Physocyclus Simon, 1893

Nolasco, Garduño & Valdez-Mondragón, Alejandro, 2022, Four new species of the spider genus Physocyclus Simon, 1893 (Araneae: Pholcidae) from Mexico, with updated taxonomic identification keys, European Journal of Taxonomy 813, pp. 173-206 : 175-185

Genus Physocyclus Simon, 1893

Type species

Pholcus globosus Taczanowski, 1874 .

Diagnosis

See Valdez-Mondragón (2010, 2013, 2014).

Distribution

The genus Physocyclus is native and endemic in North and Central America ( Valdez-Mondragón 2010), with the exception of P. globosus , which has a cosmopolitan distribution due to human activities (Valdez- Mondragón 2010, 2013, 2014). Huber & Villarreal (2020) mentioned that Caporiacco’s (1955) records of P. dugesi Simon, 1893 from Miranda (1 ♀) and Caracas (1 juv.) are dubious and presumably based on specimens of P. globosus or a species of Priscula Simon, 1893 . The natural distribution of Physocylus is mainly in arid and dry ecosystems ( Figs 9–10, 13 , 76, 77 ), such as xerophilous scrubs or deserts, although some species occur in zones with temperate and subtropical climates, such as deciduous forest ( Figs 8, 12 , 70, 72 ). The specific habitat of these spiders is between big boulders, in tree bark, or inside any hollow that provides protection ( Figs 11 , 71 ). In karstic zones, due to their trogophilic habits, they are commonly found inside caves, on the walls or between fissures ( Figs 11 , 73 , 75 ) ( Valdez-Mondragón 2010).

Composition

The genus Physocyclus is composed of two species groups: globosus and dugesi . The globosus group includes 15 species: P. bicornis Gertsch, 1971 , P. gertschi Valdez-Mondragón, 2010 , P. globosus , P. guanacaste Huber, 1998 , P. huacana Valdez-Mondragón, 2010 , P. lautus Gertsch, 1971 , P. mariachi sp. nov., P. modestus Gertsch, 1971 , P. montanoi Valdez-Mondragón, 2010 , P. paredesi Valdez- Mondragón, 2010, P. pocamadre sp. nov., P. sarae Valdez-Mondragón, 2010 , P. sikuapu sp. nov., P. validus Gertsch, 1971 , and P. xerophilus Nolasco & Valdez-Mondragón, 2020 .

The dugesi group includes 22 species: P. brevicornus Valdez-Mondragón, 2010 , P. californicus Chamberlin & Gertsch, 1929 , P. cornutus Banks, 1898 , P. darwini , P. dugesi , P. enaulus Crosby, 1926 , P. franckei Valdez-Mondragón, 2010 , P. hoogstraali Gertsch & Davis, 1942 , P. lyncis sp. nov., P. marialuisae Valdez-Mondragón, 2010 , P. merus Gertsch, 1971 , P. mexicanus Banks, 1898 , P. michoacanus , P. mysticus Chamberlin, 1924 , P. palmarus Jiménez & Palacios-Cardel, 2013 , P. pedregosus Gertsch, 1971 , P. peribanensis Valdez-Mondragón, 2014 , P. platnicki Valdez- Mondragón, 2010, P. reddelli , P. rothi Valdez-Mondragón, 2010 , P. sprousei Valdez-Mondragón, 2010 , and P. tanneri Chamberlin, 1921 .

The identification key of species of Physocyclus Simon, 1893 is updated from Valdez-Mondragón 2010 (hereafter VM 2010), using the same abbreviations.

Identification keys

Males

1. Sclerotized cones on frontal lamina of chelicerae present (VM 2010: figs 15, 22) .......................... 2

– Sclerotized cones on frontal lamina of chelicerae absent (VM 2010: figs 63, 77) ......................... 22

2. Few sclerotized cones (<10) on each frontal lamina (VM 2010: figs 42, 49) .................................. 3

– Numerous sclerotized cones (>10) on each frontal lamina (VM 2010: figs 105, 119) ..................... 5

3. Procursus pointing to front of palp, with basal half dark brown and distal half dark, with thin and long spine distally; sclerites on bulb, with oval shape in prolateral view ( Nolasco & Valdez-Mondragón 2020: figs 6–8) .................................................... P. xerophilus Nolasco & Valdez-Mondragón, 2020

– Procursus pointing to base of palp (VM 2010: figs 44, 51) .............................................................. 4

4. Embolic sclerites half-moon shaped in retrolateral view; without projection directed to base of embolus; procursus with long ventral notch and ventral protuberance on median part (VM 2010: figs 44–45)…… ............................................................................... P. globosus (Taczanowski, 1874)

– Embolic sclerites oval-shaped in retrolateral view; with conical projection directed to base of embolus; procursus with short ventral notch and without ventral protuberance on median part (VM 2010: figs 51–52)…………….. ....................................................... P. guanacaste Huber, 1998

