Leviana dimidiata (L. Koch, 1871) Framenau & Kuntner, 2022

Framenau, Volker W. & Kuntner, Matjaz, 2022, The new Australian leaf-curling orb-weaving spider genus Leviana (Araneae, Araneidae), Evolutionary Systematics 6 (2), pp. 103-133 : 103

Leviana dimidiata (L. Koch, 1871) comb. nov.

Figs 1A-D , 4A, B , 5A-C , 6A , 7A-D , 8A-F , 9

Epeira dimidiata Koch 1871: 95-96, plate 8, figs 1, 1a.

Epeira sylvicola Rainbow 1897: 518-519, plate 17, figs 4, 4a. New synonymy.

Araneus sylvicola (Rainbow).- Rainbow 1909: 222.

Araneus dimidiatus (L. Koch).- Rainbow 1911: 184; Rainbow 1916: 100; Bonnet 1955: 498; Rack 1961: 22.

Araneus sylvicolus (Rainbow).- Rainbow 1911: 194.

Aranea dimidiata (L. Koch).- Roewer 1942: 827.

Aranea sylvicola (Rainbow).- Roewer 1942: 834.

Araneus silvicola (Rainbow).- Bonnet 1955: 598 (corrected spelling; ' silva ' - Latin: wood).

Type material.

Syntypes of Epeira dimidiata L. Koch, 1871: 1 female, Mackay (21°08'S, 149°11'E, Queensland, Australia) (ZMH, ( Rack 1961)-catalogue no. 235) (examined); 2 females, same locality (L. Koch collection) (NHMUK 1915.3.5.594-595) (examined).

Syntypes of Epeira sylvicola Rainbow, 1897: 2 females, Guildford (33°50'S, 150°59'E, New South Wales, Australia) (AM KS.6679) (examined).

Other material examined.

