, 20 December 1995, G.R. Brown & G.M. Dally. Paratypes - ANIC (not registered), 1, cf, same data as holotype; NTM 1.6205, 1, cf, same data as holotype; NTM 1.6202, 1, cf, 13°14’S, 131°06’E, Adelaide River, 5 November 1996, G.R. Brown. Description of male. Body (Figs 2, 3) length 12 mm; fore wing 8 mm; hind wing 6 mm. Clypeus with obscure inverted V-shaped carina which is flattened and not sharply defined, rugosely punctate between branches of carina, finely rugosely punctate laterally; frons with short, flat, with V-shaped carina above antennal insertion, transversely carinate below mid-ocellus, rugosely punctate; vertex and gena rugosely punctate. Pronotum rugosely punctate, dorsum transversely rugose, anterior margin sharply carinate; mcsoscutum coarsely transversely rugosely punctate; mesoscutellum coarsely rugosely punctate; metanotum rugosely punctate; propodeum closely punctate becoming transversely rugosely punctate on posterior surface, dorsal and posterior surfaces delineated by carina; mesopleuron coarsely rugosely punctate dorsally becoming finely punctate ventrally; fore coxae small, conical, closely punctate. Metasoma with anterior groove on T2-5 and S2-5 scrobiculate; tergites closely and finely punctate, punctures sparser postcrolaterally and coarser on T7; epipygium (Fig. 2) rounded apically with narrow semi-membranous apical margin; stemites closely punctate, less so on disc of S2-6; SI weakly medially raised; hypopygium subparallel basally becoming sinusoidal distally without prominent lateral lobes or spines thence produced into long recurved apical spine. Genitalia (Fig. 7): parameres (in profile) long and narrow, ventral margin evenly concavcly curved over most of length, dorsal margin subparallel over basal 2/3 then abruptly converging (and slightly concave) with ventral margin over most of length, apex narrowly rounded and slightly Fig. 3. Dimorphothynnus fulvicnts, male: habitus. Scale line - 1 mm. 108 Thynnine wasps of the genus Dimorphothynnus upturned; aedeagus (Figs 5,7) short, subtriangular medially with highly modified valves sheathing the remainder of the aedeagus basally; cuspides large, subtriangular (viewed dorsally), outer margin serrated with small basally directed teeth, inner margin straight with fiat vertical inner surface; digiti small, inconspicuous and knob-like with erect cluster of setae. Colour. Black; tegulac and legs (except fore coxae) orange; setae white, and forming posterior or ventral bands on most metasomal segments especially laterally and ventrally; wings hyaline, veins black becoming orange basally. Distribution. Only known from above the northern bank of Adelaide River in the township of Adelaide River, Northern Territory. Remarks. This species, one of the smallest in the genus, can be distinguished from all the other species of Dimorphothynnus by the combination of orange legs, hyaline wings and a black metasoma. The setal banding on the metasoma suggests a relationship with D. integer from northern Queensland, but D. integer has a distinctive mesoscutellutn which is almost impunctatc and higher posteriorly with a distinct flat posterior surface that is delineated dorsally by a transverse carina. The new species differs from D. ohtusus by the presence of setal bands on the metasoma. The shape of the sheathing valve at the base of the aedeagus (Fig. 5) also distinguishes this species from D. nigricrus (Fig. 6). Etymology. This species name is derived from Latin and is a reference to leg colour. It is intended as a noun in apposition. Dimorphothynnus nigricrus sp. nov. (Figs 1,4,6, 8) Material examined. Holotype - NTM 1.6203, cT, campsite near Oenpelli reservoir. Northern Territory, 12°23’S, 133°06’E, uv light, 27 November 1996, G.R. Brown & J. Purdie. Description of male. Body (Fig. 4) length 14 mm; fore wing 10 mm; hind wing 7 mm. Fig. 4. Dimorphothynnus nigricrus , male: habitus. Scale line = 1 mm. Figs 5, 6. Dimorphothynnus species, aedeagi (and apex of basiparameres), dorsal view: 5. Dimorphothynnus fulvicrus ; 6, Dimorphothynnus nigricrus. Scale line = 0.1 mm. Figs 7, 8. Dimorphothynnus species, male genitalia, dorsal view: 7, Dimorphothynnus fulvicrus ; 8, Dimorphothynnus nigricrus. Scale line = 0.1 mm. Clypeus (Fig. 1) with obscure inverted V-shaped carina which is flattened and not sharply defined, rugosely punctate between branches of carina, finely rugosely punctate laterally; frons with short, flat, V-shaped carina above antennal insertions, transversely carinate below mid ocellus, rugosely punctate; vertex and gcna closely to rugosely punctate. Pronotum rugosely punctate, dorsum transversely rugose, anterior margin sharply carinate; mesoscutum coarsely transversely rugosely punctate; mesoscutellum coarsely and closely punctate; metanotum rugosely punctate; propodeum closely punctate, dorsal and posterior surfaces delineated by obscure carina; mesopleuron coarsely rugosely punctate dorsally becoming finely and closely punctate ventrally; fore coxae small, conical, closely punctate. Metasoma with anterior groove on T2-5 and S2-5 scrobiculate; tergites closely punctate, punctures sparser posterolaterally on T2-5 and coarser on T7; epipygium rounded apically without narrow semi-membranous apical margin; stemites closely and deeply punctate; SI weakly medially raised; hypopygium subparallel basally becoming sinusoidal distally without prominent lateral lobes or spines thence produced into long recurved apical spine. 109 G. R. Brown Genitalia (Fig. 8) with parameres (in profile) long and narrow, margins almost straight, ventral margin slightly sinusoidal, dorsal margin converging with ventral over most of length, apex narrowly rounded; aedeagus (Figs 6, 8) short, subtriangular medially with highly modified valves sheathing remainder of the aedeagus basally; cuspides large, subtriangular (viewed dorsally) with ventral lobe arising from lateral margin, outer margin (viewed dorsally) serrated with small basally directed teeth, inner margin straight with flat vertical inner surface; digiti apparently absent. Colour. Black; setae white, and forming posterior or ventral bands on most metasomal segments especially laterally and ventrally; wings hyaline, veins black. Distribution. Dimorphothynnus nigricrus is only known from the vicinity of Oenpelli, Northern Territory, from a creek flat downstream from Oenpelli reservoir. Remarks. This species is similar to D. fulvicrus in that both are smaller than most other species in the genus and have the tergites uniformly and closely punctate, but D. fulvicrus has orange legs rather than black legs. It is also distinguished from D. integer and D. obtusus as discussed for D. fulvicrus above. Etymology. This species name is derived from Latin and is a reference to leg colour. It is intended as a noun in apposition. KEYS TO THE SPECIES OF DIMORPHOTHYNNUS The majority of type specimens within Dimorphothynnus (as well as the subfamily) are male. However, two species, D. bicolor and D.fimbriatus (Smith), were described from the female only. Unfortunately both of these are a part of a complex of species that also includes D. apicalis (Smith), a species misidentified as D. haemorrhoidalis (Guerin) by Westwood (1844a), Turner (1907, 1910) and Given (1959), as well as several undescribed species. This species complex was discussed by Given (1959), who was able to demonstrate that pairs collected ‘in copula’ were not necessarily the same species, and as such the identity of the males of D. bicolor and D. fimbriatus must be considered tentative. Rhagigaster haemorrhoidalis Guerin does not belong to this genus and is here transferred to Rhytidothynnus Brown based on an examination of the holotype in MNG. Conversely, Rhagigaster obtusus Smith is a species of Dimorphothynnus because the genitalia completely match those of the type species of Dimorphothynnus. So the correct combinations for these two species are Rhytidothynnus haemorrhoidalis (Guerin) comb. nov. and Dimorphothynnus obtusus (Smith) comb. nov. The females of Dimorphothynnus barnardi (Smith), D. conjugatus, D. testaceipes, D. trunciscutis and the two new species described here, are unknown. However, it is interesting to note that Turner (1907) had a male and a female in front of him when he described D. barnardi but the female was damaged and incomplete and he therefore did not describe it. The original description of D. barnardi indicates that the clypeal carina is in the form of an inverted Y-shape and enclosing a polished almost impunctate area, but gives no mention of a transverse propodeal carina. The description of the clypeal carina excludes this species from this genus (as would the absence of a propodeal carina). It is more likely that it is a species of Rhytidothynnus. While the type has not been examined, I have seen a specimen from Pine Creek near Bundabcrg, Queensland, in ANIC that fits the description of this species and which is a species of Rhytidothynnus. This specimen was collected about 260 km southeast of the type locality, Duaringa. Its inclusion in the key is based on the original description (1 urner, 1907), and its correct placement is dependent on an examination of the holotype which is the only known specimen. Similarly, the female of D. deceptor (Smith) was not considered to be a species of Dimorphothynnus by Turner (1907), and is therefore not included in the key. Male (winged) 1 Wings fulvous; legs black [only known from the types from Geraldton, WA]. D. deceptor (Smith) Wings hyaline, fuscous or black (although veins may be orange); legs black or orange.2 2( 1) Clypeus with sharply defined inverted Y-shaped carina, branches enclosing polished triangular area [Qld] . D. barnardi (Turner) Clypeus with inverted V-shaped carina, carina not sharply defined, not enclosing a large or polished triangular area.3 3(2) Metasoma black (if lightly testaceous as in D. testaceipes, then wings black).4 Metasoma at least partially ferruginous.10 4(3) Wings hyaline to weakly infuscate.5 - Wings black.8 5(4) Mesoscutellum almost impunctate, vertically truncate posteriorly with carina sharply delineating dorsal and posterior surfaces [Qld]. D. integer (Fabricius) Mesoscutellum punctate to rugosely punctate, without distinct posterior surface clearly delineated from dorsal surfaces.6 6(5) Tergites closely and uniformly punctate, posterior margin of tergites (weakly so medially in D. fulvicrus and D. nigricrus ) and stemites and ventral margins of pleurites with dense fringe of white setae.7 Tergites punctate to closely punctate, punctures variable in size and spacing, tergites, stemites and pleurites not fringed with white setae [NSW, Vic, SA]. D. obtusus (Smith) comb. nov. 7(8) Legs orange [NT]. D. fulvicrus sp. nov. — Legs brown to black [NT]. D. nigricrus sp. nov. 110 Thynnine wasps of the genus Dimorphothynnus 8(4) Epipygium punctuate, with punctures longitudinally aligned; legs dark; mesoscutellum truncate posteriorly; mesopleura deeply and rugosely punctate [Qld, NSW, Vic, SA]. D. trunciscutus Turner Epipygium longitudinally multicarinate or irregularly longitudinally multistriate; legs orange; mesoscutellum narrowly rounded posteriorly; mesopleura punctate .9 9(8) Metasoma lightly testaceous [NSW, Vic, SA] . . D. testaceipes (Turner) - Metasoma black [NSW] . D. mono (Westwood) 10(3) Metasoma entirely ferruginous [Qld, NSW]. . D. dimidiatus (Smith) Metasoma with only apical 2 segments ferruginous .11 11(10) Wings dark; pronotum transversely carinate [Qld] . D. conjugates (Turner) Wings hyaline; pronotum punctate to rugose . .12 12(11) Mesopleurae red [WA]. D. simillimus (Smith) Mesopleurae black. 13 13(12) Fore coxae with deep, large, longitudinal, smooth and polished cavity on the inner ventral surface that is delineated by a sharp carina WA] . D. bicolor (Westwood) Fore coxae if depressed or partially concave, not deeply so and not margined by a carina.. 14 14(13) Prosternum (and adjacent areas of fore coxae) concave, concavity polished and sparsely punctuate [WA]. D. fimbriates (Smith) Prostemum not concave, punctuation more uniform [WA]. D. apicalis (Smith) Female (wingless) 1 Pygidium notched. D. apicalis (Smith) - Pygidium not notched. 2 2(1) Dorsum of T1 finely transversely carinate, not punctate; head medially depressed. . D. integer (Fabricius) T1 not transversely carinate; head not medially depressed. 3 3(2) Posterior angles of head weakly excavate. . D. bicolor (Westwood) Posterior angles of head not excavate . 4 4(3) Apex of metasoma ferruginous . . D. dimidiates (Smith) Metasoma concolorous. 5 5(4) Head and body dark brown or black . 