Bountya Townsend, 1971: 180

ROIG-JUÑENT, SERGIO, 2000, The Subtribes And Genera Of The Tribe Broscini (Coleoptera: Carabidae): Cladistic Analysis, Taxonomic Treatment, And Biogeographical Considerations, Bulletin of the American Museum of Natural History 2000 (1), pp. 1-90 : 62-70

Bountya Townsend, 1971: 180 .

TYPE SPECIES (BY MONOTYPY): Bountya insularis Townsend, 1971 .

DIAGNOSIS: Bountya belongs to the same group as Cascellius . Bountya and Anheterus are the only genera that male have foretarsomeres without adhesive vestiture. It differs from Anheterus in having 9–12 setae on lateral umbilical series.

DESCRIPTION (after Townsend, 1971): External morphology: One supraorbital seta bearing one seta at each side of head; vertex without transverse groove; antenna with antennomere 2 glabrous, 3 pubescent on apical third, 4 on apical half, 5–11 all along surface. Mandible with or without scrobal setae. Labium with submentum having eight setae; tooth of mentum simple; prementum with glossal sclerite quadrisetose. Pronotum not constricted apically, with two setae each side. Elytron with parascutellar seta on basal portion of second stria; lateral umbilical series with 9–12 setae. Abdominal sterna smooth. Male fore and middle tarsomeres without adhesive vestiture.

Male genitalia: Median lobe with dorsal surface unsclerotized and basal orifice completely open dorsally. Internal sac without sclerite X, Y, and apical plate. Left paramere asetose, with apical region thin; right setose.

Female genital tract: Subgonocoxite 9 broad; gonopod 9 short, without nematiform setae on subapical setose organ. Median oviduct and spermatheca entering separately into bursa copulatrix; helminthoid sclerite absent, bursa copulatrix long.

LARVAE: The larva was described by Townsend ( 1988).

DISTRIBUTION AND HABITAT: The genus Bountya occurs in Bounty Island, south of New Zealand, occupying a subantarctic habitat.

Creobius Guérin-Ménéville, 1838 Figures 290–297

Feronia (Creobius) Guérin-Ménéville, 1838: 4 .

Creobius : Waterhouse, 1841b: 255. Solier, 1849: 200. Semenov, 1900: 80. Csiki, 1928: 14. Blackwelder, 1944: 28. Schweiger, 1959: 3. Roig-Juñent, 1995a: 53.

Cascellius Curtis, 1839 (part): 183. Waterhouse, 1841: 254. Gemminger and Harold, 1868: 244. Waterhouse, 1881: 80. Philippi, 1887: 637. Bruch, 1911: 162. Germain, 1911: 53.

Cascelius Putzeys, 1868 (part) (lapsus): 352.

TYPE SPECIES (by monotypy): Feronia (Creobius) eydouxii Guérin-Ménéville, 1838 . DIAGNOSIS: Creobius belongs to the same group as Cascellius , and differs from all the genera of this group in having three or more supraorbital setae on each side.

DESCRIPTION: External morphology (figs. 290, 291): Three to five supraorbital punctures bearing one seta each side of head; vertex without transverse groove; temporal ridge obsolete in middle; antenna with antennomeres 1–2 glabrous, 3 setose on half or apical third, 4–11 pubescent all along surface. Mandible each with scrobal setae. Maxillary eustipes with two basal setae; maxillary palpomere 2 asetose. Labium with submentum bisetose; mentum bisetose, with tooth simple; prementum with glossal sclerite quadrisetose, ventral surface without longitudinal medial carina, with the paramedian areas deep-set; labial palpomere 2 bisetose. Pronotum not constricted apically, with five to six setae each side. Elytron with parascutellar striole joined to apical portion of stria 1; with parascutellar seta on basal portion of second stria; lateral umbilical series with 9 to 12 setae. Foretrochanter with one setae; male foretarsomeres 1–4 and middle tarsomeres 1–2 with adhesive vestiture ventrally.