Australia: New South Wales: 2 males, Beecroft, 33°45'S, 151°04'E (AM KS.53874, KS70922); 2 males, 1 female, Beecroft Reserve , 33°45'S, 151°04'E (AM KS.54073, KS.58718, KS.66205) ; 1 female, Bellevue Hill, Sydney, 33°53'S, 151°17'E (AM KS.70044); 1 male, Bonny Hills Road, off Pacific Highway, S of Port Macquarie , 31°33'S, 152°49'E (AM KS.62864) ; 1 female, Cheltenham, 33°45'S, 151°05'E (AM KS.56988); 1 female, Como, 34°01'S, 151°04'E (AM KS.32822); 1 female, Cudgen, 28°16'S, 153°33'E (QM); 12 females, 4 juv., Enfield, 33°53'S, 151°06'E (AM KS.33984); 1 male, Forster, 6 km S, 32°12'S, 152°31'E (AM KS.10204); 1 female, Frazer Reserve , Wahroonga, 33°43'S, 151°08'E (AM KS.56766) ; 1 female, Galston, Crosslands Road, 33°38'S, 151°05'E (AM KS.83701); 1 female, Gosford, 33°25'S, 151°20'E (MV K-10331); 1 female, Jamberoo Mountain , 34°43'S, 151°18'E (AM KS.81993) ; 1 female, Lake Munmorah State Recreation Reserve , Geebung, 22°12'S, 151°34'E (AM KS.17794) ; 1 female, same locality (AM KS.17811) ; 1 male, Maroota State Forest , 33°31'S, 150°59'E (AM KS.73416) ; 1 female, Medlow Bath, 33°41'S, 150°17'E (AM KS.34097); 1 female, Minnamurra Falls , 34°38'S, 150°44'E (AM KS.100355) ; 1 female, Moruya, 35°55'S, 150°05'E (AM KS.33981); 2 females, Narara, 33°23'S, 151°21'E (MV K-10332); 1 male, O'Hares Creek , 34°14'S, 150°52'E (AM KS.86691) ; 1 female, Port Stephens , south shore, 32°42'S, 152°06'E (AM KS.33983) ; 1 female, Wahroonga, 33°43'S, 151°08'E (AM KS.46010); 1 female, Wallumatta Nature Reserve , North Ryde, 33°48'S, 151°07'E (AM KS.81958) ; 1 male, Warrumbungles National Park , White Gum carpark area, 31°17'20"S, 149°02'26"E (AM KS.75194) ; 2 males, 1 female, Warrumbungles National Park , 0.9 km E of western entrance to park, 31°16'35"S, 148°57'49"E (AM KS.75238) ; 1 female, same locality (AM KS.75239). Queensland : 1 male, Belmont Hills Bushlands , 27°31'S, 153°07'E (QM S54745 ) ; 2 females, same locality (QM S54746 ) ; 1 female, same locality (QM S54744 ) ; 1 female, same locality (QM) ; 3 females, Blackdown Tableland , Stony Creek Falls, 16°53'S, 145°39'E (QM S39784 ) ; 1 female, Boondall Wetlands , 27°20'21"S, 153°04'27"E (QM S67469 ) ; 3 females, same locality (QM) ; 3 females, 1 juv., same locality (QM S67470 ) ; 1 female, same locality (QM S67468 ) ; 1 female, Boonoo Boonoo , 28°53'S, 152°06'E (QM S39797 ) ; 1 male, Bribie Island , 27°03'30"S, 153°11'32"E (AM KS.69544) ; 1 male, same locality (AM KS.69516) ; 2 males, same locality (AM KS.69609) ; 2 females, 11 juv., Brisbane , 27°28'S, 153°01'E (QM S39789 ) ; 1 female, same locality (QM S69647 ) ; 1 female, Buhot Creek , Burbank, 27°35'27"S, 153°10'19"E (QM S67807 ) ; 1 male, same locality (QM S67283 ) ; 3 males, same locality (QM S67282 ) ; 1 female, same locality (QM) ; 2 males, 1 female, same locality (QM S67628 ) ; 1 males, same locality (QM S111921) ; 1 male, 3 females, Bulimba Creek , Carindale, 27°30'9"S, 153°06'03"E (QM S67467 ) ; 1 female, same locality (QM S67466 ) ; 1 female, same