6 - Head and metasoma black, mesosoma ferruginous .7 6(5) T3 with transverse carinae mostly restricted to anterior half of sclerite. D. obtusus (Smith) comb. nov. T3 with transverse carinae more evenly distributed on sclerite. D. morio (Westwood) 7(5) T1 transversely carinate (in addition to apical margin and weak preapical carina); apex of mctasoma with a long brush of setae laterad to pygidium. . D. fimbriates (Smith) - T1 not transversely carinate (other than apical margin and weak preapical carina); apex of metasoma without long brush of setae. D. simillimus (Smith) ACKNOWLEDGEMENTS I thank John Jennings and Nigel Fortearth for refereeing the manuscript and providing useful comments. REFERENCES Ashmead, W.H. 1899. Super-families in the Hymenoptera and generic synopses of the families Thynnidae, Mymiosidae and Mutillidac. Journal of the New York Entomological Society 7: 45-60. Brown, G.R. 1993. A new species of Lestricothynnus with notes on miscoupling in Thynninae (Hymenoptera: Tiphiidae). Journal of the Australian Entomological Society 32: 197-199. Brown, G.R. 1997a. Chilothynnus, a new genus of Australian Thynninae (Hymenoptera: Tiphiidae) associated with orchids. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 13:61-71. [Original publication and reprints incorrectly dated 1996.] Brown, G.R. 1997b. Arthrothynnus , a new genus of orchid-pollinating Thynninae (Hymenoptera: Tiphiidae). The Beagle, Records of the Museums and Art Galleries of the Northern Territory 13: 73-82. [Original publication and reprints incorrectly dated 1996.] Brown, G.R. 2001. Caetrathynnus, Nitidothynnus and Pmcerothynnus , new genera of Thynninae (Hymenoptera: Tiphiidae) from northern Australia. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 16: 107-122. Brown, G.R. 2005. A Revision of Tachyphtxm Brown and description of two new genera within the Aripltron group (Hymenoptera: Tiphiidae). Journal of Natural Histoiy 39(2): 197-239. Burrell, R.W. 1935. Notes on the habits of certain Australian Thynnidae. Journal of the New York Entomological Society 43: 19-28. Dalla Torre, C.G. de 1897. Catalogus Hymenoptorum hucusque descriptorum systematicus et synonymicus. (Aus der kaiserlich- koniglichen Hof-und Staatsdruckerei, Wein) 8: 99-199. Given, B.B. 1953. General report on a search for parasites of Melolonthinae in Australia. New Zealand Journal of Science and Technology (B) 34: 322-340. Given, B.B. 1959. Notes on Australian Thynninae. II. The genera Dimorphothynnus, Rhagigaster and Eirone. Proceedings of the Linnean Society of New South Wales 83: 309-326. G. R. Brown Naumann, l.D. 1991. Hymenoptera (Wasps, bees, ants, sawflies). Pp. 916-1000. In: Naumann, l.D. (ed.). The insects of Australia. A textbook for students and research workers. Melbourne University Press, Carlton. Ridsdill Smith, T.J. 1970b. The behaviour of Hemithynnus hyalinatus (Hymenoptera: Tiphiidae), with notes on some other Thynninae. Journal of the Australian Entomological Society’ 9: 196-208. Snodgrass, R.E. 1941. The male genitalia of Hymenoptera. Smithsonian Miscellaneous Collections 99: 1-86. Turner, R.E. 1907. A revision of the Thynnidae of Australia (Hymenoptera. Pt I. Proceedings ofthe Linnean Society of New South Wales 32: 206-290. Turner, R.E. 1910. Hymenoptera Fam. Thynnidae. Genera Insectorum 105: 1-62. Westwood, J.O. 1844a. Illustrations of some species of Australian thynnideous insects. Arcana Entomologica 2: 101-110. Westwood, J.O. 1844b. Further illustrations of thynnideous insects ofAustralia. Arcana Entomologica 2: 135-146. Accepted 26 September 2011 112 The Beagle, Records of the Museums and Art Galleries of the Northern Territory\ 2011 27 : 113-121 Periclimenaeuspulitzerfinali sp. nov. (Crustacea: Decapoda: Palaemonidae), a new pontoniine shrimp from East Africa A. J. BRUCE Curator Emeritus, Museum and Art Gallery of the Northern Territory. Present address: Queensland Museum, PO Box 3300, South Brisbane OLD 4101, AUSTRALIA abruce@broad. net.au ABSTRACT A small pontoniine shrimp. Periclimenaeus pulitzerfinali, from Shelly Beach, Mombasa, Kenya, is described as new and illustrated. An appendix presents a complete list of Periclimenaeus species recorded from East Africa (Kenya, Zanzibar, Tanganika). Keywords: Periclimenaeus pulitzerfinali sp. nov., Crustacea, Decapoda, Pontoniinae, Mombasa, Kenya, sponge associate. INTRODUCTION The genus Periclimenaeus Borradaile, 1915 is the second most speciose genus of the subfamily Pontoniinae with at present 70 described species - 55 from the lndo- west Pacific region. 4 from the eastern Pacific and 11 from the Atlantic region. The various species are associated with sponge orascidian hosts, although the hosts of a number of species have not yet been reported or identified. The present report describes and illustrates a further new species from an East African sponge host, increasing the number of I ndo- west Pacific species known to 56. The single specimen is deposited in the collection of the Museum and Art Gallery of the Northern Territory, Darwin. The names Tanganyika and Zanzibar are used in their geographical sense only, as the two entities are now Politically united as Tanzania. Abbreviations used: CL refers to the postorbital carapace length; NTM, Museum and Art Gallery of the Northern Territory, Darwin (formerly Northern Territory Museum). SYSTEMATICS Family Palaemonidae Rafinesque, 1815 Subfamily Pontoniinae Kingsley, 1879 Genus Periclimenaeus Borradaile, 1915 Gender masculine. Type species, by subsequent designation of Borradaile (1917: 378), Periclimenaeus [originally spelling Periclimenaeus] robustus Borradaile, '915. Recent, Amirante Islands, Indian Ocean. Genus placed 0r i the Official List of Generic Names in Zoology under Opinion 470 dated 1957. Periclimenaeus pulitzerfinali sp. nov. (Figs 1-5) Material examined. Holotype - NTM Cr. 17287 (previously AJB # 2098), ovigerous 9, CL 5.2 mm, reef front off Shelly Beach, Mombasa, Kenya, SCUBA, 16 m, coll. J. Wood. 23 January 1974. Diagnosis. Rostral dentition 7/0, without supraorbital spines or tubercles, first abdominal segment tergite with small narrow anterior median lobe, scaphocerite tooth not exceeding lamella, first pereiopod slender, fingers stout, semi-spatulate, half palm length, major second pereiopod with feeble molar process, palm and merus ventrally spinulate, minor second pereiopod dactyl elongate, extending well beyond fixed finger, cutting edge convex, entire, tip strongly bifid, fixed finger short, cutting edge grooved, edges entire, third pereiopod dactyl robust, biunguiculate, ventral corpus minutely denticulate, telson with dorsal spines well developed, at 0.15 and 0.53 of telson length, exopod of uropod with distolateral tooth and spine only. Description. Female holotype, a small-sized pontoniine shrimp of subcylindrical, slightly compressed body form, complete with both second pereiopods. Rostrum (Fig. 1 A) slender, compressed, without distinct midrib, slightly depressed, about 0.33 of CL, reaching to about distal margin of proximal segment of antennular peduncle, with 7 slender acute teeth extending over whole of rostral length, fifth tooth longest, with numerous long setae interspersed, tip acute, ventral margin feebly convex, unarmed, non-setose. Carapace (Fig. 1B,C) slightly compressed, glabrous, without epigastric, supraorbital or hepatic spines, with feeble postorbital ‘shoulders’, antennal spine well developed, 113 A. J. Bruce Fig. 1 . Periclimenaeus pulitzerfinali sp. nov., holotype female, Mombasa, Kenya, NTM Cr. 17287: A, rostrum; B, carapace and rostrum; C, anterior carapace and appendages; D, antennule; E, antenna; F, ophthalmic somite, lateral; G, eye, dorsal; H, first abdominal tcrgite, anterior margin, dorsal. I, fifth pleuron and sixth abdominal somite; J, sixth abdominal segment, dorsal; K, telson; L, same, posterior spines, dorsal spines inset, anterior spine left, posterior spine right; M, uropod. submarginal, inferior orbital angle small rounded, curved medially, anterolateral margin of branchiostegite feebly produced, rounded. Abdomen glabrous, first segment tcrgite with small anterior median lobe (Fig. 1H), sixth segment (Fig. 1I,J) length about 1.5 times length of fifth, depressed, about 1.2 times longer than anterior width, posterolateral angle small, rounded, with small acute tooth, postero-ventral angle large, broadly acute; first to fourth pleura enlarged, rounded, fifth postero-ventrally subrectangular (Fig. II), margin feebly setose. Telson (Fig. IK) about 1.8 times sixth segment length, 0.5 of CL, 2.0 times longer than anterior width, lateral margins feebly convex, posteriorly convergent, dorsal telson spines about 0.10 of telson length, at 0.15 and 0.53 of telson length, anterior pair slightly longer than posterior, posterior margin (Fig. 1L) broadly convex without median point, 0.25 of anterior telson width, lateral posterior spines small, about 0.4 of intermediate spine length, intermediate spines short, about 0.08 of telson length, 2.5 times lateral spine length, submedian spines slightly longer and more slender than intermediate spines, densely setulose. Ophthalmic somite (Fig. IF) anteriorly with median vertical carina. Eye (Fig. IG) with large globular cornea, diameter about 0.1 of CL, well-pigmented, without obvious accessory Periclimenaeuspulitzerfinali sp. nov., from Kenya pigment spot, stalk short, about as broad as long, 0.8 of comeal diameter, length about 0.8 of corneal diameter. Antennule (Fig. ID) of normal form, damaged on right, proximal peduncular segment about twice as long as central width, distally tapering, anterolateral margin with well-developed acute tooth on left, abnormal, rounded on right, lateral margin expanded proximally, rounded, medial margin straight, thickened, sparsely setose, with well-developed ventromedial tooth at about half length, stylocerite large, broadly phylliform, distally acute, sparsely setose laterally, statocyst normal with granular statolith; intermediate segment short, about 0.16 of proximal segment length, longer than wide, distal segment similar, upper flagellum biramous, right abnormal, 2 proximal segments fused, short ramus with single segment, about 4 groups of long aesthetascs, left with 7 segments fused, shorter ramus with 2 segments, with 5 groups of aesthetascs, longer ramus slender, with 10 segments, lower flagellum similar, slender with 19+segments. Antenna (Fig. 1E) of normal form, basiceritc short, with acute tooth laterally, carpocerite subcylindrical, about 3.0 times longer than width, reaching to about 0.75 of scaphocerite length, few short setae medially; scaphocerite 2.5 times longer than wide, greatest width distally at about 0.5 of length, anterior margin bluntly angular, lateral margin straight, with well-developed distal tooth at about 0.85 of scaphocerite length, reaching level of distal margin of lamella. Thoracic stemites with third and fourth broad, unarmed, posterior stemites narrow and unarmed. Mouthparts of normal Periclimenaeus morphology. Mandible (Fig. 2A) with corpus stout, without palp; incisor process (Fig. 5A) slender, tapering, transversely truncate distally, with 4 slender subequal acute teeth, medial margin entire; molar process (Fig. 5BC) robust, subcylindrical, obliquely truncate distally, with 4 stout marginal teeth, with 2 small brushes of short simple setae. Maxillula (Fig. 2B) with feebly bilobed palp (Fig. 5D), upper lobe short, glabrous, lower lobe stouter, with small simple spinule; upper lacinia moderately broad, dorsal margin strongly convex, distal margin (Fig. 5E) with 10 short robust spines, feebly denticulate on ventral surface (Fig. 5F), lateral surface with several oblique rows of slender spiniform setae; lower lacinia short, subcylindrical, distally rounded, with numerous long terminal setae. Maxilla (Fig. 2C) with simple tapering palp, proximal lateral margin with several short plumose setae, distally rounded, shorter than basal endite, basal cndite bilobed, dorsal lobe expanded with dense simple setae, distolateral margin with several transverse rows of simple setae, ventral lobe smaller and shorter than dorsal lobe, with numerous simple setae distally; coxal endite obsolescent. Fig. 2. Periclimenaeus pulitzerfinali sp. nov., holotype female, Mombasa, Kenya, NTM Cr. 17287: A, mandible; B, maxillula; C, maxilla; D, first maxilliped; E, Second maxillipcd; F, third maxilliped. 