The wolf spider genus Artoria in New South Wales and the Australian Capital Territory, Australia (Araneae, Lycosidae, Artoriinae)

The wolf spider (Lycosidae Sundevall, 1833) genus Artoria Thorell, 1877 is revised for New South Wales and the Australian Capital Territory, Australia, to include 34 species, 21 of which are new to science: A. albopilata (Urquhart, 1893), A. alta Framenau 2004, A. beaury sp. n., A. barringtonensis sp. n., A. belfordensis sp. n., A. berenice (L. Koch, 1877), A. bondi sp. n., A. booderee sp. n., A. comleroi sp. n., A. corowa sp. n., A. equipalus sp. n., A. extraordinaria sp. n., A. flavimana Simon, 1909, A. gloriosa (Rainbow, 1920), A. grahammilledgei sp. n., A. helensmithae sp. n., A. howquaensis Framenau, 2002, A. kanangra sp. n., A. kerewong sp. n., A. lineata (L. Koch, 1877), A. maroota sp. n., A. mckayi Framenau, 2002, A. mungo sp. n., A. munmorah sp. n., A. myallensis sp. n., A. quadrata Framenau, 2002, A. slatyeri sp. n., A. strepera sp. n., A. taeniifera Simon, 1909, A. terania sp. n., A. triangularis Framenau, 2002, A. ulrichi Framenau, 2002, A. victoriensis Framenau, Gotch & Austin, 2006, and A. wilkiei sp. n. Lycosa pruinosa L. Koch, 1877, currently listed in Artoria, is considered a nomen dubium. Artoria are largely forest dwellers, although some species have preferences for more open areas such as riparian or coastal environments or grasslands. Consequently, the genus mainly occurs east and west along the Great Dividing Range, although some species can be found into the Riverina, Cobar Peneplain and Darling Riverine Plains IBRA regions to the west. Key Words


Introduction
The genus wolf spider genus Artoria Thorell, 1877 includes small, free-roaming spiders with a fairly uniform morphology. They are overall brown in colouration, with variably lighter central and marginal areas on the carapace and often with a light cardiac mark on an otherwise variably mottled abdomen ( Fig. 1A-D). These spiders are somewhat sexually dimorphic, i.e. males are slightly smaller than females and often more distinctly coloured. In addition, males may have modifications of the first pair of legs, for example brush-legged males such in China (Li et al. 2012), four on Pacific islands (New Caledonia, Vanuatu, New Hebrides, Samoa, Marquesas), and two in South-East Asia (Papua New Guinea, Philippines, Indonesia) (World Spider Catalog 2018). Three species of Artoria are currently described from Africa; however, their original descriptions suggest they are misplaced in the genus .
In Australia, species of the genus Artoria are most common in the Bassian and Torresian bioregions, i.e. along the eastern coast, in the south-east (including Tasmania), in south-western Western Australia and in the tropical northern parts of the country. Very few records are from the interior and these are often from near intermittent or permanent water bodies . Artoria species often occur in forests and woodlands, although some species appear to prefer open habitats or riparian environments Framenau et al. 2002).
Prior to this study, 13 species of Artoria were known from New South Wales (NSW) and the Australian Capital Territory (ACT). The Bush Blitz survey to Mungo National Park in 2017 (see http://www.bushblitz.org.au; accessed 20 October 2018) resulted in the discovery of a new species of Artoria which provided the opportunity to apply for funding to resolve the taxonomy of the genus in the region. Previous studies by the senior author suggested a considerable biodiversity of the genus in the eastern part of Australia. Therefore, the aim of this study was to comprehensively revise all species of Artoria in NSW and the ACT.

Material and methods
This study is based on the examination of 1,580 records (vials) including 2,760 males and 1,389 females of Artoria from NSW (1,554 records) and the ACT (26 records) in mainly the Australian Museum, Sydney, and the Australian National Insect Collection, Canberra, supplemented by material from other Australian and overseas institutions (see abbreviations of institutions below). Descriptions are based on spiders stored in 70% ethanol, preferable using recent and representative material in lieu of poorly preserved historical type specimens.
For examination and illustration, female internal genitalia were placed in a Pancreatin solution for a few hours as described in Álvarez-Padilla and Hormiga (2007) to clear soft tissue. Male pedipalps were dissected by removing the bulb from the cymbium and carefully detaching the palea region from the tegulum.
Digital images were taken using a Leica DFC 500 digital camera attached to a Leica MZ16A stereo microscope. To increase depth of field, multiple images were merged using the software package AutoMontage Pro Version 5.02 (p).
Life history data as described for each species generally refer to populations from New South Wales or the Australian Capital Territory. If species are more widespread and occur in different climatic zones (i.e. in   (male from Avon River, Victoria); B, A. mckayi (female with eggsac from Avon River, Victoria); C, A. flavimana Simon, 1909 (female from Capel, Western Australia); D, A. taeniifera Simon, 1909 (female from Ravensthorpe, Western Australia). All species occur in New South Wales; A. mckayi has also been found in the Australian Capital Territory.
This revision recognizes 34 species of Artoria in NSW, five of which currently also known from the ACT (Table 1). Eight species are known only from males and therefore require further survey and taxonomic research to identify the respective females. A number of undescribed females are still present in the collection of the Australian Museum. It was impossible to unequivocally assign these to a male described here. In total, 1,465 of the 1,580 examined vials of Artoria from NSW and ACT (= 92%) can be attributed to a described species following this study.  A. strepera sp. n.). Some occur further into central NSW (for example A. victoriensis Framenau, Gotch &A. howquaensis Framenau, 2002). Twelve species are currently known only from their type localities or from very small ranges and should be considered shortor narrow range endemic species (Harvey 2002;New and Sands 2002;Ponder and Colgan 2002) (Table 1). These are therefore of elevated conservation concern.
This regional review focussing on a single geographic state and one territory doubles the number of species of Artoria in Australia, demonstrating the high diversity of the genus in this country. At least 15 undescribed species of Artoria are known from Western Australia (V.W. Framenau, unpublished data) and similar ratios of undescribed to described species can be expected from other states. A conservative estimate for the total number of species of Artoria in Australia is 150-180 species, thereby likely representing the largest genus of Australian wolf spiders and a considerable challenge for taxonomic research.
This study documents a considerable morphological diversity of Artoria in NSW and the ACT. Some informal species groups of putatively more closely-related species can be distinguished, for example a 'lineata-group', including A. lineata, A. quadrata and A. ulrichi, in which males have a spoon-shaped tegular apophysis, or a booderee-group (A. booderee sp. n., A. corowa sp. n., A. equipalus sp. n. and A. munmorah sp. n.), having small spiders in which males have a two-lobed tegular apophysis. Other species, such as A. extraordinaria sp. n., have an unusual genital morphology for Artoria, with no known similar species. It is possible future phylogenetic research will consider these groups or species discernible at the genus level. However, as the currently described fauna of Artoria only represents a fraction of the species present in Australia based on artificial geographic boundaries, a more comprehensive taxonomic documentation is required to formulate testable phylogenetic hypotheses at the genus level. This analysis should include representatives from the total distribution range of Artoria and other genera of the Artoriinae, a subfamily that reaches into China and the Pacific Ocean (Framenau 2007;Li et al. 2012;Vink 2002;World Spider Catalog 2018). Artoria howquaensis is the morphologically closest species of this study to the type species of Artoria, A. parvula Thorell, known from China to northern Australia (Framenau 2005;Li et al. 2012).