locality (QM S67463 ) ; 1 male, same locality (QM S67461 ) ; 1 female, same locality (QM S67464 ) ; 6 males, 6 females, same locality (QM S67462 ) ; 1 female, Bundaberg Forest , 24°52'S, 152°21'E (QM) ; 1 female, Bundaberg , Baldwins Swamp, 24°52'S, 152°21'E (QM S25246 ) ; 1 female, Bundaberg, land, 24°52'S, 152°21'E (MV K-10345); 12 females, Camp Milo , Cooloola, 26°00'S, 153°05'E (QM S39750 ) ; 1 female, Chelsea Road Bushland Reserve , 27°28'58"S, 153°11'15"E (QM) ; 2 females, same locality (QM) ; 4 females, same locality (QM S65837 ) ; 2 females, same locality (QM S65840 ) ; 1 female, Closeburn , 27°20'S, 152°52'E (QM S39766 ) ; 3 females, Consuelo Tablelands, Foleys Yards, 25°01'S, 147°57'E (QM); 2 females, Cooloola , 26°12'S, 153°03'E (QM S39753 ) ; 1 female, Cooloola State Forest , 26°12'S, 153°03'E (QM S33393 ) ; 1 female, Coombabah Creek 200 m upstream Lakeside Caravan Park, 27°52'S, 153°22'E (QM S39769 ) ; 2 females, Curtis Island , 23°37'S, 151°9'E (QM S69916 ) ; 1 female, Deepwater National Park , 24°32'S, 151°58'E (QM S25402 ) ; 1 female, Downfall Creek , 27°23'S, 153°01'E (QM) ; 1 female, Drewvale, Illaweena Street, 27°38'39"S, 153°03'47"E (QM); 1 female, same locality (QM) ; 5 females, same locality (QM S65839 ) ; 1 female, Eidsvold , 25°22'S, 151°07'E (QM S39765 ) ; 1 female, Enoggera Army land, Brisbane, 27°26'S, 152°59'E (QM S22429 ) ; 1 female, Expedition Range National Park , 25°12'S, 148°59'E (QM S44180 ) ; 1 male, 5 females, 9 juv., Finch Hatton National Park , 21°9'S, 148°38'E (QM) ; 1 female, Forty Mile Scrub, SW Mt Garnet , 18°05'S, 144°51'E (QM S39778 ) ; 1 female, Girraween National Park , near, 28°50'S, 151°55'E (QM S39774 ) ; 1 female, Gold Creek Reservoir, 27°27'53"S, 152°52'32"E (QM S54749 ) ; 1 female, same locality (QM S54750 ) ; 1 female, same locality (QM) ; 1 female, same locality (QM S54751 ) ; 1 female, same locality (QM S54748 ) ; 3 females, same locality (QM S54747 ) ; 1 female, Hann Tableland , northern end, 16°48'S, 145°10'E (QM S40510 ) ; 1 female, same locality (QM S41332 ) ; 3 females, same locality (QM S40498 ) ; 2 females, Helidon , 27°33'S, 152°08'E (QM S39795 ) ; 1 female, Hurdie Gully , 14.8 km WSW Monto, 24°55'S, 150°59'E (QM S44168 ) ; 1 female, Ipswich , 27°37'S, 152°47'E (QM S39763 ) ; 2 females, Karawatha Forest , 27°37'33"S, 153°05'24"E (QM S67460 ) ; 1 female, same locality (QM) ; 2 males, same locality (QM S67459 ) ; 1 female, Kenniffs Cave , Mt Moffat, 25°01'S, 147°57'E (QM) ; 1 female, same locality (QM) ; 1 female, Kroombit Tops, 24°22'S, 151°01'E (QM); 1 female, same locality (QM) ; 4 females, same locality (QM S39751 ) ; 2 females, Kroombit Tops, Lower Dry Creek , 24°24'S, 151°01'E (QM) ; 7 females, Kroombit Tops, Upper Dry Creek , 24°25'S, 151°03'E (QM) ; 4 females, Kroombit Tops, Northern Escarpment , 24°22'S, 151°01'E (QM) ; 1 female, Kroombit Tops, Upper Kroombit Creek , 24°25'S, 151°03'E (QM) ; 1 female, Kroombit Tops, Upper TA47 Creek, 24°25'S, 151°03'E (QM); 1 