115 A. J. Bruce medial margin slightly produced, convex, non-setose; scaphognathite well developed, narrow, about 3.4 times longer than central width, anterior lobe narrow, rounded, medial margin slightly excavate, posterior lobe well developed, one third of scaphocerite length. First maxilliped (Fig. 2D) with elongate flattened subcylindrical tapering palp, about 5.0 times longer than central width, reaching anterior margin of basal enditc, with single preterminal plumose seta medially; basal endite fused with coxal endite, combined medial margin straight, distal margin produced, rounded, margins with numerous long spiniform setae; exopod with normal flagellum with 4 plumose terminal setae, some short plumose setae distolaterally, caridean lobe small; epipod lost in dissection. Second maxilliped (Fig. 2E) with normally developed endopod, dactylar segment narrow, about 3.0 times longer than central depth, medial margin straight, with numerous serrulate spines, propodal segment normal, distomedial margin not produced, with several robust spines and shorter spiniform setae, carpus, merus and ischiobasis without special features; coxa medially produced; exopod normally developed with 4 plumose terminal setae; epipod lost in dissection, small. Third maxilliped (Fig. 2F) with endopod reaching to about middle of carpocerite, ischiomerus fully fused to basis, point ofjunction on medial margin indicated by small marginal swelling, combined segment about 4.6 times longer than central width, basal region slightly expanded, medially rounded, without marginal setae, with submarginal row of 17 short plumose setae (Fig. 5H), distolateral angle (Fig. 5G) with 2 short simple submarginal spines, ischiomeral portion with numerous long slender spiniform marginal setae; penultimate segment about 0.32 of antepenultimate segment length, 3.0 times longer than wide, with numerous long simple setae medially; terminal segment about 0.75 of penultimate segment length, tapering distally, without strong terminal spine, with several transverse rows of spiniform setae medially; exopod normally developed with 4 plumose terminal setae, several short plumose marginal setae distolaterally, coxa not medially produced, with low rounded lateral plate, without arthrobranch. First pcrciopods (Fig. 3A) moderately slender, exceeding carpocerite by distal fifth of merus, chela (Fig. 3B,C) with palm subcylindrical, moderately compressed, 2.5 times longer than deep, dorsal and ventral margins feebly tuberculate, fingers (Fig. 3D) 0.43 of palm length, robust, subspatulate, with groups of short stiff setae, dactylus 2.2 times longer than basal width, dorsal margin convex, tip hooked, with stout medial tooth flanked by 2 smaller teeth, cutting edges robust, entire; carpus 1.6 of chela length, 5.5 times longer than distal width, tapering proximally, dorsal and ventral margins feebly tuberculate; merus subequal to carpus length, 5.9 times longer than maximal width, tapering slightly distally; ischium about 0.62 of merus length; basis without special featured, and coxa with small setose distoventral process. Second pereiopods grossly unequal, dissimilar. Major second pereiopod (Fig. 4A) with massive chela, about 1.33 times CL, chela (Fig. 4B) with palm minutely tuberculate (Fig. 51), oval in section, twice as long as deep, tapering slightly distally, dactylus (Fig. 4C,D) with tip missing, about 0.42 of palm length, strongly compressed, 2.5 times longer than depth, proximal cutting edge with low elongated molar process, with posterior end projecting, distal cutting edge concave, entire; fixed finger about 1.1 times longer than proximal depth, robust, with shallow fossa proximally, distally grooved, medial margin with acute tooth proximally, lateral margin with rounded setose process, tip feebly hooked, distinctly bidentate, prcterminal tooth rounded, terminal tooth acute; carpus about 0.33 of palm length, 1.3 times longer than wide, tapering strongly proximally, distally slightly excavate, unarmed; merus (Fig. 4E) 0.37 of palm length, 1.6 times longer than broad, slightly expanded centrally, ventral margin tuberculate (Fig. 5.1); ischium (Fig. 4E) subequal to meral length, distally broadened, about 2.8 times longer than distal width, ventral margin strongly tuberculate (Fig. 5K); basis and coxa robust, without special features. Minor second pereiopod (Fig. 4F) with chela about 0.6 ot CL, 0.53 of major chela length, palm oval in section, dorsally convex, smooth, ventrally straight, finely tuberculate, 2.2 times longer than deep, tapering slightly distally. projecting slightly posteriorly, sparsely setose distoventrally; dactylus (Fig. 4G,H) extending well beyond tip of fixed finger (Fig. 4G), 2.12 times longer than deep, strongly compressed, dorsal margin strongly convex, cutting edge convex, sharp, entire, tip robust, feebly hooked, distally bidentate; fixed finger short, stout, about as long as deep proximally, with small acute tip separated by small notch from cutting edge, cutting edge grooved, edges concave, entire; carpus short, stout, about 0.25 of palm length, distally expanded, tapering proximally, unarmed; merus about 0.33 of palm lcngth,1.5 times longer than deep, with ventral margin with numerous acute tubercles; ischium about 0.38 of palm length, twice as long as wide, tapering proximally, ventral margin with numerous subacute tubercles; basis and coxa robust, without special features. Ambulatory pereiopods robust; third pereiopod (Fig. 3E-G) reaching to end of scaphocerite, with dactyl (Fig. 3H) 0.29 of propod length, unguis distinctly demarcated, 2.5 times longer than basal width, curved, unarmed ventrally, corpus compressed, about 1.6 times longer than basal width, tapering strongly distally, dorsal margin moderately convex, ventral margin with acute distal slightly preterminal antcrovcrted accessory tooth, ventral margin feebly convex, with 4 minute denticles, without basal process; propod about 0.25 of CL, 3.6 times longer than proximal width, tapering slightly distally, distoventral angle armed with 2 stout spines, slightly unequal, longer spine subequal to dorsal corpus length, ventral margin with 4 spaced spines, lengths decreasing slightly proximally; carpus about 0.7 of propod length, 2.5 times longer than 116 Periclimenaeus pulitzerfinali sp. nov., from Kenya Fig. 3. Periclimenaeus pulitzerfinali sp. nov., hololype female, Mombasa, Kenya, NTM Cr. 17287: A, first pereiopod; B, same, chela; C, same, oblique; D, same, fingers; E. third pereiopod; F, same, propod and dactyl; G, same, distal propod and dactyl; H, same, dactyl; L, same, merus. central depth, tapering proximally, unarmed; merus (Fig. 31) 1.1 times propod length, 3.1 times longer than central depth, ventral margin with numerous subacute tubercles (Fig. 5L); ischium 0.9 of propod length, 0.8 of meral length, ventral margin with numerous subacute tubercles, basis and coxa robust, without special features. Fourth pereiopod similar to third, subequal in length, more slender, about 4.4 times third propod length, 4.5 times longer than proximal width, tapering slightly distally, with similar spination, spines slightly smaller. Fifth pereiopod similar, propod 1.2 times third propod length, 6.0 times longer than proximal width, with single distoventral spine only, several distal transverse rows of cleaning setae; fourth and fifth dactyls similar to third, but no ventral denticles discernible. Uropod (Fig. 1M) with protopodite posterolaterally acute; exopod about 0.8 of telson length, 2.0 times longer than broad, lateral margin slightly convex, with numerous submarginal ventral setae, with small acute tooth distally, with adjacent straight spine, about 2.0 times longer than tooth, diaeresis distinct; endopod subequal to exopod, 2.6 times longer than broad. Measurements (mm). Female holotype, postorbital carapace length, 5.2; carapace and rostrum, 6.8; total body 117 A. J. Bruce Fig. 4. Periclimenaeuspulitzerfinali sp. nov., holotype female, Mombasa, Kenya, NTM Cr. 17287: A, major second pereiopod; B, same, chela; C, same, fingers, medial; D, same, lateral; E, same, merus and ischium; F, minor second pereiopod; G. same, fingers; H, same, dactylus. length (approx.), 17.0; major second pereiopod chela, 6.7; minor second pereiopod chela, 5.7. Colouration. Uniform bright yellow. Host. Spirastrella sp., undescribed species [Porifera: Spirastrellidac], det. G. Pulitzer-Finali, 7 January 1975. Etymology. Named in honour of Gustavo Pulitzer-Finali (Trieste, October 1915 - Rapallo, 21 November 2006), in recognition of his studies on the systematics of the Porifera, of which he described 260 species (pers. comm. J.N.A. Hooper), and his assistance in the identification of sponge shrimp hosts from East African waters. Systematic position. Periclimenaeus pulitzerfinali is a typical member of its genus without major morphological differences from other species of the genus, while still being readily distinguishable from all the other species. Of the 54 lndo-west Pacific species of Periclimenaeus, only six have the carapace without supraorbital tubercles, the first pereiopod with fingers of the chela simple, cutting edges entire, the minor second pereiopod dactyl with a simple cutting edge, the third ambulatory pereiopod dactyl biunguiculate with the unguis unarmed, the ventral corpus denticulate and lacking a basal process, and the exopod of the uropod with a distolateral tooth and a mobile spine only. Of these six species that fully meet this character list, all can be distinguished from P. pulitzerfinali as follows: Periclimenaeus gartlii Bruce, 1976 scaphoccrite with distolateral tooth small, falling far short of distal lamella margin; first pereiopod fingers as long as palm, tips simple; third pereiopod dactyl with stout recurved access tooth. Periclimenaeus hebedactylus Bruce, 1970 second pereiopod fingers distally truncate, lateral posterior telson spines preterminal. Periclimenaeus lobiferus Bruce, 1978 with large antero¬ median dorsal lobe on first abdominal tergite, dactyl of first pereiopod with conspicuous dorsal tuft ol long setae. Periclimenaeus nielbrucei Bruce, 2006 rostral dentition 11/0, with distal teeth divergent, corpus of third ambulatory dactyl with robust denticles. 118 Periclimenaeus pulitzerfinali sp. nov., from Kenya Fig. 5. Periclimenaeus pulitzerfinali sp. nov., holotype female, Mombasa, Kenya, NTM Cr. 17287. Moutlipart details: A, incisor process; B, molar process, dorsal; C, same, ventral; D, maxillula, palp; E, same, distal upper lacinia; F, same, marginal spine; G, third maxilliped, distolateral ischiomerus; H, same, proximomedial ischiomerus. Ventral pereiopodal tuberculations; I, major second pereiopod, palm; J, same, merus; K, same, ischium; L, third pereiopod, merus. Periclimenaeus solitus Brace & Coombes, 1995 rostral dentition 3/0; scaphocerite with distolateral tooth small, exceeding lamella margin, first pereiopod fingers more that half palm length, palm and carpus non-tuberculate, minor second pereiopod fixed finger simple, third pereiopod merus ventrally non-tuberculate. Periclimenaeus usitatus Bruce, 1969 with ventral rostral tooth, dentition 5/1, major second pereiopod with fixed finger grooved, edges of grooves covered with minute acute tubercles, corpus of third ambulatory dactyl with corpus without ventral denticulations. Periclimenaeus pulitzerfinali most closely resembles P. solitus Bruce & Coombes, 1995 from the Cobourg Peninsula, Northern Territory, Australia, and known only from the holotype specimen. It differs in having a rostral dentition of 7/0 (versus 3/0), the scaphocerite with a strong distolateral tooth reaching to the distal margin of the lamella (versus a small tooth, slightly exceeding the lamella), the first pereiopod with the palm and propod minutely tubcrcuiate (versus non-tuberculate), the major second pereiopod with the fixed finger bidentate (versus simple), third pereiopod propod ventrally minutely tuberculate (versus non-tuberculate), distovcntral spines simple (versus minutely dentate), dactyl with unguis about 0.5 of corpus length (versus 0.8), corpus with 4 acute ventral denticles, (versus 3 more robust small teeth), distal accessory tooth more acute, slender, 0.28 of unguis length (versus more robust, 0.15 of unguis length), dorsal telson spines shorter, 0.1 of telson length (versus 0.18). Remarks. Of the 54 presently known Indo-west Pacific species of Periclimenaeus, 32 are known only from the type material. Of these, 23 are known only from the holotype specimen only. Periclimenaeus species generally are found in small sponges or ascidians as heterosexual pairs. The males, being smaller than the females, seem to be frequently overlooked. Five species are known from heterosexual pairs only, and more numerous specimens may occur in larger hosts. Periclimenaus echinimanus Duris, el ai, 2011 is unusual in that the type material consists of 16 specimens, from six sponge hosts. Where numerous specimens of a species are available they show a high degree of morphological consistency in small details. Consequently an unidentified specimen can be readily distinguished on the basis of its motphology. Indeed, the author has a ‘cheat-sheet’ atlas of the third pereiopod dactyls of the Indo- west Pacific species of Periclimenaeus which generally 119 A. J. Bruce allows, with occasional exceptions, the almost immediate identification of a specimen so that