Baalzebub youyiensis Zhao & Li, 2012 (from Guangxi, China).

The generic epithet is named after the French arachnologist Eugène Louis Simon, in recognition of his inception of Theridiosomatidae.

Simonia gen. nov. resembles Sennin Suzuki, Hiramatsu & Tatsuta, 2022 in general shape of copulatory organs. Male palps of these two genera have similar embolic divisions with at least three bristle-like, sharp embolic apophyses; epigynes of both genera have similar spoon-shaped or equicrural triangular scape. Simonia gen. nov. can be distinguished from Sennin by a combination of following characters: cymbial outgrowth (cymbial apophysis) absent (vs present), conductor axe-shaped, almost hyaline and with vein-shaped grains (vs not axe-shaped, membranous, without vein-shaped grains), embolus long, extending to the distal part of embolic division, terminally torch-shaped, with a cylindric stalk and a multiramose apex (vs shorter and apex blunt, located at the proximal part of embolic division, embolic terminal absent), all embolic apophyses are not coiled (vs strongly curved or coiled), embolic division dorsally with large, hyaline lobe (vs embolic lobe absent) (cf. Figs 1–3 and Suzuki et al. 2022: figs 7, 9); epigynal plate surface distinctly wrinkled, with a distinctive transverse fold at midlength between anterior and posterior margins (vs surface slightly wrinkled, medially without the distinctive transverse fold), the anterior part of spermathecae fused (vs not fused, just only overlapped), copulatory duct indistinct, the course of the copulatory duct simple, forming a loop in the inside of copulatory bursa (vs distinct, and course more complex, with a coiled trajectory at the basal side of the spermathecae), copulatory bursa large, nearly as long as epigyne length (vs smaller or indistinct, barely longer than 1/2 length of epigynal plate) (cf. Figs 4E–G, 5C–E, 6C–E and Suzuki et al. 2022: figs 8, 10).

Figure 1. 

Male palp of Simonia youyiensis A prolateral B retrolateral C dorsal D ventral. Abbreviations: Co = conductor; CL = cymbial lobe; Cy = cymbium; ED = embolic division; MA = median apophysis; Pc = paracymbium; SCy = cymbial setae; St = subtegulum; Te = tegulum. Scale bars: 0.2 mm (A–D).

Figure 2. 

Bulb (A) and male palp (B–D) of Simonia youyiensis, cited from Zhang and Lin (2016) A apical B ventral C prolateral D retrolateral. Abbreviations: Co = conductor; CL = cymbial lobe; Cy = cymbium; MA = median apophysis; Pc = paracymbium; SCy = cymbial setae; SD = sperm duct; St = subtegulum; Te = tegulum. Scale bars: 0.2 mm (A–D).

Figure 3. 

Male palpal sclerites of Simonia youyiensis, cited from Zhang and Lin (2016) A bulb, prolateral B embolic division, ventral C conductor, ventral D cymbium, retrolateral. Abbreviations: Co = conductor; CL = cymbial lobe; EA = embolic apophysis; ED = embolic division; EL = embolic lobe; ET = embolic terminal; MA= median apophysis; Pc = paracymbium; SCy = cymbial setae; St = subtegulum; Te = tegulum. Scale bars: 0.1 mm (A–D).

Figure 4. 

Simonia youyiensis, male habitus (A, B), female habitus (C, D) and epigyne (E–G) A dorsal B ventral C dorsal D ventral E intact, ventral F cleared, ventral (blue dashed line showing the anterior margin of the epigynal plate) G cleared, dorsal (green dashed line showing schematic course of copulatory duct). Abbreviations: CB = copulatory bursa; CD = copulatory duct; FD = fertilization duct; Sc = scape; SH = spermathecal head; SS = spermathecal stalk; Sp = spermatheca; TF = transverse fold of epigynal plate. Scale bars: 0.5 mm (A–D); 0.2 mm (E–G).