female, Kuraby , 27°37'S, 153°05'E (QM S41023 ) ; 1 male, 1 female, Mahogany Forest , Mt Moffat National Park, 24°56'S, 148°04'E (QM S111925) ; 2 females, Molangdool, 24°45'S, 151°33'E (AM KS.33985); 1 female, Moreton Island , 27°11'S, 153°24'E (QM S39782 ) ; 15 females, Mt Abbott , 20°06'S, 147°45'E (QM S38572 ) ; 1 female, Mt Coolum , 26°34'S, 153°05'E (QM S111923) ; 1 female, same locality (QM) ; 1 female, Mt Flinders , 22°32'S, 150°46'E (QM S39768 ) ; 1 female, Mt Garnet , 17°41'S, 145°07'E (QM) ; 1 female, Mt Moffat National Park , 25°01'S, 147°57'E (QM) ; 1 female, Mt Moffat , summit, 25°01'S, 147°57'E (QM) ; 1 female, 1 juv., Mt Moffat , top camp, 25°01'S, 147°57'E (QM S111924) ; 2 males, 2 females, 3 juv., Nelson, 12°38'S, 143°15'E (AM KS.32825); 7 females, Nerimberah, via Rockhampton, 23°24'S, 150°35'E (QM S9923); 1 female, North Stradbroke Island , 27°35'S, 153°27'E (QM S55768 ) ; 2 females, same locality (QM S55769 ) ; 3 females, same locality (QM S55744 ) ; 7 females, same locality (QM S55632 ) ; 2 females, same locality (QM S55743 ) ; 4 females, same locality (QM S56284 ) ; 2 females, same locality (QM S55631 ) ; 1 female, same locality (QM S55745 ) ; 1 female, same locality (MV K-10333) ; 1 female, Orchid Beach , Fraser Island, 24°58'S, 153°19'E (QM S31029 ) ; 2 females, Peel Island , 27°30'S, 153°21'E (QM S39748 ) ; 1 female, Percy Island , 21°24'S, 150°20'E (QM S39787 ) ; 2 females, Ransome Reserve , 27°29'34"S, 153°11'05"E (QM) ; 1 male, same locality (QM) ; 4 females, same locality (QM) ; 2 females, same locality (QM S65838 ) ; 3 males, 1 female, same locality (QM S65841 ) ; 1 female, Rochedale , 27°37'S, 153°9'E (QM S39757 ) ; 1 female, same locality (QM S39760 ) ; 3 males, 6 females, 2 juv., Rochedale State Forest , 27°37'S, 153°09'E (QM S39754 ) ; 1 female, same locality (QM S39759 ) ; 1 female, same locality (QM S39758 ) ; 1 female, same locality (QM S39755 ) ; 1 female, same locality (QM) ; 2 females, same locality (QM S39756 ) ; 1 female, Rundle Shale camp site, 23°39'S, 150°59'E (QM S39781 ) ; 1 female, South Percy Island , Central Valley, 21°45'S, 150°19'E (QM S33318 ) ; 1 female, South Percy Island , NW Bay, 21°45'S, 150°19'E (QM) ; 2 females, Stanthorpe , 28°40'S, 151°56'E (QM S39783 ) ; 1 male, Taringabah , 17°43'S, 145°56'E (QM S39770 ) ; 1 female, Teewah Creek , Cooloola, 26°05'S, 153°02'E (QM S39788 ) ; 1 male, 3 females, 1 juv., Teviot Brook , Boonah, 27°55'S, 152°33'E (QM S39762 ) ; 1 female, Top Moffat Camp, Mt Moffat National Park , 25°01'S, 147°05'E (QM S111926) ; 1 female, Upper Brookfield , 27°30'S, 152°55'E (QM S39796 ) ; 1 female, Warro , via Lowmead, 26°56'S, 150°55'E (QM S39779 ) ; 1 female, Woodgate , 25°06'S, 152°34'E (QM S39780 ) ; 10 females, Yeppoon , Emu Park, Rockhampton, 23°16'S, 150°05'E (QM S69646 ) ; 1 female, same locality (QM S39761 ) ; Victoria: 2 females, Bruthen, 37°42'S, 147°50'E (MV K-10335); 2 females, Dermody's Camp , Avon River State Forest, 37°48'17"S, 146°55'03"E (WAM T68049 ) .

