Nyssus Walckenaer, 1805

Nyssus Walckenaer, 1805 View in CoL View at ENA

Nyssus Walckenaer, 1805: 52 View in CoL . Type species by monotypy: Nyssus coloripes Walckenaer, 1805: 52 View in CoL , type lost, neotype here designated. The gender is masculine.

Anchognatha Thorell, 1881: 228 . Type species by original designation, Anchognatha avida Thorell, 1881 . New Synonymy.

Supunna Simon, 1897: 168 (type species by subsequent designation of Rainbow, 1911, Agroeca insularis L. Koch, 1873 ); Rainbow, 1911: 253; Petrunkevitch, 1928: 180; Roewer, 1955: 635; Bonnet, 1958: 4196; Platnick, 1989: 448. Type species discussed above in Nomenclatural Issues. New Synonymy.

Diagnosis. In species with a large paracymbial spine, males are readily distinguished from most similar genera and from males and females of Battalus in the absence of the large spines on the cheliceral fang mound, from Iridonyssus in the wider eye group and longer caput, and from Poecilipta in that the legs are all of similar size. Males of Nyssus species lacking a paracymbial spine differ from those of Leichhardteus in the extensive dorsal and ventral scutes.

Description. Carapace pear-shaped. White feathery hairs form spots or radial lines ( Figs 16 View FIGURE 16 , b, c, 79a, b, d View FIGURE 79 ) but are often lost in ethanol. Lateral profile of carapace ( Fig. 2a View FIGURE 2 ): high but more or less flat or slightly arched for its length; marginally highest point behind fovea at shallowly sloping posterior declivity; eyes set behind clypeus edge. Retrolateral face of large chelicerae smooth. Eyes ( Fig. 16b, c View FIGURE 16 ): front row slightly recurved; back strongly procurved; group occupies about half headwidth; all eyes similar in size and opacity. Clypeus low, eye region curved gently. Labium anterior border thick, pallid. Serrula long, distal. Chelicerae: fang diagonal; pallid groove in front above fang; females with shield of thick hairs over front face of fang. Legs: femora I and II similar in thickness and overall size to III and IV. Pair of long erect bristles, one basidorsal and one apicodorsal on each of patellae and tibiae I–IV; none on palp. Trochanters I–III weakly notched, slightly deeper on IV. Femora III, IV with moderate to strong but always distinct predistal constriction (most species) or unmodified ( N. coloripes , N. jaredwardeni ). Scopula: I and II weak, thin on metatarsi, full on tarsi, not obscuring cuticle; III weaker still; IV entirely absent. Claw tufts: dense, small. Spines: distinct, strong but not on raised base, on all legs, except tarsi, and palp. Tibiae I and II ventrally with three pairs, weak in males. Scutes: tracheal spiracle small, teat-shaped, with sclerotised base bear spinnerets; ridge more evident in ♂. Males with ovate shield in anterior fifth of abdomen and sclerotised shield over booklungs ( Nyssus coloripes group, most species); ventral scute present (most species) or absent ( Nyssus semifuscus , and some males of N. albopunctatus ). Spinnerets: colulus represented by setose area ( Fig. 7f View FIGURE 7 ). PLS and ALS short with domed tips; PMS digitiform; large spigots on PLS. Trichobothria: tarsi with two rows for full length. Male palp: tibia with low plate-like process prodistoventrally; strong bipartite RTA ( N. paradoxus , Fig. 88a–c View FIGURE 88 ), strong spine-like ( N. wendyae , Fig. 90a, b View FIGURE 90 ) or weak (most species) bristles retrolaterally; cymbium canoe-shaped with paracymbial spine (most species) or absent ( N. pseudomaculatus , N. jonraveni , N. yuggera , N. loureedi ); without spines or scopula; bulb a simple tear-shaped tegulum with embolus apically twisted. Epigyne: CO form pair of ovoid ridges, sometimes complete or closed, widely separated, and with additional groove and ridge below ( N. coloripes group) or medially ( N. paradoxus ) to CO. Internally of four types: in N. coloripes group, with small spermathecal head and wide ducts to FD ( Fig. 135 View FIGURE 135 ); in N. yuggera and N. loureedi , simple narrow ducts with no enlargement of head and narrow duct to FD ( Fig. 135 View FIGURE 135 ); in N. albopunctatus group, very large spermathecal head with broad lobes to FD; and in N. paradoxus and N. robertsi, CD as long as epigyne and connects through short duct to hardly enlarged spermathecal head ( Fig. 135 View FIGURE 135 ).