only confirmation by comparison with the original or more modem description is required. The number of Periclimenaeiis species known from the waters of Kenya, Zanzibar and Tanganyika is now 19 and a list is appended below, with their original references and other major descriptive reports. This region includes the type localities for six species and five species are not known from elsewhere in the Indo-West Pacific region. LIST OF CENTRAL EAST AFRICAN PER1CLIMENAEUS SPECIES (KENYA, ZANZIBAR, TANGANYIKA) Periclimenaeiis arabicus (Caiman, 1939) Periclimenes (Periclimenaeiis) arabicus Caiman, 1939: 210-211, fig. 4. Periclimenaeiis arabicus. - Holthuis 1952: 13, 130. Kenya: Mombasa (Bruce 1976b); North Kenya Banks, 82 m, Wasin Island, Wasin Channel; Mombasa, Old Port (Bruce 1976c). Zanzibar: Prison Island; Bawi Island; Chawamba Reef (Bruce 1976c). Tanganyika: Kunduchi (Bruce 1976c). Also known from Oman to Fijian Islands. Periclimenaeiis ardeae Bruce, 1970 Periclimenaeiis ardeae Bruce, 1970: 310-312; 2005: 395, 397, fig. 5. Kenya: Mombasa, Ras Iwatine (Bruce 1976b). Also known from type locality, Heron Island, Great Barrier Reef, Australia. Periclimenaeiis bidentatus Bruce, 1970 Periclimenaeiis bidentatus Bruce, 1970: 305-307; 1991: 254-255, fig. 18. Kenya: Mombasa, Shelly Beach (Bruce 1976b). Zanzibar: Bawi Island, type locality (Bruce 1970); Zanzibar Harbour; Mkokotoni Flarbour (Bruce 1976c). Also known from Papua New Guinea, New Caledonia and Australia. Periclimenaeiis djiboutensis Bruce, 1970 Periclimenaeiis djiboutensis Bruce, 1970: 307-308; 1978:256-259, figs 28-29. Zanzibar: Mangapwani, in Plakinastrella ceylonica (Dendy, 1905) (Bruce 1976c). Also known from Israel, Jibuti, Madagascar, Vietnam and Queensland. Periclimenaeiis fawatu Bruce, 2006 Periclimenaeiis fawatu Bruce, 2006a: 33^13, figs 1-6. Zanzibar: Fungu Fawatu, Unguja, type locality (Bruce 2006a). Known from type locality only. Periclimenaeiis hebedactylus Bruce, 1970 Periclimenaeiis hebedactylus Bruce, 1970: 308-310. Zanzibar: Makunduchi, 91.5 m, type locality (Bruce, 1970). Known from type locality only. Periclimenaeiis hecate (Nobili, 1904) Coralliocaris hecate Nobili, 1904: 232; 1906: 58, pi. 3, fig. 2. Periclimenaeiis hecate .-Bruce 1974: 1574-1577, figs. 11— 12, 13e; 1976b: 22-30, figs 8-11. Kenya: Watamu (Bruce 1976a). in Diplosoma cf. modestum. Also reported also from Jibouti to Tahiti. Periclimenaeiis leptodactylus Fujino & Miyake, 1968 Periclimenaeiis leptodactylus Fujino & Miyake, 1968: 90-95, figs 3-5. Zanzibar: Grave Island, Chumbe Island, Shangani (Bruce 1976c). Kenya: Wasin Channel (Bruce 1976c). Otherwise known only from the type locality in Amarni Islands, Japan. Periclimenaeiis miniitiis Holthuis, 1952 Periclimenaeiis minutus Holthuis, 1952: 134—137, figs. 57-59; Bruce 2010a: 9, fig. 6F-L. Zanzibar: Mbweni; Fuwatu Reef (Bruce 1976c). Type locality, Banda Island, Indonesia. Also reported from Somalia, Philippines and Western Australia Periclimenaeiis aff. minutus Holthuis, 1952 Periclimenaeiis minutus. — Bruce 2010a: 9 (not P. minutus Holthuis, 1952). Tanganyika: Pangani Bay (Bruce 1976c). Periclimenaeiis pulitzerjinali sp. nov. Kenya: Mombasa: present report. Periclimenaeiis quadridentatus (Rathbun, 1906) Coralliocaris quadridentatus Rathbun, 1906: 920, fig. 69, pi. 24, fig. 1. Periclimenaeiis quadridentatus. - Bruce 1976c: 473. Kenya: Mombasa, Shelly Beach (Bruce 1976b, as P. stylirostris). Type locality, Auau Channel, Hawaiian Islands, 51-79 m (Rathbun 1906). Also reported from Mariana Islands. Periclimenaeiis rhodope (Nobili, 1904) Coralliocaris (Onycocaris ) rhodope Nobili, 1904; 1906: 61, pi. 2 fig. 8. Periclimenaeiis rhodope. — Holthuis 1952: 125—129, figs 54, 55, 55bis; Bruce 1974: 1558-1562, figs 1-2, 3A,B, 7A,B. Zanzibar: Chwaka, Makunduchi, Chumbe Island (Bruce 1976c). Tanganyika: Kunduchi (Bruce 1976c). Kenya: Shimoni, Ras Iwatine, Banburi, Jadini, Mombasa Island (Bruce 1976c). Also known from Somalia, Seychelle Islands, and Queensland, possibly from China (South China Sea). Periclimenaeiis robustus Borradaile, 1915 Periclimenaeiis robustus Borradaile, 1915: 213; 1917: 324, 278, pi. 55, fig. 20; Bruce 1976c: 473; 2005: 390-395, figs 1-3. Kenya: Mombasa, Ras Iwatine, 119-141 m (Bruce 1976c). Also known from type locality, Seychelle Islands. Periclimenaeiis trispinosus Bruce, 1969 Periclimenaeiis trispinosus Bruce, 1969: 169-170; 2001: 147-152, figs 1-3. Zanzibar: Mkokotoni, type locality (Bruce 1969). Known from type locality only. 120 Periclimenaeus pulitzerfinali sp. nov., from Kenya Periclimenaeus tuamotae Bruce, 1969 Periclimenaeus tuamotae Bruce, 1969: 170-172. Kenya: Mombasa Island, Fort Jesus; Wasin Island (Bruce 1976b). Tanganyika: Kunduchi (?) (Bruce 1969). Type locality: Mururoa Atoll, Tuamotu Islands. Also reported from Northern Territory and Queensland, Australia. Periclimenaeus uropodialis Barnard, 1958 Periclimenaeus uropodialis Barnard, 1958: 18-20, fig. 6. Kenya: Mombasa, Old Port; Wasin Channel (Bruce 1976c). Zanzibar: Mkokotoni (Bruce 1976b). Tanganyika: Mafia Island (Bruce 1976c). Type locality: Dclagoa Bay, Mozambique. Also reported from Queensland, Australia. Periclimenaeus usitatus Bruce, 1969 Periclimenaeus usitatus Bruce, 1969: 172-174; 2010b: 28-29, fig. 6. Zanzibar: Off Unguja: 7°46’48’'S. 39°42’36”E, 20 m, type locality (Bruce 1969). Known from type locality only. Periclimenaeus zanzibaricus Bruce, 1969 Periclimenaeus zanzibaricus Bruce, 1969: 174-175; 2006b: 15-20, figs 8-9. Kenya: Ras Iwatine; Mombasa (Bruce 1976b). Zanzibar: Uroa, type locality (Bruce 1969); Marumbi (Bruce 1976c). Otherwise reported only from Queensland, Australia. ACKNOWLEDGEMENTS I am most grateful to Prof. Maurizio Pansini, University of Genova, Italy, for his information on Gustavo Pulitzer- Finali. This study was facilitated by support from the Australian Biological Resources Study. REFERENCES Barnard, K.H. 1958. Further additions to the Crustacea fauna list of Portuguese East Africa. Memories do Museu Dr. Alvaro de Castro 4: 3-23. Borradaile, L.A. 1915. Notes on carides. Annals and Magazine of Natural History (8) 15: 205-213. Borradaile, L.A. 1917. On the Pontoniinae. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership of Mr J. Stanley Gardiner. Transactions of the Linnean Society of London, Zoology’ (2) 17: 323-396, pis. 52-57. Bruce, A.J. 1969. Preliminary descriptions of ten new species of the genus Periclimenaeus Borradaile, 1915 (Crustacea, Decapoda Natantia. Pontoniinae). Zoologische Mededelingen, Leiden 44 (12): 159-175. Bruce, A.J. 1970. Further preliminary descriptions of new species of the genus Periclimenaeus Borradaile, 1915 (Crustacea, Decapoda Natantia. Pontoniinae). Zoologische Mededelingen, Leiden 44 (21): 305-315. Bruce, A.J. 1974. Observations upon some specimens of the genus Periclimenaeus Borradaile (Decapoda Natantia, Pontoniinae) originally described by G. Nobili. Bulletin du Museum national d’Histoire naturelle, 3‘ serie 258, Zool. 180: 1557-1583. Bruce, A.J. 1976a. A report on a small collection of pontoniine shrimps from the northern Indian Ocean..Journal of the Marine Biological Association of India 16 (2)( 1974): 437-545. Bruce, A.J. 1976b. A report on a small collection of shrimps from the Kenya National Marine Parks at Malindi, with notes on selected species. Zoologische Verhandelingen, Leiden 145: 1-72. Bruce, A.J. 1976c. A synopsis of the pontoniinid shrimp fauna of central East Africa. Journal of the Marine Biological Association of India 16 (2)( 1974): 462-490. Bruce, A.J. 1978. A report on a collection of pontoniine shrimps from Madagascar and adjacent seas. Zoological Journal of the Linnean Society 62: 205-290. Bruce, A.J. 1991. Shallow-water palaemonoid shrimps from New Caledonia (Crustacea: Decapoda). Pp. 221-279, figs 1-31. In: Richer de Forges, B. (edj. Le benthos desfonds meubles des logons de Nouvelle-Caledonie. [Benthos of the soft bottoms in New Caledonian lagoons.] Editions de l’ORSTOM: Paris. Bruce, A.J. 2001. A re-examination of Periclimenaeus trispinosus Bruce and P. orbitospinatus Bruce (Crustacea: Decapoda: Pontoniinae). Zoologische Mededelingen, Leiden 75(9): 147-158. Bruce, A.J. 2005. A re-description of Periclimenaeus robustus Borradaile, the type species of the genus Periclimenaeus Borradaile, 1915 (Crustacea: Decapoda: Pontoniinae). Cahiers de Biologic Marin 46: 389-398. Bruce, A.J. 2006a. Periclimenaeus fawatu spec. nov. (Crustacea: Decapoda: Pontoniinae), from Zanzibar. Zoologische Mededelingen, Leiden 84(3): 33-43. Bruce A.J. 2006b. Periclimenaeus nielbrucei sp. nov. (Crustacea: Decapoda: Pontoniinae), a new sponge associate from the Capricorn Islands, Queensland, with notes on related Periclimenaeus species. Zootaxa 1224: 1-22. Bruce, A.J. 2010a. Periclimenaeus devaneyi sp. nov., from Oahu, Hawai'i (Crustacea: Decapoda: Pontoniinae). Zootaxa 2372: 339-388. Bruce, A.J. 2010b. More pontoniine shrimps (Crustacea: Decapoda: Palaemonidae) from the CReefs 2009 Heron Island expedition. Zootaxa 2604: 20-36. Bruce, A.J. & Coombcs, K.E. 1995. The palaemonoid shrimp fauna (Crustacea: Decapoda: Caridca) of the Cobourg Peninsula, Northern Territory. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 12: 101-144. Caiman, W.T. 1939. Crustacea: Caridea. The John Murray Expedition 1933-1934, Scientific Reports 6: 183-224. Fujino, T. & Miyake, S. 1968. Description of two new species of pontoniid shrimps (Crustacea, Decapoda, Palaemonidae) commensal with sponges. Ohmu. Occasional papers of Zoological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan 1: 85-96. Holthuis, L.B. 1952. The Decapoda of the Siboga Expedition. Part XL The Palaemonidae collected by the Siboga and Snellius Expeditions with remarks on other species. 11. Subfamily Pontoniinae. Siboga Expedition Monograph 39a 10: 1-252, figs 1-110, tab. 1. Nobili, G. 1904. Diagnoses prcliminairc de vingt-huit especes nouvelles de stomatopodes et decapodes macroures de la Mer Rouge. Bulletin du Museum national d’Histoire naturelle 10: 228-238. Nobili, G. 1906. Faune carcinologique dc la Mer Rouge. Decapodes et stomatopodes. Annales des Sciences naturelles, Zoologie (9) 4(1-3): 1-347. Rathbun. M.J. 1906. The Brachyura and Macrura of the Hawaiian Islands. US Fish Commission Bulletin 23(3): 827-930. Accepted 20 October 2011 121 ' "• . > The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 21: 123-133 A new species and new records of Oncinopus (Crustacea: Brachyura: Inachidae) from northern Australia PETER J. F. DAVIE Queensland Museum, PO Box 3300, South Brisbane OLD 4101, AUSTRALIA peter.davie@qm.qld.gov.au ABSTRACT A new species of the Indo-west Pacific majoid spider crab genus Oncinopus is described from Darwin Harbour (Northern Territory), as well as the North West Shelf (Western Australia) and the Gulf of Carpentaria (Queensland). It can be distinguished from other species by differences in the morphology of the male first gonopod, rostral shape, the setation of the dactyl of the first pereiopod, and the length of the last two pairs of pereiopods. Oncinopus angustifrons Takeda & Miyake, 1969 is recorded from the North West Shelf, and is the first record of this species from Australia. Oncinopus aranea (Dc Haan, 1839) is recorded from northwestern Australia for the first time. Five species of Oncinopus are now recognised, with four of these now known from Australia. A key to distinguish them is provided. Keywords: Inachidae, northern Australia, North West Shelf, Darwin, taxonomy, new species, new records. INTRODUCTION Species in the majoid genera Oncinopus and Achaeus (family Inachidae) are distinctive in having the dactyli and propodi of their fourth and fifth pereiopods subchelate, very much like the ‘carrying crabs’ of the families Dorippidae and Dromiidae. Unlike those crabs however, inachids do not use these structures for carrying objects over their back, but for clinging tightly to the substrate in areas with strong currents, with their carapaces, chelipeds and other legs extended in the current to gather food (Ng et al. 2008). In particular, the first two pairs of ambulatory legs of these genera have remarkably long comb-like spines used to trap food particles. Oncinopus has been considered to contain four species: 0. aranea (De Haan, 1839); O. neptunus Adams & White, 1848; O. angustifrons Takeda & Miyake, 1969; and 0. postillonensis Griffin & Tranter, 1986. Ng et al. (2008) also listed, in error, O. subpellucidus Stimpson, 1857 as a valid species, with O. angulatus Haswell, 1880 as a junior synonym. However, both species were described from Port Jackson, New South Wales, and both were treated as junior synonyms of O. neptunus by Griffin & Tranter (1986), who found only O. neptunus to be present in Port Jackson. Griffin & Tranter (1986: 38) also indicated that they had seen a female specimen from