5. Palp femur with a ventral conical apophysis in the middle (VM 2010: figs 86, 121) ...................... 6

– Palp femur without ventral conical apophysis in the middle (VM 2010: figs 128, 135) .................. 8

6. Palp femur thin, with long conical apophyses; embolus short, curved and thin; lateral apophyses of chelicerae absent; chelicerae with big cones on basal and prolateral parts of frontal lamina, except on anterior half of region with upside-down U-shape (VM 2010: figs 84–86) .................................. ............................................................................................................. P. mysticus Chamberlin, 1924

– Palp femur wide, with small conical apophyses; embolus large and wide; lateral apophyses of chelicerae present; chelicerae with small cones scattered (VM 2010: figs 98–100) ........................ 7

7. Chelicerae with large and curved lateral apophyses, without oval protuberance on basal part of frontal lamina; sclerotized cones on prolateral part of frontal lamina and toward prolateral part of lateral apophyses of chelicerae; procursus with short and thin apical spine; embolic sclerites small, oval apically; embolus rounded in dorsal part (VM 2010: figs 98–100) ..... P. reddelli Gertsch, 1971

– Chelicerae with small and trapezoidal lateral apophyses, with oval protuberance on basal part of frontal lamina; sclerotized cones along frontal lamina and on frontal protuberance of chelicerae; procursus with large and wide apical spine; embolic sclerites large, triangular apically; embolus straight and triangular (VM 2010: figs 119–121) .................... P. franckei Valdez-Mondragón, 2010

8. Embolus pointing in perpendicular position to longitudinal axis of palp femur (VM 2010: figs 10, 135) ................................................................................................................................................... 9

– Embolus pointing to base of palp femur (VM 2010: figs 31, 107) ................................................. 14

9. In retrolateral view, palp with inconspicuous notch between embolic sclerites and embolus; embolus partially covering embolic sclerites; dorsal protuberance present on bulb; chelicerae with lateral apophyses slightly curved; sclerotized cones of chelicerae on half basal, on prolateral part of frontal lamina, and prolateral part of lateral apophyses (VM 2010: figs 133–136) ....................................... ............................................................................................ P. brevicornus Valdez-Mondragón, 2010

– In retrolateral view, palp with conspicuous notch between embolic sclerites and embolus; embolus not covering embolic sclerites; dorsal protuberance absent on bulb; chelicerae with lateral apophyses straight; sclerotized cones of chelicerae in other position (VM 2010: figs 8, 10, 22, 24) .............. 10

10. Embolic sclerites thin; circular notch between embolic sclerites and embolus; embolus with circular dorsal part; sclerotized cones of chelicerae on prolateral part of frontal lamina and prolateral part of lateral apophyses (VM 2010: figs 8–11) ...................... P. californicus Chamberlin & Gertsch, 1929

– Embolic sclerites wide; oval notch between embolic sclerites and embolus; embolus with curved dorsal part; sclerotized cones of chelicerae scattered (VM 2010: figs 24–25, 178–179) ................11

11. Chelicerae with sclerotized cones on basal half of frontal lamina and few scattered cones on distal part; lateral apophyses of chelicerae long; procursus pale in basal half and dark in distal half, with a short apical spine; embolus wide, oval basally ( Jiménez & Palacios-Cardiel 2013: figs 5–8) .......... .................................................................................. P. palmarus Jiménez & Palacios-Cardiel, 2013

– Chelicerae with sclerotized cones on ¾ of total length of frontal lamina and prolateral part of lateral apophyses; lateral apophyses of chelicerae short; procursus totally dark, with a long apical spine; embolus with other shape (VM 2010: figs 22, 24, 176, 178) ......................................................... 12

12. Chelicerae without sclerotized cones on small central region of frontal lamina; in retrolateral view, procursus with long ventral notch, starting at level of dorsal apophyses of procursus (VM 2010: figs 176–178) ............................................................................ P. darwini Valdez-Mondragón, 2010

– Chelicerae with sclerotized cones totally covering ¾ of length of frontal lamina; in retrolateral view, procursus with small ventral notch at middle (VM 2010: figs 22) ................................................. 13

13. Chelicerae with long cones on frontal lamina; procursus shorter than in P. peribanenis ; distal spine slightly wider than in P. peribanensis ; in retrolateral view, procursus with conspicuous notch between embolic sclerites and embolus; embolus wide and straight (VM 2010: figs 22– 24) .................................................................................................................... P. dugesi Simon, 1893

– Cheliceraewithsmallconesonfrontallamina;procursuslongerthanin P.dugesi ;withdistalspinethinner than in P.dugesi ; procursus with inconspicuous notch between embolic sclerites and embolus; embolus wide and sinuous ( Valdez-Mondragón 2014: figs 6, 8) .... P. peribanensis Valdez-Mondragón, 2014

14. Embolus with distal concavity, ending in a sharp tip (VM 2010: figs 31–32) ............................... 15

– Embolus without distal concavity, ending in a rounded or curved tip (VM 2010: figs 17–18) ...... 19