Type species: Nyssus coloripes Walckenaer, 1805 View in CoL , by monotypy.

Distribution and Habitat. Nyssus occurs throughout Australia and both N. coloripes and N. albopunctatus have been presumably introduced to New Zealand on shipping containers. Nyssus is common in open habitats including heath, open eucalypt forest, grassland and savannah woodland, and may be found in sunny patches on the fringes of rainforest and vine thickets.

Included species: Supunna albopunctatus ( Hogg, 1896) , Anchognatha avida Thorell, 1881 , Nyssus coloripes Walckenaer, 1805 , N. emu sp. nov., Agroeca insularis L. Koch, 1873 , N. jaredwarden sp. nov., N. jonraveni sp. nov., N. loureedi sp. nov., N. luteofinis sp. nov., N. paradoxus sp. nov., N. pseudomaculatus sp. nov., N. robertsi sp. nov., N. semifuscus sp. nov., N. wendyae sp. nov. and N. yuggera sp. nov.

Remarks. The gender of Supunna is feminine and that of Nyssus is masculine ( Bonnet 1958). One of the two most common species of Nyssus is the highly ornate Nyssus coloripes ( Fig. 73a–d View FIGURE 73 , commonly known as Supunna picta ); the other is Nyssus albopunctatus ( Fig. 79b View FIGURE 79 ). N. albopunctatus is quite distinctive in the black legs and white spots on a jet black abdomen. A number of other Nyssus species have the same or a very similar pattern but all are morphologically quite distinct.

Species Groups

Nyssus is here broken into four species groups ( N. coloripes group, N. albopunctatus group, N. avidus group and N. paradoxus group) based upon genitalic characters and patterns of hair on the carapace. Initially, it seemed a genus for each may be warranted but the defining characters are intermediate and that decision is reserved for a cladistic and molecular analysis.

Key to species of Nyssus View in CoL

Males (of N. insularis and N. robertsi View in CoL unknown)

1. Palp. Large retrolateral tibial apophysis ( Fig. 88a–c View FIGURE 88 )............................................ Nyssus paradoxus Palp. No View in CoL tibial apophysis ( Fig. 90a, b View FIGURE 90 ) or tibial apophysis subtle ( Fig. 90a, b View FIGURE 90 )...................................... 2

2(1). Femora III and IV distally normal ( Fig. 75b View FIGURE 75 )................................................................ 3 Femora III and IV with distinct distal constriction (as in Iridonyssus kohouti View in CoL , Fig. 6d View FIGURE 6 )............................... 4

3(2). Paracymbial spine on large basal process, within one diameter of cymbial base ( Fig. 74a, g View FIGURE 74 )................. N. coloripes Paracymbial View in CoL spine on small subbasal process but not very basal ( Fig. 75d, e View FIGURE 75 )........................... N. jaredwardeni View in CoL

4(2). Paracymbial spine minute or absent ( Fig. 85a, b View FIGURE 85 )............................................................. 5 Paracymbial spine large, distinct (e.g., Fig. 90 View FIGURE 90 , 91a View FIGURE 91 ........................................................... 8 View FIGURE 8

5(4). Embolus apically with corkscrew-like tip ( Figs 85a View FIGURE 85 , 87c View FIGURE 87 )...................................................... 6 Embolus very long or straight ( Fig. 81a, g View FIGURE 81 )................................................................. 7

6(5). Embolus apically with very short, corkscrew-like tip ( Fig. 85a View FIGURE 85 )......................................... N. yuggera Embolus View in CoL apically longer, screw-like tip ( Fig. 87c View FIGURE 87 ).................................................... N. loureedi View in CoL

7(5). Embolus, darkly sclerotised tip very long ( Fig. 81g View FIGURE 81 ); white submedial bands on carapace small and separate ( Fig. 81f View FIGURE 81 )................................................................................................. N. jonraveni Embolus View in CoL , darkly sclerotised tip short ( Fig. 81a View FIGURE 81 ); white submedial bands on carapace long and almost continuous ( Fig. 81e View FIGURE 81 )........................................................................................ N. pseudomaculatus View in CoL

8(4). Embolus spiral large, open ( Figs 90a View FIGURE 90 , 91a View FIGURE 91 ).................................................................. 9 Embolus spiral smaller ( Fig. 80a, b View FIGURE 80 )...................................................................... 10

