Tomarus

López-García, Margarita M. & Deloya, Cuauhtémoc, 2019, Five New Species of the Dynastine Genus Tomarus Erichson (Coleoptera: Scarabaeidae), with an Illustrated Key to Species, The Coleopterists Bulletin 73 (1), pp. 127-141 : 135-139

KEY TO ADULTS OF THE SPECIES OF THE GENUS TOMARUS

1. Pronotum without fovea or tubercle .......... 2

1 ʹ. Pronotum with fovea and tubercle, occasionally very small. If fovea absent, then pronotum nearly smooth, with minute, sparse punctures .......................................... 9

2. Apex of metatibia crenulate ( Fig. 24 ). Male protarsal inner claw wide and bifid, protarsomere 5 enlarged ........................... 3

2 ʹ. Apex of metatibia entire ( Fig. 25 ). Male protarsal inner claw protarsus simple, protarsomere 5 subcylindrical .................... 5

3. Protibia with 5 teeth: 3 large and 2 small denticules (one basal and one before second large tooth) ( Fig. 26 ). Mexico and USA.... ................ Tomarus sallaei (Bates, 1888)

3 ʹ. Protibia with 4 teeth: 3 large and a small basal denticule ( Fig. 27 ) .............................. 4

4. Pygidium with dense rugosity on entire surface. Apex of metatibia with 9–10 spinules. Parameres short, with an acute lateral tooth on each side (Fig. 45). USA and Canada........ ................... Tomarus relictus (Say, 1825)

4 ʹ. Pygidium with large, sparse punctures. Apex of metatibia with 14–15 spinules. Parameres long, without acute lateral tooth (Fig. 46). Ecuador and Peru................................. ...... Tomarus peruvianus (Endrödi, 1970)

5. First apical tooth of mandible acute ( Fig. 28 ). Frontoclypeal region with a thick, straight carina ( Fig. 30 ). Mesotibia strongly narrowed before apex. Mexico and Central America ................... Tomarus nasutus (Burmeister, 1847)

5 ʹ. First apical tooth of mandible truncate ( Fig. 29 ). Frontoclypeal region with a pair of cariniform tubercles ( Fig. 31 ). Mesotibia not or only slightly contracted before apex ................. 6

6. Parameres with a large, dorsal tooth on each side, apex straight (Fig. 47). Females identifiable only by association with males. El Salvador, West Indies, and South America... ............... Tomarus fossor (Latreille, 1813)

6 ʹ. Parameres without dorsal teeth, with a ventral tooth on each side, apex variable ........... 7

7. Parameres with acute, triangular apex ( Figs. 18–19 ). Metatibia strongly narrowed before apex. Apex of metatibia with 19 spinules. Brazil................... Tomarus paranaensis López-Garc´ıa and Deloya, new species

7 ʹ. Parameres with apex truncate to rounded (Figs. 46–47). Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia with 25–29 spinules ............................ 8

Figs. 45–68. Tomarus species , parameres. 45) T. relictus ; 46) T. peruvianus ; 47) T. fossor ; 48) T. spinipenis ; 49) T. burmeisteri ; 50) T. neglectus ; 51) T. gibbosus ; 52) T. rostratus ; 53) T. cicatricosus ; 54) T. ebenus ; 55) T. similis ; 56) T. bituberculatus ; 57) T. selanderi ; 58) T. maimon ; 59) T. discrepans ; 60) T. gyas ; 61) T. laevicollis ; 62) T. colombianus ; 63) T. subtropicus ; 64) T. maternus ; 65) T. adoceteus ; 66) T. pullus ; 67) T. pumilus ; 68) T. roigjunenti .