15. Embolus with dorsal part slightly curved, almost straight, and with small distal ventral concavity (VM 2010: figs 72–73) ................................................................................................................... 16

– Embolus with dorsal part strongly curved, distal ventral concavity large (VM 2010: figs 163– 164) ................................................................................................................................................. 17

16. In retrolateral view, lateral apophyses of chelicerae short and conical, in frontal view without protruding laterally of chelicerae; embolus straight and long (VM 2010: figs 29–31) ...................... ...................................................................................................................... P. enaulus Crosby, 1926

– In retrolateral view, lateral apophyses of chelicerae wide and rounded apically, in frontal view without protruding laterally of chelicerae; embolus straight and shorter than in P. enaulus and P. sprousei (VM 2010: figs 70–72) .................................................................................. P. merus Gertsch, 1971

– In retrolateral view, lateral apophyses of chelicerae strongly wider than in P. enaulus and P. merus , in frontal view protruding laterally of chelicerae; embolus long and sinuous (VM 2010: figs 168, 170) .......................................................................................... P. sprousei Valdez-Mondragón, 2010

17. Sclerotized cones of chelicerae scattered along frontal lamina; embolic sclerites partially covered by embolus; in dorsal view, embolic sclerites sigmoid; lateral apophyses of chelicerae absent; chelicerae with basal-frontal protuberance; palp femur with ventral-distal conical apophysis, rounded apically (VM 2010: figs 161–163) .................................................. P. marialuisae Valdez-Mondragón, 2010

– Sclerotized cones of chelicerae not scattered along frontal lamina; embolic sclerites not covered by embolus; in dorsal view, embolic sclerites straight; lateral apophyses of chelicerae present; chelicerae without basal-frontal protuberance; palp femur without ventral-distal conical apophysis (VM 2010: fig. 105) ........................................................................................................................ 18

18. Embolic sclerites short; embolus with claw-shape apically, with deep distal-ventral concavity; lateral apophyses of chelicerae short and straight, ending in rounded tip; sclerotized cones of chelicerae on prolateral part and basal half of frontal lamina (VM 2010: figs 105–108) ......................................... ................................................................................................................ P. tanneri Chamberlin, 1921

– Embolic sclerites long, spine-shaped; embolus wide, with shallow distal ventral concavity; lateral apophyses of chelicerae long and straight, ending in a sharp tip; sclerotized cones of chelicerae on prolateral part of frontal lamina, first basal ⅓ of chelicerae, and on prolateral part of lateral apophyses (VM 2010: figs 126–129) ............................... P. michoacanus Valdez-Mondragón, 2010

19. In retrolateral view, embolus apically ending in a rounded or curved shape; procursus thin and short, with inconspicuous distal spine; sclerotized cones of chelicerae scattered along frontal lamina (VM 2010: figs 15, 17, 183, 185) ................................................................................................... 20

– In retrolateral view, embolus apically ending in a rounded shape in ventral part, and dorsal part ending in a sharp tip; procursus wide and long, with conspicuous distal spine; sclerotized cones of chelicerae on prolateral-basal part of frontal lamina and prolateral part of lateral apophyses (VM 2010: figs 56, 58, 91, 93) ....................................................................................................... 21

20. Embolus apically ending in a rounded shape; embolic sclerites long and wider than in P. rothi , without distal notch dorsally; dorsal apophysis of procursus shorter than in P. rothi ; sclerotized cones of chelicerae along of frontal lamina and on prolateral part of lateral apophyses; lateral apophyses of chelicerae wide and long, in lateral view conical and curved (VM 2010: figs 15–17) ...................... ...................................................................................................................... P. cornutus Banks, 1898

– Embolus apically ending in a curved shape; embolic sclerites long and thinner than in P. cornutus , with small distal notch dorsally; dorsal apophysis of procursus larger than in P. cornutus ; sclerotized cones distributed until ⅔ part of length of frontal lamina and on lateral apophyses; lateral apophyses of chelicerae wide and short, in lateral view conical, ending in two tips with different sizes (VM 2010: figs 183–185) ................................................................................. P. rothi Valdez-Mondragón, 2010

21. Embolic sclerites long and wide, rounded apically; embolus without apical notch; procursus with apical spine thinner than in P. hoogstraali ; sclerotized cones on basal half, prolateral part of frontal lamina and prolateral part of lateral apophyses; lateral apophyses of chelicerae shorter than in P. hoogstraali , projected at almost same length of chelicerae (VM 2010: figs 91–94) .......................... ................................................................................................................ P. pedregosus Gertsch, 1971

– Embolic sclerites long and thin, spine-shaped apically; embolus with apical notch; procursus with apical spine wider than in P. pedregosus ; sclerotized cones on basal and prolateral part of frontal lamina of chelicerae but missing on lateral apophyses; lateral apophyses of chelicerae longer than in P. pedregosus , projected at same length of chelicerae (VM 2010: figs 56–59) .................................. ................................................................................................. P. hoogstraali Gertsch & Davis, 1942

22. Two lateral apophyses in each chelicera ( Figs 17–19 ) ................................................................... 23