Small sized with body length 1.50–1.65 in males and 1.90–2.45 in females; carapace 0.83–0.96 long in males and 0.91–1.08 in females. Carapace nearly pyriform, in profile highest just behind ocular area, gradually sloping to pedicel, c. 1.35–1.45 times longer than high; carapace smooth, with long, sparse setae, yellow brown to dark brown, slightly darker anteriorly; cervical groove V-shaped, radial grooves and fovea indistinguishable. Sternum yellowish brown to dark, distinctly darker than carapace, clothed with dense setae, heart-shaped, anterior edge truncate, anterior and lateral margins with brown extensions fitting intercoxal concavities; posterior region strongly protruding between coxae IV. Female palp distally with erect, thin, dark bristles. Chelicerae slightly darker than carapace. Labium triangle shaped. Maxillae nearly trapezoidal, anterior edge straight, anterior and lateral margins slightly curved, slightly convergent posteriorly, with dense scopulae on inner margins. Legs long, uniformly coloured, slightly lighter than carapace, with darker femora and coxae I. Leg formula 1243. Abdomen ovoid, nearly as long as wide, abdominal colours and patterns variable; marginally clothed with sparse long setae, ventrally covered by fine short setae. Spinnerets brown. Male palp: same as in type species. Epigyne with small, hyaline scape (SC); epigynal plate surface distinctly wrinkled, with distinctive transverse fold (TF) at midlength between anterior and posterior margins; scape (SC) extending from posterior margin of epigynal plate, translucent; copulatory openings indistinct; copulatory duct (CD) indistinct, with simple course, forming loop in inside of copulatory bursa; spermathecae (Sp) consist of relatively large head (SS, anterior part) and slightly narrower stalk (SS, posterior part), and in addition with distinctly small base (SB, distal part) in S. sumatra sp. nov.; spermathecal heads fused, located centrally and juxtaposed; fertilization ducts (FD) acicular, membranous, located on basal surface of spermathecae; copulatory bursae (CB) represented spherical or oval sacs, large, nearly as long as epigyne length, surface hyaline, wrinkled and ribbed, bursae touching each other.

Simonia youyiensis (Zhao & Li, 2012) from Laos, Vietnam and Guangxi in China, S. steineri sp. nov. from Laos, and S. sumatra sp. nov. from Sumatra.

A preliminary genus-level taxonomic molecular analysis of Theridiosomatidae from Southeast Asia was carried, based on five targeted genes (two mitochondrial genes: 16S and COI; three nuclear genes: 18S, 28S, and H3). According to the results (unpublished): (1) the monophyly of the genus Simonia gen. nov. is supported; (2) this new genus is related to two genera exclusively distributed in SE Asia, Karstia and Sennin. Morphologically, the new genus is also similar to Karstia, but can be distinguished by the absence of cymbial outgrowth, presence of torch-shaped embolic terminal and large, hyaline embolic lobe, and by the fused anterior parts of the spermathecae, large copulatory bursa, as well as a set of other characters of the copulatory organs (see diagnosis of the genus above and key to theridiosomatid genera endemic to Oriental Realm below).

The type species of Simonia gen. nov., S. youyiensis was originally assigned to the Baalzebub, although it did not show typical Baalzebub features. Baalzebub is definitely not monophyletic. There is a strong possibility that Baalzebub sensu stricto contains only two species from the Neotropical Realm: B. baubo and B. albonotatus (Petrunkevitch, 1930). These Neotropical Baalzebub species share the following distinctive suite of characters, here contrasted with the corresponding condition in Simonia gen. nov.: 1) embolic apophyses spatulate, thick and blunt (vs bristle-like and slender, apically sharp); 2) embolus claw-shaped, not branched (vs torch-shaped, with a cylindric stalk and a multiramose apex); 3) embolic lobe absent (vs present); 4) conductor not axe-shaped, surface smooth (vs axe-shaped, surface with many vein-shaped grains); 5) scape large, at least longer than 1/2 length of epigynal plate (vs small, no more than 1/4 length of epigynal plate); 6) epigynal plate surface smooth (distinctly wrinkled, with a distinctive transverse fold); and 7) copulatory bursa surface smooth, small, less than 1/2 length of epigynal plate (surface wrinkled and ribbed, large, as long as length of epigyne) (cf. Coddington 1986: figs 161–164, 183, 184, 186, 187 and Figs 1–3, 4E–G, 5C–E, 6C–E). In view of the above-mentioned facts, it is currently impossible to discern any obvious derived features that could indicate a close relationship between S. youyiensis and the genus Baalzebub sensu stricto, leaving no doubts that our new combination and the establishment of a new genus are correct.

Holotype ♀ and paratypes 3♀ (IZCAS), China: Guangxi, Pingxiang City, Youyi Town, Bantou Vill., Niuyan Cave, 22°05.666'N, 106°45.439'E, 251 m, 18.I.2011, Z. Chen and Z. Zha leg. Examined.

