Psednocnemis, West, Rick C., Nunn, Steven C. & Hogg, Stephen, 2012

West, Rick C., Nunn, Steven C. & Hogg, Stephen, 2012, A new tarantula genus, Psednocnemis, from West Malaysia (Araneae: Theraphosidae), with cladistic analyses and biogeography of Selenocosmiinae Simon 1889, Zootaxa 3299, pp. 1-43 : 4-7

Lyra :

5. Female lyra : (0) rudimentary ( Fig. 47 ) or absent; (1) oval with proximally truncate form ( Fig. 49a ); (2) almost circular (Fig. 52a); (3) reinform ( Fig. 51a ); (4) 1–3 rows of heavy clavates (not reniform) (Fig. 50a). Length = 6, CI = 0.66, RI = 0.86. In those selenocosmiine species that show loss of prolateral maxillary lyra , several issues arise, namely this trait (loss of lyra ) is the singular defining feature of Yamia ( Schmidt & Haupt 2004, Raven 2005, Zhu & Zhang 2008). Yet, many undescribed Orphnaecus species (West & Nunn, pers. obs.) and some Australian species (R. Raven, pers. comm. 2008) also show losses of maxillary lyra . Secondly (and most critical to character state coding), Phlogiellus species show wide ranges of lyrate bacillae counts, but no distinct division between presence/absence (some species such as P. baeri show 1–3 bacillate setae). Therefore, instead of using the ambiguous state “absent”, due to overly subjective character delineation combined with false multi-state character coding ( Sereno 2007), it becomes logical to include the rudimentary condition within the same transformational character state (0). This resolves those subjective character state tendencies, as noted by Jenner (2004), Sereno (2007, 2009), Assis (2009) and Vogt et al. (2010) which can threaten the value of morphological characters. This was so critical to Jenner (2004) and Sereno (2009), they stated that such artificial/weak character state delineation can “ threaten the very future of morphology based cladistics as a vigorous discipline ” (surely a strong enough argument as to support our reasoning). Other genera display unique maxillary lyrate arrangements, Selenocosmia have an almost circular patch of very short bacillae (Fig. 52a), some Orphnaecus possess a reniform patch ( Fig. 51a ), while Chilobrachys lyra comprises 1–3 rows of heavy claviform bacillae (Fig. 50a).

6. Setal fringe ventrally to prolateral maxillary suture: (0) absent; (1) fine pallid setae ( Figs 49a , 50a); (2) scopulate, dark setae ( Fig. 47 ). Length = 3, CI = 0.66, RI = 0.8. The genera Poecilotheria , Haplocosmia and some Chilobrachys possess a setal fringe just ventral to the prolateral maxillary suture, but dorsal to the lyra . In Poecilotheria this fringe is dark, almost scopulate ( Fig. 47 ), while in Haplocosmia ( Fig. 49a ) and Chilobrachys (Fig. 50a) the fringe is formed from fine, pallid setae.

7. Tubercles on prolateral maxillary surface: (0) absent; (1) present ( Fig. 47 ). Length = 1, CI = 1, RI = 0. All Poecilotheria possess 1–12 tubercles immediately distad to patch of rudimentary bacillae on prolateral maxillary surfaces ( Fig. 47 ). These tubercles are not known in any other selenocosmiine genus.

Sexual morphology (female): Pérez-Miles et al. (1996: characters 13, 14) used spermathecal morphology in their cladistic analysis of Theraphosinae . Guadanucci (2005: characters 11, 12) also used spermathecal morphology in his analysis of Ischnocolinae. Pérez-Miles et al. (2008: characters 8–11) used spermathecal morphology to define their new theraphosine genus Ami Pérez-Miles 2008 ; West et al. (2008: characters 23, 24) used spermathecal morphology in their analysis of Theraphosidae . Five characters were used based on spermathecal morphology, all of which (with the exception of character 8, states 0 and 1 = West et al. 2008: character 23, states 2 and 0) are new.

8. Spermathecae: (0) fused ( Figs 28 b, 30); (1) not fused ( Fig. 31 b). Length = 2, CI = 0.5, RI = 0.83. Within Selenocosmiinae examined, both fused ( Smith 2004: fig. 6; 2006: fig. 6) and unfused ( Kulczyński 1908: fig. 1; Schmidt 1999: fig. 5; Haupt & Schmidt 2004: figs 7–9; Zhu & Zhang 2008: figs 4 I, 6 I, 8 I, 9 F; West & Nunn 2010a: figs 6, 23, 51, 70, 101; 2010b: figs 17, 37, 84, 141, 160, 181, 200, 201) spermathecae were noted.

