Novakiella trituberculosa (Roewer, 1942)

Framenau, Volker W., Vink, Cor J., Scharff, Nikolaj, Baptista, Renner L. C. & Castanheira, Pedro de S., 2021, Review of the Australian and New Zealand orb-weaving spider genus Novakiella (Araneae, Araneidae), Zoosystematics and Evolution 97 (2), pp. 393-405 : 393

Novakiella trituberculosa (Roewer, 1942)

Figs 1 , 2 , 3 , 7 , 8

Epeira tri-tuberculata Urquhart 1887: 78-79, pl. 7, fig. 2; pl. 8, fig. 1 (preoccupied by Epeira trituberculata Lucas, 1846; currently listed as junior synonym of Cyclosa insulana (Costa, 1834).

Epeira tri-tuberculata Urquhart 1888: 120-121, pl. 11, figs 7, 8.

Aranea trituberculosa Roewer 1942: 834 (replacement name).

Novakia trituberculata Court and Forster 1988: 119-124, figs 359, 553-562.

Novakiella trituberculosa Court and Forster, in Platnick 1993: 457.

Type material.

Syntypes of Epeira tri-tuberculata Urquhart, 1887, 2 females, Karaka, New Zealand (37°06'S, 174°53'E), A.T. Urquhart (CMNZ 2005.135.112). Examined.

Other material examined.