8. Parameres wide, with parallel sides and minute spines on apicolateral third; ventral teeth short, triangular (Fig. 48). Females identifiable only by association with males. Argentina........... ................. Tomarus spinipenis Neita and Ratcliffe, 2017

8 ʹ. Parameres slender, apex narrowed, without spines on apicolateral third; ventral teeth long (Fig. 49). Females identifiable only by geography or association with males. Bolivia, Argentina, and Brazil............................. ..... Tomarus burmeisteri (Steinheil, 1872)

9. Antennal club 1.5–2.0 times longer than antennomeres 2–7 ( Fig. 32 ). Ocular canthus acute .............................................. 10

9 ʹ. Antennal club shorter, subequal in length to antennomeres 2–7 ( Fig. 33 ). Ocular canthus rounded .......................................... 16

10. Mandibles with 3 acute teeth. Clypeus subtrapezoidal, base 1.5 times wider than apex. Apex of metatibia with 11–15 spinules...11

10 ʹ. Mandibles with 2 acute apical teeth and a widely rounded basal tooth. Clypeus usually narrowed toward apex, base ±2 times wider than apex. Apex of metatibia with more than 25 spinules ...................................... 12

11. Clypeal teeth triangular to transverse, located on rounded anterior angles of clypeus ( Fig. 16 ). Parameres without ventral, apical tooth ( Figs. 14–15 ). Southern Florida, West Indies, Mexico (Tabasco), and Atlantic Coast of South America from Colombia to Brazil (Amapá to Espiritu Santo) ...................... ...... Tomarus cuniculus (Fabricius, 1801)

11 ʹ. Clypeal teeth conical, located in middle of apical margin ( Fig.13 ). Anterior angles of clypeus strongly acute and elevated. Parameres with ventral apical tooth ( Figs. 11–12 ). Atlantic Coast of Brazil (Rio de Janeiro to Santa Catarina) ..................................... .............. Tomarus moroni López-Garc´ıa and Deloya, new species

12. Frontoclypeal region with a bilobed carina or 2 low, transverse tubercles separated by 1 tubercle diameter or more ( Figs. 34–36 ) .. .. ...................................................... 13

12 ʹ. Frontoclypeal region with a high, straight carina ( Fig. 30 ) ................................. 15

13. Frontal tubercle short, separated by more than 2 tubercle diameters ( Fig. 34 ). Pronotum and elytra nearly smooth, with minute, sparse punctures. Pronotal fovea absent ( Figs. 39, 42 ). Pronotal tubercle like a swelling ( Fig. 39 ). Peru, Chile, and Argentina............... ....... Tomarus villosus (Burmeister, 1847)

13 ʹ. Frontal tubercles longer, separated by about a tubercle diameter ( Figs. 35–36 ). Pronotum and elytra strongly punctate, with small to large punctures. Pronotal fovea small to large. Pronotal tubercle large, conical to triangular ( Figs. 40–41 ) ......................... 14

14. Pronotal fovea large, deep, about 1.2 times interocular width ( Fig. 43 ). Clypeus short ( Fig. 35 ) and with lateral margins perpendicularly elevated ( Figs. 37, 40 ). Clypeal teeth triangular to transverse, separated by a tooth diameter ( Fig. 37 ). Brazil, Argentina, and Uruguay........................................ ........ Tomarus rubripes (Boheman, 1858)

14 ʹ. Pronotal fovea small, shallow, 0.25 times interocular width ( Fig. 44 ). Clypeus long, strongly narrowed towards apex ( Fig. 36 ), lateral margins not elevated ( Figs. 38, 41 ). Clypeal teeth long, acute, not separated ( Fig. 38 ). Argentina...................................... .... Tomarus bidentulus ( Fairmaire, 1892)

15. External surface of protibia (in frontal view) nearly smooth, with only minute, sparse punctures. Pronotum and elytra deeply punctate, punctures large, round. Apical half of parameres wide and with parallel sides, ventral tooth wide and located apically (Fig. 50). USA............................................ ........ Tomarus neglectus (LeConte, 1847)

15 ʹ. External surface of protibia (in frontal view) deeply or at least obviously punctate. Pronotum and elytra with minute to small punctures. Apical half of parameres slender and slightly narrowed, ventral tooth triangular and located medially (Fig. 51). USA and Mexico......................................... ......... Tomarus gibbosus (De Geer, 1774)

16. Clypeus trapezoidal. Clypeal teeth small, transverse and separated by more than 2 tooth diameters ........................................ 17

16 ʹ. Clypeus triangular. Clypeal teeth large, usually triangular and acute, separated by less than 2 tooth diameters or not separated ........... 20

