Phryganoporus Simon, 1908

Gray, Michael R., 2002, The Taxonomy and Distribution of the Spider Genus Phryganoporus Simon (Araneae: Amaurobioidea: Desidae), Records of the Australian Museum 54 (3), pp. 275-292 : 276-280

publication ID	https://doi.org/ 10.3853/j.0067-1975.54.2002.1367
persistent identifier	https://treatment.plazi.org/id/C22187DE-FC59-5053-A46C-F91723A1FD1B
treatment provided by	Felipe
scientific name	Phryganoporus Simon
status

Treatment

Genus Phryganoporus Simon View in CoL

Phryganoporus Simon, 1908: 367 View in CoL (type species Amaurobius gausapatus Simon, 1906 ). Roewer, 1954: 1370. Lehtinen, 1967: 259, 324.

Ixeuticus (part).– Main, 1971: 119. Brignoli, 1983: 495.

Badumna View in CoL (part).– Gray, 1983: 248.

Diagnosis. Cribellate spiders. Body silvery grey to brown with light and dark brown patterning on abdomen. In solitary or communal webs. Separated from Badumna by the strong presence of white setae on the body; the possession on the male palp of a large, prolaterally directed, basal tegular bulge; extension of the conductor fold across to the retrolateral side of the bulb; and a basal RTA. From Forsterina and Reinga by the prolateral origin of the embolus and the presence of a transverse epigynal ridge. From Namandia by the sinuous course of the embolus across the ventral bulb and the presence of fused paracribellar spigots on the PMS; and from Paramatachia by these characters plus the presence of a median apophysis.

Description. Medium-sized cribellate spiders (CL 1.9–4.8) that live in solitary and communal sheet webs and run on top of the sheet. Overall colouration varies from silvery grey to brown, depending on the density of white hairs present (juveniles are often lighter in colour than adults). Body setae plumose, comprising shorter, broader, more recumbent hairs, and longer bristle-like hairs ( Fig. 1f View Figure 1 ). Legs with several “plumose” setal types ( Fig. 1a–c,i,j View Figure 1 ); long, vertical, plumose hairs sparsely scattered on lateral and dorsal surfaces. Feathery hairs absent. Carapace cuticle dark brown with a cover of white plumose hairs, giving a silvery grey to brown appearance (older specimens can appear browner through loss of hairs, cf. Fig. 3d View Figure 3 ). Chelicerae dark brown. Labium and maxillae dark brown with pale apices. Sternum silvery grey to brown, white hairs numerous to few. Abdomen light silvery grey to brown, anterior dorsum usually with a short, dark brown, mid-dorsal stripe, often delimited by a border of white hairs. Posterior dorsum with variable patterns of light and dark brown chevrons that may be more or less fused posteriorly, often with lateral tufts of white hairs. Lateral abdomen with brown flecks, often coalescing into a pair of irregular, longitudinal, brown stripes bordering the venter. Ventral abdomen with a light to dark grey-brown mid-ventral stripe (often enclosing paler patches) flanked by lateral white-pigmented stripes. Spinnerets brown. Legs light brown with darker brown pigment rings often giving a banded appearance. Caput prominent, gently arched; carapace highest near mid-caput. Eyes in two rows, anterior eye row slightly recurved, posterior eye row straight to moderately procurved; AME or ALE largest, lateral eyes slightly protuberant. Tapeta of indirect eyes canoe-shaped. Clypeus low, less than twice as wide as AME. Chilum divided. Chelicerae robust, with boss, fang with serrate retroventral margin; teeth 2–4 retromarginal, 3–5 promarginal; 4–5 moderately long modified setae on retromargin near fang base, about 15 along promargin, longest apically. Maxillae broad, lateral margins more or less convex. Labium short and wide, bluntly coniform, with short basal notches. Sternum longer than wide with a short extension between coxae IV. Legs 1243 or 1423. 3 tarsal claws, superior claws with 7–13 curved teeth, inferior claws with 2–3 teeth. Claw tufts and scopulae absent. Trochanters shallowly notched. Trichobothria in single row on tarsi and metatarsi, rather short but lengthening distally, 5–6 on tarsus I, 4–5 on metatarsus I, 3–4 in short, subdistal row on tarsus and metatarsus IV; trichobothria present on tibiae and on palpal tarsus and cymbium; bothria collariform, proximal plates with fine, longitudinal ridges. Tarsal organ placed subdistally, distal to trichobothria; capsulate with small pyriform pore ( Fig. 1a–e View Figure 1 ). Male palp. Tibia short and wide, with two retrolateral apophyses: an apical ventrad RTA, ventrally protuberant with a narrow, semi-translucent distal margin; and a basal dorsad RTA, robust and spine-like. Inner surface of basodorsal apophysis with 1–2 small trichobothria. Patella tarsus I; b, tarsus IV. c, trichobothria, metatarsus IV, dorsal. d,e, tarsus I: d, tarsal organ; e, distal bothrium. f, setae, dorsal abdomen. g,h, metatarsus IV and calamistrum, dorsal. i,j, leg setae, ventral: i, metatarsus; j, tarsus.