9. Lateral bilobe spermathecal morphology: (0) apically swollen ( Fig. 38 b); (1) not apically swollen ( Fig. 6 ). Length = 2, CI = 0.5, RI = 0.75. In all Psednocnemis ( West & Nunn 2010b, figs 37, 64, 141; herein, ( Figs 6 , 40 ) as well as L. robustus ( Smith 1988; West 1991; West & Nunn 2010a, fig. 101), lateral bilobes lack apical swelling, differing from all other selenocosmiine species with bilobed spermathecal morphology.

10. Bilobed spermathecae with medial and lateral lobes of similar length ( Fig. 43 ): (0) absent ( Figs 38 b, 45); (1) present ( Figs 40, 43, 44 b). Length = 2, CI = 0.5, RI = 0.91.

11. Spermathecal lobes with mushroom cap ends (mushroomiform): (0) absent; (1) present. Length = 2, CI = 0.5, RI = 0.8. All Selenocosmia within this analysis ( Figs 44 b, 45), L. fuscus and L. lessunda ( West & Nunn 2010a: figs 51, 70), respectively, herein: ( Fig. 38 b) possess at least one lobe with mushroom cap (mushroomiform) ends.

12. Spermathecae tombstone-shaped ( Fig. 36 ): (0) absent; (1) present. Length = 1, CI = 1, RI = 0. This trait was noted in an undescribed Orphnaecus species used in our analysis as well as Orphnaecus philippinus ( Schmidt 1999: fig. 5).

Sexual morphology (male): Male palpal bulb characters are well established synapomorphies of and within Theraphosinae ( Raven 1985, Pérez-Miles et al. 1996, Bertani 2000). Bertani (2000) identified homologous keels within Theraphosinae , we follow him in exploring embolus morphology in Selenocosmiinae , utilizing six characters, herein.

13. Terete embolus lanceolate (LTE) ( Figs 41, 44 a): (0) absent ( Figs 28 a, 31a, 32, 34, 35a, 37a, 38a); (1) present ( Figs 39, 41, 44 a). Length = 1, CI = 1, RI = 0.

14. Single retrolateral keel on embolus (R): (0) absent; (1) strong and heavy ( Figs 34, 35 a); (2) weak ( Figs 37 a, 38a). Length = 2, CI = 1, RI = 0. In all Phlogiellus ( Fig. 34 ) and Orphnaecus ( Fig. 35 a) examined, a strong and heavy single keel (similar position to R, or retrolateral keel of Bertani 2000) runs along the entire embolus, twisting 90 degrees distally. Lyrognathus (except L. robustus ) also possess a single retrolateral (RR) keel ( West & Nunn 2010a: figs 5, 40 and, herein, Figs 37 a, 38a), although it is far less pronounced.

15. Basal embolus with distinct heavy retrolateral lobe (BL): (0) absent; (1) present ( Figs 34, 35 a). Length = 1, CI = 1, RI = 0. In all Phlogiellus ( Zhu & Zhang 2008: figs 8 B, 9 I and, herein, ( Fig. 34 ) and Orphnaecus ( Fig. 35 a), retrolateral embolus keel terminates basally as an enlarged lobe (BL). A similar lobe is present in Lyrognathus (RBL) ( Figs 37 a, 38a), but is far smaller, not so distinct. Additionally, an apophysis is present in Haplocosmia himalayana , however, this structure originates from the tegulum, not from the embolus and thus not applicable to this character.

16. Embolus with distal (A) and sub-distal (SA) keels ( Figs 31 a, 32): (0) absent; (1) present. Length = 1, CI = 1, RI = 0. All Chilobrachys and Haplocosmia used in this analysis ( Figs 31 a, 32, respectively) possess both A and SA keels.

17. Distal spiral curl on embolus (AC): (0) absent; (1) present ( Fig. 39 ). Length = 1, CI = 1, RI = 0. In all known male Psednocnemis (herein, Fig. 39 ; West & Nunn 2010b: figs 58, 116, 135), the embolus terminates distally with distinct distal “spiral curl” (AC).