Taxonomy
Family Lycosidae Sundevall, 1833 Subfamily Artoriinae Framenau, 2007 Genus Artoria Thorell, 1877 Australian Forest Runners Artoriella Roewer, 1960(synonymy established in Framenau, 2002 Lycosula Roewer, 1960(synonymy established in Framenau 2007 Trabaeola Roewer, 1960(synonymy established in Framenau 2002 Type species. Artoria parvula Thorell, 1877, by original designation (Thorell 1877). Diagnosis. In NSW and the ACT, five genera of the subfamily Artoriinae have been found: Artoria, Artoriopsis Framenau, 2007, Diahogna Roewer, 1960, Kangarosa Framenau, 2010and Tetralycosa Roewer, 1960. The row of the anterior eyes in the latter three genera is generally wider than the row of the posterior median eyes and therefore these genera are thought to form a natural group (Framenau 2006(Framenau , 2010Framenau and Hudson 2017). In contrast, species in Artoria ( Fig. 2A-E) and Artoriopsis generally have the anterior eye row narrower than the posterior median eye row. Artoriopsis differs from Artoria by a more elongated carapace that is adorned by distinct light lateral bands, and a distinct colouration of the abdomen with a central diamond-shaped black spot (with the exception of Artoriopsis whitehouseae Framenau, 2007, which has a black abdomen with irregular light central band (Framenau 2007)). There are also genus-specific differences between the male pedipalps of all genera, specifically with respect to the basoembolic and tegular apophyses. For example, the tegular apophyses in Diahogna, Kangarosa and Tetralycosa are generally wider at the base than apically, whereas in Artoria they are narrow at the base and widen apically. The tegular apophysis of Artoriopsis is variable and in general most similar to that of Artoria, but the genera are easily distinguished by the somatic characters above.
Description (after Framenau 2002). Small to medium-sized wolf spiders (total length ca. 2.5-10 mm); males slightly smaller than females; carapace brown to black with darker radial pattern; light median and lateral bands sometimes present; abdomen brown to dark grey, often with mottled pattern and mostly with a light lanceolate heart mark; carapace longer than wide, dorsal profile straight in lateral view; head flanks in frontal view steep in most males but may be a gentle slope in females.
Chelicerae with three (rarely one or two) promarginal and three (rarely one or two) retromarginal teeth; labium as long as or slightly longer than wide; leg formula IV>I>I>III; tegular apophysis located apically at tegulum and of variable shape; basoembolic apophysis broad, heavily sclerotised and bent ventrally; embolus of varying shape (slim to very thick); terminal apophysis functioning as conductor, sometimes forming a groove for the resting embolus; most species with varying number of macrosetae at tip of male cymbium; a scopulate patch of setae occasionally present dorsally on cymbium. Epigyne variable, a simple opening of varying shape or covered by a sclerotised ovoid plate.  A. barringtonensis sp. n. (AM KS122794) straight, distance between AME/AME at least twice of AME/ALE.; C, A. kanangra (KS45008) slightly procurved, evenly spaced; D, A. belfordensis sp. n. (AM KS127759) eyes strongly procurved, evenly spaced.; E, A. extraordinaria sp. n. (AM KS128074) eyes strongly procurved, distance between AME/ALE at least twice AME/AME. Scale bars: 0.1 mm. 7 Basoembolic apophysis (BA) rounded (Fig. 46F)  Distance between AME/ALE at least twice of AME/AME (Fig. 2E)  Palea about as long as wide (Fig. 47H)  17 Anterior spinnerets ventrally pale (Fig. 19B)  39 Distance between AME/ALE at least twice of AME/AME (Fig. 2E)  Lycosa albo-pilata Urquhart, 1893: 123-125.-Rainbow 1911: 1911. Lycosa albopilata, Urquhart.-Roewer 1955Bonnet 1957Bonnet : 2632McKay 1973: 378;McKay 1985: 74. Artoria albopilata (Urquhart) Diagnosis. Artoria albopilata is most similar to A. gloriosa, a species endemic to Lord Howe Island. Males differ in the shape of the tegular apophysis, which ends in three lobes in A. albopilata (Fig. 3E) and two tips in A. gloriosa (Fig. 20E). The tip of the embolus is sharp in A. albopilata, but broad and blunt in A. gloriosa. Females differ in the shape of the epigyne, in particular the anterior border, which is semicircular in A. albopilata, but undulating in A. gloriosa. The median septum fills the atrium in A. albopilata, but is truncated anteriorly in A. gloriosa.
Description. Artoria albopilata has been described in detail (Framenau 2005). A diagnosis and diagnostic images (Figs 3A-H, 46F) are provided here to facilitate identification.
Life history and habitat preferences. Artoria albopilata is a forest species occurring in leaf litter. In NSW and the ACT, it has been found in open to closed dry to wet sclerophyll forests, with one record from a Hoop Pine plantation.
Mature males and females appear in spring in October with the highest number of records in November. There is a much smaller peak around March. A single female with eggsac was found in January. Mature spiders can be found until May. There are also few records of mature spiders from July.
Distribution. In NSW and ACT A. albopilata is particularly found east of the Great Dividing Range (Fig. 4). It has also been found in south-eastern Queensland, south-eastern Victoria, southern South Australia (incl. Kangaroo Island) and Tasmania (Framenau 2005). Diagnosis. The tegular apophysis in male A. alta has a distinctive shape with a terminal part that resembles, in ventral view of the pedipalp, a bicycle seat (Fig. 5E). The female here associated with A. alta has a distinct waved anterior border of the epigyne and two posterior dark lobes (Fig. 5G).

