Liphistius Schiödte, 1849

Peter J. Schwendinger, 2017, A revision of the trapdoor spider genus Liphistius (Mesothelae: Liphistiidae) in peninsular Malaysia; part 1, Revue suisse de Zoologie 124 (2), pp. 391-445 : 393

Liphistius Schiödte, 1849

Type species: Liphistius desultor Schiödte, 1849 , described from female holotype deposited in the Natural History Museum of Denmark (= Zoological Museum of the University of Copenhagen).

Diagnosis and characterisation: See

Platnick

&

Sedgwick (1984); habitus as in Fig. 2 A-B.

Key to the species-groups in peninsular Malaysia:

1 Embolus proper deeply split, distal portion of sclerotised and membranous part disconnected from each other on both sides (on retrodorsal side more widely so than on proventral side); ventrolateral wall of sclerotised part and of membranous part basally running into a short but very wide, mostly dorsad-directed fold (called “dorsal lamella” in Schwendinger, 1990: figs 60-62) ( Fig. 3A, C, E, G ); distal margin of tegulum (corresponding to marginal tegular apophysis of Heptathelinae; see Schwendinger & Ono, 2011) drawn into a more or less distinctly prominent edge; subtegulum with or without apophysis. Poreplate with small CDO and rather small and simple receptacular cluster; mediolateral processes on ventral rim of poreplate present or absent (e.g. Schwendinger, 1990: figs 37-39, 43-45, 47-49, 53-56) .......................................................................................................................................... trang -group

- Embolus proper much less distinctly split, sclerotised and membranous part touching each other on both sides and forming a closed tube; connecting fold mostly long and narrow, indistinct and running prolaterad ( Fig. 3B, D, F, H ) (in L. endau and L. negara sp. nov. bent prodorsad instead of running straight proventrad or prolaterad; Figs 6F, H ; 8F-G); distal margin of tegulum indistinct, never developed as an edge; subtegular apophysis never present. Poreplate with very large quadrangular, pentangular or trapezium-shaped CDO (see Fig. 5 ), or (rarely) with small or medium-sized circular CDO (see Fig. 24F ), or with indistinct slit-like CDO (see Figs 16A , C-D; 22A, C, E, G); receptacular cluster large or medium-sized, always complex, never simple and digitiform; mediolateral processes on ventral rim of poreplate always absent ................................................................................................................ 2

2 Contrategulum with distal edge proventrally ending in a downward-curved, U-shaped or V-shaped row of denticles ( Figs 4C ; 6A, C; 8A, D); para-embolic plate short or absent. Vulval plate strongly sclerotised; CDO large, of various shapes; receptacular cluster large, always undivided ....................................................... malayanus -group

- Distal edge of contrategulum proventrally not ending in a U-shaped or V-shaped row of denticles; para-embolic plate short or long. Vulval plate completely or only partially scerotised; CDO small to large, rounded, angular or slit-like, entire, medially divided by a longitudinal septum ( Fig. 13G ), or completely divided into two paramedian openings ( Figs 13A , C-D; 14I); receptacular cluster entire or divided into two paramedian halves ( Figs 13B , E-F, H; 14J) ...................................................................................................................................................................... 3

3 Contrategulum with more or less distinct ventral process ( Figs 10B , 11F , 14 C-D). Vulval plate completely scerotised; CDO rounded or angular, entire ( Fig. 12A, C, F ), medially divided by a longitudinal septum ( Fig. 13G ), or completely divided into two paramedian openings ( Figs 13A , C-D; 14I); receptacular cluster always divided into two paramedian halves ( Figs 12B, D, G ; 13B, E-F, H; 14J) ........................................................ tioman -group

- Contrategulum always without ventral process. Posterior portion of vulval plate only partially scerotised; CDO and receptacular cluster never divided longitudinally.............................................................................................. 4

4 Para-embolic plate long, its distal margin coarsely serrate ( Figs 15 B-D, 17B-D); dorsal apex of contrategulum narrow and pointed ( Figs 15 E-F, 17G-H); distal margin of cymbium with two indistinct, subequal lobes ( Figs 15B , 17B ); tibial apophysis divided into a long part carrying three megaspines and a short part with only one megaspine, all these megaspines short and bent ( Figs 15A , G-I; 17A, I-K). Vulval plate with posterior stalk reduced to a small sclerite at posterior margin, far apart from poreplate ( Figs 16 A-E, 18A-F) ......................................... linang -group