18. Embolus with multiple distal keels: (0) present; (1) absent. Length = 1, CI = 1, RI = 0. Among all species used in this analysis, those with more than one keel (or what could be interpreted as a keel) were scored with the state “present”. The outgroup taxon Reichlingia annae ( Reichling 1997: fig. 1) possesses a ventral crest (VC) and prolateral inferior (PI) keel. In Poecilotheria ( Smith 2004: fig. 13, 14; Smith 2006: figs 13, 14; herein, Fig. 28 a) three keels are distinct, one reverse twisting retrolateral (RCR) keel, as well as prolateral superior (PS) and prolateral inferior (PI) keels, corkscrewing along the entire embolus (in the opposite direction to other selenocosmiine species with embolus keels). Chilobrachys displays prolateral superior (PS) and inferior (PI) keels, sub-distal (SA) and distal (A) keels ( Fig. 31 a) and in Haplocosmia , male embolus is flattened, twisted, with strong distal (A), sub-distal (SA), and retrolateral (R) keels ( Fig. 32 ).

Legs:

19. Scopula division on tar. III: (0) divided; (1) undivided. Length = 2, CI = 0.5, RI = 0.89. The conditions “divided/undivided tarsal scopula” have been used extensively throughout history of classification of Mygalomorphae but, herein, is applied only to tar. III. Raven discussed tarsal scopula division in regard to “Anterior/Posterior Gradations” ( Raven 1985); Guadanucci (2005) followed from Raven, both concluded a dependency for tarsal scopula division with most developed scopulation in anterior legs, grading to weaker conditions in hind legs. Furthermore, Guadanucci (2005) concluded tarsal scopula division is not dependant upon size and is useful in phylogenetic analysis (contra Pérez-Miles 1994).

20. Scopula division on met. IV: (0) divided; (1) undivided. Length = 1, CI = 1, RI = 0. West & Nunn (2010b, figs 19, 69, 74), following Pocock (1895a; 1900a) used undivided met. IV scopula to diagnose most Lyrognathus species. Gallon (2003) considered undivided met. IV scopula apomorphic for Eumenophorinae. Guadanucci (2005) concluded, in his work on tarsal scopula division, that dependency of tarsal scopula condition between legs means the more entire scopulate condition, the more apomorphic states it will show. This would also apply to metatarsal scopula.

21. Scopula on met. IV retrolaterally stronger: (0) absent; (1) present. Length = 1, CI = 1, RI = 0. West & Nunn (2010b) first noted the presence of scopula on retrolateral side of division on met IV as almost entire, with prolateral side only extending 1/3 to 2/3 basally in Coremiocnemis (group 1) species ( West & Nunn 2010b: figs 21, 87, 163, 206).

22. Met. IV scopula divided by strong diagonal line: (0) absent; (1) present. Length = 1, CI = 1, RI = 0. Both Coremiocnemis kotacana and the Coremiocnemis sp. from Borneo, used within our data matrix, possess met IV scopula divided and separated distally by strong dividing lines; there is no gradual diminishing of scopula ( West & Nunn 2010b: figs 165, 206).

23. Scopula on tarsi laterally extended: (0) absent ( Figs 29, 35 c, 38c, 42,); (1) present ( Fig. 46 ). Length = 1, CI = 1, RI = 0. Following West et al. (2008: character 9, state 3). Raven (1985: table 3, character 17) utilized a similar character in his cladistic analysis of Theraphosidae subfamilies, “ tarsi broader than metatarsi: absent/ present ”, a trait he found synapomorphic for Aviculariinae . This character is noted in all Poecilotheria considered for analyses.

Leg morphology:

24. Width of Tib. IV in relation to Tib. I: (0) similar to, or smaller than ( Fig. 17 ); (1) incrassate, much thicker than ( Fig. 38 c). Length = 1, CI = 1, RI = 0. Pocock (1895a; 1900a) used incrassate tib. IV to diagnose Lyrognathus . Smith (1988), West (1991) and West & Nunn (2010a: fig. 38c) followed Pocock (1895a). Pérez-Miles et al. (1996: character 16) used this character in their analysis of Theraphosinae .

25. Leg spines on tibia: (0) present; (1) absent. Length = 1, CI = 1, RI = 0. Raven (1985) and West et al. (2008: character 11) discussed leg spination in Barychelidae and considered reductions in leg spination to be a derived condition in Theraphosidae .

26. Number of clavate trichobothria on tar. IV: (0) under 25; (1) 25 and over. Length = 2, CI = 0.5, RI = 0.91. Raven (1985: table 1, character 37) considered reduction/loss of tarsal trichobothria a synapomorphy for Atypoidina.