with retrodorsal bristle; retrodorsal patellar apophysis usually absent or rudimentary but sometimes well developed; spatulate white hairs present or absent. ( Figs. 5e,g,l View Figure 5 , 6f View Figure 6 ). Cymbium broad with short, broadly conical apex. Tegulum broad with a strong, prolaterally directed, basal tegular bulge. Tegulum and conductor ridged and moderately sclerotised. “S”-shaped, falciform conductor arises from the middle and prolateral tegulum; broadest basally, wrapping across the entire width of the bulb; marginally grooved and distally tapering ( Fig. 5d,e,l View Figure 5 ). Embolus an elongate, sinuous rod, tapering near apex; arising prodorsally from the base of the tegulum, it curves ventrad over the tegular bulge and follows an “S” shaped course with the conductor across the ventral bulb. Median apophysis subapical, membraneous and spoon shaped, directed apically to retrolaterally ( Fig. 6d–f View Figure 6 ). Epigynum with a deep, central fossa bounded posteriorly by a sclerotised transverse ridge of variable width; lateral teeth present, just posterolateral to transverse ridge ( Figs. 5c View Figure 5 , 7d View Figure 7 ). Internal genitalia with two wide, membraneous copulatory ducts leading to the paired spermathecae. Each spermatheca consists of a glandulate central lobe connected via a sclerotised interlobar duct of variable length to a small, sometimes indistinct, distal lobe ( Figs. 5f View Figure 5 , 6g View Figure 6 ). Spiracle opening anterior to and half as wide as cribellum ( Fig. 2a,g View Figure 2 ). Tracheal system with four branched tubes confined to the abdomen. Calamistrum in central half to two-thirds of metatarsus IV in females, weakly developed in males. Spinning organs ( Fig. 2a–j View Figure 2 , female). Cribellum entire or divided ( Fig. 2f,j View Figure 2 ); spinning field(s) narrow; divided fields well separated (by about one fifth of a field width); males with non-functional, undivided cribellar plate, about three quarters width of functional cribellum. Spinnerets short; ALS broad, 2-segmented, each bordered anteriorly by a row of thick plumose hairs; PMS shortest, 1-segmented; PLS 2- segmented, as long as ALS with a coniform terminal segment. In female, ALS with 2 adjacent major ampullate gland spigots medially and about 35 piriform spigots; PMS with 1 minor ampullate, 1 cylindrical and 8 aciniform spigots, plus 4 fused paracribellar bases each with 5–14 spigots; PLS with c. 12 spigots, identified as 8 aciniform, 1–2 cylindrical, 1 modified PLS spigot (base truncate apically) with 2 associated paracribellar spigots, the base of the proximal spigot being partially or completely fused with the modified PLS spigot base ( Fig. 2a–e,g–i View Figure 2 ).

Included species. Phryganoporus candidus (L. Koch) , Phryganoporus vandiemeni (Gray) n.comb., P. nigrinus Simon , P. davidleei n.sp., P. melanopygus n.sp.