– One single apophyses in each chelicera (VM 2010: figs 77–78) .................................................... 27

23. Chelicerae with both lateral apophyses close to each other, in parallel position, pointing downwards; lateral apophyses of chelicerae with 3-4 sclerotized cones ( Figs 17–19 ) ....................................... 24

– Chelicerae with lateral apophyses pointing in perpendicular position, forming different angles; lateral apophyses of chelicerae without sclerotized cones (VM 2010: figs 1–2) ............................ 25

24. In frontal view, longest lateral apophyses of chelicerae wide and dark, with stridulatory ridges on it, without sclerotized cones; frontal-distal apophyses of chelicerae conical, with 3-4 sclerotized cones; procursus straight, with ventral notch at median part, with a long spine distally; in retrolateral view, embolic sclerites short; in dorsal view, embolus thin basally and becoming wider distally and in retrolateral view ending in a rounded tip ( Figs 17–19 , 21–22 , 24–27 ) .............. P. mariachi sp. nov.

– In frontal view, longest lateral apophyses of chelicerae short and pale, with 3-4 apical sclerotized cones; frontal-distal apophyses of chelicerae conical, without sclerotized cones; procursus curved, with ventral notch at basal part, without distal spine; in retrolateral view, embolic sclerites long and wider than in P. mariachi sp. nov.; in dorsal view, embolus straight and in retrolateral view slightly curved, ending in “J” shape (VM 2010: figs 197, 200) .......... P. platnicki Valdez-Mondragón, 2010

25. Chelicerae with basal-lateral apophyses short, ending in small rounded tip; chelicerae with distalfrontal conical apophyses, sclerotized, protruding from the distal margin of chelicerae; procursus with long and thin distal spine; embolic sclerites rounded dorsally; embolus square in retrolateral view, with a small spine sub-distally on dorsal view (VM 2010: figs 63–66) ....... P. lautus Gertschi, 1971

– Chelicerae with paired lateral apophyses, conical, ending in sclerotized and sharp tips; chelicerae with small distal-frontal conical apophyses or lateral-distal apophyses; procursus with short distal spine or long and wide; embolic sclerites curved or square-shaped dorsally (VM 2010: figs 1–3; Valdez-Mondragón 2014: figs 23–25) ............................................................................................ 26

26. In retrolateral view, chelicerae with one lateral wide apophysis and the other small and triangular, forming a 90° angle between them; procursus conical, with rounded ventral notch in retrolateral view; embolic sclerites wide and square; embolus without distal spine (VM 2010: figs 1–4) .......... .................................................................................................................... P. bicornis Gertschi, 1971

– In retrolateral view, chelicerae with one lateral apophysis wide and ax-shaped and the other thin and conical, forming a <90° angle between them; procursus straight, with oval ventral notch in retrolateral view; embolic sclerites short, slightly oval; embolus with a wide distal spine (Valdez- Mondragón 2014: figs 21, 23–26) ........................................... P. paredesi Valdez-Mondragón, 2010

27. Stridulatory ridges occupying full length of lateral apophyses of chelicerae (VM 2010: figs 112– 113) ................................................................................................................................................. 28

– Stridulatory ridges occupying part of total length of lateral apophyses of chelicerae (VM 2010: figs 140–141) .................................................................................................................................. 32

28. Procursus wide and straight, ending in a short spine ( Fig. 44 ; VM 2010: fig. 79) ......................... 29

– Procursus thin and conical, ending in a long spine (VM 2010: figs 156, 192) ............................... 30

29. Procursus wider in distal half than basal half, with curved projection ventrally; embolic sclerites with square notch in median part; embolus ending in a rounded small ventral projection, without ventral projections and without small spine dorsal-apically but with a conspicuous triangular projection in dorsal view (VM 2010: figs 79–80) .......................................................... P. modestus Gertsch, 1971

– Procursus wider in basal half than distal half, without curved projection ventrally; embolic sclerites square-shaped and without square notch in median part; embolus ending in square shape, with a small ventral projection pointing to base of femur and with small spine dorsal-apically ( Figs 44–47 , 50–51 ) ................................................................................................................... P. sikuapu sp. nov.

30. Palp femur straight ventrally; lateral apophysis of chelicerae wide, shield-shaped; embolic sclerites slightly curved in retrolateral view; embolus wide, with apical wide and curved concavity (VM 2010: figs 190–192) ................................................................................ P. sarae Valdez-Mondragon, 2010

– Palp femur curved ventrally (VM 2010: fig. 114) .......................................................................... 31

31. Palp femur thinner than in P. montanoi ; lateral apophysis of chelicerae wide and triangular-shaped in frontal view, with half length of chelicerae; embolic sclerites short, not protruding embolus; embolus long and inverted S-shaped (VM 2010: figs 112–114) ....................................................................... ...................................................................................................................... P. validus Gertsch, 1971