AUSTRALIA: Australian Capital Territory: 1 female, 1 juvenile, Canberra , 35°18'S, 149°08'E (SAM) ; 1 female, Kaleen, Canberra , 35°17'S, 149°13'E, 25.iv.1990 ( SAM 31192 ) ; 1 male, 1 female, Kaleen, Canberra , 35°17'S, 149°13'E, 21.v.1988 (SAM) ; 1 male, Red Hill, 14 Pera Place , 35°20'S, 149°08'E, 24.ii.1982, M. S. Harvey leg. (WAM T73527 ) ; 1 male, same locality, 01.iv.1983, M. S. Harvey leg. (WAM T73571 ) ; New South Wales: 1 male, Beecroft , 33°45'S, 151°04'E, 25.i.1999, J. Noble leg. (AM KS58630) ; 1 female, Botany , 33°57'S, 151°12'E, 04.vii.1961, R. Mascord leg. (AM KS32651) ; 1 female, same locality, 26.vii.1966, R. Mascord leg. (AM KS32652) ; 1 female, Coonabarabran, ' Smokey Hole' , 31°16'S, 149°17'E, 04.x.1978, E. Edmondson leg. (AM KS7545) ; 1 male, Dulwich Hill , 33°54'S, 151°08'E, 17.ii.1977, H. Ehmann leg. (AM KS0741) ; 1 male, Epping Strip , 33°46'S, 151°05'E, 10.i.1996, J. Noble leg. (AM KS49920) ; 1 male, Hillsdale, Sydney , 33°57'S, 151°13'E, 03.ii.1972, R. Mascord leg. (AM KS34148) ; 1 male, Khancobin , 26.iv.1990 (SAM) ; 1 female, Lord Howe Island, Erskine Valley Transect, 31°34'58"S, 159°04'45"E, pitfall traps, 01.xi.1978, T. Kingston leg. (AM KS88175) ; 1 male, Lord Howe Island, Goat House Cave , 31°33'54"S, 159°05'18"E, 10.ii.1971, M. Gray leg. (AM KS20999) ; 1 female, Mount Colah , 33°40'S, 151°07'E, M. Gray leg. (AM KS48820) ; 1 male, Park Beach, Coffs Harbour , 30°18'S, 153°07'E, 23-24.v.1986 (SAM) , 1 male, Punchbowl , 33°56'S, 151°03'E, 02.xii.1940, Ms Levitt leg. (AM KS33550) ; 1 male, The Rock Nature Reserve , 30km SW Wagga Wagga, 35°16'S, 147°05'E, sweeping/beating, 13.xii.2000, C. A. Car leg. (AM KS93847) ; Queensland: 1 male, Bardon, Brisbane , 27°27'S, 152°58'E (QM) ; 1 female, Burleigh Heads , 28°06'S, 153°26'E (QM) ; 1 male, 1 juvenile, Endfield Station , 27°55'S, 149°43'E (QM) ; 1 female, Eurimbula , 24°11'S, 151°50'E, C. Horseman leg. (AM KS12771) ; 1 female, Gatton, Queensland Agricultural College , 27°34'S, 152°20'E, S. Pearce leg. (QM S66755 ) ; 1 male Hurdle Gully , 13km WSW Monto, 24°54'00"S, 150°59'55"E, 23.ix-20.ix.1997, G. Monteith leg. (QM) ; 1 male, Jerons St Park , 27.xi.2009, R. Whyte leg. (WAM S84676 ) ; 1 male, Jevons , 26.xii.2009, R. Whyte leg. (QM S84670 ) ; 1 female, same locality, 01.i.2010, R. Whyte leg. (QM S84672 ) ; 1 male, Masthead Island, Great Barrier Reef , 23°32'S, 151°44'E, C. Hedley leg. (AM KS32650) ; 1 male, Mt Gavial , 1km S, 23°36'S, 150°29'E, 17.xii.1998, D. J. Cook leg. (QM S69354 ) ; 1 male, Oakey , 27°27'S, 151°42'E (QM) ; 1 male, Teewah Creek, Cooloola , 26°02'S, 153°03'E (QM) ; 1 female, Walton Bridge Reserve , 20.xi.2009, R. Whyte leg. (QM S84666 ) ; South Australia: 1 male, Baird Bay , 33°09'S, 134°22'E, 12.i.1995, J. M. Waldock & P. Payne leg. (WAM T73553 ) ; 1 male, 1 female, Beautiful Valley Caravan Park, adjacent, near Wilmington , 32°39'S, 138°06'E, 14.iv.1993 (SAM) ; 1 male, Belalie Creek, Jamestown , 33°12'S, 138°36'E, 10.iv.1993, D. Hirst leg. (SAM) ; 2 males, Cape Gantheaume , 1km N Point Tinline, Kangaroo Island, 35°59'S, 137°37'E, 10.xi.1987, D. Hirst leg. (SAM) ; 1 male, Caracoorte Cave Reserve , 37°05'S, 140°47'E, 25.iv.1979, D. Lee leg. (SAM) ; 1 female, Caroline