Artoria alta Framenau, 2004
Description. The male of A. alta has been described in detail (Framenau 2004). A diagnosis and diagnostic images (Figs 5A-H, 46L) are provided here to facilitate identification. A putative female of the species (the only female of Artoria currently known from alpine habitats near the type locality), is illustrated here (Fig. 4), although conspecifity remains unclear. The body colouration of the female is much more distinct than that of the male and this female may represent a different species. We decided to include the female here as potential candidate of the female A. alta to have a public documentation of this morphotype.
Life history and habitat preferences. Artoria alta is known only from subalpine or alpine habitats in Kosciuszko National Park in New South Wales. Mature males and females have so far only been found in summer, between end of November and end of December.
Distribution. Known only from Kosciuszko National Park, NSW, in the Australian Alps (AUA) IBRA region (Fig.4). Other material examined. Known only from type specimen.
Etymology. The specific name is an adjective in apposition derived from the type locality, Barrington Tops National Park.
Diagnosis. The tegular apophysis in males of A. barringtonensis sp. n. is distinct amongst the known species of the genus due to the presence of a row of apical teeth, which are visible in lateral view (Fig. 6D).
Anterior eye row. Straight, distance between AME/ AME at least twice of AME/ALE.
Legs. Femora and tibiae of leg I very dark; other legs yellow brown, with no annulations (Fig. 6A).
Female unknown. Life history and habitat preferences. The single male of this species was found at the edge of an upland swamp in summer (January).
Distribution. Artoria barringtonensis sp. n. is known only from its type locality, the Barrington Tops National Park in the south-eastern NSW North Coast (NNC) IBRA region (Fig. 4).    (Framenau, 2004)    Etymology. The specific epithet is a noun in apposition and refers to the type locality, Beaury State Forest.
Life history and habitat preferences. Artoria beaury sp. n. is a forest species most commonly collected in litter from dry sclerophyll and subtropical forests or rainforests. Records also include Bunya and Hoop Pine plantations.
Mature males have only been found in November and December, mature females from November to May, with peaks in December and April. A single female with eggsac was found in March.
Distribution. In NSW, Artoria beaury sp. n. occurs in the north-east, principally in the NSW North Coast (NNC), New England Tablelands (NET) and South Eastern Queensland (SEQ) IBRA regions with a single isolated record also from the South East Coastal (SEC) region (Fig.  8). Artoria beaury sp. n. has also been found in south-eastern Queensland (V.W. Framenau, unpublished data). Etymology. The specific name is an adjective in apposition derived from the type locality, Belford National Park.
Diagnosis. Males of A. belfordensis sp. n. differ from all other Artoria species, except A. taeniifera, by the base of the basoembolic apophysis, which is drawn out into an apical tip (Fig. 47C). It differs from A. taeniifera by the shape of the tegular apophysis, distinctly two-pronged in A. taeniifera, but three-lobed in A. belfordensis sp. n. There is some variation in this character, but also in the shape of the tegular apophysis within this species. Too little material is available to judge if this represents intra-or interspecific variation. We therefore consider this variation capable of occurring within one species. Females are most similar to females of A. taeniifera but differs in the shape of the atrium which is inverted U-shaped in A. taeniifera (Fig. 38G) and obscure in A. belfordensis (Fig. 9G).
Prosoma. Length 2.5, width 1.8; carapace light yellow-brown with indistinct dark radial and black V-shaped pattern between cephalic and thoracic region; covered with dark setae centrally and marginally with a band of lighter setae (Fig. 9A); sternum medium brown, dusted dark grey (Fig. 9B).
Life history and habitat preferences. Artoria belfordensis sp. n. was collected in undisturbed or moderately disturbed forest. Mature males were collected in June, October and November and the single female in October, indicating winter to spring maturity.
Description. Artoria berenice has been described in detail 2005). A diagnosis and diagnostic images (Figs 10C, D, G, H, 11A-H, 46H) are provided here to facilitate identification, especially in differentiation from similar newly-described species.
Life history and habitat preferences. In NSW males and females appear from late August with highest numbers throughout October to January. Males are rarely found in March but then appear again in April and May. Females occur throughout summer and autumn until May, with two activity peaks in November/December and April and can sometimes be found into the winter