27. Position of clavates on tar. IV: (0) proximal 2/3 to 3/ 4 in straight line; (1) proximal 1/2 or less in straight line; (2) In proximal 1/ 3 in "U" shape. Length = 3, CI = 0.66, RI = 0.91. Raven (1985) discussed variation of leg trichobothria in Mygalomorphae and used those variations in his systematic revision of the infraorder. Among Selenocosmiinae examined, several clavate trichobothrial conditions were noted. Additionally, Guadanucci (in prep.) noted the clavate arrangement in Poecilotheria (Guadanucci, in prep.: fig. 167) as our state (2).

28. Brush of penicillate setae on retrolateral tib. IV ( Fig. 38 c): (0) absent; (1) present. Length = 1, CI = 1, RI = 0. West & Nunn (2010a) used the presence of penicillate setae along retrolateral surfaces of tib. IV to diagnose all Lyrognathus ( West & Nunn 2010a: figs 8, 20, 53, 71, 98). We introduce this character for analysis for the first time.

29. Brush of recurved setae along retrolateral met. IV ( Fig. 42 ): (0) absent; (1) present. Length = 1, CI = 1, RI = 0. West & Nunn noted in Coremiocnemis (group 1) the presence of recurved setae in brushform along retrolateral surfaces of met. IV ( West & Nunn 2010b: figs 21, 87, 165, 206). The description of Psednocnemis gen. nov. indicates this characteristic is diagnostic to Coremiocnemis (except C. tropix Raven 2005 ) which is tested here for the first time.

30. Leg width formula IV, III, I, II: (0) absent; (1) present. Length = 2, CI = 0.5, RI = 0.83.

Chelicerae:

31. Intercheliceral setal fringe proximodorsally: (0) absent ( Fig. 14 ); (1) present ( Fig. 49 b). Length = 1, CI = 1, RI = 0. In Haplocosmia , a distinct fringe of long pallid setae is present, intermixed with intercheliceral pegs. This fringe has not been seen outside Haplocosmia , and is considered here for the first time.

32. Intercheliceral pegs with prolateral keeled ridge ( Fig. 49 b): (0) absent; (1) present. Length = 1, CI = 1, RI = 0. In Haplocosmia , and only within that selenocosmiine genus, intercheliceral pegs possess a distinct prolateral keel. As in character 31, this trait is tested for the first time.

33. Spine cluster on retrolateral proximomedial chelicerae ( Fig. 15 ): (0) absent; (1) present. Length = 2, CI = 0.5, RI = 0.94. West & Nunn first noted retrolateral cheliceral spines in Lyrognathus ( West & Nunn 2010a: figs 26, 60, 80, 96), Coremiocnemis ( West & Nunn 2010b: figs 9, 47, 69, 107, 170, 189, 215) and in Selenocosmia ( West & Nunn 2010a). The same feature is present in Psednocnemis ( Fig. 15 ).

Figures

FIGURES 46 – 48. Poecilotheria species. 46. P. metallica Pocock 1899, female, in life, Nallamala Range, Pradesh, India, dorsal view. 47. P. f o r m o s a Pocock 1899, holotype female, NHM 1898.10.10.1 – 2, right maxillae (inverted), prolateral view. 48. P. metallica, female, QM S 88462, left strikers, retrolateral view. Photos: 46 R. West; 47 D. Court; 48 S. Nunn.

FIGURES 49 a – 50 b. 49 a, b. Haplocosmia himalayana (Pocock 1899), holotype female, NHM 1899.8. 19.3. 49 a. Right maxillae (inverted), prolateral view. 49 b. Right chelicerae, prolateral view. 50 a, b. Chilobrachys sericeus (Thorell 1895), syntype male, NHM 1895.9. 21.21. 50 a. Left maxillae, prolateral view. 50 b. Left strikers, retrolateral view. Photos: R. Raven.

FIGURES 51 a – 52 b. 51 a, b. Orphnaecus species, female, QM S 83783. 51 a. left maxillae, prolateral view. 51 b. Orphnaecus species, female, QM S 83783, strikers, retrolateral view. 52 a. Selenocosmia javanensis (Walckenaer 1837), female, QM S 83754, left maxillae, prolateral view. 52 b. S. javanensis female QM S 83754, strikers, retrolateral view. Photos: S. Nunn.