Forest, ' Snowgum Reserve' , 37°56'S, 140°56'E, 20.iv.1979, G. Grass leg. (SAM) ; 1 male, Coromandel Valley, Mt Lofty Ranges , 32°05'S, 138°38'E, 02.iii.1996, L. N. Nicolson leg. (SAM) ; 1 female, Dog Lake Road, SE Langhorne Creek, 35°17'S, 139°02'E, J. Eckert leg. (SAM) ; 1 female, Eucla , 77km E, 31°28'S, 129°37'E, 23.ii.1978, B. Y. Main leg. (WAM T87424 ) ; 1 female, Forestville , 34°57'S, 138°35'E (SAM) ; 1 male, Gluepot Reserve , 11.3km W Gluepost Homestead, 33°45'16"S, 139°59'58"E, 26.xi.-06.xii.2000 (SAM NN19454) ; 1 female, Kangaroo Island, 35°45'S, 137°37'E (SAM) ; 1 male, Kangaroo Island, Western River Wilderness Protection Area, Waterfall Creek near waterfall, 35°41'44"S, 136°54'37"E, beating, 09-10.v.2010, M. G. Rix & D. Harms leg. (WAM T102789) ; 1 male, Lake Gilles Conservation Park , 33°05'S, 136°39'E, 21.xi.1995 (SAM) ; 1 female, Langhorne Creek , 35°18'S, 139°02'E, C. Wilson leg. (SAM) ; 1 female, 1 juvenile, Marino Rocks to Halletts Cove , near railway line, 35°04'S, 138°30'E, 05.ix.1967, H. M. Cooper leg. (SAM) ; 2 females, Melrose , camping area, 32°49'S, 138°11'E, 14.iv.1987, D. Hirst leg. (SAM) ; 1 male, Mitcham , 34°59'S, 138°37'E, 21.i.1979, R. V. Southcott leg. (SAM) ; 2 females, Muston , Kangaroo Island, 35°49'S, 137°44'E, 03.vii.1967, H. M. Cooper leg. (SAM) ; 1 female, Nappyalla , 35°20'S, 139°07'E, J. Eckert leg. (SAM) ; 1 female, Port Wakefield , 34°11'S, 138°09'E, 14.iii.2004, B. S. Pavey leg. (SAM) ; 1 female, Port Wakefield , T-junction W, 34°11'S, 138°09'E, B. Pavey leg. (SAM) ; 1 male, Pyap , 34°27'S, 140°29'E, 02-09.vi.1990, L. N. Nicolson leg. (SAM) ; 1 female, Sellicks-Aldinga Scrub , 35°17'S, 138°27'E, 22.ix.1987, D. Hirst leg. (SAM) ; 1 male, Serpentine Lakes , 28°30'S, 129°00'E, 16.iv.1994, D. Hirst leg. (SAM) ; 1 female, St Peters, Adelaide , 33°55'S, 151°11'E, 27.i.1975, P. Walker leg. (AM KS32104) ; 1 male, Walkerville, Adelaide , 34°53'S, 138°37'E, 20.vi.1984, J. Thurmer & D. Hirst leg. (SAM) ; 1 male, Windsor Gardens, Adelaide , 34°52'S, 138°39'E, 20.vi.1988, D. Hirst leg. (SAM) ; 2 males, East Risdon , 42°50'S, 147°21'E, 27.iv.1961, V. V. Hickman leg. (AM KS28582) ; Tasmania: 3 males 1 juvenile, Launceston , 41°27'S, 147°10'E, 06.iv.1928, V. V. Hickman leg. (AM KS28538) , 1 female, same locality, 03.ix.1929, V. V. Hickman leg. (AM KS28547) ; 1 female, Liberty Creek Track, S side Macquarie Heads , 42°12'S, 145°12'E (QM) ; 3 males, 2 females, 2 juveniles, New Town , 42°53'S, 147°19'E, 10.ii.1934, V. V. Hickman leg. (AM KS28699) ; 1 female, St. Columba Falls , 41°19'17"S, 147°55'34"E, 07.iii.2006, G. Hormiga leg. (NHMD) ; Victoria: 1 female, Balwyn, 24 Yandilla Street , 37°48'S, 145°04'E, 05.iv.1981, M. S. Harvey leg. (WAM T73551 ) ; 1 male, same locality, 31.iii.1981, M. S. Harvey leg. (WAM T73572 ) ; 1 male, same locality, 12.i.1981, M. S. Harvey leg. (WAM T73573 ) ; 1 male, same locality, 05.iv.1981, M. S. Harvey leg. (WAM T73574 ) ; 1 male, same locality, 28.xii.1982, M. S. Harvey leg. (WAM T73577 ) ; 1 male, Beaconsfield , 38°03'S, 145°15'E, 07.xi.1893 (MV K9424) ; 1 female, Cann River , 20km N, 37°17'S, 149°12'E, 17.vi.1987, R. J. Raven leg. (QM S13154 ) ; 1 female, Canterbury, 7 Quantock St , 