Hypogena vacca (Fabricius, 1801)

(Figs 1, 2, 3, 38, 39)

Trogosita vacca Fabricius, 1801: 153 .

Ulosonia vacca (Fabricius, 1801) . (Shultz, 1901).

= Ulosonia parvicornis Fairmaire, 1892: 250 (Synonymy in Kulzer 1962: 93).

Type Material: LECTOTYPE (male) labeled: (a) “Amer. Mer. / Schmidt”; (b) “ Mus de Sehestedt ”; (c) “ Trogosita vacca Fabr. ”; (d) on red paper “Type.”; (e) “ LECTOTYPE / Hypogena / vacca / Grey & Smith, 2020” (ZMUK). Designated here . PARALECTOTYPE (female) labeled: (a) on a red label “Type”; (b) “Amer. Mer. / Schmidt”; (c) “Mus de Sehestedt”; (d) “ Trogosita ♀ vacca Fabr. ”; (e) “ PARALECTOTYPE / Hypogena / vacca / Grey & Smith, 2020” (ZMUK). (See Fig. 38). This image was taken by Dr. Michael Kuhlmann at the Zoological Museum of Kiel University. In the original description, Fabricius (1801) did not designate a holotype. The specimen designated here is intended to fix the taxonomic status of this species. The designated specimen was chosen from the syntype series because it had the most complete locality information .

Additional material examined: (139 females, 172 males, 2 sex unknowns): ARGENTINA: 8 females, 9 males . TB# 21182, 21206 (NMNH); TB# 18130-18131 (CASC); TB# 17928, 17935, 18133 (NHMB); TB# 18020, 18025, 21762 (NMNH); TB# 18134, 21169-21174 (BMNH). ARGENTINA: Buenos Aires: 7 females . TB# 17936-17937 (NHMB); TB# 18036-18039 (NMNH); TB# 21208 (BMNH). ARGENTINA: Chaco: 2 males . TB# 18138 (NHMB); TB# 18136 (BMNH). ARGENTINA: Jujuy: 1 female . TB# 18137 (NHMB). ARGENTINA: Mendoza: 1 male . TB# 17934 (NHMB). ARGENTINA: Salta: 1 female . TB# 21209 (CASC). ARGENTINA: Tucumán: 2 females, 6 males . TB# 18132 (CASC); TB# 18728-18729 (NHMB); TB# 18518, 18520-18523 (BMNH). BOLIVIA: 5 females, 8

males. TB# 21153-21162, 21165-21166 (NABF); TB# 18141 (NMNH) . BRAZIL: 27 females, 31 males. TB# 21216 (ALRC); TB# 21167-21168 (NABF); TB# 18352-18353, 18360-18363 (CASC); TB# 18731-18732 (unknown); TB# 21215 (CASC); TB# 21133-21151 (NMPC); TB# 17920, 17927, 17930, 17932-17933, 17938, 17941-17950, 17958, 18730, 18733-18734 (NHMB); TB# 18026-18028, 18030, 21181 (ADSC); TB# 21177 (BMNH); TB# 18029 (RLAC) . BRAZIL: Espírito Santo: 1 female, 3 males. TB# 21211-21214 (BMNH) . BRAZIL: Minas Gerais: 1 male. TB# 18139 (ADSC) . BRAZIL: Rio Grande do Sul: 1 female. TB# 18144 (NHMB) . BRAZIL: Santa Catarina: 19 females, 26 males. TB# 21178 (NMNH); TB# 17963, 17966-17967, 18333-18335, 18349, 18354, 18372-18373, 18377-18378, 18387, 18506-18508, 21760 (CASC); TB# 17926, 17931, 17939-17940, 18706-18719, 18721-18725 (NHMB); TB# 17968-17970 (NMNH); TB# 18524 (BMNH) . BRAZIL: Sao Paulo: 1 male. TB# 18135 (BMNH) . MEXICO: 1 female, 1 male. TB# 21203, 21205 (BMNH) . PARAGUAY: 2 females, 3 males. TB# 17951-17955 (NHMB) . USA: New Mexico: Santa Fe Co.: 2 males. TB# 18726-18727 (NHMB) . country not specified: 64 females, 78 males, 2 sex unknowns. TB# 17847-17856, 17960-17962, 17964, 18331-18332, 18336-18348, 18350-18351, 18355-18359, 18364-18371, 18374-18376, 18379-18386, 18388-18428, 18456-18469, 18505, 18510, 18513-18517, 21758-21759, 21761 (CASC); TB# 21210 (CASC); TB# 17921-17925, 17929, 18143 (NHMB); TB# 17971-17972, 18031-18035, 18140, 18142 (NMNH); TB# 21164, 21175-21176, 21204, 21207, 21763-21764 (BMNH) .