FIGURES 28 – 36. 28 a – b: a. Poecilotheria tigrinawesseli Smith 2006, male, embolus, retrolateral view. b. P. tigrinawesseli, female, spermatheca, dorsal view. 29. Phlogiellus watasei (Kishida 1920), female, dorsal habitus. 30. Chilobrachys fimbriatus Pocock 1899, holotype female, NHM 1899.11. 2.22, spermatheca, dorsal view. 31 a – b: a. Chilobrachys andersoni (Pocock 1895), paratype male, NHM 1895.9. 21.21, embolus, retrolateral view. b. Chilobrachys andersoni, holotype female, NHM 1891.10. 15.6, spermatheca, dorsal view. 32. Haplocosmia nepalensis Schmidt & von Wirth 1996, male, QM S 83827, embolus, retrolateral view. 33. Phlogiellus atriceps Pocock 1897, paratype female, NHM 1897.11. 1.6 – 7, spermatheca, dorsal view. 34. Phlogiellus baeri (Simon 1877), male, QM S 83789, embolus, retrolateral view. 35 a – c: a. Orphnaecus pellitus Simon 1892, syntype male, MNHN AR 4678, embolus, retrolateral view. b. Orphnaecus pellitus, paratype female, MNHN AR 4678, spermatheca, dorsal view. c. Orphnaecus pellitus, female, dorsal habitus. 36. Orphnaecus philippinus (Schmidt 1999), female, QM S 83833, spermatheca, dorsal view. Abbreviations: A—apical keel, BL—basal lobe, PI—prolateral inferior keel, PS—prolateral superior keel, Rretrolateral keel, RCR—reverse corkscrew retrolateral keel, SA—sub-apical keel. Illustrations: S. Nunn.

FIGURES 37 – 45. 37 a – b: a. Lyrognathus crotalus Pocock 1895, male, QM S 88361, embolus, retrolateral view. b. Lyrognathus crotalus, female, QM S 88362, spermatheca, dorsal view. 38 a – c: a. Lyrognathus lessunda West & Nunn 2010, paratype male, QM S 74066, embolus, retrolateral view. b. Lyrognathus lessunda, holotype female, QM S 74067, spermatheca, dorsal view. c. Lyrognathus lessunda, holotype female, QM S 74067, habitus, dorsal view. 39. Psednocnemis davidgohi sp. nov., paratype male, QM S 88022, embolus, retrolateral view. 40. Psednocnemis jeremyhuffi (West & Nunn 2010), paratype female, QM S 88071, spermatheca, dorsal view. 41. Coremiocnemis obscura West & Nunn 2010, male, QM S 88360, embolus, retrolateral view. 42. Coremiocnemis hoggi West & Nunn 2010, paratype female, QM S 74074, dorsal habitus. 43. Coremiocnemis kotacana West & Nunn 2010, holotype female, QM S 83759, spermatheca, dorsal view. 44 a – b: a. Selenocosmia javanensis (Walckenaer 1837), male, NHMW 173 embolus, retrolateral view. b. Selenocosmia javanensis, female, QM S 88355, spermatheca, dorsal view. 45. Selenocosmia species (A), female, RMBR ZRC. ARA. 02, spermatheca, dorsal view. Abbreviations: AC—apical spiral curl, LTE—lanceolate terete embolus, RBL—reduced basal lobe, RR—reduced retrolateral keel, TE—terete embolus. Illustrations: S. Nunn.

FIGURES 2 – 7. Psednocnemis davidgohi sp. nov., holotype female, QM S 88303. 2. Carapace, dorsal view. 3. Eyes, dorsal view. 4. Labium, ventral view. 5. Sternum, labium, maxilla and coxae, ventral view. 6. Spermatheca, ventral view. 7. Abdomen, dorsal view. Scale bar = 10 mm for 2, 5 and 7; 5 mm for 4; 2 mm for 3; 1 mm for 6. Photos: S. Nunn.

FIGURES 14 – 20. Psednocnemis davidgohi sp. nov., holotype female, QM S 88303. 14. Left intercheliceral pegs, prolateral view. 15. Left proximomedial retrolateral cheliceral spine cluster. 16. Close up of intercheliceral pegs. 17. Left patella and tibia IV, dorsal view. 18. Left metatarsus and tarsus IV, prolateral view. 19. Left metatarsus and tarsus IV, retrolateral view. 20. Left metatarsus and tarsus IV, ventral view. Scale bar = 10 mm for 17 – 19; 5 mm for 20; 2 mm for 14 and 15; 1 mm for 16. Photos: S. Nunn.