China: 1♂ 12♀ (IZCAS), Guangxi Prov., Baise City, Pingguo Co., Liming Vill., Ganmoyan Cave, 23°48.330'N, 107°31.526'E, 22.IX.2015, J. Wu and Z. Chen leg.; 1♂ 7♀ (NHMSU), Guangxi Prov., Hechi City, Donglan Co., Sanshi Town, Gongping Vill., a nameless cave, 24°21.347'N, 107°23.190'E, 368 m, 9.II.2015, Y. Li and Z. Chen leg. Laos: 3♂ 17♀ (NHMSU), Bolikhamsai Prov., Lak Sao City, Transiten, Lang Cave, 18°23.318'N, 104°32.675'E, 318 m, VIII.2012, P. Jäger leg. Vietnam: 1♂ 3♀ (IZCAS), Phú Tho Prov., Thanh Son Dist., Xuan Son Commune, Xuan Son National Park, Lang Cave, 21°06.600'N, 104°57.600'E, 375 m, 26.X.2012, H. Zhao and Z. Chen leg.

Females of S. youyiensis are most similar to those of S. sumatra sp. nov. by having similar habitus and general shape of the vulva. Simonia youyiensis differs from S. sumatra sp. nov. in 1) anterior margin of the epigynal plate slightly concaved (c. 140°) (vs concaved c. 120°) (cf. Fig. 4E, F and Fig. 6C, D), 2) spermathecae comma-shaped (vs spermathecae shaped like the whole hind leg of a frog) (cf. Fig. 4G and Fig. 6E), and 3) bursae spherical (vs egg-shaped) (cf. Fig. 4G and Fig. 6E).

Male (Fig. 4A, B): Carapace brown, slightly darker in ocular region, without distinct pattern; cervical groove and radial grooves indistinct. AER distinctly recurved, PER slightly procurved. Sternum yellowish brown. Mouthparts reddish brown. Legs uniformly yellowish. Abdomen round, dorsum basically light brown, with 3 pairs of black diagonal bands on sides, forming 3 V-shaped stripes; venter centrally with black trapezoidal speckle, marginally with several discontinuous streaks. Measurements: Total length 1.62. Carapace 0.83 long, 0.81 wide. Clypeus 0.21 high. Sternum 0.52 long, 0.49 wide. Abdomen 1.14 long, 1.08 wide. Length of legs: I 3.32 (1.01, 0.39, 0.88, 0.63, 0.41); II 2.51 (0.81, 0.32, 0.58, 0.51, 0.29); III 1.84 (0.52, 0.21, 0.38, 0.41, 0.32); IV 2.22 (0.49, 0.32, 0.52, 0.58, 0.31).

Palp (Figs 1–3): Femur unmodified. Patella round and small, nearly as long as tibia, dorsally bears strong and long macroseta slightly longer than tibia. Tibia cup-shaped, about 1/6–1/5 length of cymbium, with several sparse setae. Cymbium (Cy) broad, navicular, c. 2.1 × longer than wide, dorsally clothed with dense setae, basolaterally with lobe (CL) and row of longitudinally arranged setae (SCy) along suture between cymbium and lobe. Cymbial lobe represented by small and semi-transparent sheet, just like outline of thumb. Paracymbium (Pc) L-shaped, about 1/5–1/4 length of cymbium, with sharp and spine-like tip. Tegulum (Te) capacious and inflated, c. 1.5 × longer than wide, surface with countless delicate texture; sperm duct (SD) indistinct and sinuate. Subtegulum (ST) located prolaterally to tegulum, represented by large and surface smooth tubercle, slightly longer than 1/2 length of tegulum. Median apophysis (MA) small, about 2/5 length of tegulum, shaped like tadpole, consisting of relatively wide head (distal part) and narrow tail (proximal part); distal process blunt, apex round and rough, with many tiny and scale-like tooth; proximal process triangular, gradually narrowing toward its apex, apex sharp. Conductor (Co) large and axe-shaped, aligned transversely on anterior part of bulb; almost all of conductor hyaline and with vein-shaped grains, except membranous distal margin. Embolic division nearly as long as conductor, hidden behind conductor, consisting of broad embolic lobe (EL), embolic terminal (ET), and at least 3 apophyses (EA); embolic terminal short, less than 1/2 length of embolic lobe, shaped like a torch, with cylindric stalk and multiramose apex; all embolic apophyses bristle-like and slender, slightly longer than embolic terminal; some apexes of embolic terminal and embolic apophyses overpass distal margin of conductor.