37°49'S, 145°04'E, 13.vi.1981, M. S. Harvey leg. (WAM T73556 ) ; 3 females Croydon , 37°47'S, 145°16'E, 28.ii.1999, S. W. Fulton leg. (MV K10088 ) ; 1 male, 1 female, 1 juvenile, Echuca , 36°08'S, 144°45'E, 01.xi.1955 (MV K9427) ; 1 female, Emerald , 37°56'S, 145°27'E, 13.v.1948, C. Oke leg. (AM KS32504) ; 1 female, Frankston , 38°08'S, 145°08'E, 30.iv.1994 (SAM) ; 1 male, Nyah to Kooloonong (no exact location), B. Harvey leg. (MV K9451) ; Western Australia: 1 female, Attadale , 32°01'S, 115°48'E, 20.vii.1962, A. R. Main leg. (WAM T87192 ) ; 1 female, Bannister , 32°39'S, 116°33'E, 15.vi.1985, B. Y. Main leg. (WAM T73564 ) ; 1 female, Bannister , 32°39'S, 116°33'E, 15.vi.1985, B. Y. Main leg. (WAM T73565 ) ; 1 female (WAM T73566 ) ; 1 male, Boolathana Station , 24°24'49"S, 113°44'41"E, pitfall trap, 15.i.-31.v.1995, J. M. Waldock et al. leg. (WAM T73548 ) ; 1 female, Boya, Helena Valley , 31°54'S, 116°03'E, 06.x.1982, B. Y. Main leg. (WAM T73561 ) ; 1 female (WAM T73562 ) ; 1 female, same locality, 08.x.1982, P. Hussey leg. (WAM T73563 ) ; 1 male, Commonwealth Road, West , 32°44'13"S, 118°16'16"E, wet pitfall trap, 30.x.1997 - 15.v.1998, N. A. Guthrie leg. (WAM T74852 ) ; 1 female, Durokoppin Nature Reserve , 31°24'S, 117°46'E, 03.vi.1989, B. Y. Main leg. (WAM T73567 ) ; 1 female, same locality, 05.v.1987. B. Y. Main leg. (WAM T73568 ) ; 1 male, Grass Patch, Fitzg. Loc. 41, 32°13'56"S, 121°46'00"E, 29.xi.1978, A. F. Longbottom leg. (WAM T73578 ) ; 1 male, Hurlstone Nature Reserve , 32°32'32"S, 119°22'42"E, wet pitfall trap, 30.x.1997 - 20.v.1998, P. van Heurck et al. leg. (WAM T74854 ) ; 1 male, Jarrahdale (Alcoa) Mine area , 31°16'S, 116°06'E, K. E. C. Brennan leg. (WAM T48214 ) ; 1 female, vacuum collector, M. L. Moir leg. (WAM T48215 ) ; 1 female, N of Lake King-Norseman Road , 33°04'54"S, 119°59'53"E, wet pitfall trap, 15.x.1999 - 25.x.2000, N. A. Guthrie leg. (WAM T74863 ) ; 1 male, Lake Morgan, Helms Arboretum , 33°43'09"S, 121°48'29"E, wet pitfall trap, 15.x.1999 - 01.xi.2000, P. van Heurck et al. leg. (WAM T74853 ) ; 1 male, Mount Gibson iron-ore mine, 29°36'02"S, 117°12'25"E, pitfall trap, 15-30.iv.2005, S. Thompson leg. WAM T67918 ); 1 male, Roe Plain, between Mundrabilla and Madura , 32°04'S, 126°31'E, 26.ii.1990, B. Y. Main leg. (WAM T87431 ) ; 1 female, Stirling Range National Park, Moingup Spring , 34°24'S, 118°06'E, 10.vi.1993, J. M. Waldock & A. Sampey leg. (WAM T74423 ) ; 1 female, Torbay Hill, Lot 40, 35°04'S, 117°37'E, 07.x.1983, B. Y. Main leg. (WAM T73569 ) ; 1 female, same locality, 06.x.1993, B. Y. Main leg. (WAM T73570 ) ; 1 female, Two Peoples Bay Nature Reserve , track near coast, 34°59'25"S, 118°09'35"E, 01.v.2008, M. Rix & M. S. Harvey leg. (WAM T81707 ) . NEW ZEALAND: North Island: 1 male, Paranui Scenic Reserve , 35°05′S 173°27′E, at night, 16.ii.2000, G. Hall leg. (NZAC 03038710) ; 1 male, Karaka , 37°06'S, 174°53'E, A.T. Urquhart leg. (CMNZ 2005.135.112) ; 1 male, Hamilton , 37°48′S 175°18′E, 2014, B.N. McQuillan leg. (LUNZ 00012963) ; 1 female, Hamilton , 37°48′S 175°18′E, 2014, B.N. McQuillan leg. (LUNZ 00012964) ; South Island: 1 female, Kaituna Valley , 43°44′S 172°42′E, 24.v.1975, R.R. Forster leg. (NZAC 03038712) .