Male genitalia (figs. 292–296): Sternum 9 with complete ring, having a portion not sclerotized, and lateral margins narrow (figs. 292, 293). Median lobe with dorsal surface unsclerotized and basal orifice completely open dorsally (fig. 296). Internal sac without sclerite X, Y, and apical plate; armature of small spiculae (fig. 296). Left paramere asetose (fig. 294); right paramere with long hairs from the middle (fig. 296).

Female genital tract (fig. 297): Rami of gonocoxite 9 short. Subgonocoxite 9 broad, without setae; gonopod 9 short, without nematiform setae on subapical setose organ. Spermatheca broad (fig. 297, sp); median oviduct and spermatheca enter separately into bursa copulatrix; helminthoid sclerite absent. Bursa copulatrix long, with apical accessory gland (fig. 297), without basal sclerite; ligula not sclerotized.

DISTRIBUTION AND HABITAT: The genus Creobius has a single species restricted to the northern Nothofagus forest of Argentina and Chile, being one of the most common carabid species in this area.

MATERIAL EXAMINED: Creobius eydouxii (AMNH, CM, CUIC, IADIZA, MCZ, USNM) .

Promecoderus Dejean, 1829 Figures 298–325

Promecoderus Dejean, 1829: 25 . Waterhouse, 1842: 205. Lacordaire, 1854: 244. Putzeys, 1868: 328; 1873: 319. MacLeay, 1873: 330. Sloane, 1890: 190; 1920: 124. Csiki, 1928: 24. Lawrence et al., 1987: 109.

Cnemacanthus Gray, 1832: 276 (type species Cnemacanthus gibbosus Gray, 1832 by monotypy).

TYPE SPECIES (by monotypy): Promecoderus brunnicornis Dejean, 1829 .

DIAGNOSIS: Promecoderus belongs to the same group as does Cascellius , and is characterized by the following combination of characters: submentum with four to six setae, lateral umbilical series with four setae, and fore and middle male tarsomeres with adhesive vestiture.

DESCRIPTION: External morphology (figs. 298, 299, 305, 317–319): One supraorbital puncture bearing one seta on each side of head; vertex with or without transverse groove; temporal ridge complete or absent from middle; antenna with antennomeres 1– 2 glabrous, 3 with apical ring or setose on apical half, 4 setose on apical half or complete setose, 5–11 pubescent all along surface. Mandibles with scrobal setae. Maxillary eustipes with two basal setae; maxillary palpomere 2 asetose. Labium with four to six setae on submentum; mentum bisetose, with tooth simple, small; prementum with glossal sclerite quadrisetose, ventral surface without longitudinal medial carina, with paramedian areas deep-set; labial palpomere 2 bisetose. Pronotum not constricted apically, with two setae each side. Elytron with parascutellar striole joined to apical portion of stria 1; with parascutellar seta on basal portion of second stria; lateral umbilical series with four setae. Abdominal sterna 4–7 smooth, with two lateral foveae each side (fig. 317) or extended mediad as a groove (fig. 318). Foretrochanter bisetose; male foretarsomeres 1–4 or 1–3 and middle tarsomeres 1–2 with adhesive vestiture ventrally.

Male genitalia (figs. 300–303, 306–310, 312–315, 320–321, 324): Sternum 9 with complete sclerotized ring (fig. 313, 314) or incomplete (figs. 300, 306, 307) with lateral margins narrow. Median lobe with dorsal surface unsclerotized and basal orifice completely open dorsally (figs. 303, 312, 315, 321, 324). Internal sac without sclerite X, Y, and apical plate; armature of groups of small spiculae (figs. 310, 320), in some species with a sclerotized plate near the middle (figs. 324). Left paramere asetose (figs. 301, 308); right setose (figs. 302, 309). Some species exhibit an inversion of the aedeagus (figs. 321) that renders the left paramere setose and the right asetose.