the Mortensen Pacific Expedition to Mindoro, Puerto Galera, the Philippines, which they believed to be a new fifth species, but refrained from describing it because they lacked a male. The male first gonopod of Oncinopus species is very unusual amongst majoids in being strongly twisted about three times. This was critically described by Takeda & Miyake (1969), who first recognised the importance of this structure in species discrimination. The present study was stimulated by the discovery of the new Oncinopus species during identifications of beam- trawl catches, following a environmental impact survey of Middle Arm, Darwin Harbour, Northern Territory, for the development of the new city of Weddell. Further investigation revealed that this new species was also present in other unidentified material from northern Australia in the collections of the Queensland Museum. Also amongst this latter material were two other Oncinopus species representing new distributional records, and these are also reported here. Oncinopus aranea and O. neptunus are widespread Indo-west Pacific species, and well known from Australian waters (Griffin & Tranter 1986; Davie 2002; Poore 2004). However the other two species, O. postillonensis and O. angustifrons , appear to have a more limited range, so far being only known from southern Japan, the East China Sea, the Philippines and Indonesia (see Griffin & Tranter 1986; Marumura & Takeda 2009). Thus, it was of interest to discover O. angustifrons amongst the present material from north-western Australia. Poore et al. (2008) have already tentatively identified O. cf. angustifrons from somewhat further south in Western Australian waters, and thus, given the present material, it is more likely that their identification was correct, although their material was not re-examined as part of this study. Institutional abbreviations: NTM, Museum and Art Gallery of the Northern Territory (formerly Northern Territory Museum), Darwin; QM, Queensland Museum, Brisbane. 123 P. J. F. Davie Fig. 1. Oncinopus kathae sp. nov., dorsal views. A, holotype female (NTM-Crl7096); B, paratype male (NTM-Cr 17097). 124 Oncinopus (Inachidae) from northern Australia Abbreviations: cb, carapace breadth; Gl, male first gonopod; P2-P5, pereiopods 2-5. Measurements are of carapace breadtli at the widest point followed by carapace length. SYSTEMATICS Family Inachidae Macleay, 1838 Oncinopus De Haan, 1839 Gender masculine. Type species, by monotypy, Inachus (i Oncinopus ) aranea De Haan, 1839. Recent. Japan (no specific locality given). Inachus ( Oncinopus ) De Haan, 1839: 87. Oncinopus. - Miers 1879: 645; 1886: 20; Alcock 1895: 182; Sakai 1938: 204 (key), 206; 1965: 66; 1976: 155; Griffin 1966: 274 (key); Takeda & Miyake 1969: 476; Griffin & Tranter 1986: 34; Davie 2002: 295; Poore 2004: 362-364; Ng, Guinot & Davie 2008: 112. Diagnosis. Carapace rounded triangular, depressed, without spines; rostrum obtusely bilobate. Eyes non- retractile. Basal antennal article very slender, free distally, not reaching anterior margin of eyestalk. Weakly developed interantennular septum, no interantennular spine. Last pair of pereiopods in subdorsal position, last two pairs subchelate. Abdomen distinctly 7-segmented in both sexes. Male first gonopod twisted about 3 times. Oncinopus kathae new species (Figs 1-3, 4A, B, 5E ) Material examined. Holo type - NTM Cr. 17096, female (7.1 x 9.8 mm, 8.8 mm postrostral), West Arm, Darwin Harbour, Stn 35, 12°34.997’S, 130°53.668’E, beam trawl, 8.9 m, coll. N. Smit & party, 28 Mar 2011. Paratpes - NTM Cr. 17097, male (6.6 * 9.9 mm, 8.8 mm postrostral), West Ann, Darwin Harbour, Stn 37, 12°35.518’S, 130°53.335’E, beam trawl, 12.6 m, coll. N. Smit & party, 28 March 2011. QM W29099, male (6.0 x 8.8 mm, 8.0 mm postrostral), North West Shelf, Western Australia, CSIRO Stn 02B06BT, 19°04.4’S, 118°47.3’E, beam trawl, 82 m, coll. T. Ward & party, 28 April 1983. QM W23654, male (5.9 x 7.8 mm), North West Shelf, Western Australia, CSIRO Stn 02B05BT, 19°04.3’S, 118°50.5’E, beam trawl, 83 m, coll. T. Ward & party, 27 April 1983. QM W23657, male (5.9 x 7.1 mm), North West Shelf, Western Australia, CSIRO Stn 02B09BT, 19°28.5'S, 118°55.3’E, beam trawl, 40 m, coll. T. Ward & party, 26 April 1983. ADDITIONAL (NON-TYPE) MATERIAL - QM W23656, ovig. female (6.1 x 7.6 mm), North West Shelf, Western Australia, CSIRO Stn 03B03BT, 19°55.9’S, 117°55.5’E, beam trawl, 43 m, coll. T. Ward & party, 26 June 1983. QM-W173I4, male (5.8 x 8.0 mm), female (6.3 x 7.4 mm), Gulf of Carpentaria, 12°00.5’S, 141 0 11,9’E, dredged, R.V. Southern Surveyor, CSIRO/QM party, 37 m, 3 December 1990. Description. Carapace sub-triangular, more widely flanged posteriorly adjacent leg bases; carapace width approx. 0.75 (male) to 0.8 (female) postrostral carapace length. Surface smooth, but obscured by epibenthic fouling when alive; regions somewhat defined, with long swollen postrostral gastric elevation with slightly elevated mesogastric tubercle; rounded cardiac elevation armed with a pair of indistinct blunt lateral tubercles; intestinal region smooth. Rostrum of 2, narrow, triangular, flattened, blunt spines; outer lateral margins subparallel, separated by a broad V-shaped hiatus; length approx 0.1 times postrostral carapace length; inner margins with long curled stout setae; outer margins with shorter straight setae. Supraorbital eave not obviously broadened, but distinctly elevated, eyestalks clearly visible dorsally for most of their length; eyestalk short and stout, cornea enlarged. Hepatic margin produced laterally as broad rounded triangle, often quite pronounced. Branchial region with longitudinal rows of curled hairs on dorsal margin. Basal antennal article slender, smooth, distally free; second segment subequal or slightly shorter than basal segment, similarly slender; third segment about half length second; antennal flagellum long. Frontal margin deflexed below rostrum as a broad rounded lobe, not visible in dorsal view; interantennular septum obsolete. Antennules slender; palps folding transversely. Third maxilliped smooth; ischium with a low, longitudinal ridge below articulation with merus, inner margin produced as a long broad rounded lobe extending about two-thirds length of merus; merus longer than wide, anterolateral angle not produced, inner lateral margin with raised rim. Pterygostomian region narrow, smooth; margin weakly indicated, smooth. Sternum generally smooth; sternites 3-4 deeply laterally excavated, with high narrow medial longitudinal ridge. Male cheliped stout, relatively short, smooth; ischium and merus with longitudinal row of long bristle-like setae ventrally on outer face and a row of shorter hooked setae dorsally; carpus elongate, inner margin broadly rounded, with row of long bristle-like setae, outer face with shorter hooked setae; chela with short hooked setae dorsally and ventrally; palm approx. 2.3 times as long as high; fingers approx. 0.4 times as long as palm, weakly incurved; a narrow, smooth gape between fingers in proximal two thirds, finely dentate distally. First ambulatory leg (measured from attachment to tip of dactyl) about 2.5 times (approx. 2.6) postrostral carapace length (female holotype), smooth, propodus and daetylus dorsoventrally flattened, length of propodus about 4.4 times width. P2 and P3 similar; anterior and posterior margins of carpus, propodus and daetylus with fringing rows of long strong setae; a row of short curled setae also present dorsally on merus, carpus and propodus; daclyli with a thick brush of long, softer setae on inner margin. P4 and P5 similar, subchelate, sexually dimorphic being obviously shorter in female; P5 shortest, angled dorsally, length (measured from attachment to base of dactyl) approx. 1.7 times postrostral carapace length in male, and 1.3 times in female; merus of P5 6.5 times 125 P. J. F. Davie Fig. 2. Oncinopus katliae sp. nov., ventral views. A, holotype female (NTM-Crl 7096); B, paratype male (NTM-Cr 17097). longer then wide in male versus 5.1 times in female; merus, carpus and propodus with sparse, long, plumose setae; carpi with cluster of plumose setae disto-ventrally; propodus arched, bearing thick setae on inner margin; dactyl strongly recurved, armed with row of 6-8 small teeth ventrally along its length. Male abdomen broad, 7 free segments, third somite with lateral margin strongly produced, convex; somites 4-6 longitudinally ridged, with a medial tubercle anteriorly; sixth somite about two thirds as wide as third segment, width about 3.5 times length; tclson broadly rounded, approx. 2.8 times wider than long. Female abdomen with fused somites; 126 Oncinopus (Inachidae) from northern Australia a large medial tubercle on each of somites 3-5; a low medial ridge on sixth somite and telson. Female gonopore with simple ventral opening, not elevated; holotype with 9 large eggs. Male first gonopod twisted, aperture terminal and abdominal; first major twisted section very wide and stout, second twist much narrower; distal portion beyond last twist tubular, extended about 0.3 times total length of Gl, slightly recurved apically, tip truncate, not developed into prominent snout. Remarks. Oncinopus kathae sp. nov. can be distinguished from other species of Oncinopus using the key provided. As pointed out by Takeda & Miyake (1969), gonopod morphology appears to be very good for species discrimination (see Fig. 5). By this feature, O. kathae is probably closest to O. neptunus but differs from that species in having a long, untwisted, tubular distal portion (Fig. 5E), and noticeably shorter pereiopods 3 and 4 (cf. Fig. 1B and 6A). The next most similar species is O. angustifrons, but again, the Gl of O. kathae differs by its simple tubular tip, and enlarged first twisted section which is much broader than the second twist (cf. Fig. 5B and 5E). Also the rostral lobes of O. angustifrons are more widely divergent, and the outer lateral margins are also divergent in that species (Fig. 4D, E), versus subparallel in O. kathae (Fig. 4A, B). Oncinopuspostillonensis differs from all others by the long, Fig. 3. Oncinopus kathae sp. nov.. A, B, D: paratype male (NTM-Crl7097); C: holotype female (NTM-Crl7096); A, dorsal view of male pereiopods 4 and 5; B, third maxilliped; C, detail of female second pereiopod showing carpus, propodus and dactylus; D. ventral view of propodus and dactylus of male pereiopods 4 and 5. 127 P. J. F. Davie Fig. 4. A, B, Oncinopus kathae sp. nov., paratype male (NTM-Crl 7097), dorsal and ventral views of frontal region. C-F, Oncinopiis angustifrons Takeda & Miyake, 1969: C, male (4.3 mm cb) (QM-W23655); D. carapace of holotype; E, ventral view of frontal region of allotype female; F, second perciopod of holotype. (Figures D-F, after Takeda and Miyake, 1969: fig. 4). 128 Oncinopus (Inachidae) from northern Australia dorsoventrally flattened rostral lobes (Fig. 6D. E) and by the broad interorbital region which conceals most of the eyestalk (Fig. 6E). Finally, O. aranea also differs in G1 morphology by its shorter distal section ending in a ‘duck’s head’, and by having the second twist of the G1 at least as large and prominent as the first (Fig. 5C). Poore et al. (2008) reported a single specimen of Oncinopus aranea from off the Western Australian coast (27°08’S) from 414 m depth. The carapace shows a marked resemblance to O. kathae sp. nov. however, as it appears to be a female and missing most limbs, it can not be unambiguosly attributed to the present new species, at least not without direct examination. The presence of a thick brush of long, softer setae on the inner margin of the dactyli of P2 and P3 (Figs 1 A, 3C), has not been noted or illustrated for other species (cf. Fig. 6) and appears to be a unique character for O. kathae sp. nov. It is not present in the specimens of O. angustifrons or 0. aranea studied here. Oncinopus kathae sp. nov. also appears to have fewer, and relatively larger, eggs than the ovigerous female of 0. angustifrons, and it would be interesting to compare this character with other species. Habitat. Although field notes were made on the type of bottom for most of the West Ann stations sampled during the Weddell Survey, they were not recorded specifically for stations 35 or 37. However, it can be inferred to have been a mixture of low rocky clumps on a muddy sand substrate. Nearby sites were noted as being gravelly mud with some shell grit, or gravelly sandy mud. The North West Shelf sites were located in deeper, much cleaner, offshore areas, and there were small amounts of clean coral sand associated with some of the specimens. The depth range for Oncinopus kathae sp. nov. is 8 to 83 m. Etymology. Named for my beautiful wife Kathleen - because the promise of a new species is long overdue, and not because of any