17. Protibia tridentate. Apex of metatibia with 6–8 spinules, margin dentate. Parameres not strongly elongated and with apicolateral or medial teeth ..................................... 18

17 ʹ. Protibia quadridentate, basal tooth small and acute. Apex of metatibia with 13–15 spinules, margin slightly crenulate. Parameres strongly elongated and without lateral teeth (Fig. 52). Desert coast of Peru and Chile... ............ Tomarus rostratus Dupuis, 2014

18. Pronotum, elytra, and pygidium with minute to small punctures. Male protarsal inner claw bifid, protarsomere 5 enlarged. Apex of sternite 8 not emarginate in females ............ 19

18 ʹ. Pronotum, elytra, and pygidium with large, deep, round punctures. Male protarsal inner claw simple, acute, protarsomere 5 subcylindrical. Apices of parameres strongly expanded outwards (Fig. 53). Apex of sternite 8 emarginate in females. Mexico to Colombia........ ............. Tomarus cicatricosus (Prell, 1937)

19. Parameres wide, basal third widened, apical third strongly contracted and apex dilated (Fig. 54). Females identifiable only by geography or association with males. Mexico to Brazil............................................ .. .... .... Tomarus ebenus (De Geer, 1774)

19 ʹ. Parameres slender and long, apical third not strongly contracted, sides nearly parallel (Fig. 55). Females identifiable only by geography or association with males. Costa Rica to Brazil....................................... ............ Tomarus similis (Endrödi, 1968)

20. Pronotal tubercle large, usually acute. Pronotal fovea deep and wide .................. 21

20 ʹ. Pronotal tubercle minute to small, rounded. Pronotal fovea shallow and narrow .......... 25

21. Pronotal fovea transversely oval and as wide as interocular distance. Pronotal punctation minute. Parameres as in Fig. 56. Mexico to Bolivia............. Tomarus bituberculatus (Palisot de Beauvois, 1811)

21 ʹ. Pronotal fovea rounded and narrower, about 1/2–2/3 times interocular width. Pronotal punctation small to large .................... 22

22. Metasternum glabrous, without setae. Pronotal fovea 1/3 wider than interocular width. Parameres with or without a short ventral tooth on each side (Fig. 57–58) ........... 23

22 ʹ. Metasternum with short, scattered setae on the outer sides. Pronotal fovea 2/3 wider than interocular width. Parameres with a long dorsal tooth on each side .................... 24

23. Pronotal fovea shallow, sometimes as a rugose surface. Parameres with apical third triangularly dilated (Fig. 57). Pacific Coast of Mexico ..................... Tomarus selanderi (Cartwright, 1959)

23 ʹ. Pronotal fovea deep. Parameres with apical fourth roundly dilated (Fig. 58). Ecuador, Peru, Chile, Guianas, and Brazil.............. ........... Tomarus maimon Erichson, 1847

24. Apex of parameres wide (Fig. 59). Females identifiable only by geography or association with males. Mexico to Venezuela............... ................. Tomarus discrepans (Escalona and Joly, 2006)

24 ʹ. Apex of parameres narrowed, ending in a triangular or rounded point (Fig. 60). Females identifiable only by geography or association with males. Costa Rica to Argentina............. Tomarus gyas Erichson, 1848 (in part)

25. Frontoclypeal tubercles conical to triangular .......................................... 26

25 ʹ. Frontoclypeal tubercles transverse, low, sometimes difficult to see .............................. 30

26. Pronotum punctate, with small to large, dense punctures. Apex of metatibia with 23–25 spinules .................................................. Tomarus gyas Erichson, 1848 (in part)

26 ʹ. Pronotum nearly smooth, with minute to small, sparse punctures. Apex of metatibia with 8–12 spinules ............................ 27

27. First interval of elytra smooth or with sparse punctures. Parameres as in Fig. 61. Mexico to Costa Rica........................................... ........... Tomarus laevicollis (Bates, 1888)

27 ʹ. First interval of elytra strongly punctate, punctures similar in size or smaller than in other intervals .................................. 28