– Palp femur wider than in P. validus ; lateral apophysis of chelicerae wide and shield-shaped in frontal view, almost with same length as chelicerae; embolic sclerites long, with an S-shape, protruding embolus; embolus small and square (VM 2010: figs 154–156) ......................................................... ................................................................................................ P. montanoi Valdez-Mondragón, 2010

32. In lateral view, chelicerae with lateral-basal apophysis projected toward frontal part, and thin lateraldistal apophysis, ending in serrated tip; procursus curved, with curved ventral notch and a rounded distal-dorsal projection; embolic sclerites wide and conspicuous, with small rounded distal notch; in retrolateral view, embolus wider than in P. huacana , without notch between bulb and embolus (VM 2010: figs 140–143) ......................................................... P. gertschi Valdez-Mondragón, 2010

– In lateral view, chelicerae with only one lateral apophysis wide and square-shaped; procursus conical, with ventral notch “V”-shaped; embolic sclerites conical and slightly curved, without small rounded distal notch; in retrolateral view, embolus thinner than in P. gertschi , with rounded notch between bulb and embolus (VM 2010: figs 147–150) ........................................................................ ................................................................................................. P. huacana Valdez-Mondragón, 2010

Females

1. Epigynum in ventral view, bell-shaped, with lateral constrictions strongly marked in median part, with anterior half smaller than posterior half (VM 2010: figs 19, 26) ........................................................... 2

– Epigynum in ventral view, with other shape, without constrictions in median part, or if present, barely visible (VM 2010: figs 60, 67) .............................................................................................................. 19

2. Ventral apophyses of epigynum long and conical or long and flat (VM 2010: figs 19, 21, 130, 132). 3

– Ventral apophyses of epigynum small, with other shape (VM 2010: figs 95, 97, 137, 139) .............. 15

3. Pore plates long and thin (VM 2010: figs 39–41, 75) ............................................................................ 4

– Pore plates with other shape (VM 2010: figs 20, 27) ............................................................................. 9

4. Epigynum with light ventral region, rounded, close to epigastric furrow (VM 2010: figs 35, 74) ...... 5

– Epigynum with light ventral region, triangular, close to epigastric furrow (VM 2010: figs 109, 130) 8

5. Epigynum with ventral apophyses long or flat ( Fig. 61 ; VM 2010: fig. 74) ......................................... 6

– Epigynum with ventral apophyses short, conical and rounded distally (VM 2010: figs 33–38) .......... 7

6. Epigynum with ventral apophyses long and flat, rounded distally; epigynum rounded in lateral view; wide light region in median part triangular; pore plates wide and oval ( Figs 61–63 ) ............................. ......................................................................................................................................... P. lyncis sp. nov.

– Epigynum with ventral apophyses thin and sharp apically, slightly curved in lateral view; epigynum slightly curved in lateral view; wide light region in median part circular; pore plates thin and long (VM 2010: figs 74–76) ....................................................................................... P. merus Gertsch, 1971

7. Epigynum with ventral apophyses rounded apically, with upside-down chair shape (type I), or with ventral apophyses long and conical, ending in a rounded tip (type II); in frontal view, anterior rounded protuberances on ventral apophysis absent; pore plates shorter than in P. sprousei (VM 2010: figs 33– 41) ...................................................................................................................... P. enaulus Crosby, 1926

– Epigynum with ventral apophyses wide and rounded basally, ending in a conical tip; in frontal view, anterior rounded protuberances on each ventral apophysis; pore plates longer than in P. enaulus (VM 2010: figs 172–175) ............................................................. P. sprousei Valdez-Mondragón, 2010

8. Epigynum with ventral apophyses straight in ventral view, short and conical in lateral view; epigynum with light ventral region close to epigastric furrow shorter than in P. michoacanus ; pore plates short and thin, straight, slightly wider in posterior part (VM 2010: figs 109–111) .......................................... ...................................................................................................................... P. tanneri Chamberlin, 1921

– Epigynum with ventral apophyses wide and curved in ventral view, wide and curved in lateral view; epigynum with light ventral region close to epigastric furrow longer than in P. tanneri ; pore plates long, slightly curved, wider in anterior part (VM 2010: figs 130–132) ............................................................ ............................................................................................... P. michoacanus Valdez-Mondragón, 2010

9. Epigynum with ventral apophyses wide in basal part, with small tips (VM 2010: figs 26, 180) ....... 10

– Epigynum with ventral apophyses long and conical, with wide tips (VM 2010: figs 19, 165) .......... 11

10. Epigynum with tips of ventral apophyses separated from each other, tips longer than in P. darwini ; pore plates thin, curved, “V”-shaped; light region close to epigastric furrow smaller than in P. darwini (VM 2010: figs 26–28) ......................................................................................... P. dugesi Simon, 1893

– Epigynum with tips of ventral apophyses very close to each other, tips smaller than in P. dugesi ; pore plates wide and straight; light region close to epigastric furrow longer than in P. dugesi (VM 2010: figs 180–182) .................................................................................. P. darwini Valdez-Mondragón, 2010