months. A female with eggsac was found in November and one with spiderlings in December.
Artoria berenice is a forest species. Habitat descriptions found on collection labels include 'wet sclerophyll', 'semi-cleared woodland near Pinus plantation', 'grassy forest', 'Eucalyptus botryoides-Allocasuarina torulosa, open forest', 'Angophora costata woodland ridge top, open forest', but also 'gravel pit', and 'cave entrance, cave in doline'.
Distribution. In NSW, A. berenice is particularly found east of the Great Dividing Range, i.e. in the coastal South Eastern Queensland (SEQ), NSW North Coastal (NNC), Sydney Basin (SYB), and South East Corner (SEC) IBRA bioregions. It has been found west of the Great Dividing Range, with one single isolated record from central NSW (Fig.12). Artoria berenice has also been found in Queensland, eastern Victoria and throughout Tasmania and occurs in New Caledonia and Vanuatu (Framenau 2005). Banksia (AM KS128073). Paratypes: 3 males, same as holotype (AM KS116188).
Other material examined. Known only from type material.
Etymology. The specific name is a noun in apposition referring to the type locality, Bondi State Forest.
Diagnosis. Similar to other species in the lineata-group, the tegular apophysis of the male pedipalp is spoon-shaped (Fig. 6G) in A. bondi sp. n., but is much smaller than in A. lineata, A. quadrata or A. ulrichi. In that, A. bondi sp. n. males resemble A. strepera sp. n. males in which the embolus and terminal apophysis protrude much further behind the tegulum than in A. bondi sp. n.
Female unknown. Life history and habitat preferences. The type material of Artoria bondi sp. n. has been found in Banksia woodland in November.

Artoria booderee
Etymology. The specific name is a noun in apposition referring to the type locality, Booderee National Park.
Diagnosis. Males of A. booderee sp. n. share with A. corowa sp. n., A. munmorah sp. n. and A. equipalus sp. n. a distinctly bi-lobed tegular apophysis. They differ by having a more elongated cymbium with ca. 2-3 distinct macrosetae on its tip, with 4-5 present in A. equipalus sp. n. and absent in both other species of this group. The epigyne of female A. booderee sp. n. is incised posteriorly and most similar to that of A. corowa sp. n., but the anterior edge of the epigyne in A. booderee sp. n. is distinctly semi-circular, not so in A. corowa sp. n.
Life history and habitat preferences. Artoria booderee sp. n. appears to be a coastal species, as it has so far only been found on the northern and southern headlands around Jervis Bay and near Sydney. No habitat data were included on the locality labels.
All mature males at Jervis Bay were collected in August. The one from Sydney specimen was collected in December. Mature females were found in August, December and February suggesting that A. booderee sp. n. is a not tightly synchronised with any specific season.
Distribution. Known only from around Jervis Bay and the Mt Kembla Basin in the Sydney Basin (SYB) IBRA region (Fig. 12). Other material examined. Known only from type material.
Etymology. The specific name is a noun in apposition referring to the type locality, Comleroi Road.
Diagnosis. Males of A. comleroi sp. n. are most similar to those of A. slatyeri sp. n. based on the elongated spoonshaped tegular apophysis. They differ by an apical small lobe being present in A. comleroi sp. n. Females of A. comleroi sp. n. display an epigyne shape with the atrium forming two distinct ovoid depressions in ventral view.
Epigyne about as long as wide, strongly sclerotized at posterior rim, atrium formed by two distinct ovoid depressions (Fig. 16G); spermathecal heads globular about their diameter apart, spermathecal stalks attached medially and centrally bent (Fig16H).
Life history and habitat preferences. No habitat data were found with the type specimens. The male and female types were collected in June suggesting Artoria comleroi sp. n. to be winter mature.

Artoria corowa
Etymology. The specific name is a noun in apposition referring to the type locality, NW of Corowa.
Diagnosis. Males of A. corowa sp. n. share with A. booderee sp. n., A. munmorah sp. n. and A. equipalus sp. n. a distinctly bi-lobed tegular apophysis. They differ from these by the tegular apophysis not protruding past the cymbium edge in ventral view and the presence of a disto-ventral cluster of macrosetae. The epigyne of female A. corowa sp. n. is incised posteriorly and therefore most similar to that of A. booderee sp. n., but lacking the distinct semi-circular anterior edge of the epigyne of A. booderee sp. n.
Epigyne: slightly wider than long, strongly sclerotized lateral posterior tips, atrium shaped like an in-verted love-heart with posterior incision (Fig. 14G); spermathecal heads globular and around their diameter apart, spermathecal stalks attached laterally and strongly S-shaped (Fig. 14H).
Life history and habitat preferences. There was little but varied habitat data available with the specimens, including 'roadside', 'Eucalyptus largiflorens patch', 'Eucalyptus camaldulensis patch', Casuarina cristata patch' and 'native grassland', are insufficient to form a clear account of the habitat preferences of A. corowa sp. n. remain unclear. All specimens were collected between November and December, suggesting the species is largely summer mature. The single coastal record of the species, a mature male and female, is from May (autumn). Other material examined. Known only from type material.
Etymology. The specific epithet is a compound noun in apposition derived from the Latin equus (= horse) and palus (= swamp) and refers to the type locality, near Horse Swamp Track.

Diagnosis.
Males of A. equipalus sp. n. share with A. booderee sp. n., A. corowa sp. n. and A. munmorah sp. n. a distinctly bi-lobed tegular apophysis, but differ from these by the basal lobe of the tegular apophysis being more sclerotized and oriented more 'horizontally' in ventral view. The cymbium tip in A. equipalus has 4-5 macrosetae, which are absent in A. corowa sp. n. and A. munmorah sp. n. Artoria booderee sp. n. has 2-3 macrosetae on the cymbium tip which is more elongate than that of A. equipalus sp. n. The epigyne of female A. equipalus sp. n. is most similar to that of A. mungo sp. n., but the shape of the spermathecal heads is very different in both species with in particular with respect to the insertion of the spermathecal stalks (postero-medial in A. equipalus sp. n., lateral in A. mungo sp. n.).
Legs. Tibiae, metatarsi and tarsi of leg I very dark to almost black; other legs light brown, annulated (Fig. 17A).
Life history and habitat preferences. The habitat description with the type material is 'upland swamp'. Spi- ders were found in January suggesting Artoria equipalus sp. n. to be summer mature.
Distribution. Artoria equipalus sp. n. is known only from its type locality in Barrington Tops National Park in the NSW North Coast (NNC) IBRA region (Fig. 15). Etymology. The specific epithet is an adjective in apposition derived from Latin (extraordinaria -unusual) and refers to the unusual shape of the basoembolic apophysis of the male pedipalp.