Notes on synonymies. Phryganoporus Simon is revalidated and removed from its synonymy with Badumna Thorell ( Gray, 1983) . Phryganoporus nigrinus Simon is removed from synonymy with B. candida (L. Koch) . Phryganoporus gausapatus (Simon) is returned to synonymy with P. candida . Phryganoporus g. gausapatus Simon and P. tubicola Simon are retained in synonymy with P. candida (L. Koch) .

Distribution. Australia and Norfolk Island.

Subgroups. Within Phryganoporus , two subgroups are recognised.

P. candidus – vandiemeni: Cribellum divided ( Fig. 2f View Figure 2 ). Basal segments of modified PLS spigot and proximal paracribellar spigot joined proximally ( Fig. 2e View Figure 2 ). Calamistrum occupies central half to three-fifths of metatarsus IV (0.48–0.57:1), retrodorsal glabrous zone narrow, distal end of calamistrum usually delimited by a retrodorsal, subdistal spine ( Fig. 1g View Figure 1 ). Transverse epigynal ridge usually broad, lateral teeth broadly pointed ( Fig. 5c,h,j View Figure 5 ). Interlobar spermathecal ducts short, curving laterally to the distal spermathecal lobe ( Fig. 5f,i,k View Figure 5 ). Male palp with central limb of the S-shaped embolus/conductor with a strong diagonal slope so that the upper section of the “S” is much smaller than the lower section ( Fig. 5d View Figure 5 ).

Patellar apophysis present, vestigial or absent ( Fig. 5e,g,l View Figure 5 ).

P. davidleei – nigrinus – melanopygus: Cribellum entire ( Fig. 2j View Figure 2 ). Basal segments of modified PLS spigot and proximal paracribellar spigot completely fused ( Fig. 2i View Figure 2 ). Calamistrum occupies central two-thirds of metatarsus IV (0.60–0.70:1), retrodorsal glabrous zone wide, subdistal delimiting spine absent ( Fig. 1h View Figure 1 ). Transverse epigynal ridge usually slender ( Figs. 6h View Figure 6 , 7d View Figure 7 ). Interlobar spermathecal ducts elongate, acutely bent (“elbowed”) laterally, doubling back to join the small, rounded distal spermathecal lobes ( Fig. 6g View Figure 6 ). Male palp with the central limb of the S-shaped embolus/conductor horizontally oriented or gently sloping so that the upper section of the “S” is more than half the size of the lower section ( Figs. 6d View Figure 6 , 9a View Figure 9 ). Patellar apophysis absent ( Fig. 6f View Figure 6 ).

Remarks

A previous electrophoretic study of the Badumna candida species group” (Colgan & Gray, 1992) supported the validity of Phryganoporus Simon and the synonymy of P. gausapatus with P. candidus , as well as recognising several more or less distinct genetic populations. Of these, the two most distinct groups proved to be morphologically distinct and are given specific recognition here ( P. davidleei n.sp., P. melanopygus n.sp.). All representatives of B. vandiemeni were consistently associated in analyses as a distinct basal or near basal clade within the main candidus group. Consequently, despite its close morphological similarity to P. candidus , P. vandiemeni is maintained here.

Spinnerets. Simon (1908) recorded the presence of an entire cribellum as a characteristic of his new genus Phryganoporus . This observation may have been based upon the female of P. nigrinus which he described at that time—the divided cribella of his Amaurobius gausapata and P. gausapata occidentalis females perhaps going unnoticed. Main (1971) noted the presence of a divided cribellum in her concept of Ixeuticus candidus . In fact, as demonstrated here, Phryganoporus possesses both entire and divided cribellum states ( Fig. 2f,j View Figure 2 ). These states are correlated with differences in metatarsus IV and calamistrum structure ( Fig. 1g,h View Figure 1 ). Given that the immediate outgroups of Phryganoporus ( Badumna , Forsterina ) have divided cribellum plates, the direction of change here appears to be from the divided to the entire state. Bond & Opell (1997) also record the presence of both cribellar states in the American dictynid genus Mallos . In dictynid spiders, including Mallos , the cribellum is typically entire. However, divided cribella characterise representatives of a derived clade in Mallos , within which two sister species show reversion to the entire state. The polarity of cribellar state changes are of considerable phylogenetic interest. The “entire” state seen in the basal araneomorphs (hypochilids and austrochiloids) is considered plesiomorphic for the group, while the basal groups of the Haplogynae and Entelegynae ( Filistatidae and Eresoidea) show the derived “divided” state ( Griswold et al., 1999). Ramirez (2000) makes the interesting observation that the early instars of the austrochiloid spider Thaida peculiaris have a “bilobate cribellum”. The possibility that this reflects the presence of a divided cribellum state in an ancestral austrochiloid remains open.