Diagnosis: The male of this species has two large supraorbital horns and a small tuberculate clypeal horn. The supraorbital horns are usually very thick, perpendicular to the head, and often diverge away from each other. The clypeal horn is variable, sometimes appearing very short, or sometimes larger with the middle portion of the horn being somewhat thinner than the apical portion. The punctures on this species are of two sizes and the coarse punctures are primarily located on the lateral portions of the pronotum and are widely spaced away from one another. The posterior edge of the pronotum is raised. This species is small, usually between 5 - 7 mm long. The female is similar in every aspect except the presence of the horns (Fig. 38).

Redescription: Male. Size: 5—7 mm; 2.5— 3 mm.

Head. Clypeal horn short and similar in thickness at base to one supraorbital horn (4:1; 5:1; 7:1). Clypeal horn projected forward, almost smooth (8:0; 9:0); may be tuberculate or thinner in middle and widen slightly apically. Supraorbital horns long and thick (10:1; 11:1); from dorsal view, may not bend or may bend outwards (12:0); produced perpendicularly to head (13:1); bent slightly forward (14:1); finely punctate (15:1). Antennae with stellate sensoria (22:0). Third antennomere similar size as fourth and more than 1.5 times size of second (18:0; 19:2). Vertex and frons not punctate, almost smooth (26:0; 28:0). Gena not extending laterally past eye (36:0). Clypeus not forming circular arch due to clypeal horn (29:2). Mentum trapezoidal, punctate and setose (40:0; 42:1; 41:1). Labrum fully setose (39:1). Groove on mandible present, continuing to connection point to cranium (44:1).

Thorax. Pronotal puncturing biordinal in size (54:1). Coarse punctures numerous; concentrated on lateral portions of pronotum (57:0). Posterior edge of pronotum raised. From lateral view, pronotum not arched (58:0). Lateral margins of pronotum subparallel (61:1). Margins of pronotum expanded wider than diameter of one coarse puncture (62:2). Anterior edge of pronotum extending to middle of eye but not past eye (63:1). Hypomeron laterally rugulose (75:2); not punctate. Prosternum finely punctate (67:1); separated by less than one diameter (68:1); no pubescent (66:0). Prosternal process not continuing past posterior margin of pronotum (69:0). Elytral striae punctate (46:1; 47:1); punctures separated by less than one diameter (48:0). Elytral interstices convex and punctate (49:1); punctures as large or larger than one fourth size of strial puncture size (50:0); separated by more than one diameter (51:1). Mesepisternum with punctures as large as or larger than eye facets (71:2); separated by less than one diameter (72:1). Row of cuticular teeth present on posterior edge of metasternum (77:1).

Legs. Tarsi pubescent. Cuticular outgrowths present on proximal two protarsomeres (86:1). Tibiae bearing two apical spurs (85:1). Lateral edge of protibia with more than seven socketed spines on distal half of tibia (81:1; 82:1). Spines on medial edge of protibia along entire length (83:1).