Artoria extraordinaria sp. n.
Diagnosis. The male pedipalp of A. extraordinaria sp. n. is highly unusual within the genus owing to the shape of the basoembolic apophysis, which forms an almost 360-degree circle (Fig. 47I). The female epigyne is poorly defined with two almost transparent lateral lobes (Fig. 18G).
Total length 3.6. Prosoma. Length 2.1, width 1.4; carapace light reddish-brown with dark radial pattern and black V-shaped pattern between cephalic and thoracic region; indistinct and irregular broad lighter central and marginal band (Fig. 18A); sternum pale, with darker margin (Fig. 18B).
Anterior eye row. Strongly procurved, distance between AME/ALE at least twice AME/AME.
Anterior eye row. Strongly procurved, distance between AME/ALE at least twice AME/AME.
Life history and habitat preferences. Based on locality data, Artoria extraordinaria sp. n. is a forest species. Detailed habitat data were not available on locality labels of the specimens. Most specimens were collected between February and April, with one male recorded from June, suggesting Artoria extraordinaria sp. n. to be autumn to winter mature.
Distribution. This species was found some distance from the coast at scattered localities in the NSW North Coast (NNC), Sydney Basin (SYB) and South Easter Corner (SEC) IBRA bioregions (Fig. 15).

Figs 1C, 15, 19 A-H, 46D
Yellow-handed Forest Runner Artoria flavimanus Simon, 1909: 192, fig. 9 (2002)).  . The gaping shape of the tegular apophysis of the male pedipalp (Fig. 46D) is distinctive. Males can also be identified be the distinctive leg formula, with leg I longer than leg IV (I>IV>II>III). The shape of the epigyne of A. flavimana has a semi-circular to pentagonal atrium which includes an isolated ovoid median septum (Fig. 19G).

Material examined. Holotype female of
Description. Artoria flavimana has been described in detail  Life history and habitat preferences. Artoria flavimana occurs predominantly in eucalypt forests. In NSW and ACT, most males were recorded in May and August and females also into November. Reproductive activity appears to occur mostly in winter.
Distribution. Artoria flavimana occurs in south-eastern NSW and the ACT in the South East Corner (SEC), South Eastern Highlands (SEH) and Australian Alps (AUA) IBRA regions (Fig. 15). The species is also known from Victoria, Tasmania, south-eastern South Australia and south-western Western Australia (Framenau , 2005. Diagnosis. Artoria gloriosa is similar to A. albopilata; however, the tegular apophysis of the male pedipalp of A. gloriosa lacks the apical lower tip present in A. albopilata. The tip of the embolus of A. gloriosa is blunt (Fig. 46E), whereas it is sharp and sickle-shaped in A. albopilata (Fig.  46F). The median septum of the female epigyne of A. gloriosa does not fill out the whole atrium as in A. albopilata (Fig. 3G) but is truncated anteriorly (Fig. 20G).
Life history and habitat preferences. Artoria gloriosa is a forest species dwelling in litter in a variety of forest types on Lord Howe Island. Mature males and females have largely been found between November and February, with two records of males from May and a fe-male with spiderlings in July. Two females with spiderlings were found in December.
Life history and habitat preferences. Habitat descriptions on collection labels with Artoria grahammilledgei sp. n. suggest the species to be a woodland and forest dweller in litter, i.e. 'dry sclerophyll forest', 'Angophora costata woodland ridge top', 'Eucalyptus botryoides-Allocasuarina torulosa open forest' and 'litter & soil beneath Casuarina'.
Mature male Artoria grahammilledgei sp. n. have been found almost throughout the year, except November, February and March with two distinct activity periods in April and July. Mature females have been found in all months with a first peak from April to August, and a second in November. A female with eggsac was found in October/ November and females with spiderlings in January, April, November and December. Therefore, the species appears largely autumn to winter mature, but with wide seasonal variation in relation to its reproductive behaviour.
Distribution. In NSW, Artoria grahammilledgei sp. n. appears common east of the Great Dividing Range but has also been found along its west (Fig. 23). The species is also known from Queensland (V.W. Framenau, unpublished data). Other material examined. 175 males, 100 females (2 with spiderlings) and 1 juvenile 134 records. AUSTRALIA: New South Wales: 2 males, 4 km NE of Mt Wog Wog, 17 km SE Bombala, 37°04'30"S, 149°28'00"E (AM KS98791, KS98810); 1 male, Bar-Etymology. The specific epithet is a matronym honouring Dr Helen Smith of the Australian Museum, Sydney, for her ongoing support of our spider research.
Diagnosis. Artoria helensmithae sp. n. males and females are very similar to those of A. beaury sp. n. (see diagnosis above).
Prosoma. Length 2.2, width 1.5; carapace light yellow-brown with dark radial pattern and black V-shaped pattern between cephalic and thoracic region; indistinct and irregular lighter central and marginal band (Fig.  24A); sternum light, margin dusted dark grey (Fig. 24B).
Legs. Femora, tibiae and metatarsi of leg I very dark to almost black; other legs brown, with darker annulations; tarsi and metatarsi lighter brown (Fig. 24A).
Life history and habitat preferences. Habitat descriptions found on the collection labels of A. helensmithae sp. n. include 'under tall shrubs', 'sand dunes', open forest, litter', 'rainforest' and 'dry forest'. A single record is from the dark zone of a cave. Therefore, A. helensmithae sp. n. seems to show some variation in its habitat preferences. Most records also state 'litter'.
Artoria helensmithae sp. n. appears to have three peaks of reproductive activity, the first around April/May, then in July and again from October to December. Females with spiderlings were found in December. This is an unusual phenology for wolf spiders, i.e. this species appears to be reproductively active in autumn, winter and summer.
Distribution. In NSW, A. helensmithae sp. n. has generally been found east of the Great Dividing Range, with records from the NSW North Coast (NNC), Sydney Basin (SYB), South East Corner (SEC) and South East Highlands (SHE) IBRA regions (Fig. 23). The species has also been found in eastern and central Victoria (V.W. Framenau, unpublished data). Diagnosis. Amongst Australian Artoria, A. howquaensis is most similar to A. parvula, a tropical species not yet found in NSW or the ACT. This species most easily diagnosed by its body colouration, being very dark brown to black with distinct white lateral bands on the carapace (Fig. 25A, B), this colour pattern being more distinct in live specimens. The tegular apophysis is similar to those of the booderee-group (i.e. two-lobed), but in contrast to those species, the basal lobe is pointy (not rounded) and directed ventrally (Fig. 25E). The epigyne is poorly sclerotised with the Y-shaped internal genitalia visible through the cuticle (Fig. 25G, H).