The presence of a “modified PLS spigot” and associated paracribellar spigots has been noted in many cribellate entelegyne araneomorphs by Griswold et al. (1999). Griswold (1990) noted that the large modified PLS spigot has a cylindrical but apically tapered base and a cylindrical distal section. In Phryganoporus the base is cylindrical and widely truncate apically, with a distal section that is more or less cylindrical in P. candidus and weakly clavate in P. nigrinus . Additionally the bases of the modified PLS spigot and the proximal paracribellar spigot are basally fused in P. candidus and completely fused in P. nigrinus . The pattern seen in P. candidus is also present in Badumna spp. The fusion of these spigots, as well as their proximity, indicates a close functional relationship.

Relationships. Phryganoporus is most closely associated with the Badumna – Forsterina (“badumnine”) generic group which is characterised by the sinuous (“S or E-shaped”) course of the conductor and embolus across the bulb; and less closely with the Namandia – Matachia (“namandiine”) generic group, which shares with the previous group the “spoon-shaped” membranous median apophysis (or derivative hook-like form or absent), the plate-like (or derivative form) apicoventral RTA and the almost universal presence of epigynal teeth. Morphological trends in Namandia support the derivation of the sinuous embolus/conductor conformation typical of the “badumnine” group from the circular to semicircularfalciform conformation typical of the “namandiine” group. The PMS of genera representing these generic groups differ considerably. In Phryganoporus , Badumna and Forsterina the paracribellar spigot bases are fused while in Namandia , Matachia and Paramatachia they are free, the presumed plesiomorphic condition.

Like Matachia , many New Zealand genera placed in the Desinae of Forster (1970) and the Amphinectidae of Forster & Wilton (1973) can be associated with these generic groups. For example, almost all of Forster’s desine genera, exemplified by Matachia , Goyenia , Panoa and Manawa , can be related to the “namandiine” group on the basis of palpal (and spinneret) conformation. Of the New Zealand amphinectids, Reinga fits within the “badumnine” group near Forsterina (E-shaped embolus with retrolateral origin) while Mamoea also has “badumnine” affinities (S-shaped embolus with prolateral origin). “Amphinectid-desine” associations have also been suggested by Griswold et al. (1999) and Davies & Lambkin (2000).

The placement of Phryganoporus here with the Desidae remains provisional. Badumna (syn. Ixeuticus ) has been variously associated with the Desidae or Desinae by Lehtinen (1967), Forster (1970) and Griswold et al. (1999). The nominal genus, Desis , is a highly derived taxon whose relationships are unclear. Some of the distinctive characters of Desis include the greatly increased size of the spigot field on the posterior median spinnerets, a double row of trichobothria on the tarsi, apically tapered maxillae, absence of a serrula. Others, like the strongly porrect jaws (also seen in “namandiine” groups and more widely) and reduction of leg spines could be simply specialisations related to living and feeding in an intertidal habitat. The general morphology of the male palp is rather similar to the “namandiine” pattern noted above, but the presence of a “secondary tegular process” is an important difference. Davies (1998) has suggested that this could represent the “primary” conductor (sensu Lehtinen, 1967) whose function has been taken over by the larger falciform “secondary” conductor. This raises doubts over the homology of the functional conductors in Desis and the “badumnine” and “namandiine” groups. All of this suggests that these groups cannot be readily associated with Desis or the Desinae.