11. Epigynum with ventral and conical paired projections on posterior margin, ventral apophyses conical and slightly curved, separated by a notch on anterior margin; pore plates wide and curved (VM 2010: figs 165–167) ........................................................................... P. marialuisae Valdez-Mondragón, 2010

– Epigynum without ventral and conical paired projections on posterior margin (VM 2010: fig. 19) 12

12. Epigynum with ventral apophyses thin and long, with a sharp tip; epigynum with wide and light region on median part; epigastric furrow triangular-shaped; pore plates with half-circle shape ( Jiménez & Palacios-Cardiel 2013: figs 10–13) ............................... P. palmarus Jiménez & Palacios-Cardel, 2013

– Epigynum with ventral apophyses wide and long, with a rounded tip (VM 2010: figs 19–21) ......... 13

13. Epigynum with ventral apophyses close to each other; epigynum with thin and triangular light region in median part; pore plates wide and oval, slightly curved, close to each other (VM 2010: figs 19– 20) ...................................................................................................................... P. cornutus Banks, 1898

– Epigynum with ventral apophyses widely separated from each other; epigynum with wide light region in median part (VM 2010: figs 187, 189) ............................................................................................. 14

14. Epigynum with ventral apophyses pointing downward; light region in median part with rhomboid shape; pore plates thinner than in P. rothi , forming a 90° angle ( Valdez-Mondragón, 2014: figs 15– 17) ......................................................................................... P. peribaniensis Valdez-Mondragón, 2014

– Epigynum with ventral apophyses pointing toward front; light region in median part oval; pore plates wider than in P. peribanensis , forming a>90° angle (VM 2010: figs 187–189) .................................... .............................................................................................................. P. rothi Valdez-Mondragón, 2010

15. Chelicerae with stridulatory ridges; epigynum with ventral apophyses wide and flat; pore plates wide, oval-shaped (VM 2010: figs 12–14, 137–139) .................................................................................... 16

– Chelicerae without stridulatory ridges; epigynum with ventral apophyses small and conical; pore plates with other shape (VM 2010: figs 95–97, 123–125) ............................................................................. 17

16. Epigynum with ventral apophyses flat and rhomboid-shaped, with porosities; epigastric furrow forming a>90° angle; pore plates wide and oval, without a sclerotized region in anterior margin (VM 2010: figs 12–14) ......................................................................... P. californicus Chamberlin & Gertsch, 1929

– Epigynum with ventral apophyses rounded, without porosities; epigastric furrow curved; pore plates wide and oval, with a sclerotized region in anterior margin (VM 2010: figs 137–139) ......................... .................................................................................................. P. brevicornus Valdez-Mondragón, 2010

17. Anterior part of epigynum circular; ventral apophyses of epigynum inconspicuous; epigynum with dark central region forming upside-down “T” shape; pore plates small and triangular (VM 2010: figs 123–125) ................................................................................. P. franckei Valdez-Mondragón, 2010

– Anterior part of epigynum square; ventral apophyses of epigynum conspicuous; epigynum without dark central region with upside-down “T” shape; pore plates elongated (VM 2010 figs 95–97) ...... 18

18. Ventral apophyses of epigynum close to each other, smaller than in P. reddelli ; epigynum with conspicuous triangular dark region in median part; pore plates large and oval, with a small constriction in middle, without sclerotized region in anterior margin (VM 2010: figs 95–97) .................................. ..................................................................................................................... P. pedregosus Gertschi, 1971

– Ventral apophyses of epigynum separated from each other, longer than in P. pedregosus ; epigynum without triangular dark region in median part; pore plates markedly curved at posterior part, with sclerotized region in anterior margin (VM 2010: figs 102–104) .................... P. reddelli Gertschi, 1971

19. Epigynum longer than wide (VM 2010: figs 60, 88) ........................................................................... 20

– Epigynum wider than long (VM 2010: figs 46, 53) ............................................................................. 22

20. Epigynum with two ventral apophyses on anterior margin, small and cylindrical; a third large and conical apophysis located on basal ⅓ part, ending in a rounded tip; pore plates markedly thin and long, almost in parallel position to each other (VM 2010: figs 88–90) ........... P. mysticus Chamberlin, 1924

– Epigynum without two ventral apophyses on anterior margin; epigynum without sharp and conical apophyses, and without a third large and conical apophysis ventrally (VM 2010: fig. 60; Figs 67– 68 ) .......................................................................................................................................................... 21

21. Epigynum with two anterior, circular, wide apophyses, short and circular in ventral view; two ventral protuberances, flat and curved on middle part; epigynum without long and light bellshaped region in median part; pore plates thin and long, wider in posterior half (VM 2010: figs 60– 62) ................................................................................................. P. hoogstraali Gertsch & Davis, 1942

– Epigynum with two small anterior apophyses, oval in ventral view; without two ventral protuberances on middle part; epigynum with long and light bell-shaped region in median part; pore plates markedly thinner and longer than in P. hoogstaali ( Figs 67–69 ) ......................................... P. pocamadre sp. nov.