- Para-embolic plate short, its distal margin smooth; dorsal apex of contrategulum wider, tounge-shaped, with rounded or pointed tip ( Figs 19D , 21 I-L, 23D-G); distal margin of cymbium with prodorsal lobe distinctly longer than proventral one ( Figs 19C , 21H , 23J ); tibial apophysis undivided, its megaspines long or short, curved but not bent. Posterior stalk wide, close to poreplate, not or only partially connected to poreplate ( Figs 20 , 22 , 24 ) .......... .......................................................................................................................................................... batuensis -group

Figures

Fig. 2. Habitus of two Liphistius species from peninsular Malaysia and southern Thailand. (A) Liphistius endau, female from Kota Tinggi (Malaysia), dorsal view. (B) Liphistius indra sp. nov., male paratype from the Sankalakhierie Mountains (Thailand), same view.

Fig. 3. SEM-micrographs of embolus proper of left palpal organ of two males: Liphistius desultor from Penang Hill (A, C, E, G) and Liphistius malayanus from Fraser’s Hill (B, D, F, H). (A) Retroventral-distal view. (B) Dorsal-distal view. (C) Distal view (dorsal side to the right). (D) Distal view (prolateral side to the right). (E) Retrodorsal view. (F) Prodorsal view. (G) Proventral view. (H) Prolateral view. Abbreviations: f - major embolic fold; m - membranous part of embolus proper; s - sclerotised part of embolus proper.

Fig. 4. Liphistius malayanus, details of palp of four males: Templer Park (A, C-D, H-I); holotype of L. malayanus cameroni (B, E, G); Cameron Highlands, ZRC (F, J), Fraser’s Hill, MHNG (K-L). (A) Paracymbium and tibial apophysis of left palp, retroventral view. (B) Tibial apophysis of right palp, ventral view. (C) Distal part of left palpal organ, proventral view (arrow indicating V-shaped row of denticles at proventral end of distal edge of contrategulum). (D, F) Same, retrodorsal and slightly proximal view. (E) Same, retrodorsal view. (G-H) Left palpal organ, distal view (dorsal side up). (I, K) Distal edge of contrategulum of right palp, distal view (dorsal side to the left). (J) Same, distal and slighly prolateral view. (L) Same of left palp, distal and slighly prolateral view (dorsal side to the right). Abbreviations: a - dorsal apex of contrategulum; de - distal edge of contrategulum. Scale lines: 1.0 mm (A; B; C-D; E, G; F; H-L).

Fig. 5. Liphistius malayanus, vulval plate of eight females: paratype of L. m. cameroni from Berinchang, Cameron Highlands (A- B); exuvia of female from Ringlet, Cameron Highlands (C); female (SMF 64093) from Fraser’s Hill (D); female (moulted 24. VI. 2002) from Fraser’s Hill; poreplate slightly deformed by brief immersion in cold KOH (E); female (SMF 56206) from Fraser’s Hill (F); female (SMF 60037) from Genting Highlands (G); female (moulted 8. XII. 2005) from Genting Highlands (H); exuvia of female from Templer Park (I-J). (A, C-I) Dorsal view. (B, J) Ventral view. Scale lines: 1.0 mm (A-B; C, F; D; E, G; H; I-J).

Fig. 6. Liphistius endau, details of left palp of eight males: from type locality, matured 27. II. 2002 (A-F); from type locality, matured 1. II. 2002 (G-I); from unknown locality, died 22. III. 2012 (J); from unknown locality, killed 30. IX. 2010 (K); from Gunung Belumut (L); from Kota Tinggi, matured 22. X. 2014 (M); same place, matured 20. VIII. 2014 (N); same place, matured 6. XI. 2014 (O). (A) Distal part of palp, ventral view. (B) Paracymbium and tibial apophysis, retroventral view. (C) Palpal organ, proventral view (arrow indicating U-shaped row of denticles at proventral end of distal edge of contrategulum). (D) Same, retrodorsal and slightly proximal view. (E) Same, retrodorsal view. (F, H) Same, distal view (dorsal side up). (G) Embolus complex, proventral and slightly distal view. (I) Distal edge of contrategulum, distal and slighly prolateral view (dorsal side to the right). (J-O) Same, distal view. Abbreviations: dw - dorsal wall of sclerotized part of embolus proper; t - tooth on distal edge of contrategulum. Scale lines: 1.0 mm (A-B; C-E, G-H; F, I-O).

Fig. 10. Liphistius johore, female holotype (A); Liphistius panching, male from type locality (B). (A) Dorsal view of vulval plate (redrawn from Platnick & Sedgwick, 1984: fig. 79) (arrows indicating anterolateral invaginations in margin of poreplate). (B) Distal view of left palpal organ (dorsal side up) (redrawn from Sedgwick & Platnick, 1986: fig. 5).