Diagnosis.

Male N. trituberculosa can easily be distinguished from N. boletus sp. nov. by the much stronger apico-prolateral spur on tibia of leg II and the morphology of key pedipalp sclerites (Figs 1C-E , 2A, B , 4D-F , 5 ): median apophysis with a longer basal portion and a smaller pointed curved and acute tip in N. trituberculosa (projection to the tip flattened, much longer, reaching beyond the radix base and tip rounded in N. boletus sp. nov.), conductor lobe two-lobed in N. trituberculosa (mushroom-shaped in N. boletus sp. nov.), and embolus almost straight in N. trituberculosa (with tip strongly bent, thin and very acute in N. boletus sp. nov.). Females of N. trituberculosa differ from those of N. boletus sp. nov. by details in the epigyne plate (Figs 3C , 6C ), specifically its subtriangular shape in N. trituberculosa (trapezoidal in N. boletus sp. nov.); the transverse and short wrinkles, mainly laterally visible in N. trituberculosa (more pronounced and crossing the plate in N. boletus sp. nov.); copulatory openings less conspicuous in N. trituberculosa (clearly visible in N. boletus sp. nov.); and bridge thin and longer in N. trituberculosa (subtriangular, much wider at posterior margin in N. boletus sp. nov.).

Redescription.

Male (WAM T73571 [images]; WAM T73573 [measurements]): Total length: 5.3. Carapace (Fig. 1A ) 2.7 long, 2.1 wide, light brown with large black bands on lateral margins and yellowish setae throughout except from fovea to pedicel; cephalic area subquadrate; fovea longer than wide and bearing a long black spot. Eyes ringed in black, lateral ones located on small tubercles (Fig. 1A ). AME 0.22, ALE 0.11, PME 0.14, PLE 0.11; row of eyes: AME 0.58, PME 0.40, PLE 0.86. Chelicerae with paturon light brown and fangs reddish-brown; three promarginal teeth, central one largest, two or three retromarginal teeth, basal one largest. Legs (Fig. 1A, B, E ) yellow with dark brown spots on joints; tibiae II with strong apico-prolateral spur that carries a stout spine; metatarsi and tarsi of leg II slightly curved; leg formula IV> I> II> III; length of segments (femur + patella + tibia + metatarsus + tarsus = total length): I - 3.97 + 1.30 + 3.25 + 2.86 + 0.97 = 12.35, II - 2.73 + 1.17 + 2.21 + 0.90 + 0.97 = 7.98, III - 2.34 + 0.71 + 1.23 + 1.30 + 0.72 = 6.30, IV - 3.51 + 0.97 + 2.14 + 2.14 + 0.84 = 9.62. Labium wider than long, subtriangular, brown (Fig. 1B ); endites rounded, light brown (Fig. 1B ). Sternum slightly longer than wide, light brown, with thick black edges and large anterior guanine white spot (Fig. 1B ). Abdomen (Fig. 1A, B ) 3.06 long, 2.47 wide; subtriangular, longer than wide, abdominal humps strong and posterior end reaching over spinnerets; dorsum beige with posterior darker folium pattern; sparsely covered with long brown setae; sides dark olive-grey; venter beige and irregularly mottled with brown spots. Pedipalp (Figs 1C, D , 2A, B ) length of segments (femur + patella + tibia + cymbium = total length): 0.58 + 0.26 + 0.26 + 0.78 = 1.88; conductor lobe bilobed, basal lobe rounded and apical lobe pointed and connected to a wide and concave lateral expansion, whose ectal border has a dense black field of scale-like structures; terminal apophysis with an inflated and membranous body, ending in curved and pointed well-sclerotised tip; embolus slightly sinuous in mesal view, tapering apically towards conductor when expanded (Fig. 2A, B ); conductor very conspicuous and subquadrate with a projected and sclerotised tip; median apophysis stout, heavily sclerotised, forming an arch over the radix, medially curved and extended into a basally pointing tip.

Female (WAM T73556): Total length: 9.0. Carapace 3.5 long, 2.8 wide, eyes, chelicerae, legs, labium, endites and sternum essentially as in male (Fig. 3A, B ). Eye measurements: AME 0.18, ALE 0.09, PME 0.14, PLE 0.14; row of eyes: AME 0.63, PME 0.47, PLE 1.46. Pedipalp length of segments (femur + patella + tibia + cymbium = total length): 1.12 + 0.48 + 0.48 + 1.20 = 3.28. Leg formula IV> I> II> III; length of segments (femur + patella + tibia + metatarsus + tarsus = total length): I - 3.92 + 1.60 + 3.20 + 2.96 + 0.96 = 12.64, II - 3.68 + 1.52 + 2.88 + 0.90 + 0.88 = 9.86, III - 2.40 + 0.88 + 1.44 + 1.44 + 0.80 = 6.96, IV - 4.08 + 1.36 + 2.56 + 2.48 + 1.04 = 11.52. Abdomen (Fig. 3A, B ) 5.00 long, 4.80 wide, similar to male in shape and colour, but slightly more rounded, with less projected humeral humps. Epigyne plate subtriangular, with smooth anterior portion near the basis of scape and lateral wrinkles; scape broken off (Fig. 3C-E ) (as in all specimens examined by us). If present, scape triangular with indistinct transverse wrinkles ( Court and Forster 1988, figs 554, 555). Spermathecae spherical and very large, occupying most of the genital area, medially connected to copulatory openings (Fig. 3D, E ).