Female genital tract (figs. 304, 311, 316, 322, 323, 325): Rami of gonocoxite 9 small. Subgonocoxite 9 broad, with ensiform setae; gonopod 9 short and broad, without nematiform setae on subapical setose organ. Spermatheca broad (figs. 311, 316, 322, 325, sp); median oviduct and spermatheca enter together (figs. 311, 322) or separated into bursa copulatrix; helminthoid sclerite absent. Bursa copulatrix long; ligula sclerotized (figs. 323) or not.

LARVAE: The larva of P. concolor was described by Moore (1964).

HABITAT: The species of Promecoderus occupy different habitats, ranging from the rainforest ( Nothofagus ) to dry open woodland (Darlington, 1961; Lawrence et al., 1987) or dry sclerophyll zone and open scrub (Sloane, 1890; Lawrence et al., 1987).

GEOGRAPHICAL DISTRIBUTION: Promecoderus occurs in Tasmania and southeastern Australia.

REMARKS: The genus Promecoderus is one of the most diverse broscine genera, with 44 species, and there is no clear synapomorphy of the group. For this reason, I treat the genus as five different groups of species in attempting to prove the monophyly of Promecoderus . This genus needs a full revision. I consider only those species in which males have an adhesive vestiture ventrally on fore and middle tarsomeres 1 and 2 as belonging to Promecoderus . The species considered as a Tasmanian group by Sloane (with adhesive vestiture on foretarsomeres and without it on the middle tarsomeres) are considered here as belonging to Acallistus . The species without adhesive vestiture on fore and middle tarsomeres are considered here as belonging to Anheterus .

MATERIAL EXAMINED: The species studied, arranged by natural groups proposed by Sloane (1890), are as follows:

Group A, brunnicornis group (figs. 300– 306): P. brunnicornis (MCZ) . This group is characterized by the presence of a row of setae on male sterna 4–7 and two setae each side of sternum 8.

Group B, concolor group (figs. 307–314): P. concolor (AMNH) . This group is characterized by elytra smooth, and male foretarsomeres 1-3 with adhesive vestiture ventrally. Other characters are: no row of setae on male sterna 4–7 and sternum 8 with one setae each side.

Group C, clivinoides group (figs. 315– 318): P. comes (AMNH) , P. dyschiroides (MCZ) and P. aff. clivinoides (AMNH) . This group is characterized by: no row of setae on male sterna 4–7; one setae each side of sternum 8; elytra striate; male foretarsomeres 1– 4 slender, with adhesive vestiture ventrally.

Group D, albaniensis group (figs. 326– 327): P. albaniensis ( AMNH; MCZ). This group is characterized by no row of setae on male sterna 4–7; one setae each side of sternum 8; elytra striate; abdominal sterna 4–7 with deep rounded lateral foveae, without a transverse sulcus; male foretarsomeres 1–4 broad, with adhesive vestiture ventrally.

Group E, gibbossus - subdepressus - inornatus group (figs. 321–325): P. subdepressus (AMNH) , P. gibbosus (MCZ, USNM) , P. aff. elegans (AMNH) , P. inornatus (MCZ) . This group is easily recognized because its species have foveo-sulci on abdominal sterna 4–7, a prominent sulcus dorsally posterior to the eyes, and a sclerotized plate in the internal sac. Also some species of this group have the aedeagus inverted (fig. 321), where the left paramere has the shape and setae of the right, and the right is asetose and with the shape of the left.

Acallistus Sharp, 1886 Figures 326–335

Acallistus Sharp, 1886: 362 . Broun 1893: 983. Sloane, 1920: 127.

Promecoderus : Britton, 1949: 537. Lawrence et al. (part), 1987: 109.

TYPE SPECIES (by monotypy): Acallistus simplex Sharp, 1886 (junior synonym of Promecoderus tasmanicus Castelnau, 1867 ).

DIAGNOSIS: Acallistus belongs to the same group as does Cascellius . It differs from South American creobiines in having submentum quadrisetose, from Bountya and Anheterus in having male foretarsomeres with adhesive vestiture, and from Promecoderus in lacking adhesive vestiture on middle tarsomeres.