Abdomen. First three visible abdominal segments setose and laterally rugulose (89:1; 90:1; 91:1). All abdominal segments are finely punctate. setae arising from punctures (94:3).

Female. Similar to male, except as follows: Cuticle on intraocular space raised near eye and extending transversely inwards (23:2). Intraocular space impinging on eye (24:1). Vertex and frons punctate; punctures as large or larger than facets of eye (25:2; 27:2). Clypeus punctate; punctures smaller than eye facets and spread evenly across clypeus (31:1; 32:2). Anterior margin of clypeus flat, but with corners rounded (30:1). Clypeus tumid (33:1).

Distribution (Fig. 39): Paraguay, Brazil (Minas Gerais, Sao Paulo, Espirito Santo, Rio Grande do Sul, Santa Caterina) Bolivia, Argentina (Tucumán, Jujuy, Salta, Buenos Aires, Chaco, Mendoza).

Key to Hypogena species

1) Mesepisternum setose; prosternum and hypomeron setose with white to golden setae; coarse pronotal punctures scattered across pronotal surface; clypeal horn short, pointed anteriorly from clypeus; supraorbital horns short, tuberculate, and pointed anteriorly (Fig. 27); Ecuador, Brazil, Costa Rica, Honduras, Panama, Venezuela, Mexico, Cuba, Guatemala (Fig. 28)........................................................................................... H. hirsuta sp. nov.

- Mesepisternum not setose; pronotal punctures occasionally as above, but not in combination with setation on the mesepister- num; hypomeron occasionally setose, setae always absent on mesepisternum; clypeal and supraorbital horns variable..... 2

2) Line of setae present on all femurs; coarse punctures scattered across pronotum; clypeal horn short, tuberculate; supraorbital horns short, tuberculate, not pointing inwards as in H. marginata (Fig. 10); Brazil .................... H. akuma sp. nov.

- Line of setae not present on any femur; pronotal punctures and horns may be similar to those in H. akuma, but not in combination with line of setae on femora......................................................................... 3

3) Pronotum and elytra with long distinct setae arising from coarse punctures; body elongate; coarse pronotal punctures concentrated on lateral portions of pronotum (Fig. 34); [male specific characters unknown] Columbia........ H. reburra sp. nov.

- Pronotum and elytra not bearing long porrect hairs; body usually not as elongate; pronotal puncture concentration variable; if pronotal punctures concentrated on lateral portions of pronotum, then the elytra and pronotum not bearing long setae..... 4

4) Pronotal punctures small, of one size, or with at most 4 larger punctures (Fig. 17).................................. 5

- Pronotal punctures of two sizes with numerous large punctures (Fig. 21)......................................... 8

5) Mentum medially and longitudinally produced; hypomeron without punctures, smooth; horns long, thin, straight and subequal in length; horns usually not as well developed as those of H. tricornis (Fig. 17); Brazil, Costa Rica, Cuba, Guatemala, Nicaragua, Mexico, Panama (Fig. 18)................................................... H. canaliculata (Champion)

- Mentum not raised; hypomeron usually punctate; horn thickness variable, but bent to point anteriorly, usually not equal in length.............................................................................................. 6

6) Mentum medially depressed; hind femur weakly bent; clypeal horn long, and arising perpendicular to head; clypeal horn thinnest at base and widening apically and flat on top; supraorbital horn strongly bent both inwards and forward (Fig. 35); Peru ................................................................................... H. triceratops Steiner

- Mentum not depressed; hind femur not bent; horns variable.................................................... 7

7) Anterior portion of first two abdominal segments laterally rugulose; length: 9—10 mm; clypeal horn conical, projecting forward from clypeus; supraorbital horns large, thinnest basally and widening apically; supraorbital horns strongly bent to point anteriorly (Fig. 29); Brazil, Peru (Fig. 30)................................................ H. laevicollis (Kulzer)