Artoria howquaensis Framenau, 2002
Description. Artoria howquaensis has been described in detail Framenau et al. 2006). A diagnosis and diagnostic images (Figs 25A-H, 46J) are provided here to facilitate identification, in particular from similar newly-described species.
Life history and habitat preferences. Artoria howquaensis is a riparian species which has previously been found in open habitats along the margins of rivers and springs. The records in NSW are consistent with this habitat preference as the species was found in open, grassy and swampy areas.
Mature males and females were found in December, with further records of males in March.
Distribution. In NSW, A. howquaensis has been found in scattered records in the northern half of the state east and west of the Great Dividing Range in the Nandewar (NAN), NSW North Coast (NNC) and Sydney Basin (SYB) bioregion (Fig. 23). The species is likely more widespread in the state, but riparian habitats are rarely sampled. The species is known to occur throughout Victoria and South Australia 2005;Framenau et al. 2006). Other material examined. AUSTRALIA: New South Wales: 1 male, Barren Grounds Nature Reserve, 14 km NW Jamberoo, Illawarra Escarpment, 34°40'28"S, 150°42'45"E (AM KS63088).

Artoria kanangra
Etymology. The specific epithet is a noun in apposition referring to the type locality, Kanangra-Boyd National Park.
Diagnosis. Males of A. kanangra sp. n. are most similar to those of the booderee-group (A. booderee sp. n., A. corowa sp. n., A. munmorah sp. n. and A. equipalus sp. n.), but differ in the shape of the tegular apophysis, which is birdhead-shaped in A. kanangra sp. n. (Fig. 26C).
Legs. Femora and tibiae of leg I very dark; other legs brown, femora and tibia with darker annulations, particularly ventrally; tarsi and metatarsi lighter yellow-brown (Fig. 26A).
Female unknown.
Life history and habitat preferences. The holotype was found in creek sphagnum at 1,200 m altitude suggesting the species to be riparian. Adult males were found in August and December.
Distribution. Artoria kanangra sp. n. is currently known only from two localities in eastern-central NSW, from the Sydney Basin (SYB) and South Eastern Highlands (SHE) IBRA bioregions (Fig. 27). Other material examined. Known only from type material.
Etymology. The specific epithet is a noun in apposition referring to the type locality, Kerewong State Forest.
Diagnosis. The pedipalp morphology of males of A. kerewong sp. n. is distinct within the genus, with the terminal section of the tegular apophysis having both a pointed apical and a pointed basal tip. The basoembolic apophysis is the longest amongst the known Artoria species. The female of A. kerewong sp. n. is currently unknown. Description.
Pedipalps. Tibia as long as broad; cymbium tip with distal cluster of macrosetae (Fig. 26G, H); dorsal scopula patch present but sparse; tegular apophysis with narrow stalk and bent triangular tip, tip reaching margin of cymbium, prolaterally strongly pointed (Fig. 26G); palea about twice as long as wide, basoembolic apophysis twice as long as broad, bent posteriorly and broadly rounded; embolus extremely broad, widely semicircular; terminal apophysis not visible (Fig. 48A).
Female unknown. Life history and habitat preferences. No habitat data were on the collection label of the holotype, which was found in November/December suggesting it is summer mature.
Distribution. Artoria kerewong sp. n. is known only from the type locality, the Kerewong State Forest in the NSW North Coast (NNC) IBRA region (Fig. 27). Diagnosis (after Framenau 2002). Artoria lineata forms part of the lineata-group, that includes those species in which the male pedipalp has a spoon-shaped tegular apophysis and the median septum of the female epigyne is often inverted T-shaped. In NSW and the ACT this group also includes A. quadrata and A. ulrichi. Identification of the species in this group is notoriously difficult and, in males, is best achieved by dissecting the pedipalp for an examination of the palea. Males of both A. lineata and A. ulrichi have an oval tegular apophysis (Fig. 46A, B), which is more rectangular or apically truncated in A. quadrata (Fig. 46C). Males of A. lineata and A. ulrichi can be distinguished by the position of the embolus and terminal apophysis. The tip of both is visible apically of the tegulum in A. lineata but totally concealed by the tegulum in A. ulrichi. In the dissected palea, the basal edge of the embolus is straight in A. lineata (Fig. 46A), but distinctly curved in both A. ulrichi (Fig. 46B) and A. quadrata (Fig. 46C).
The female epigyne in both A. lineata and A. ulrichi has an inverted, T-shaped median septum, however, the posterior border is light or transparent along its whole length in A. lineata, whereas the lateral tips of the medium septum are dark all around in A. ulrichi. Description. Artoria lineata has been described in detail  Life history and habitat preferences. Artoria lineata is a forest species and has been found in the leaf litter of dry and wet sclerophyll forests and rainforest. Mature males and females can be found all year round with peaks of activity in August and October. Females can also be found in high numbers through summer and into autumn.
Distribution. Artoria lineata appears to be widespread in all IBRA regions immediately east and west of the Great Dividing Range (Fig. 27). The species otherwise occurs in Queensland, throughout Victoria, south-eastern South Australia and Tasmania 2005) Artoria maroota sp. n. Etymology. The specific name is a noun in apposition referring to the type locality, Maroota State Forest.