DESCRIPTION: External morphology (figs. 326–328, 330): One supraorbital puncture each side bearing one seta on each side of head; vertex with shallow transverse groove; temporal ridge complete or absent from middle; antenna with antennomere 1 and 2 glabrous, 3 with apical ring of setae, 4 setose on apical half, 5–11 setose all along surface. Mandibles each with scrobal setae. Maxillary eustipes with two basal setae; maxillary palpomere 2 asetose. Labium submentum quadrisetose; mentum tooth reduced to a small convexity ( A. tasmanicus , A. plebius ) (fig. 327) or distinctly developed ( A. longus ) (fig. 330); prementum with glossal sclerite quadrisetose, ventral surface without longitudinal medial carina, with the paramedian areas deep-set (fig. 328); labial palpomere 2 bisetose. Pronotum not constricted apically, with two setae each side. Elytron with parascutellar striole joined to apical portion of stria 1; with parascutellar seta on basal portion of second stria; lateral umbilical series with four setae. Abdominal sterna 4–7 smooth, with a shadow of lateral fovea. Foreleg trochanter bisetose; male foretarsomeres 1–3 or 1–4 with adhesive vestiture ventrally; middle tarsomeres without it.

Male genitalia (figs. 331–335): Sternum 9 with a complete ring, a portion not sclerotized, and lateral margins narrow (figs. 331, 332). Median lobe with dorsal surface unsclerotized and basal orifice completely open dorsally (fig. 335). Internal sac without sclerite X, Y, and apical plate. Left paramere asetose (fig. 333); right setose on two apical thirds (fig. 334).

Female genital tract (fig. 329): Rami of gonocoxite 9 short. Subgonocoxite 9 broad, with ensiform setae; gonopod 9 short, without nematiform setae on subapical setose organ. Median oviduct and spermatheca entering separately into bursa copulatrix; helminthoid sclerite absent. Bursa copulatrix long; without ligula.

HABITAT: The species inhabit tall forest (Lawrence et al., 1987).

GEOGRAPHICAL DISTRIBUTION: The genus Acallistus occurs in Tasmania. Sharp (1886) reported its occurrence in New Zealand, but Britton (1949) stated that this record was erroneous.

REMARKS: The genus was recognized originally for a single species, Acallistus simplex Sharp 1886 . It was later considered by Britton (1949) as junior synonym of Promecoderus tasmanicus Castelnau, 1867 . Sloane (1890) revised the genus Promecoderus , and stated that P. tasmanicus constitutes a natural group together with other species. In this contribution, I consider this group of species as genus Acallistus , different from Promecoderus .

MATERIAL EXAMINED: Acallistus tasmanicus (MCZ) , A. plebius ( NEW COMBINATION) ( USNM), A. longus ( NEW COMBINATION) ( MCZ), A. cuprescens ( NEW COMBINATION) ( MCZ), and A. aff. longus (ANIC) .

Anheterus Putzeys, 1868 Figures 336–342

Anheterus Putzeys, 1868: 345 ; 1873: 337. Csiki, 1928: 23.

Promecoderus : Sloane, 1890 (part): 192; Lawrence et al. (part), 1987: 109.

TYPE SPECIES (by subsequent monotypy in Putzeys, 1868): Promecoderus gracilis Germar, 1848 .

DIAGNOSIS: Anheterus belongs to the same group as does Cascellius . Anheterus and Bountya are the only genera of this group without adhesive vestiture on male foretarsomeres. Anheterus differs from Bountya in having four setae on lateral umbilical serie.

DESCRIPTION: External morphology (figs. 336, 337): One supraorbital puncture bearing one seta on each side of head; vertex without transverse groove; temporal ridge complete; antenna with antennomeres 1–2 glabrous, 3 with apical ring of setae; 4 setose on apical half, 5–11 pubescent all along surface. Mandibles each with scrobal setae. Maxillary with eustipes having two basal setae; maxillary palpomere 2 asetose. Labium with submentum quadrisetose; tooth of mentum simple; prementum with glossal sclerite quadrisetose apically, ventral surface without longitudinal medial carina, with the paramedian areas deep-set; labial palpomere 2 bisetose. Pronotum not constircted apically, with two setae each side. Elytron with parascutellar striole joined to apical portion of stria 1; with parascutellar setae on basal portion of second stria; lateral umbilical series with four setae. Abdominal sterna 4–7 with a lateral foveae each side. Foreleg trochanter bisetose; male fore and middle tarsomeres without adhesive vestiture.