28. Apex of pygidium strongly dilated in females. Parameres with small accessory tooth posteriorly joined to base of large lateral tooth ( Fig. 3 ). Panama........................... .......... Tomarus laticaudus López-Garc´ıa and Deloya, new species

28 ʹ. Apex of pygidium regularly rounded. Parameres without small accessory tooth at base of large lateral tooth (Fig. 59) ............. 29

29. Parameres with apicolateral tooth acute and shorter than basolateral tooth (Fig. 62). Protibia with a fourth small, basal tooth. Colombia (Pacific Coast) ................ Tomarus colombianus López-Garc´ıa and Gasca-Álvarez, 2014

29 ʹ. Parameres with 2 lateral teeth equal in length and shape ( Fig.8 ). Protibia tridentate with a small, basal undulation. Northern Venezuela............. ....... Tomarus maracaiboensis López-Garc´ıa and Deloya, new species

30. Pronotum with minute to small punctures. Metatibia subrectangular, sides nearly parallel. Apex of metatibia with 9–17 spinules .............................................. 31

30 ʹ. Pronotum with large, deep, punctures. Metatibia subtriangular, strongly widened towards apex. Apex of metatibia with 21–30 spinules .......................................... 33

31. Pronotum nearly smooth, with sparse, minute punctures. Parameres with large lateral tooth strongly curved backwards (Fig. 63). USA..................... Tomarus subtropicus (Blatchley, 1922)

31 ʹ. Pronotum punctate, with small, dense punctures. Parameres with large lateral tooth not strongly curved back, subapical tooth rounded or obtuse ............................. 32

32. Parameres with large lateral tooth short and wide, subapical tooth rounded (Fig. 64). Scutellum with 2 lines of punctures parallel to lateral borders. Apex of metatibia with 14–17 spinules. Honduras to Bolivia.................. ............. Tomarus maternus (Prell, 1937)

32 ʹ. Parameres with large lateral tooth long and slender, subapical tooth obtuse (Fig. 65). Scutellum with sparse punctures. Apex of metatibia with 9–11 spinules. Bahamas and Cayman Islands.................................... .................. Tomarus adoceteus Ratcliffe and Cave, 2010

Figures

Figs. 24–44. Tomarus species. Metatibia: 24) T. sallaei; 25) T. nasutus. Protibia: 26) T. sallaei; 27) T. relictus. Mandible: 28) T. nasutus; 29) T. fossor. Head: 30) T. nasutus; 31) T. burmeisteri. Antenna: 32) T. villosus; 33) T. maimon. Head, dorsal view: 34) T. villosus; 35) T. rubripes; 36) T. bidentulus. Head, frontal view: 37) T. rubripes; 38) T. bidentulus. Head and pronotum, lateral view: 39) T. villosus; 40) T. rubripes; 41) T. bidentulus. Pronotum: 42) T. villosus; 43) T. rubripes; 44) T. bidentulus.

Figs. 17–19. Tomarus paranaensis. 17) Habitus; 18) Parameres, frontal view; 19) Parameres, lateral view. Scale bars = 2 mm (Fig. 17), 0.5 mm (Figs. 18–19).

Figs. 10–16. Tomarus moroni: 10) Habitus; 11) Parameres, frontal view; 12) Parameres, lateral view; 13) Head, dorsal view. Tomarus cuniculus: 14) Parameres, frontal view; 15) Parameres, lateral view; 16) Head, dorsal view. Scale bars = 2 mm (Figs. 10, 13, 16), 0.5 mm (Figs. 11–12, 14–15).

Figs. 2–6. Tomarus laticaudus: 2) Habitus; 3) Parameres, frontal view; 4) Parameres, lateral view; 5) Female pygidium, ventral view. Tomarus laevicollis: 6) Female pygidium, ventral view. Scale bars = 2 mm (Figs. 2, 5–6), 0.5 mm (Figs. 3–4).

Figs. 7–9. Tomarus maracaiboensis. 7) Habitus; 8) Parameres, frontal view; 9) Parameres, lateral view. Scale bars = 2 mm (Fig. 7), 0.5 mm (Figs. 8–9).