22. Epigynum with ventral paired concavities on median part (VM 2010: figs 67, 151) ......................... 23

– Epigynum without ventral concavities on median part (VM 2010: figs 46, 81) ................................. 29

23. Epigynum with oval and large concavities on median part (VM 2010: figs 5, 151) ........................... 24

– Epigynum with small concavities on median part, “U”-shaped, close to epigastric furrow (VM 2010: figs 67, 144) ........................................................................................................................................... 26

24. Chelicerae with stridulatory ridges; ventral apophyses of epigynum wide, forming a “T” shape with median region between central concavities; pore plates wide, slightly curved, pointing to each other (VM 2010: figs 151–153) ............................................................ P. huacana Valdez-Mondragón, 2010

– Chelicerae without stridulatory ridges; ventral apophyses of epigynum thin, not forming a “T” shape with median region between central concavities (VM 2010: figs 5–7, 194–196) ............................... 25

25. Ventral apophyses of epigynum wider than in P. sarae , located on anterior part; in ventral view, apophyses pointing to each other; pore plates wide and oval, without oval translucid structures below them (VM 2010: figs 5–7) ............................................................................... P. bicornis Gertsch, 1971

– Ventral apophyses of epigynum thinner than in P. bicornis , located on central part; in ventral view, apophyses pointing downwards; pore plates small, slightly curved in posterior part, located above two oval translucid structures (VM 2010: figs 194–196) ........................ P. sarae Valdez-Mondragón, 2010

26. Ventral concavities of epigynum close to each other (VM 2010: fig. 67) ........................................... 27

– Ventral concavities of epigynum separated from each other (VM 2010: fig. 144) ............................. 28

27. Epigynum with ventral apophyses small and triangular in ventral view; ventral concavities on posterior part, with “W” shape; pore plates oval, smaller than in P. paredesi , above translucid structures (VM 2010: figs 67–69) ....................................................................................... P. lautus Gertsch, 1971

– Epigynum with apophyses small and conical in ventral view; ventral concavities large and oval, on central part; pore plates semicircular, longer than in P. lautus , without oval translucid structures below ( Valdez-Mondragón 2014: figs 31–33) ......................................... P. paredesi Valdez-Mondragón, 2010

28. Epigynum with ventral apophyses wide in ventral view, close to each other, ending in a rounded tip, located on median part; in ventral view, two deep and elongated anterior concavities absent; pore plates small and oval, above oval translucid structures (VM 2010: figs 144–146) ........................................... ........................................................................................................ P. gertschi Valdez-Mondragón, 2010

– Epigynum with ventral tiny apophyses, widely separated from each other, ending in a small sharp tip, located on anterior margin of epigynum; in ventral view, two deep and elongated anterior concavities present; pore plates wide and oval, with a small contraction in middle, without oval translucid structures below (VM 2010: figs 201–203) ................................................. P. platnicki Valdez-Mondragón, 2010

29. Epigynum with ventral apophyses small, conical, on median part (VM 2010: figs: 81, 83) .............. 30

– Epigynum with ventral apophyses with different size, shape and position, or even absent (VM 2010: figs 116, 118) ......................................................................................................................................... 33

30. Chelicerae without stridulatory ridges; epigynum corrugated ( Figs 55–56 ) ....................................... 31

– Chelicerae with stridulatory ridges; epigynum not corrugated (VM 2010: fig. 81) ............................ 32

31. Pore plates circular, small, widely separated from each other, without translucid structures below them; in lateral view, epigynum without curved concavity (VM 2010: figs 158–160) ..................................... ...................................................................................................... P. montanoi Valdez-Mondragón, 2010

– Pore plates oval, longer than in P. montanoi , close to each other, with translucid oval structures below; in lateral view, epigynum with curved concavity ( Figs 55–57 ) ................................ P. sikuapu sp. nov.

32. Epigynum with ventral apophyses small and conical in ventral view, thin and conical in lateral view; pore plates elongated, above oval translucid structures (VM 2010: figs 81–83) .................................... ......................................................................................................................... P. modestus Gertsch, 1971

– Epigynum with ventral apophyses small and slightly elongated in ventral view, wide and conical in lateral view; pore plates oval, above oval translucid structures ( Nolasco & Valdez-Mondragón 2020: figs 14–16) .............................................................. P. xerophilus Nolasco & Valdez-Mondragón, 2020

33. Epigynum with rounded shape, without ventral apophyses; with dark central spot; pore plates short and triangular, above oval translucid structures (VM 2010: figs 116–119) ................................................... ............................................................................................................................. P. validus Gertsch, 1971

– Epigynum with bell shape, with ventral apophyses; without dark central spot; pore plates oval or elongated, without oval translucid structures below ( Figs 31–33 ) ...................................................... 34

34. Pore plates elongated, wider distally; conical apophyses located on two dark regions; epigynum with paired concavities close to epigastric furrow ( Figs 31–33 ) ..................................... P. mariachi sp. nov.