Male genitalia (figs. 338–341): Sternum 9 with sclerotized ring complete with lateral margins narrow (fig. 338). Median lobe with dorsal surface unsclerotized and basal orifice completely open dorsally (figs. 341). Internal sac without sclerite X, Y, and apical plate. Left paramere asetose (fig. 339); right setose on apical two thirds (fig. 340).

Female genital tract (figs. 342): Rami of gonocoxite 9 short. Gonopod 9 short, without nematiform setae on subapical setose organ. Spermatheca broad (fig. 342, sp); median oviduct and spermatheca entering together into bursa copulatrix; helminthoid sclerite absent. Bursa copulatrix long; ligula small, unsclerotized.

HABITAT: The species inhabit open scrub and low open woodland (Lawrence et al., 1987).

GEOGRAPHICAL DISTRIBUTION: The genus Anheterus occurs in South Australia (Adelai- de) and ‘‘ Northern Territory of Southern Australia ’’ (Sloane, 1892).

REMARKS: This genus, as redefined herein, contains three species, A. gracilis , A. distinctus (Sloane, 1890) , and A. ambiguous (Sloane, 1892), considered by Sloane (1890, 1892) as a natural group of species within the genus Promecoderus .

MATERIAL EXAMINED: Anheterus gracilis (AMNH, ANIC, MCZ) .

Figures

Figs. 290–297. Creobius eudouxy. (290) Head and prothorax; (291) mouthparts, ventral view; (292) sternum 9, ventral view; (293) lateral view; (294) left paramere; (295) right paramere; (296) median lobe, lateral view, showing internal sac; (297), female genital tract, ventral view.

Figs. 298–318. Promecoderus. (298–304), group A, Promecoderus brunnicornis: (298) head and prothorax; (299) mouthparts, ventral view; (300) sternum 9, ventral view; (301) left paramere; (302) right paramere; (303) median lobe, lateral view; (304) female genital tract, ventral view. (305–312) Group B, Promecoderus howitti: (305) mouthparts, ventral view; (306) sternum 9, ventral view; (307) lateral view; (308) left paramere; (309) right paramere; (310), apex of median lobe, lateral view showing internal sac; (311) female genital tract, ventral view; (312) Promecoderus lucidicollis, median lobe, lateral view. (313–316) Group C, Promecoderus dischiroides: (313) sternum 9, ventral view; (314) lateral view; (315) median lobe, lateral view; (316) female genital tract, ventral view. (317) Sternites of Promecoderus clivinoides; (318) sternites of Promecoderus gibbosus.

Figs. 319–325. Promecoderus. (319–323), group E, Promecoderus gibbosus: (319) head and prothorax; (320) median lobe, lateral view showing internal sac; (321) median lobe, lateral view showing inversion of parameres, (322) female genital tract, ventral view; (323) ligula of female genital tract. (324, 325) Group D, Promecoderus albaniensis: (324) median lobe, lateral view showing the internal sac; (325) female genital tract, ventral view.

Figs. 326–342. Acallistus and Anheterus. (326–342) Acallistus plebius: (326) head and prothorax; (327) mouthparts, ventral view; (328) ligula, ventral view; (329) female genital tract, ventral view. (330– 335) Acallistus longus: (330) mouthparts, ventral view; (331) sternum 9, ventral view; (332) lateral view; (333) left paramere; (334) right paramere; (335) median lobe, lateral view. (336–342) Anheterus gracilis: (336) head and prothorax; (337), mouthparts, ventral view; (338) sternum 9, ventral view; (339) left paramere; (340) right paramere; (341) median lobe, lateral view; (342) female genital tract, ventral view.