Female. Somatic features as in Fig. 4C, D and coloration distinctly lighter than in male. Measurements: Total length 2.40. Carapace 0.91 long, 0.90 wide. Clypeus 0.20 high. Sternum 0.53 long, 0.51 wide. Abdomen 1.56 long, 1.27 wide. Length of legs: I 3.81 (1.31, 0.42, 0.78, 0.89, 0.41); II 3.17 (0.87, 0.43, 0.81, 0.68, 0.38); III 2.03 (0.63, 0.28, 0.37, 0.44, 0.31); IV 2.32 (0.62, 0.31, 0.49, 0.57, 0.33).

Epigyne (Fig. 4E–G). Epigynal plate large, c. 1.4 × wider than long, anteriorly slightly concaved by c. 140°, arrangement of various parts of vulva indistinctly visible through tegument. Scape (Sc) T-shaped, small, about 1/5 length of epigynal plate. Spermathecae (Sp) comma-shaped, with bean-shaped head (SH) and slightly curved stalk (SS); anterior surface of spermathecal heads touching anterior margin of epigynal plate; spermathecal stalks extending obliquely, separated by about 2–3× diameters of stalks. Copulatory bursae (CB) spherical.

China (Guangxi), Laos, Vietnam (Fig. 7).

Holotype ♀ (IZCAS) and paratype 1♀ (IZCAS), Laos: Hua phan Pro., Guesthouse cave, 20°24.176'N, 104°13.818'E 13.I.2008, H. Steiner leg.; paratype 1♀ (NHMSU), Hua phan Pro., Tham Kuong Tai, 20°27.470'N, 104°09.850'E, 12.I.2014, H. Steiner leg.

The specific name is a patronym after Mr. Helmut Steiner (Hessen, Germany), collector of the type series.

The females of S. steineri sp. nov. can be easily distinguished from other congeners by the following characters: 1) dorsum of abdomen uniformly coloured (vs with pattern) (cf. Fig. 5A and Figs 4C, 6A); 2) spermathecae separated from anterior margin of epigynal plate by more than 1.5 × diameters of spermathecal head (vs nearly touching anterior margin) (cf. Fig. 5E and Figs 4G, 6E); and 3) spermathecae hammer-shaped, spermathecal heads globular, spermathecal stalks straight (vs spermathecae not hammer-shaped, spermathecal heads bean-shaped, spermathecal stalks slightly curved) (cf. Fig. 5E and Figs 4G, 6E).

Figure 5. 

Holotype female of Simonia steineri sp. nov., habitus (A, B) and epigyne (C–E) A dorsal B ventral C intact, ventral D cleared, ventral (blue dashed line showing the anterior margin of the epigynal plate) E cleared, dorsal (green dashed line showing schematic course of copulatory duct). Abbreviations: CB = copulatory bursa; CD = copulatory duct; FD = fertilization duct; Sc = scape; SH = spermathecal head; SS = spermathecal stalk; Sp = spermatheca; TF = transverse fold of epigynal plate. Scale bars: 0.5 mm (A, B); 0.2 mm (C–E).

Figure 6. 

Holotype female of Simonia sumatra sp. nov., habitus (A, B) and epigyne (C–E) A dorsal B ventral C intact, ventral D cleared, ventral (blue dashed line showing the anterior margin of the epigynal plate) E cleared, dorsal (green dashed line showing schematic course of copulatory duct). Abbreviations: CB = copulatory bursa; CD = copulatory duct; FD = fertilization duct; SB = spermathecal base; Sc = scape; SH = spermathecal head; SS = spermathecal stalk; Sp = spermatheca; TF = transverse fold of epigynal plate. Scale bars: 0.5 mm (A, B); 0.2 mm (C–E).