– Pore plates oval; conical apophyses not located on two dark regions; epigynum without paired concavities close to epigastric furrow (VM 2010: figs 46–47, 53–54) ................................................ 35

35. Epigynum with one long and curved apophysis, slightly bifurcated at tip; pore plates with half oval shape, close to each other (VM 2010: figs 46–48) .............................. P. globosus (Taczanowski, 1874)

– Epigynum with three ventral apophyses, one on anterior part and two in median, not bifurcated distally; pore plates oval, separated by their own width (VM 2010: figs 53–55) ................................................. ........................................................................................................................ P. guanacaste Huber, 1998

Figures

Figs 8–13. Typical habitats of the spider genus Physocyclus Simon, 1893 from Mexico. 8. Deciduous forests (Nayarit). 9–10. Oasis in desert (Baja California Sur). 11. Cave entrance located in a deciduous forest (Guerrero). 12. Thorny scrub with columnar cacti (Guanajuato). 13. Xerophilous scrub (Baja California Sur).

Figs 74–77. Habitat and microhabitat of Physocyclys lyncis sp. nov. (74–75) and Physocyclus pocamadre sp. nov. (76–77). 74. Pine-oak forest, predominant vegetation in the Ejido La Primavera, Ecotourist Park, Zapopan, Jalisco, Mexico. 75. “Cueva del Lince” (type locality), inside of the Ejido la Primavera, Ecotourist Park, Zapopan, Jalisco, Mexico. 76. Habitat close to the beach “El Tecolote”, La Paz, Baja California Sur. 77. Xerophytic scrub, close to the beach “Punta San Francisquito” (type locality), Baja California.

Figs 70–73. Habitat and microhabitat of Physocyclus mariachi sp. nov. (70–71) and Physocyclus sikuapu sp. nov. (72–73). 70. Disturbed deciduous forest near Municipality of Hostotipaquillo (type locality), Jalisco, Mexico. 71. Big boulders on the ground where the specimens were collected (red arrows indicate the specific microhabitat). 72. Deciduous tropical forest, Municipality of Aquila, Costa Aquila Michoacan, Mexico. 73. Cave at the viewpoint of Costa Aquila (type locality).

Figs 14–19. Physocyclus mariachi sp. nov. Male (holotype). 14–16. Habitus in dorsal, lateral and ventral views, respectively. 17. Carapace, frontal view. 18, 19. Chelicerae in frontal and lateral views, respectively. Scale bars: 14–16 = 1 mm; 17–19 = 0.5 mm.

Figs 20–23. Physocyclus mariachi sp. nov. Male (holotype). 20–22. Left palp, prolateral, retrolateral and dorsal views, respectively. 23. Bulb of male palp, dorsal view. Scale bars: 0.5 mm.

Figs 24–27. Physocyclus mariachi sp. nov. Male (holotype). 24–25. Detail of chelicerae, frontal and lateral views, respectively. 26. Details of procursus, embolus and embolic sclerites of left palp, retrolateral view. 27. Detail of bulb, dorsal view. Scale bars: 0.5 mm.

Figs 44–47. Physocyclus sikuapu sp. nov. Male (holotype). 44–46. Right palp, prolateral, retrolateral and dorsal views, respectively. 47. Bulb of male palp, dorsal view. Scale bars= 0.5 mm.

Figs 48–51. Physocyclus sikuapu sp. nov. Male (holotype). 48–49. Detail of chelicerae, frontal and lateral views, respectively. 50. Details of procursus, embolus and embolic sclerites of right palp, in retrolateral view. 51. Detail of bulb, dorsal view. Scale bars= 0.5 mm.

Figs 58–63. Physocyclus lyncis sp. nov. Female (holotype). 58–60. Habitus in dorsal, lateral and ventral views, respectively. 61–63. Epigynum in ventral, lateral and dorsal views, respectively. Scale bars: 58– 60 = 1 mm; 61–63 = 0.5 mm.

Figs 64–69. Physocyclus pocamadre sp. nov. Female (holotype). 64–66. Habitus in dorsal, lateral and ventral views, respectively (red and blue arrows indicate the dorsal protuberance on carapace and dorsal patch on opisthosoma). 67–69. Epigynum in ventral, lateral and dorsal views, respectively. Scale bars: 64–66 = 1 mm; 67–69 = 0.5 mm.

Figs 52–57. Physocyclus sikuapu sp. nov. Female (paratype). 52–54. Habitus in dorsal, lateral and ventral views, respectively. 55–57. Epigynum, ventral, lateral and dorsal views, respectively (red arrow indicates translucid bag-shaped structure). Scale bars: 52–54 = 1 mm; 55–57 = 0.5 mm.

Figs 28–33. Physocyclus mariachi sp. nov. Female (paratype). 28–30. Habitus in dorsal, lateral and ventral views, respectively; red arrows indicate a dorsal patch in the opisthosoma (28) and the dorsal protuberance in carapace (29). 31–33. Epigynum, ventral, lateral and dorsal views, respectively. Scale bars: 28–30 = 1 mm; 31–33 = 0.5 mm.