Diagnosis.
Males of Artoria maroota sp. n. are able to be distinguished from all other species of the genus by the black setal brushes on the tibia of leg I and the inverted L-shaped tegular apophysis (Fig. 29C).
Female unknown. Life history and habitat preferences. No detailed habitat data were available from the locality labels. As the species was mainly collected during a forest survey it appears to be a forest dweller.
Mature males were collected from October to December suggesting it is spring/summer mature.
Distribution. This species was found in the Sydney Basin (SYB) IBRA region in eastern central NSW (Fig. 30). Diagnosis. Artoria mckayi is most similar to A. albopedipalpis , a species which has only been recorded from Victoria . The ventrally-pointing tegular apophysis of the male pedipalp ( Fig. 31E) is somewhat similar to that of A. berenice (Fig. 11E) but is much slimmer. The female epigyne (Fig. 31G) may resembles that of A. berenice (Fig. 11G) but is overall much rounder and less constricted posteriorly.

Artoria mckayi Framenau, 2002
Description. Artoria mckayi has been described in detail . A diagnosis and life and diagnostic images (Figs 1A-B, 31A-H, 46G) are provided here to facilitate identification.
Life history and habitat preferences. Artoria mckayi is a riparian habitat specialist and can mainly be found along the edges of rivers and creeks on both muddy and rocky banks. In NSW and the ACT mature spiders have been found between October and May. In Victoria, the adult spiders (species  can be found throughout the year with lower numbers mainly in spring (March).
Distribution. Artoria mckayi occurs at rivers and creeks of the Great Dividing Range (Fig. 30). The species otherwise occurs in Queensland, Victoria, south-eastern South Australia and Tasmania 2005). Etymology. The specific name is a noun in apposition referring to the type locality.
Diagnosis. Based on the shape of the tegular apophysis, males of A. mungo sp. n. are most similar to A. helensmithae sp. n. and A. beaury sp. n. Artoria beaury sp. n. differs distinctly in the shape of the basoembolic apophysis which is much broader than that of A. mungo sp. n. In A. helensmithae sp. n. the apical edge of the tegular apophysis is much more indented than in A. mungo sp. n. Female A. mungo sp. n. are most similar to A. wilkiei sp. n. based on the shape of the epigyne in ventral view, but the spermathecal heads of the latter are much larger and touching medially.
Life history and habitat preferences. It appears that this species has affinities to at least intermittently flooded areas, as it has been found near swamps and creeks in dry sclerophyll bushland and scrubland. Mature spiders have been found in August, September and January suggesting it is spring-to summer-mature.
Legs. Femora and tibiae of leg I, II dark; other legs yellow-brown, with slightly darker annulations (Fig. 33A).
Life history and habitat preferences. There is only a single record with habitat information, 'canopy of trees with grass under'; therefore, habitat preferences of A. munmorah sp. n. remain unclear.
Mature spiders have been found between October and January, therefore this species appears to be spring-to summer-mature.
Distribution. Artoria munmorah sp. n. is currently known from eastern central NSW along both sides of the Great Dividing Range and occurs in the NSW North Coast (NNC), Sydney Basin (SYB), South Eastern Highlands (SHE), NSW South Western Slope (NSS) and Brigalow Belt South (BBS) IBRA regions (Fig. 34).  (Fig. 29G). It appears most similar to A. belfordensis sp. n. (Fig. 9E), but it does not have the drawn out and pointed bases of the basoembolic apophysis of that species.
Pedipalps. Pedipalp globular, tibia as long as broad; cymbium tip with cluster of disto-ventral macrosetae (Fig. 29G, H); dorsal scopula patch sparsely present, short; tegular apophysis distally wide, deeply indented, basally narrowed to 1/3, retrolateral part widely scooped with two tips not reaching margin of cymbium (Fig. 29G); basoembolic apophysis about as long as broad, broadly rounded; not visible and not dissected as the holotype has only one pedipalp (Fig. 29G). Female unknown. Life history and habitat preferences. Habitat preferences of A. myallensis sp. n. are not known. The holotype was found at the beginning of spring (September).
Distribution. Artoria myallensis sp. n. is known only from the type locality, the coastal Myall Lakes in the southern NSW North Coast (NNC) IBRA region (Fig. 34).
Remarks. The transfer of Lycosa pruinosa to Artoria was based on the morphology of the male pedipalp as illustrated in the original description (L. Koch 1877; fig. 2a), which displays a clear artoriine structure (Framenau 2005).
However, the body colouration of the holotype illustrated (L. Koch 1877; fig. 2) with a light median band on a uniformly black abdomen does not match the genus description. As part of our comprehensive revision of NSW Artoria we could not identify this species, originally described from Sydney. The most similar species is Artoriopsis whitehouseae Framenau, 2007; however, the type locality, Sydney, for A. pruinosa falls far outside the range of that species, which so far has only been found west of the Great Dividing Range (Framenau, 2007;unpublished data). Considering the uncertainty identifying A. pruinosa due to the lack of the type specimen, we here consider the species-group name Lycosa pruinosa L. Koch, 1877 a nomen dubium. Distribution. Artoria quadrata is common east and west of the Great Dividing Range, from coastal areas into the Brigalow Belt South (BBS) and NSW South Western Slopes (NSS) IBRA regions (Fig. 34). The species otherwise occurs in Queensland and Victoria 2005 Diagnosis. The shape of the tegular apophysis in males of A. slatyeri sp. n. is distinctive within Artoria (Fig. 36C), most closely resembling that of A. grahammilledgei sp. n. (Fig. 22E); however, it is much more elongated than the tegular apophysis of that species. The basoembolic apophysis is much narrower and longer in A. slatyeri sp. n. than in A. grahammilledgei sp. n.
Female unknown. Life history and habitat preferences. Artoria slatyeri sp. n. is apparently a forest species, the type material being found in a forest reserve. Mature males were found in pitfall traps in December and February suggesting the species is summer-mature.
Distribution. Artoria slatyeri sp. n. is currently known only from the Bondi State Forest and Wadbilliga National Park in southern NSW in the South Eastern Highlands (SEH) IBRA region (Fig. 37). Other material examined. Known only from type material.
Etymology. The specific epithet refers to the scientific genus-group name of the Australian Currawong birds, Strepera Lesson, 1831. Currawong is also the name of the type locality.
Diagnosis. Males of A. strepera sp. n. most closely resemble A. bondi sp. n. as both have a small, spoon-shaped tegular apophysis; however, the embolus and terminal apophysis in A. strepera sp. n. are much longer and protruding distinctly apically from the tegulum (Fig. 48F), in contrast to A. bondi sp. n. in which these are inconspicuous (Fig. 47D).
Female unknown. Life history and habitat preferences. The habitat preferences of A. strepera sp. n. are unknown. The male types were found in spring (October).
Distribution. Artoria strepera sp. n. is currently known only from the type locality, Currawong, north of Sydney in the Sydney Basin (SYB) IBRA region (Fig. 37).
Life history and habitat preferences. The habitat of A. taeniifera in NSW is unknown. In Western Australia, from where most records of this species have been reported, A. taeniifera prefers open habitats such as coastal dunes, gardens and open woodlands. The single female from NSW was found in March suggesting the species to be autumn-mature. In south-western Western Australia, mature spiders have been found in winter and early spring.
Distribution. Artoria taeniifera has only been found once in NSW, centrally in the Cobar Plain (COP) IBRA region (Fig. 37). There is a second isolated record of the species in South Australia (Framenau, unpublished data). It is unknown if these represent persistent populations.
The species occurs commonly in south-western Western Australia, from where it was initially described 2005;unpublished data).   with isolated records around Sydney (Sydney Basin -SYB) and in the south-east of the state (South East Corner -SEC) (Fig. 37). This species has also been found in south-eastern Queensland (V.W. Framenau, unpublished data). Description. Artoria ulrichi has been described in detail . A diagnosis and diagnostic images (Figs 42A-H, 46B) are provided here to facilitate identification.