Female (Fig. 5A, B): Carapace, uniformly brown except a V-shaped dark brownish band along cervical groove; cervical groove and radial grooves indistinct. AER distinctly recurved, PER almost straight in dorsal view. Sternum deep brown. Mouthparts proximally reddish brown, distally light brown, inner margins white. Legs uniformly yellowish. Abdomen round, marginally clothed with sparse long setae, uniformly yellowish white, without any markings or distinct pattern. Measurements: Total length 2.12. Carapace 1.03 long, 0.86 wide. Clypeus 0.21 high. Sternum 0.55 long, 0.52 wide. Abdomen 1.46 long, 1.37 wide. Length of legs: I 3.12 (0.98, 0.32, 0.93, 0.58, 0.31); II 3.03 (0.92, 0.28, 0.81, 0.69, 0.33); III 2.42 (0.73, 0.27, 0.58, 0.63, 0.21); IV 2.93 (0.89, 0.32, 0.77, 0.64, 0.31).

Epigyne (Fig. 5E–G). Epigynal plate c. 1.2 × wider than long, anteriorly concaved by c. 140°, copulatory bursae distinctly visible through integument. Scape (Sc) shaped like an acute triangle, relatively large, about 1/4 length of epigynal plate. Spermathecae (Sp) hammer-shaped, with globular-shaped heads (SH) and columnar stalk (SS); anterior surface of spermathecae separated from anterior margin of epigynal plate by more than 1.5 × diameters of spermathecal head; spermathecal stalks straight, extending obliquely, separated by about 3–4× diameters of stalks. Bursae (CB) subglobular.

Known only from the type locality, Guesthouse cave, Huapan Province, Laos (Fig. 7).

Figure 7. 

Distribution records of Simonia gen. nov. species.

Holotype ♀ and paratype 1♀ (IZCAS), Indonesia: West Sumatra Prov., Kab Agam Co., Kecamatan Matur district, 0°15.739'S, 100°18.491'E, 01.XII.2013, H. Zhao leg. Paratypes 1♀ and 1juv. (NHMSU), West Sumatra Prov., Kab Agam Co., Simarasok Vill., Cua Luwuang, 0°14.902'S, 100°28.993'E, 710 m, 11.I.2014, H. Zhao leg.

The species name is derived from the type locality; noun in apposition.

The female of the new species resembles those of S. youyiensis in having similar habitus and general appearance of the epigyne (cf. Fig. 6 and Fig. 4C–G), but can be easily distinguished by 1) anterior margin of epigynal plate concaved c. 120° (vs c. 140°) (cf. Fig. 6C, D and Fig. 4E, F), 2) spermathecae shaped like the whole hind leg of a frog, consisting of head, stalk and base (vs comma-shaped, only with head and stalk) (cf. Fig. 6E and Fig. 4G), and 3) bursae egg-shaped (vs spherical) (cf. Fig. 6E and Fig. 4G).

Female (Fig. 6A, B): Carapace brownish posteriorly, distinctly darker anteriorly and marginally, with distinct pattern on pars cephalica consisting of pair of dark lateral bands and Ψ-shaped markings behind posterior eyes, markings starting from behind PME and PLE almost reaching dark fovea. AER distinctly recurved, PER distinctly recurved in dorsal view. Sternum uniformly black. Mouthparts coloured as sternum. Legs uniformly yellowish white. Abdomen round, covered with sparse long setae; dorsum basically black, centrally with 2 pairs of muscular depressions, anteriorly with pair of large, nearly fan-shaped patches, posteriorly with pair of √-shaped band; venter slightly darker than dorsum, without distinct pattern. Measurements: Total length 2.28. Carapace 0.96 long, 0.88 wide. Clypeus 0.22 high. Sternum 0.48 long, 0.50 wide. Abdomen 1.56 long, 1.28 wide. Length of legs: I 2.71 (1.02, 0.33, 0.57, 0.51, 0.28); II 2.42 (0.83, 0.27, 0.52, 0.49, 0.31); III 1.74 (0.48, 0.21, 0.41, 0.41, 0.23); IV 2.29 (0.91, 0.27, 0.42, 0.41, 0.28).

Epigyne (Fig. 6E–G). Epigynal plate c. 1.2 × wider than long, anteriorly concaved by c. 120°. Scape (Sc) spoon-shaped, small, about 1/6 length of epigynal plate. Spermathecae (Sp) shaped like the whole hind leg of a frog, with bean-shaped head (SH), conical stalk (SS) and distinctly narrowed base (SB); anterior surface of spermathecae touching anterior margin of epigynal plate; spermathecal stalks and bases extending obliquely, widely separated by c. 1/2 width of epigynal plate. Bursae (CB) egg-shaped.

Known only from the type locality, West Sumatra, Indonesia (Fig. 7).