Diagnosis.

Leviana dimidiata comb. nov. is most similar to L. folium sp. nov. in both somatic (size, colouration) and genitalic morphology. However, they can be clearly separated by the abdominal pattern, particularly by the shape of the black anterior shoulder marks and lateral lines. In L. dimidiata comb. nov., the anterior black marks and lateral lines form an obtuse internal angle (e.g. Figs 7A , 8A - arrows), whereas in L. folium sp. nov. this angle is acute (e.g. Figs 12A , 13A - arrows). Male pedipalp morphology differs subtly in some key sclerites; for example, the terminal apophysis has a sclerotised tip in L. dimidiata comb. nov. which is absent in L. folium sp. nov. (Fig. 7D vs Fig. 13D ), the conductor is more prominent and more elongated and the spine in the basal arch of the median apophysis is comparatively much larger (Fig. 7C vs Fig. 12C ). The base of the scape of the female epigyne protrudes further anteriorly in L. dimidiata comb. nov. than in L. folium sp. nov. (Fig. 8C vs Fig. 13C ).

Description.

Male (based on QM S111921). Total length 6.6. Carapace (Fig. 7A ) 3.4 long, 2.3 wide; light brown centrally with dark brown lateral flanks; white setae mainly in cephalic and centrally lighter area; fovea indistinct roundish depression. Eyes: AME 0.20, ALE 0.13, PME 0.14, PLE 0.14; row of eyes: AME 0.48, PME 0.44, PLE 1.12. Sternum (Fig. 7B ) 1.6 long, 1.1 wide; orange-brown; heavy dark grey pigmentation towards margins; few white setae; few brown bristles marginally. Labium subtriangular, 0.48 long, 0.51 wide; basally dark reddish-brown. Maxillae yellow-brown. Chelicerae dark-brown; few white setae in basal half; three retromarginal teeth of equal size, four promarginal teeth with the second-basal one largest. Legs: leg formula I> II> IV> III; dark brown with distinct yellow-brown annulations; lengths of segments (femur + patella + tibia + metatarsus + tarsus = total leg length): pedipalp 0.8 + 0.2 + 0.2 + - + 1.0 = 2.2, I 3.5 + 1.5 + 3.2 + 3.4 + 1.3 = 12.9, II 2.9 + 1.4 + 2.4 + 2.9 + 1.1 = 10.7, III 1.9 + 0.8 + 1.1 + 1.3 + 0.8 = 5.9, IV 2.4 + 1.1 + 2.8 + 2.9 + 0.8 = 8.1. Abdomen (Fig. 7A, B ) 3.7 long, 2.4 wide; elongate ovoid; dorsally yellow-grey becoming darker posteriorly; two black central marking anteriorly that continue laterally as black lines almost to spinnerets; four pairs of dark brown sigilla centrally; covered with long brown bristles and white setae; venter centrally dark grey with two longitudinal lateral lighter bands, covered with brown setae; spinnerets light brown.

Pedipalps (Figs 7C, D ; also 4A, B, 5A-C): median apophysis apically broadly lobed, basally with thorn opposing a hooded base; terminal apophysis fleshy, elongated with double-tip; embolus with broad base, curved and short tip; conductor wider than high in ventral view.

Female (based on QM S67628; epigyne variation QM S25402). Total length 7.1. Carapace (Figs 8A ) 3.5 long, 2.6 wide; light yellowish-brown, cephalic area somewhat darker and with a dark brown median band; few white setae in particular in cephalic area; fovea shallow and poorly demarcated. Eyes: AME 0.18, ALE 0.15, PME 0.13, PLE 0.11; row of eyes: AME 0.46, PME 0.35, PLE 1.23. Sternum (Fig. 8B ) 1.8 long, 1.3 wide; yellowish-brown; dark brown pigmentation laterally; covered with few white setae; few long brown bristles marginally. Labium 0.64 long, 0.44 wide, colouration as male. Maxillae as male. Chelicerae dark brown; few white setae in basal half, black setae mesally mainly in apical half; four promarginal teeth, with the apical and third one largest; three retromarginal teeth of similar size. Legs: leg formula I> II> IV> III; light brown with indistinct darker annulations, most distinct on tibiae and metatarsi of leg III and IV; lengths of segments (femur + patella + tibia + metatarsus + tarsus = total length of leg): pedipalp 1.0 + 0.3 + 0.2 + - + 0.8 = 2.3, I 2.9 + 1.5 + 2.3 + 2.4 + 1.1 = 10.2, II 2.6 + 1.1 + 1.9 + 2.1 + 0.8 = 8.5, III 1.6 + 1.0 + 1.5 + 1.1 + 0.6 = 5.8, IV 2.7 + 1.1 + 1.8 + 1.9 + 0.8 = 8.3. Abdomen (Fig. 8A, B ) 3.5 long, 3.2 wide; ovoid, slightly dorsoventrally compressed; dorsal colour pattern and setae largely as male, however lighter; venter as male, but with narrow light guanine line laterally; spinnerets brown.

Epigyne (Fig. 8C-F ) ovoid wider than long, atrium bulging; central division broadly oval with narrow base; scape with almost parallel lateral border, almost translucent; often broken off anteriorly; spermathecae ovoid, almost touching centrally; fertilization duct very slightly curved and attaching laterally at atrium (Fig. 6A ).

Variation.

Males total length 4.8-6.6 (n = 5); females total length 6.4-9.9 (n = 7). There is little colour variation in males and females of this species, although the posterior part of the abdomen might be darker than in the specimens illustrated. Two out of seven females measured for this study had their epigyne scape intact.

Remarks.