Artoria triangularis Framenau, 2002
Life history and habitat preferences. In NSW, A. ulrichi has been found in forest habitats, including rainforests. Mature spiders were found from November through to June.
Distribution. In NSW, A. ulrichi has been found east and west of the Great Dividing Range in the NSW North Coast (NNC), Sydney Basin (SYB) and South Eastern Highlands (SHE) IBRA regions (Fig. 41). There is also a single record from Victoria . Diagnosis (after Framenau et al. 2006). Males of A. victoriensis can be distinguished from all other Australian Artoria by the shape of the median apophysis which resembles an upside-down sock in ventral view. The female epigyne is distinctively oval, with a white center and a sclerotized posterior rim reaching medially into this center.

Artoria victoriensis Framenau, Gotch & Austin, 2006
The specimens illustrated here vary somewhat from those originally described, in particular with respect to the internal female genitalia. It is possible, that A. victoriensis may include more than one species pending a more detailed evaluation of its intraspecific variation across its wider range.
Description. Artoria victoriensis has been described in detail . A diagnosis and diagnostic images (Figs 43A-H, 46I) are provided here to facilitate identification.
Life history and habitat preferences. Artoria victoriensis can typically be found in open, moderately moist habitats, including suburban garden and parks.
Males of the species were predominantly found from October to December, with a single record from April. Female activity is similar, although mature females are not uncommon in the later summer months.
Etymology. The specific epithet is a patronym in honour of Mr Lance Wilkie, who collected the holotype.
Diagnosis. The pedipalp morphology in males of A. wilkiei sp. n. is most similar to that of A. barringtonensis sp. n. and A. bondi sp. n. based on the small, spoon-shaped tegular apophysis. However, A. wilkiei sp. n. is the only species amongst these in which the tegular apophysis has a small tooth pointing ventrally, visible in retrolateral view (Fig. 45F). The female epigyne of A. wilkiei sp. n. is similar to that of A. mungo sp. n., but the spermathecal heads are much larger and touch medially.
Description. Male (based on holotype AM KS128072). Total length 4.3. Prosoma. Length 2.3, width 1.7; carapace yellow-brown dusted with grey and indistinct light radial pattern; lateral margin and central band pale yellow, broader in cephalic area (Fig. 45A); sternum pale, with darker margin (Fig. 45B).  This paper was completed with support from the Australian Government's Australian Biological Resources Study (ABRS) Bush Blitz Strategic Taxonomy Grants Scheme Grant Nr: TTC216-05 to study the new wolf spiders of Artoria (Lycosidae) from ACT and NSW: integrating Bush Blitz specimens into taxonomic research on an abundant predator.
We are thankful to Robert Whyte and Robert Raven for constructive suggestions that greatly improved the quality of the manuscript.
Volker Framenau acknowledges the continuing support by Mark Harvey and Julianne Waldock (WAM) for his taxonomic studies on Australian arachnids. The foundations of this study were laid by a research grant (2002)(2003)(2004)(2005) on Australian wolf spiders funded by the Australian Biological Resources Study (ABRS) to Mark Harvey (Western Australian Museum) and Andy Austin (The University of Adelaide) (grant number 202-041). VWF also appreciates the unwavering support by his partner