- First two abdominal segments completely laterally rugulose; length: 7-9 mm; clypeal horn long, flat, widening apically and flat to emarginate on top; supraorbital horns long, produced perpendicular to head and strongly curved forward (Fig. 23); Mexico, Costa Rica (Fig. 24)................................................................ H. dejeani (Champion)

8) Coarse pronotal punctures densely clustered on disc; all horns subequal in length; supraorbital horns bending slightly toward each other (Fig. 36); USA, Mexico, Guatemala, Belize, El Salvador, Honduras, Costa Rica, Panama, Columbia, Venezuela, Ecuador, Cayman Islands, Cuba, Jamaica (Fig. 37)........................................ H. tricornis (Dalman)

- Coarse pronotal punctures not concentrated primarily on disc; horns usually not equal in length or as thin as in H. tricornis .. ................................................................................................... 9

9) Coarse pronotal punctures concentrated on lateral portions of pronotum......................................... 10

- Coarse pronotal punctures scattered throughout pronotum.................................................... 12

10) Hypomeron and prosternum setose with distinct golden setae; clypeal horn short, conical and arising forward from clypeus; supraorbital horns long, bending slightly inward toward each other; supraorbital horns projecting perpendicularly from head (Fig. 13); Nicaragua, Mexico, Peru, Columbia, Panama, Haiti, Brazil, Ecuador, Honduras, Guatemala, Dominican Republic, Costa Rica, El Salvador (Fig. 14)......................................................... H. brasilica (Perty)

- Hypomeron and prosternum without distinct setae; horns not similar to above, either much thicker or bent to point anteriorly .................................................................................................. 11

11) Posterior edge of pronotum with raised edge; supraorbital horns thick and either not bending laterally or bending away from each other; clypeal horn short, tuberculate; clypeal horn often conical but may be thinner in the middle and widening apically (Fig. 38); Brazil, Bolivia, Paraguay, Argentina (Fig. 39)..................................... H. vacca (Fabricius)

- Posterior edge of pronotum without raised edge; clypeal horn long, flattened and truncated; clypeal horn widening apically and then becoming flat to slightly emarginate on top; supraorbital horns strongly curved forward to point anteriorly (Fig. 19); Peru, Colombia, Panama, Mexico (Fig. 20)........................................................... H. cat Steiner

12) Coarse pronotal punctures more than twice the size of fine punctures; male with three long horns; clypeal horn three times the diameter, at base, of supraorbital horns; Hind horns perpendicular, not bending (Fig. 15); Brazil, Colombia (Fig. 16)........................................................................................ H. brasiliensis (Kulzer)

- Coarse pronotal punctures two times the size or less of fine punctures; clypeal horn less than three times the size of one supraorbital horn........................................................................................ 13

13) Lateral edges of pronotum gradually raised, weakly sloping upwards from margins of pronotum; clypeal horn not short, of similar size to supraorbital horns; horns similar to H. tricornis but not as long (Fig. 25); Honduras, Mexico, Bahamas, Costa Rica, Guatemala, Brazil, Cuba, USA, Jamaica (Fig. 26).................................. H. depressa (Champion)

- Lateral edges of pronotum not as depressed as above; after flat margin, lateral edges of pronotum steeply raised and flattens out near disc; clypeal horn usually short; supraorbital horns variable............................................... 14

14) Male with cuticular spike on profemur; conical and produced forward; supraorbital horns long, not curved toward each other, but slightly bent forward (Fig. 11); Brazil, Peru (Fig. 12).................................. H. amazonica (Kulzer)

- Male without cuticular spike on profemur; supraorbital horns variable in size but not bent forward.................... 15

15) Supraorbital horns short, tuberculate, projected forward from head (Fig. 32); Mexico, Costa Rica, Guatemala, Peru, USA (Fig. 33)............................................................................. H. marginata (LeConte)

- Supraorbital horns long, projected perpendicularly from head (Fig. 21); USA, Mexico, Guatemala (Fig. 22).................................................................................................. H. cryptica sp. nov.