morphological similarities! Oncinopus angustifrons Takeda & Miyake, 1969 (Figs 4C-F, 5B) Oncinopus angustifrons Takeda & Miyake, 1969: 478- 481 figs 4, 5c, d; Takeda 1973: 31: Sakai 1976:156; Takeda & Kurata 1976: 124; Griffin & Tranter 1986: 34-35; Sakai et al. 2004: 255, 4 figs; Ng, Guinot & Davie 2008: 112. ? Oncinopus angustifrons. - Poore et al. 2008: 59, unnumbered fig. Material examined. QM W23655, 2 males (3.1, 4.3 mm cb) 3 females (4.0, 5.1, 5.4 mm cb), North West Shelf, Western Australia, CS1RO Stn 02B06BT, 19°04.4’S, 118°47.3’E, beam trawl, 82 m, coll. T. Ward & party, 28 April 1983. QM W29100 (ex W23654), male (4.8 mm cb), North West Shelf, Western Australia, CSIRO Stn 02B05BT, 19°04.3’S, 118°50.5’E, beam trawl, 83 m, coll. T. Ward & party, 27 April 1983. QM W29101 (cxW23657), ovig. female (4.8 mm), North West Shelf, Western Australia, CSIRO Stn 02B09BT, 19°28.5’S, 118°55.3’E, beam trawl, 40 m, coll. T. Ward & party, 26 April 1983. Diagnosis. Carapace strongly tapered anteriorly from broad hepatic region to constriction behind rostral lobes; gastric region circular, convex, median posterior part faintly demarcated from greater remaining part; cardiac region isolated, distinctly bifid at summit; rostral lobes weak, triangular, not dorso-ventrally flattened, separated by broad hiatus, lobes divergent; interorbital region only concealing basal part of unretracted eyestalk. Chelipeds slender, merus fringed with row of long, stout, setae along both borders; palm not swollen, slightly shorter than fingers. Male G1 strongly widened subapically; female gonopore weakly elevated, opening more posteriorly than medially. (After Sakai et al. 2004). Remarks. The Australian material agrees very well with the detailed description of Takeda & Miyake (1969), and in particular the male G1 is a perfect match with their figure (see Fig. 5 B). Interestingly, at some sites on the North West Shelf, it was collected sympatrically, in the same beam trawls, with specimens of the new species, O. kathae. Poore et al. (2008) tentatively identified O. cf. angustifrons from somewhat further south in Western Australian waters and although that material has not been studied as part of the present report, the present specimens add weight to their identification being correct. Distribution. Japan - Ogasawara-shoto (Takeda & Miyake 1969, Takeda 1973); East China Sea (Takeda & Miyake 1969); Philippines - San Bernadino Strait (Griffin & Tranter 1986), Sulu Archipelago - off Jolo (Griffin & Tranter 1986); Indonesia - Kepulauan Kai and Aru (Griffin & Tranter 1986). Now extended to the North West Shelf, Western Australia. The bathymetric range for Oncinopus angustifrons is from 36 to 200 m. Oncinopus aranea De Haan, 1839 (Figs 5C, 6B, C) Inachus (Oncinopus) aranea DeHaan, 1839: 100, pi. 29, fig. 2, pi. H; Yamaguchi 1993: 585. Oncinopus aranea. - Adams & White 1848: 3; Miers 1886: 20; Ortmann 1893: 37; Alcock 1895: 183; Rathbun 1902: 133; 1906:879; 1911: 247; Yokoya 1933: 137; Sakai 1936: 80, pi. 18, fig. 3; 1938:206, fig. 3, pi. 21, fig. 3; 1976: 155, pi. 48, fig. 1; Takeda & Miyake 1969: fig. 5e-f; Takeda 1979: 153; Griffin & Tranter 1986: 35; Yamaguchi & Baba 1993: 331, fig. 109; Muraoka 1998: 22; Davie 2002: 295; Poore 2004: 364, fig. 107b; Sakai et al. 2004: 256, 6 figs; Poore et al. 2008: 59, unnumbered fig. Oncinopus araneus. - Sakai 1935: 65; 1965: 66, pi. 27, fig. 1; Serene & Vadon 1981: 123; Ng, Guinot & Davie 2008:112. Nomcnclatural note. The specific epithet aranea is a Latin noun meaning a spider. Dc Haan (1839) introduced it intentionally as a noun in aposition and his spelling has been followed by the majority of authors. Sakai (1935) and 129 P. J. F. Davie Fig. 6. A, Oncinopus neptumis Adams & White, 1848 (original figure from Adams & White, 1848: pi. 2, fig. 1); B, C, O. aranea (De Haan, 1839) (B, from De Haan, 1839, pi. 29 fig. 2; C, lcctotype male, following Yamaguchi & Baba, 1993: fig. 109): D-H, O. postillonensis Griffin & Tranter, 1986 (After Griffin & Tranter 1986): D, carapace, dorsal view; E, frontal region, ventral view; F, second left pereiopod, dorsal view; G, fourth left pereiopod; H, fifth left pereiopod. 130 Oncinopus (Inachidae) from northern Australia Fig. 5. Sternal and abdominal views of male first gonopods from known Oncinopus species. A, (). neptunus Adams & White, 1848 (from Port Jackson, NSW; holotype G1 is figured by Takeda and Miyake, 1969: fig. 3); B, O. angustifrons Takeda & Miyake. 1969 (holotype); C, O. aranea (De Haan, 1839)(from Amakusa Is., Japan); D, O. postillonensis abdominal view (holotype); E, O. kathae sp. nov. (paratype male NTM-Crl7097). A-C after Takeda & Miyake (1969); D after Griffin & Tranter (1986). a minority of subsequent authors incorrectly interpreted it as an adjective. Material examined. QM W23658, male (2.9 mm eb), North West Shelf, Western Australia, CSIRO Stn 01B16BT, 19°59.3’S, 117°03.5’E, beam trawl, 52 m, coll. T. Ward & party, 22 February 1983. QM W23660, male (3.1 mm cb). North West Shelf, Western Australia, CSIRO Stn 01B17BT, 20°00.2’S, 117°00.5’E, beam trawl, 53 m, coll. T. Ward & party. 22 February 1983. Diagnosis (after Sakai et cil. 2004). Carapace moderately tapered from hepatic region to constriction behind rostral lobes; gastric region circular; convex, median posterior part faintly demarcated from remaining greater part; cardiac region isolated, distinctly bifid at summit; rostral lobes weak, triangular - not dorso-ventrally flattened, lobes divergent, intcrorbital region only concealing basal part of unretracted eyestalk. Chclipeds slender, meats fringed with row of long, stout setae along both borders; palm not swollen, slightly shorter than fingers. Male G1 strongly widened distally, with subapical beak-like lateral lobe; female gonopore distinctly elevated, opening medially. Remarks. The two specimens (both male) available to me from northwestern Australia agree with previous descriptions, and with the figure of the male G1 given by Takeda & Miyake (1969). Yamaguchi & Baba (1993) designated a lectotype and provided photographs of the male lectotype and female paralectotype. Distribution. Seychelles and Cargados Carajos (Rathbun 1911); Laccadives (Alcock 1895); Maldives - (Alcock 1895), Nallandu (Rathbun 1902); Sri Lanka (Alcock 1895); Andamans (Alcock 1895); Malay Peninsula (Alcock 1895); Japan - (De Haan 1839), Kagoshima (Ortmann 1893), west of Suno-saki (Yokoya 1933), Shimoda (Sakai 1935), Tatcyama Bay, Misaki, Shimoda, Nanki Shirahama, Ibusuki, and Nagasaki (Sakai 1938), Sagami Bay (Sakai 1965), Tokyo Bay, Sagami Bay, Shimoda, Kii Nagashima, and Kii Minabe (Sakai 1976), Shiono-misaki (Takeda 1979), Tosa Bay (Muraoka 1998); Philippines - north of Lubang Island (Serene & Vadon 1981); Sulu Archipelago - off North Ubian and off Jolo (Griffin & Tranter 1986); Indonesia - Moluccas Passage (Miers 1886), Lesser Sunda Islands, Selat Sunda, Selat Bali, Flores Sea, Scram, and Kepulauan Kai (Griffin & Tranter 1986); Arafura Sea (Miers 1886); Hawaiian Islands- Kauai and Modu Manu (Rathbun 1906). Bathymetric range: 3-400 m; Australia - southern Australian coast (Miers 1886; Poore 2004), Moreton Bay and Saumarez Reef, Queensland (Griffin & Tranter 1986a); range now extended to northwestern Australia. KEY TO SPECIES OF ONCINOPUS 1. Rostral lobes broad, dorsoventrally flattened, separated by a narrow hiatus, inner margins of lobes parallel, outer margins curved, converging distally; intcrorbital region broad, concealing most of unretracted eyestalk. . O. postillonensis Griffin & Tranter - Rostral lobes weak and narrow or triangular, not dorsoventrally flattened, separated by a broad hiatus, lobes divergent or outer margins parallel; intcrorbital region only concealing basal part of unretracted eyestalk.2 2. Male G1 tubular or only slightly widened before apex; female gonopore a simple ventral opening, not elevated.3 Male G1 strongly widened before apex, or with a lateral lobe; female gonopore at least weakly elevated.4 131 P. J. F. Davie 3. Male G1 with apex moderately flanged, last twist extending almost to tip; pereiopods 4 and 5 very long, merus of male P5 approx. 12 times longer than wide. . O. neptunus Adams & White - Male G1 with apex simple, truncate, distal portion tubular beyond last twist, extended about 0.3 times total length of Gl; pereiopods 4 and 5 relatively shorter, merus of male P5 approx. 6.1 times longer than wide. O. kalhae sp. nov. 4. Male Gl strongly widened subapically; female gonopore on a weak elevation, opening more posteriorly than medially. . O. angustifrons Takeda & Miyake - Male Gl with a beak-like lateral lobe subapically; female gonopore distinctly elevated and opening medially.. O. aranea (De Haan) ACKNOWLEDGEMENTS I am pleased to thank Chris Glasby of the Museum and Art Gallery of the Northern Territory for organising my visit to Darwin to examine the Weddell collection, and also Sue Horner of the same museum for sharing her room, and cheerful day-to-day assistance during my visit. 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Rathbun, M.J. 1906. The Brachyura and Macrura of the Hawaiian Islands. Bulletin of the United States Fish Commission 23(3): i-viii, 827-930, figs 1-79, pis 1-24. Rathbun, M.J. 1911. Marine Brachyura. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the Leadership of Mr J. Stanley Gardiner. Vol. Ill, part II, No. XI. Transactions of the Linnean Society' of London (Zoology) (2)14(2): 191-261. figs 1-2, pis 15-20. Sakai, T. 1935. List of marine animals around Shimoda area. Biological Report of Shimoda Marine Biological Station, Tokyo University’ of Literature and Science 1: 23-85, 3 pis, 2 maps. [In Japanese] Sakai, T. 1936. Crabs of Japan: 66 plates in life colours with descriptions. Sanseido: Tokyo. [In Japanese] Sakai, T. 1938. Brachygnatha, Oxyrhyncha. Studies on the crabs of Japan 3. Yokendo: Tokyo. Sakai, T. 1965. The Crabs ofSagami Bay collected by His Majesty the Emperor of Japan. 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Systematic and biogeographic notes on the crabs obtained by dredging at the sea around Cape Shiono-misaki, K.ii Peninsula. Memoirs of the National Science Museum, Tokyo 12: 151-157, figs 1-3. [In Japanese with English summary] Takeda, M.& Kurata. Y. 1976. Crabs of the Ogasawara Islands. 2. First report on the species obtained from stomachs of fishes. Researches on Crustacea 7: 116-137. figs 1-6, tables 1-2. [In Japanese with English summary] Takeda, M. & Miyake, S. 1969. Crabs from the East China Sea III. Brachygnatha Oxyrhyncha. Journal of the Faculty of Agriculture, Kyushu University 15(4): 469-521, figs 1-12, pis 17, 18. Yamaguchi, T. 1993. A list of species described in the Crustacea Volume of Fauna Japonica as belonging to the Japanese Fauna. Pp. 571 -598, figs 1 -2. In: T. Yamaguchi (cd.) Ph. F. von Siebold and Natural Histoiy of Japan. Crustacea. Shimoda Printing Inc.: Japan. Yamaguchi, T. & Baba, K. 1993. Crustacean specimens collected in Japan by Ph. F. von Siebold and H. Burger and held by the National Natuurhistorisch Museum in Leiden and other museums. Pp. 145-563. In: Yamaguchi, T. (ed.) Ph. F. von Siebold and Natural History of Japan Crustacea. Shimoda Printing Inc.: Japan. Yokoya, Y. 1933. On the distribution of decapod crustaceans inhabiting the continental shelf around Japan, chiefly based upon the materials collected by S.S. Soyo-Maru, during the years 1923-1930. Journal of the College of Agriculture, Tokyo Imperial University 12(1): 1-226, figs 1-71, tables 1-4. Accepted 11 November 2011 133 The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 27 : 135-142 Euprymnapardalota sp. nov. (Cephalopoda: Sepiolidae), a new dumpling squid from northern Australia AMANDA REID Australian Museum, Malacology, 6 College Street, Sydney NSW2010, AUSTRALIA Honorary Research Associate, University of Wollongong, Northfields Avenue, Wollongong NSW 2522, AUSTRALIA mandy. reid@austmus.gov.au ABSTRACT Examination of specimens of Euprymna Steenstrup. 