Rack (1961) considered a female in the ZMH the holotype of Epeira dimidiata , however, two further females from the type locality lodged in the NHMUK clearly bear the typical labels of L. Koch which suggests these to by syntypes. The original description does not state the number of specimens on which the description was based, and we therefore consider all specimens from Port McKay in the ZMH and NHMUK as syntypes.

The syntypes of Araneus sylvicolus (Rainbow, 1897) are clearly conspecific with those of L. dimidiata comb. nov. based on genitalic and somatic morphology, considering the intraspecific variation as described here. We therefore consider E. sylvicola Rainbow, 1897 a junior synonym of L. dimidiata (L. Koch, 1871) comb. nov.

Life history and habitat preferences.

Adult males were mainly found between September and January, with a peak in November. Mature females were mainly found between September and April, with two peaks in December and February.

Records of L. dimidiata comb. nov. are from a variety of open and closed forests and woodlands, including those of eucalypt, Casuarina and Melaleuca . However, there are also some records from heathlands and vine scrubs.

Distribution.

Eastern Australia from far north Queensland to eastern Victoria (Fig. 9 ).

Figures

Figure 1. Leviana dimidiata comb. nov. natural history (at Whites Hill, Queensland): A. Female vertical orb-web with spider in curled leaf retreat connected with hub via signal line; B. Same web, close up; note V-shaped open sector with signal line, closed hub with abrupt transition between hub loops and sticky spiral, and all radii originating at hub; C. Female in defensive pose after her leaf-retreat had been torn open; D. Leaf retreat with female (not seen) feeding on dragonfly prey.

Figure 4. Leviana dimidiata comb. nov., interpretation of the male pedipalp anatomy (QM S 54745). A. Ventral view; B. Dorsal view. Scale bar: 0.2 mm.

Figure 5. Leviana dimidiata comb. nov., expanded male pedipalp (QM S 111921). A. Ventral view; B. Apical view; C. Dorsal view. Scale bar: 0.2 mm.

Figure 6. Leviana gen. nov., cleared epigynes, posterior view. A. L. dimidiata comb. nov. (AM KS. 81993); B. L. cincinnata sp. nov. (QM S 111931). Scale bars: 0.05 mm.

Figure 7. Leviana dimidiata comb. nov., male (QM S 111921). A. Habitus, dorsal view; B. Habitus, ventral view; C. Left male pedipalp, ventral view; left male pedipalp, dorsal view. Scale bars: 2 mm (A, B); 0.2 mm (C, D). Arrow in (A) points to diagnostic character distinguishing L. dimidiata comb. nov. and L. folium sp. nov.

Figure 8. Leviana dimidiata comb. nov., female (A, B, F QM S 67628; C-E QM S 25402). A. Habitus, dorsal view; B. Habitus, ventral view; C. Epigyne, ventral view; D. Epigyne, lateral view; E. Epigyne, posterior view; F. Epigyne, ventral view (scape broken off). Scale bars: 2 mm (A, B); 0.2 mm (C, D). Arrow in (A) points to diagnostic character distinguishing L. dimidiata comb. nov. and L. folium sp. nov.

Figure 9. Distribution records of Leviana dimidiata comb. nov. and L. cincinnata sp. nov.

Figure 12. Leviana folium sp. nov., male holotype (QM S 111932). A. Habitus, dorsal view; B. Habitus, ventral view; C. Left male pedipalp, ventral view; left male pedipalp, dorsal view. Scale bars: 2 mm (A, B); 0.2 mm (C, D). Arrow in (A) points to diagnostic character distinguishing L. folium sp. nov. and L. dimidiata comb. nov.

Figure 13. Leviana folium sp. nov., female (A, B, F QM S 111922; C-E QM S 116638). A. Habitus, dorsal view; B. Habitus, ventral view; C. Epigyne, ventral view; D. Epigyne, lateral view; E. Epigyne, posterior view; F. Epigyne, ventral view. Scale bars: 2 mm (A, B); 0.1 mm (C, D). Arrow in (A) points to diagnostic character distinguishing L. folium sp. nov. and L. dimidiata comb. nov.