9(8). Palpal tibia distally with long strong bristles, especially retrolaterally ( Fig. 90a, b View FIGURE 90 ).......................... N. wendyae Palpal View in CoL tibia with similar bristles for length ( Fig. 91a View FIGURE 91 )..................................................... N. emu View in CoL

10(8). Embolus apically with very short, corkscrew-like tip ( Fig. 82a View FIGURE 82 ).......................................... N. avidus Embolus View in CoL apically long spiral ( Fig. 80a View FIGURE 80 )................................................................... 11

11(10). Embolus distally long, slender tube-like ( Fig. 80a, b View FIGURE 80 ); black spider with bold white markings on carapace and abdomen ( Fig. 79b View FIGURE 79 )................................................................................... N. albopunctatus Embolus View in CoL short, converging quickly ( Fig. 78a, b View FIGURE 78 ); brown spiders with no strongly contrasting markings................. 12

12(11). Triangular paracymbial spine on distinctly raised basal process ( Fig. 78a, b View FIGURE 78 )............................. N. semifuscus Digitiform View in CoL paracymbial spine not on raised process ( Fig. 77b View FIGURE 77 )......................................... N. luteofinis View in CoL

Females (of N. emu View in CoL , N. jonraveni View in CoL and N. wendyae View in CoL unknown)

1. Copulatory orifices large, almost contiguous or within one diameter ( Figs 74c View FIGURE 74 , 80d View FIGURE 80 )................................. 2 Copulatory orifices at least one of their diameters apart ( Fig. 81c View FIGURE 81 )............................................... 3

2(1). Abdomen black with median stripe and two pairs of pale spots posteriorly ( Fig. 73a–d View FIGURE 73 ); femora III and IV normal distally (as in N. jaredwardeni View in CoL , Fig. 75b View FIGURE 75 )................................................................... N. coloripes View in CoL Abdomen dorsally black with 3–4 pairs of white spots; carapace black with white submarginal band ( Figs 79b View FIGURE 79 , 80c View FIGURE 80 ); femora III and IV with distinct distal constriction (as in Iridonyssus kohouti View in CoL , Fig. 6d View FIGURE 6 )............................ N. albopunctatus View in CoL

3(1). Carapace with radial lines of white hair on black to brown cuticle (as in Figs 77e View FIGURE 77 , 79d View FIGURE 79 , 83a View FIGURE 83 , 84a View FIGURE 84 ).......................4 Carapace with submarginal brackets of white hair (as in Fig. 81e, f View FIGURE 81 ).............................................. 8

4.(3). Spermathecae with enlarged head (as in Fig. 82d View FIGURE 82 )............................................................ 5 Spermathecae without enlarged head ( Fig. 85d View FIGURE 85 ); Queensland ........................................... N. yuggera View in CoL

5(4). Spermathecae with very large head (as in Fig. 82d View FIGURE 82 )........................................................... 6 Spermathecae with small head (as in Figs 77d View FIGURE 77 , 87b View FIGURE 87 )..........................................................7

6(5). Copulatory orifices large, spiralled ( Fig. 82c View FIGURE 82 )........................................................ N. avidus Copulatory View in CoL orifices very small ( Fig. 87e View FIGURE 87 ).......................................................... N. insularis

7(5). Spermathecae contiguous for their length ( Fig. 77d View FIGURE 77 ); Queensland ....................................... N. luteofinis Spermathecae View in CoL predistally curve ectally then contiguous distally ( Fig. 87a, b View FIGURE 87 ); Lord Howe Island............... N. loureedi View in CoL

8(3). Epigyne with rounded V-shaped ridge between copulatory orifices ( Fig. 89a View FIGURE 89 )...................................... 9 Space between copulatory orifices smooth (as in Fig. 76a View FIGURE 76 ).................................................... 10

9(8). Median epigynal groove ends anterior to epigastric ridge ( Fig. 89a, b View FIGURE 89 ).................................. N. paradoxus View in CoL Median epigynal groove extends to epigastric ridge ( Fig. 89d View FIGURE 89 ).......................................... N. robertsi View in CoL

10(8). Femora III and IV with distinct distal constriction (as in Iridonyssus kohouti View in CoL , Fig. 6d View FIGURE 6 ).............................. 11 Femora III and IV distally normal ( Fig. 75b View FIGURE 75 ).................................................... N. jaredwardeni View in CoL

11(10). Abdomen dorsally black with 3–4 pairs of white spots; carapace black with white submarginal band ( Fig. 81e View FIGURE 81 )................................................................................................. N. pseudomaculatus View in CoL Abdomen brown in posterior half, without spots................................................... N. semifuscus View in CoL