1887 from northern Australia led to the discovery of a new species. It is described here as Euprymna pardalota sp. nov. It is distinguished from all but one other nominal species of Euprymna (E. phenax Voss, 1962) in having two rows of suckers on the arms, rather than four rows. It differs from E. phenax in a number of traits, including: the possession of large spots over most of the body, the shape of the funnel organ, the modification of the hectocotylus, and spermatophore length and structure. In addition, the discovery among the Australian Museum collection of a specimen of E. phenax, previously known only from the single male holotype, supports the validity of this taxon, which was previously described as unresolved. Keywords: Cephalopoda, Sepiolidae, sepiolid, Euprymna , Euprymna phenax, Euprymna pardalota, dumpling squid. INTRODUCTION The most recent review of the genus Euprymna Steenstrup, 1887 is that of Norman & Lu (1997). In that work, 12 nominal species were recognised, with five species considered valid: E. benyi Sasaki, 1929; E. hoylei Adam, 1986; E. morsei (Vcrrill, 1881); E. scolopes Berry, 1913; E. tasmanica (Pfeffer, 1884) and an undescribed species, E. sp. 1. Two species, E. schneehageni (Pfeffer, 1884) and E. pusilla (Pfeffer, 1884), were considered to be nomena dubia, and the status of E. albatrossae Voss, 1962; E. bursa (Pfeffer, 1884), E. phenax Voss, 1962 and E. stenodactyla (Grant, 1833) was deemed to be unresolved. In the same year, an additional species, E. hyllebergi Nateewathana, 1997, from the Andaman Sea, Thailand, was described. More recently, another species, Euprymna megaspadicea Kubodera & Okutani, 2002 has been recognised from Japan. In Norman & Lu’s (1997) paper, only one species, E. tasmanica, was reported to occur in Australia. It is found in shallow waters around southern Australia from Moreton Bay, Queensland, through to Bass Strait, Victoria. However, two unresolved taxa, one from northern Australia and one from south-western Australia, were mentioned by Norman & Lu (1997). The morphology of the northern unresolved taxon (treated as E. stenodactyla in Lu & Phillips (1985)) is described as very similar to that of E. tasmanica, although DNA analyses has demonstrated that it is distinct (M. Nishiguchi unpublished data). The second unresolved taxon, also very similar to E. tasmanica, is described by Norman & Lu (1997) as represented by insufficient material to fully resolve. (For this reason, the distribution of E. tasmanica in Reid & Jercb (2005) is described as extending north to Shark Bay in Western Australia.) All the abovementioned Australian species have four transverse rows of suckers on the arms. A visit to the Museum and Art Gallery of the Northern Territory in 2008 led to the discovery of a distinct species of Euprymna collected in northern Australian waters. More specimens were subsequently found among the collections of the Western Australian Museum and the Australian Museum. Unlike nearly all other nominal Euprymna species, this one has two transverse rows of suckers on the normal (non-hectocotylised arms). The only described species with two rows of suckers is E. phenax from the Philippines. Comparison of the Australian material with the E. phenax holotype (previously the only known specimen) indicated that the northern Australian form belongs to a new species and can be distinguished from E. phenax by a number of traits. This new species is described as E. pardalota. MATERIALS AND METHODS Terminology, measurements, indices, and abbreviations for anatomical structures follow Roper & Voss (1983), with a few minor differences (Table 1). In the current paper, ASC refers to the total number of suckers on each designated arm (this abbreviation refers to the number of suckers on the basal half of each arm in Roper & Voss (1983), with ASCT 135 A. Reid Table 1. Explanation of measurements and counts. Definitions largely follow Roper and Voss (1983). New or modified definitions are indicated by an asterisk (*). Indices (shown in squar e brackets) are calculated by expressing each measure as a percentage of mantle length Anterior Mantle Join - AMJ: width of attachment of mantle to head [AMJI]. Arm Length - AL: length of each designated (i.e. 1,2 etc.) arm measured from first basal (proximal-most) sucker to distal tip of arm (Arm 1, dorsal; 2, dorso-lateral; 3, ventro-lateral; 4, ventral) [ALI], Arm Sucker Count * ASC: total number of suckers on each designated amt (e.g. ASC2). Arm Sucker diameter - AS: diameter of largest normal sucker on each designated (i.e. 1 , 2 etc.) arm [ASIn]. Club Length - C1L: length of tentacular club measured from proximal-most basal suckers (carpus) to distal tip of club [C1LI]. Club Row Count - CIRC: number of suckers in transverse rows on tentacular club. Club Sucker diameter - CIS: diameter of largest sucker on tentacular club [C1SI]. Egg Diameter -EgD: diameter of largest egg present in the ovary or oviduct [EgDI], Eye Diameter - ED: diameter of eye [EDI]. Fin Insertion - FI: length of fin as joined to mantle [FII]. Fin Insertion anterior * - Fla: anterior origin of fin measured from mantle margin to anterior-most junction of fin and mantle [FHa], Fin Width - FVV: greatest width of single fin [FWI], Free Funnel length - FFu: the length of the funnel from the anterior funnel opening to the point of its dorsal attachment to the head [FFuI], Funnel Length - FuL: the length of the funnel from the anterior funnel opening to the posterior margin measured along the ventral mid¬ line [FuLI]. Head Length - HL: dorsal length of head measured from point of fusion of dorsal arms to anterior tip of nuchal cartilage [HU], Head Width - HW: greatest width of head at level of eyes [HWI]. Mantle Length - ML: dorsal mantle length. Measured from anterior-most point of mantle to posterior apex of mantle. Mantle Width - MW: greatest straight-line ventral width of mantle [MWI], Ventral Mantle Length - VML: length of ventral mantle measured along midline [VMLI]. used for the total number of arm suckers). All measurements are in millimetres (mm). Measurements and counts for individual mature specimens are presented in Table 2; the range of values for each character is expressed in the description as: minimum -mean- maximum (SD). The values for each sex arc given separately. Unless otherwise stated, ranges refer to the four mature male and two mature female specimens available for study. The E. pltenax holotype was measured by the author. In some cases the measurements differ slightly from those tabulated in Voss (1963). Other abbreviations: AM, Australian Museum; MV, Museum Victoria; NTM, Northern Territory Art Gallery and Museum; WAM, Western Australian Museum. For scanning electron microscopy, arm and club suckers (and in one case a whole arm) was removed from the middle of designated anns and the tentacular club, mounted, then air dried and examined in a Zeiss Evo LSI5 SEM using a Robinson Backscatter detector. As only two mature female specimens were available for examination, the female reproductive tract was not removed for examination. Similarly, the digestive tract was not removed to avoid excessive damage to type material. Some difficulty was experienced dissecting out the male reproductive tract intact and pinning it out for examination and illustration, despite attempting to do so with valuable type material. Should more, particularly fresh material, become available, it would be useful to examine and fully describe these internal structures. It was not possible to do so for this work. TAXONOMY Euprymna Steenstrup, 1887. Gender feminine. Type species, by original designation, Inioteuthis morsei Verrill, 1881. Recent. Western Pacific and eastern Indian Oceans. Diagnosis. Broad ligament between head and mantle; commissure greater than one-third of head width. Transverse suckers in 2 or more rows on normal (non-hectocotylised) arms. Stalked suckers in 6 or more transverse rows on tentacular clubs. Left arm 1 hectocotylised in mature males; distally with 2-4 rows of small suckers on elongate, columnar pedicels; suckers with narrow openings and chitinous rims; basal part of hectocotylised arm with normal suckers and 1-2 papillae in ventral sucker row, sometimes bearing tiny sucker(s). Enlarged arm suckers usually present in male (not markedly so in E. phenax and E. pardalota sp. nov.). Paired light organs in mantle cavity, ventral and closely adherent to ink sac. Gladius absent. Euprymna pardalota sp. nov. (Figs 1-9; Tables 1, 2) Material examined. FIolotype-NTM P.15796, 1 cf (8.6 mm ML, mature), Western Australia, Timor Sea, Cartier Reef, rotenone station on slope, 12°32'S, 123°33'E, 13 m, coll. H.K. Larson and M. Selway, 15 March 1990. Paratypes - NTM P.2496, 19 (7.8 mm ML, mature), Western Australia, Timor Sea, Cartier Reef, southern reef flat, rotenone station, 12°32.8'S, 123°32.3'E, 0.2-0.6 m, coll. R.C. Willan, 5 May 1992; NTM P.2497, 9 (9.0 mm ML, mature), Western Australia, Timor Sea, Cartier Reef, 136 A new species of dumpling squid from northern Australia SW corner, spur and groove region on coral reef at low tide, 12°32.8'S, 123°32.4'E, 10-20 m, coll. B.C. Russell, 8 May 1992; WAM S.34548, lcf (8.5 mm ML, mature), Western Australia, Kimberley, northwest end of Long Reef, 13°48'S, 125°47'E, 12 m, coll. F.E. Wells and C.W. Bryce, 17 August 1991; AM C.303908, 1 cf (9.5 mm ML, mature), rotenone station, Queensland, 10°59.980'S, 144°1.220'E, 6 m, coll. Aust. Museum party, Stn Qld 669, 15January 1993; AM C.456837, lcf (9.4 mm ML, mature), Queensland, Great Barrier Reef, E of Lizard Island, Yonge Reef, 1 ml N of platform, coral and sand, back reef, 14°35.00'S, 145°37.00'E, 1-15 m, coll. D.F. Hoese, 1 December 1978. Non-type material - NTM R 17968,1 juvenile (2.8 mm ML, immature), Australia, Western Australia, Timor Sea, boat anchorage off West Island, Ashmore Reef, attracted to light at night, 12°14'S, 122°56'E, 0.5 m, coll. H.K. Larson, 15 September 1987; NTM R15788, 2 juveniles (3.2 mm, 3.5 mm ML, immature), Western Australia, Timor Sea, Ashmore Reef, surface, attracted to light, 12°25'S, 122°57'E, 0.5 m, coll. H.K. Larson, 13 September 1987; NTM P.2494, 1 juvenile (5.2 mm ML, immature), Western Australia, Timor Sea, western end of Cartier Reef, in surge channel on coal reef, 12°32.2'S, 123°31.80'E, 10-14 m, coll. B.C. Russell, 4 May 1992; NTM R.2495, 19 (4.3 mm ML, immature). Western Australia, Timor Sea, SW corner of Cartier Reef, 12°32.6'S, 123°32.2'E, 8-15 m, coll. B.C. Russell, 9 May 1992; AM C.453180, lcf (7.0 mm ML, immature), Queensland, Great Barrier Reef, Linnet Reef, W side, 14° 47.00'S, 145° 20.00'E, 3-15 m, coll. Australian Museum, 22November 1975; AM C.456838, lcf (7.3 mm ML, immature), Queensland, Great Barrier Reef, Capricorn Group, One Tree L, fine sandy sediment, 23° 30.00'S, 152° 28.00'E, 4 m, coll. G. Anderson, 7 February 1975. Other material examined. Euprymna phenax Voss 1962: Holotype- USNM 575328, 1 cf (11.0 mm, mature), Philippines, Panay Island, Naso Point [not Nogas Point as stated in Voss, 1962], 11°10'N 122°30'E, dip net, coll. FV Albatross, 3 February 1908; AM C.453179, lcf (8.1 mm ML, immature), Philippines, Luzon L, Lingayen Gulf, coral, rotenone, 16°26.00'N 119° 55.00'E, 2-15 m, coll. D.F. Hoese and party, 16 April 1980. Diagnosis. Small species, up to 9.5 mm mantle length. Funnel organ dorsal element spade-shaped. Left dorsal arm hectocotylised in male: basally, 2 pairs of normal suckers; third sucker pedicel in ventral row modified into distally directed, enlarged, fleshy papilla, without terminal sucker; following 2-3 rows suckers normal; distal end of arm bears 9-15 pairs of suckers on enlarged columnar sucker pedicels. Spermatophores with discrete cement body, SLI 46.5-75.8. Preserved specimens cream with large, deep purple irregularly-shaped pigment spots on dorsal and ventral head and mantle. Fins with 1-3 large spots close to junction with mantle. Aboral surface of arms with medio- longitudinal row of large dark purplish blotches and smaller spots between base of each sucker and extending on to Fig. 1. Euprymna pardalota sp. nov.: A, dorsal view, male holotype, NTM P.15796, 8.6 mm ML, scale bar 3 mm; B, ventral view, same specimen, scale bar 3 mm. Euprymna plienax Voss: C, dorsal view, male holotype, USNM 535328, 11.0 mm ML, scale bar 3 mm; L), ventral view, same specimen, scale bar 3 mm. sucker pedicels. Aboral surface of club with row of large blotches close to keel and smaller blotches or bars and spots toward club margin. Description. Counts and indices for individual specimens are given in Table 2. Only mature specimens were measured. Species small: ML mature male 8.5-9.0-9.5 (SD, 0.5), female 7.8-8.4-9.0 (SD, 0.8). Mantle short, broad, dome- shaped, rounded posteriorly; MWI male 84.2- 90.0 -101.2 (SD, 7.8), female 70.0-79.9-89.7 (SD, 14). Dorsal mantle joined to head, ventral mantle margin straight or distinct wide ‘m’ shape. Fins rather small, rounded; fin length approx. 50% ML, FI la male 23.5- 30.1- 39.5 (SD, 6.8), female 34.6-35.1-35.6 (SD, 0.7); fin width about 20% ML, FWI male 36.8-40.2^7.1 (SD, 4.7), female 38.5-38.7-38.9 (SD, 0.3), attached dorso-laterally towards posterior half of mantle; posterior margins curved; anterior margins with well-developed lobes, lateral lobes crescentric. Anterior edges of fins do not project to level of anterior mantle margin (Figs 1A, B). 