Remarks.

The study was conducted over many years and at different institutions and therefore imaging and descriptive work based on variable specimens (plural) availability at the time. This explains why the male N. trituberculosa is here redescribed based on two specimens; one imaged many years ago, but not measured, and the measurements added for a second specimen more recently.

Habitat preferences and life history.

In Australia, mature males of N. trituberculosa were found between November and June, with peaks in January and April. Mature females were found all year round with the lowest numbers of records in November and December. Here, the species is mainly found in "pastoral habits" and constructs a horizontal orb-web amongst low grasses or weeds, with the centre pulled up by stabilizing threads. The webs are up to 0.1 m above ground. Additionally, habitat descriptions on specimen labels include “woodland”, "open forest", "shrubs near ground", "in long grass", but the species also seems to occur in more disturbed habitats such as "among garden rubbish", "ex toilet", "walking on wall at night", "inside house on wall", "stationary on door knob". In New Zealand it is mostly found in pastoral habitats ( Court and Forster 1988), which suggests that it is introduced.

Distribution.

Novakiella trituberculosa has been recorded from all Australian states, except Northern Territory, south of ca. 22°S Latitude (Fig. 7 ). In New Zealand the species is more frequently found in the North Island but it has also been found in some South Island localities ( Court and Forster 1988) (Fig. 8 ).

Figures

Figure 1. Novakiella trituberculosa, male (WAM T 73571). A. Dorsal habitus; B. ventral habitus; C. Left pedipalp, mesal view; D. Left pedipalp, ventral view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Scale bars: A, B, 1 mm; C, D, 0.5 mm; E, 1.2 mm.

Figure 2. Novakiella trituberculosa, male (WAM T 73571), expanded left pedipalp. A. Ventral view; B. Retrolateral view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Scale bars: 0.5 mm

Figure 3. Novakiella trituberculosa, female (WAM T 73556). A. Dorsal habitus; B. Ventral habitus; C. Epigyne, ventral view; D. Cleared epigyne, ventral view; E. Cleared epigyne, dorsal view. Abbreviations: CO, copulatory openings; S, scape; Sp, spermatheca. Scale bars: A, B, 2 mm; C-E, 0.1 mm.

Figure 4. Novakiella boletus sp. nov., male holotype (MV K 9867). A. Dorsal habitus; B. Ventral habitus; C. Left tibia, ventral view; D. Left pedipalp, mesal view; E. Left pedipalp, ventral view, F. left pedipalp, apical view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; P, paracymbium; Ra, radix; TA, terminal apophysis. Scale bars: A, B, 2 mm; C, 1 mm; D-F, 0.5 mm.

Figure 5. Novakiella boletus sp. nov., male holotype (MV K 9867), expanded left pedipalp. mesal view. Abbreviations: C, conductor; CL, conductor lobe; E, embolus; MA, median apophysis; Ra, radix; TA, terminal apophysis. Scale bar: 0.5 mm.

Figure 6. Novakiella boletus sp. nov., female (SAM). A. Dorsal habitus; B. Ventral habitus; C. Epigyne, ventral view. Abbreviations: CO, copulatory openings; L, lips; S, scape. Scale bars: A, B, 2 mm, C, 0.2 mm.

Figure 7. Distribution records of Novakiella trituberculosa (green circles) and Novakiella boletus sp. nov. (red stars) in Australia.

Figure 8. Distribution records of Novakiella trituberculosa in New Zealand; red circle points to material examined for this study and yellow diamonds point to records literature records (Court and Forster 1988).