Fig. 11. Liphistius tioman, details of left and right palp of a male (matured 3. I. 2002) from the type locality; left palp (A-C, E-H) and right palp (D, I). (A) Distal part of tibia and base of tarsus, retroventral view. (B) Palpal organ and tips of bristles on cumulus, retrolateral view. (C) Distal part of cymbium and palpal organ, prolateral view. (D) Palpal organ, retrolateral view. (E) Same, retroventral view. (F) Same, distal view (dorsal side at 1 o’clock). (G) Tibial apophysis, ventral view. (H) Same, retroventral view. (I) Same, retrolateral and slightly proximal view. Abbreviations: e - embolus proper; pe - para-embolic plate; rd - retrodorsal part of tibial apophysis; rv - retroventral part of tibial apophysis; vp - ventral process of contrategulum. Scale lines: 1.0 mm (A, G-H; B-E, I; F).

Fig. 12. Liphistius tioman, vulval plate of three females from the type locality: specimen moulted 17. XII. 2001 (A-B); exuvia, moult 7. VI. 2003 (C-E); exuvia, moult 16. II. 2002 (F-G). (A, C, F) Entire structure, dorsal view. (B, D, G) Same, ventral view (vesicles in G omitted). (E) Receptacular cluster, ventral and slightly posterior view. Scale lines: 1.0 mm (A-B; C-G).

Fig. 13. Liphistius panching, vulval plate of four female specimens from the type locality: specimen moulted 16. X. 2002 (A-B), moulted VIII. 2005 (C), built egg case 16. VII. 2001 (D-F), moulted 30. XII. 2005 (G-H). (A, C-D, G) Entire structure, dorsal view. (B, E, H) Same, ventral view. (F) Receptacular cluster, ventral and slightly posterior view. Scale lines 1.0 mm: (A-C, F-H; D-E).

Fig. 14. Liphistius negara sp. nov., details of left palp of male holotype (A-C, E-H) and of male paratype (D); vulval plate of female allotype (I-J). (A) Distal part of palp, ventral view. (B) Distal part of tibia and base of tarsus, retroventral view. (C-D) Palpal organ, distal view (dorsal side up). (E) Same, proventral view. (F) Same, retrodorsal view. (G) Tibial apophysis, retrolateral and slightly proximal view. (H) Distal part of cymbium, prolateral view. (I) Vulval plate, dorsal view. (J) Same, ventral view. Abbreviation: t - tooth on distal edge of contrategulum. Scale lines: 1.0 mm (A; B, G-H; C-D; E-F; I-J).

Fig. 15. Liphistius linang sp. nov., details of left palp of two males; holotype (A-E, G-H), paratype matured 14. IX. 1999 (F, I). (A) Distal part of palp, ventral view. (B) Distal part of cymbium and palpal organ, prolateral view (arrow indicating gap in distal edge of contrategulum). (C) Palpal organ, proventral and slightly proximal view. (D) Same, retrodorsal view. (E-F) Same, distal view (dorsal side up). (G) Paracymbium and tibial apophysis, retrolateral view. (H-I) Tibial apophysis, retrolateral and slightly proximal view. Abbreviations: a - dorsal apex of contrategulum; pe - para-embolic plate. Scale lines: 1.0 mm (A, G; B-F; H-I).

Fig. 16. Liphistius linang sp. nov., vulval plate of three female paratypes (A-E): specimen moulted 30. VII. 1999 (A-B), moulted 10. III. 2000 (C), exuvia, moult 14. IX. 1999 (D-E); undissected genital sternite of allotype (F). (A, C-D) Dorsal view. (B, E, F) Ventral view. Scale lines: 1.0 mm (A-E; F).

Fig. 17. Liphistius indra sp. nov., details of left (A-I) and right (J-K) palp of three males; holotype (A, C-D, G, I), paratype matured 4. X. 2012 (B, F, H, K), paratype matured 24. VIII. 2011 (E, J). (A) Distal part of palp, ventral view. (B) Distal part of cymbium and palpal organ, prolateral view. (C) Palpal organ, retrodorsal view. (D) Same, proventral and slightly proximal view. (E-F) Tegulum, retrodorsal view. (G-H) Palpal organ, distal view (dorsal side up). (I) Paracymbium and tibial apophysis, retrolateral view. (J-K) Tibial apophysis, retrolateral and slightly proximal view. Scale lines: 1.0 mm (A, I; B-F, H; J-K), 0.5 mm (G).