137 A. Reid Fig. 2. Euprymna pardalota sp. nov., male paratype, NTM P.15796, 8.6 mm ML: A, funnel, scale bar 2 mm; B, funnel organ, scale bar 0.5 mm. Funnel long, conical, base broad, tapered, projecting anteriorly beyond junction of ventral arms 4 (Figs IB, 2A); FuLl male 55.3-62.4-69.8 (SD, 6.0), female 66.7- 66.7 - 66.7 (SD, 0); free for most of its length, FFul male 32.7- 38.5-41.2 (SD, 3.0), female 35.6-37.0-38.5 (SD, 2.1). Funnel valve small, flaplike, rounded anteriorly. Funnel organ dorsal element broad, spade-shaped, rounded anteriorly broadening posteriorly with small projection medially (Fig. 2B) and indistinct median ridge; ventral elements ovoid with acute anterior tips. Funnel locking cartilage long, narrow, parallel-sided with shallow groove ot uniform depth (Fig. 2A). Mantle cartilage compliments funnel member: short straight ridge. Flead broader than long in both sexes, HLI male 42.1-5L9-64.7 (SD, 9.4), female 53.8-55.3-56.7 (SD, 12.6); HWI male 91.6-95.7-100.0 (SD, 3.4), female 93.6- 94.0-94.4 (SD, 0.6), only very slightly narrower than mantle width. Eyes large, EDI male 17.0-21.4-25.6 (SD. 4. 7), female 19.2 -22,4- 25.6 (SD, 4.5); ventral eyelids free. Distinct, large photosensitive vesicle on latero-posterior surface of head, posterior and ventral to eye, close to mantle opening. Small pore of unknown function close anterior margin of each eye. Amis broad basally, tapered distally; order 2:3:1:4 or 2:3:4:1 (Table 2). Ann length index of longest ann in male (ALI2) 61.2-73.1-87.2 (SD, 10.9), female (ALL) 72.2- 74.6 - 76.9 (SD, 3.3). All anns similar in shape, U-shaped in section; indistinct keels present on median aboral sides of arms 4 only. Sucker pedicels large, each with fine, curved bilobed lappets on posterior margin. Arms suckers biserial; Fig. 3. E. pardalota sp. nov. male paratype, AM C.456837, 9.4 mm ML: A, SEM, arm 3, right side, scale bar 2 mm: B, SEM, enlargement of sucker rim, scale bar 10 pm: C, hectocotylus, ventral side view, scale bar 1 mm. spherical throughout (Fig. 3A). Chitinous rim of arm suckers without teeth on inner margin. Infundibulum with 3-4 rows of stalked processes, with broader, ovoid, outermost rims; processes contain tufts of finger-like papillae (Fig. 2B), peripheral sucker rim processes rectangular, radially arranged, without finger-like papillae. Suckers on arms 2 and 3 larger than those on amis I and 4 in both sexes; suckers largest medially, basal few and extreme distal few rows of suckers abruptly smaller. Male arms suckers larger than female arm suckers (Table 2) (but not greatly enlarged). Sucker counts range from 20-32 on each ann, mean sucker counts on arms 1-4 in male 23,27, 25 and 22 respectively; mean sucker counts on anns 1-4 in female 27, 30, 30 and 24 respectively. All amis connected by relatively shallow webs, protective membranes absent. Left dorsal ann of male hectocotylised: basally, 2 pairs of normal suckers (not enlarged or modified); third sucker pedicel in ventral row modified into distally directed, enlarged, fleshy papilla, without terminal sucker; following 2-3 rows suckers normal; distal end of ann bears 9-15 pairs of suckers on enlarged columnar sucker pedicels (Fig. 3C); proximal-most sucker pedicels in modified 138 A new species of dumpling squid from northern Australia Fig. 4. Euprymna pardalota sp. nov. male paratype, AM C.303908, 9.5 mm ML: A. club, scale bar 2 mm; B, SEM, scale bar 400 pm; C, enlargement of club suckers, scale bar 35 pm; D, enlargement of individual club sucker, scale bar 10 pm. portion broadest and separate; distal-most columnar sucker pedicels closely packed. Hectocotylised amt slightly longer than corresponding unmodified right dorsal arm. Tentacles long, slender, stalks naked, semicircular in section; oral surface convex. Club relatively short, between 0.25x to 0.3x mantle length; C1L1 male 22.3 -27.5 -34.9 (SD, 5.3), female 25.6- 28,4— 31.1 (SD, 3.9), crescent shaped, tapers to pointed end distally (Fig. 4A, B). Sucker-bearing face of club convex. Suckers approx. 0.1 mm diameter in centre of club; arranged in 6-9 oblique rows. Swimming keel on aboral side of carpus broad, extends posteriorly Fig. 5. Euprymna pardalota male holotypc, NTM P. 15796, 8.6 mm ML: lo, light organ; ap, anal papilla; r, rectum; scale bar 0.5 mm. beyond carpus. Keel forms groove on oral side (Fig. 4A, B). Club sucker dentition (Fig. 4C, D): inner ring without teeth; infundibulum with 3 rows of processes; inner 2 rows sub-rectangular bearing rows of comb-like papillae. At periphery, processes narrower and more elongate, without papillae. Well-developed paired light organs present overlying and joined to ink sac (Fig. 5). Individual lobes rectangular medially, with large, rounded, shoulder-like bulb anteriorly, slightly enlarged, rounded, posteriorly and angled outer margin. Gills with 22-23 lamellae per demibranch (it = 5). Buccal membrane with six lappets and fringed inner margin; suckers absent. Radula with 7 transverse rows of teeth (Fig. 6A). Rhachidian teeth simple, without cusps, triangular, sightly concave laterally and ventrally. First lateral teeth similar in size and shape to rhachidian teeth with pointed cusps displaced laterally and directed towards midline of radula. Second and third laterals with elongate bases, longer than the first, curved. Third laterals with scythe-like teeth, longer than second laterals. Upper beak (Fig. 6B) with long, pointed rostrum, hood curved, high above crest posteriorly; jaw angle only slightly acute (angle almost 90°); lateral wall edge with slight indentation. Lower beak (Fig. 6C, D) with blunt pointed protruding rostrum, rostral edge obtuse with distinct inner angle; hood pointed posteriorly (Fig. 6D); hood notch absent, wings almost straight, widely spread. Distinct dark pigmentation restricted to rostrum and hood of upper and lower beaks. Gladius absent. Fig. 6. Eupiynvia pardalota: A, portion of radula, female paratype NTM P.2497,9.5 mm ML, scale bar 100 pm; B, upper beak, lateral view, male paratype AM C.303908, 9.5 mm ML, scale bar 0.5 mm (broken line indicates damaged margin); C, lower beak, lateral view, male paratype, AM C.456837,9.4 mm ML, scale bar 0.3 mm; D, lower beak ventral view, same specimen, scale bar 0.3 mm. 139 A. Reid Male reproductive tract similar in structure to congeners (Fig. 7). Accessory spermatophoric gland large, well developed. Spermatophores (17 in spermatophore storage sac of specimen AM C.456837) approx. 0.5x to 0.75x mantle length; SpLI 46.5- 61.3 -75.8 (SD, 14.6, /; = 3). Sperm reservoir contains coiled sperm cord. Cement body unipartite; aboral end cup-shaped, connects to sperm reservoir via a narrow duct; oral end broadens toward junction with ejaculatory apparatus (Fig. 8A). Oral end of ejaculatory apparatus with 3-4 simple coils. Female reproductive tract: Ovary large, occupies large proportion of posterior end of mantle cavity; displaces other organs when mature. Opens via single thick-walled oviduct at anterior end on left side. Nidamental glands paired, broad, located ventral to ovary toward anterior end. Inverted cream coloured U-shaped accessory nidamental glands located toward distal end of nidamental glands. Eggs spherical, 0.4 mm diameter (n = 1); EgDI 5.1 (n = 1). Colour in alcohol cream with large deep purple irregularly shaped spots, sometimes with darker centres, on dorsal and ventral head and mantle; spots larger on dorsal surface of mantle and head than those on ventral surface (Fig. 1A-B). Shiny bluish iridophores on head around eyes. Fins with 1-3 large spots (usually one large and 1-2 smaller) close to junction with mantle, otherwise chromatophores absent from fins dorsally and ventrally. Aboral surface of arms with mcdio-longitudinal row of large dark-purplish blotches and smaller spots between base of each sucker and extending on to sucker pedicels. These latter, smaller chromatophores are predominant on ventral side of arms 2-4, with fewer on arms 1. Spots sometimes present between dorsal arm suckers, if present, usually on distal portion of arms. Arms appear barred. Aboral surface of club with row of large blotches close to keel and smaller blotches or bars and spots toward club margin (Fig. 8C). Distinct patterning also visible on juveniles. Habitat. Coral reef. Type locality. Western Australia, Timor Sea, Cartier Reef, 12°32'S, 123°33'E, 13 m. Distribution. Australia: Western Australia, from 12°14'S, 122°56'E to Queensland, 23° 30.00'S, 152° 28.00'E, depth range 0.2 m to 20.0 m (Fig. 9). While the scale of the distribution map would suggest that this species occurs in the open ocean, all specimens collected are, in fact, associated with reefs. Etymology. The species name, pardalota, is derived from the Greek, pardalotus, meaning ‘spotted like a leopard’ and refers to the large and prominent spots that are distinctive in this species. It is adjectival. DISCUSSION The discovery of a new Euprymna in Australian waters is an exciting one. Unlike E. tasmanica, this northern Australian taxon clearly differs in having two rather than Fig. 7. Euprymna pardalota'. Male reproductive tract, male paratype, AM C.456837, 9.4 mm ML, scale bar 2 mm. asg, accessory spermatophoric gland; mg, mucilaginous gland; sg, spermatophoric gland; ss, spermatophore storage sac (sac partially broken). Testis (joined to mucilaginous gland) not shown. Fig. 8. Euprymna pardalota: A, spermatophore cement body, male paratype, AM. C.303908, 9.5 mm ML, scale bar 0.2 mm. Euprymna phenax: B. spermatophore cement body, male holotype, USNM.575328,11.0 mm ML, scale bar 0.1 mm. Euprymna pardalota: C, aboral view of tentacular club, female paratype, NTM P.2496, scale bar 0.5 mm. four transverse rows of arm suckers. The only other nominal species with two rows of ami suckers is E. phenax from the Philippine islands. Comparison of the E. phenax holotype with the northern Australian Euprymna , together with a single additional non-type E. phenax specimen (also from the Philippines), indicates that the taxon from Australia is distinct. A number of morphological characters distinguish the specimens from these two localities. Euprymna phenax has smaller, and many fewer chromatophores than E. pardalota (Fig. 1). The fins of the E. phenax (Fig. 1C, D) are smaller and are positioned more anteriorly on the body than those observed in the Australian specimens. The dorsal member of the funnel organ is spade-shaped in E. pardalota and inverted V-shape in E. phenax. Differences are also apparent in the hectocotylus. While both species have the third sucker pedicel from the base of the arm enlarged into 140 A new species of dumpling squid from northern Australia a fleshy papilla, E. pardalota has only 2-3 pairs of normal arm suckers distal to this papilla (proximal to the series of columnar suckers), while E. phenax has approx, seven pairs of normal suckers distal to the fleshy papilla, followed by about 10 pairs of columnar suckers (versus 9-15 in E. pardalota). The fleshy papilla bears a sucker in the E. phenax holotype (but not in specimen AM C.453179), but none of the male E. pardalota examined had a sucker on this papilla. In addition, based on Voss’s measurements, and measurement of the arm suckers of C.453179, the arm suckers are smaller in E. phenax males than in E. pardalota (Table 2). Variation in the size and arrangement of arm suckers has been used to diagnose other Euprymna species (Norman & Lu, 1997). Many of the suckers are absent in the holotype, a fact noted by Voss (1963: 46) who stated, “one cannot overlook the possibility that one or more may be enlarged in a complete specimen”. However, the more recently collected non-type E. phenax specimen (albeit not fully mature) has intact suckers that arc sightly enlarged Table 2. Euprymna pardalota sp. nov. Measurements (mm), counts and indices of mature specimens of both sexes. Measurements for the E. phenax holotype are included for comparison (some of Voss’ 1963 data were used, for example sucker diameters, as most suckers have now fallen off the holotype). Museum Reg. no. Sex NTM P.2496 paratype female NTM P.2497 paratype female WAM S.34548 paratype male NTM P.15796 holotype male AM C.456837 paratype male AM C.303908 paratype male USMN 575328 E. plienax holotype male ML 7.8 9.0 8.5 8.6 9.4 9.5 11.0 MWI 89.7 70.0 89.4 101.2 85.1 84.2 75.5 AMJI 48.7 44.4 57.6 52.3 47.9 52.6 45.5 VMLI 105.1 105.6 111.8 98.8 98.9 94.7 102.7 FWI 38.5 38.9 47.1 39.5 37.2 36.8 30.0 Flla 34.6 35.6 23.5 39.5 28.7 28.4 30.0 FII 34.6 35.6 41.2 31.4 34.0 29.5 27.3 FuLI 66.7 66.7 61.2 69.8 55.3 63.2 59.1 FFul 38.5 35.6 41.2 40.7 37.2 34.7 32.7 HLI 53.8 56.7 64.7 52.3 56.4 42.1 45.5 HWI 93.6 94.4 100.0 95.3 95.7 91.6 83.6 EDI 19.2 25.6 17.6 25.6 17.0 25.3 14.5 ALII 44.9 55.6 50.6 75.6 63.8 54.7 45.5 ALI2 76.9 72.2 61.2 87.2 69.1 74.7 65.5 ALI3 66.7 66.7 56.5 86.0 63.8 68.4 65.5 ALI4 57.7 50.0 44.7 69.8 53.2 57.9 54.5 ASInl 4.49 4.44 4.12 5.81 2.66 3.68 2.73 ASIn2 5.13 6.11 5.88 8.14 5.32 5.26 3.18 ASIn3 5.13 5.56 5.88 7.56 5.32 5.26 2.27 ASM 4.49 5.56 4.12 5.81 3.62 3.16 2.27 ASCI 28 27 20 24