Fig. 19. Liphistius batuensis, details of left palp of two males from type locality: specimen collected 24. VII. 1969 (A-B, F, I), specimen matured 8. II. 2002 (C-E, G-H). (A) Distal part of palp (palpal organ not indicated), ventral view. (B) Paracymbium, retrolateral view. (C) Distal part of cymbium, prolateral view. (D) Palpal organ, distal view (dorsal side up; arrow indicating proventral angle at base of para-embolic plate). (E) Palpal organ, proventral view (arrow indicating same structure as in D). (F) Embolus complex, proventral view. (G) Palpal organ, retroventral and slightly proximal view (arrow indicating same structure as in D-E). (H) Same, retrodorsal view. (I) Same, prolateral view. Abbreviation: T - tegulum. Scale lines: 1.0 mm (A, C; B) and 0.5 mm (D-I).

Fig. 20. Liphistius batuensis, vulval plate of five females from the type locality: exuvia, moult 24. I. 2002 (A-B); specimen SMF 13907 (C); exuvia, moult 12. VII. 2001 (D-E); specimen leg. XI. 1976 (F); exuvia, moult 13. I. 2002 (G-H). (A, C-D, F-G) Dorsal view. (B, E, H) Ventral view. Scale lines: 1.0 mm (A-B, D-E, G-H; C, F).

Fig. 21. Liphistius tempurung, details of left palp of four males: specimen from type locality, matured 13. XI. 2005 (A-F, I); from Gua Cicak, matured 23. I. 2005 (G, J); from type locality, matured 26. IX. 2004 (H, K); from type locality, matured 25. IX. 2004 (L). (A) Cymbium and palpal organ, ventral view. (B) Palpal organ, retrodorsal and slightly distal view. (C) Distal margin of cymbium and palpal organ, prolateral view. (D) Paracymbium, retrolateral view. (E) Distal part of palpal tibia, ventral view. (F) Tibial apophysis, retrodorsal and slightly proximal view. (G) Same, retrolateral and slightly proximal view. (H) Distal part of cymbium, prolateral view. (I-L) Palpal organ, distal view (dorsal side up). Abbreviations: T - tegulum; vp - ventral process of contrategulum. Scale lines: 0.5 mm (A, D-G; B-C, H-L).

Fig. 22. Liphistius tempurung, details of copulatory organ and genital sternite of six females: Gua Cicak, specimen moulted 8. I. 2005 (A- B, N); Gua Tempurung, exuvia, moult 22. I. 2007 (C-D, M); Gua Tempurung, exuvia, moult 9. XII. 2004 (E-F); Gua Tempurung, exuvia, moult 21. X. 2004 (G-H); medium-sized female from Gua Cicak (I-J); small female from Gua Tempurung (K-L). (A, C, E, G, I, K) Vulval plate, dorsal view. (B, D, F, H) Same, ventral view. (J, L) Receptacular cluster, ventral view. (M-N) Posterior margin of genital sternite, ventral view. Scale lines: 1.0 mm (A-D, I-L; E-H; M-N).

Fig. 23. Liphistius priceae sp. nov., details of palp of six males; holotype (A-D, H-K), male paratype collected mature (E), paratype matured 20. VII. 2004 (F), paratype matured 1. VIII. 2004 (G), paratype matured 18. IX. 2004 (L-M) and paratype matured 11. VIII. 2005 (N-O). (A) Left cymbium and palpal organ, ventral view. (B) Distal part of left tibia, ventral view. (C) Left paracymbium, retrolateral view. (D-E) Left palpal organ, distal view (dorsal side up). (F-G) Dorsal apex of contrategulum of right palpal organ, distal view (dorsal side to the left). (H) Distal part of left palpal organ, prolateral view. (I) Same, retrodorsal and slightly distal view. (J) Distal part of cymbium, prolateral view. (K-L, N) Left tibial apophysis, retrolateral and slightly proximal view. (M, O) Right tibial apophysis, same view. Abbreviations: T - tegulum; vp - ventral process of contrategulum. Scale lines: 1.0 mm (A-C, K-O), 0.5 mm (D-I; J).

Fig. 24. Liphistius priceae sp. nov., vulval plate of four female paratypes (all from exuviae): paratype, moult 1. I. 2006 (A-B), allotype, moult 24. XI. 2004 (C-D), paratype, moult 13. XI. 2005 (E), paratype, moult 2. IV. 2005 (F). (A, C, E-F) Dorsal view. (B) Lateral view. (D) Ventral view. Scale lines: 0.5 mm (A-F).