Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae)

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1 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) Tomáš LACKNER The Hokkaido University Museum, N10 W8, Sapporo, Japan, Abstract. The genus Terametopon Vienna, 1987, consisting of two subgenera Terametopon and Psammoprinus Gomy & Vienna, 1996, is revised herein. It now contains seven species, one of which, T. (T.) foveatus sp. nov. is described as new. The species T. truncatus Gomy & Vienna, 1999, known from a single male with severely damaged body (head and prothorax missing), has not been incorporated into this study and is retained as Terametopon incertae sedis. Male genitalia and SEM photographs of all known species (except for T. truncatus) are given. A key to the species is provided. Taxonomy, new species, Coleoptera, Histeridae, Saprininae, Terametopon, Afrotropical INTRODUCTION VIENNA (1987) established the genus Terametopon Vienna, 1987 in a paper dealing with several new Afrotropical species of histerids from the collections of the Transvaal Museum (Pretoria, South Africa). He included two new species in the newly established genus: Terametopon levissimestriatus Vienna, 1987 (type species of the genus) and T. endroedyi Vienna, Both taxa were known from only two specimens per species. Later MAZUR (1993) described another new species, T. namibiensis Mazur, 1993, based on a single female. GOMY & VIENNA (1996) revised the genus, and, based on the specimens from the Paris and Transvaal museums, they depicted male genitalia of T. namibiensis for the first time, and described another new species, T. deserticola Gomy & Vienna, In their revision, they divided the genus into two subgenera; this division being made on the basis of the presence/ absence of a peculiar frontoclypeal projection. Species possessing such a projection (unique with the subfamily Saprininae) remained in the nominotypical subgenus, whilst species without it were placed in a newly established subgenus Psammoprinus Gomy & Vienna, 1996, with T. namibiensis as its type species. Later, these two authors (GOMY & VIENNA 1999) added two more species into this genus; one, T. (T.) prominentinotus Gomy & Vienna, 1996 has been included in the nominotypical subgenus, but the other one, T. truncatus Gomy & Vienna, 1996, owing to the severe damage of its body, has not been placed into either of the subgenera. During my visit to South Africa in 2008 I have had the chance to visit the South African National Collection of Insects as well as the Transvaal Museum, both located in the capital, Pretoria. Almost all type specimens were studied, some borrowed and additional undetermined material has been discovered, mostly at the South African National Collection of Insects. Furthermore, the holotype of T. (P.) namibiensis has been borrowed from Stuttgart (Germany) and studied. This paper presents a revision of all known species included in the genus, as well as description of a new species. However, species T. truncatus has been omitted from this study, because (although it is a male) its thorax and head are missing and the key -43-

2 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) characters for the subgeneric/species placement are found on head and prothorax. It is herein retained as Terametopon incertae sedis. Species of the genus Terametopon are generally very rare in collections and are not readily collected, even in their preferred habitat (see GOMY & VIENNA 1999: 5). They are known almost exclusively from the dunes of the Namib Desert, Namibia, with the exception of a one specimen of T. (P.) namibiensis captured in Botswanan part of Kalahari Desert and T. (T.) foveatus sp. nov. from the Namibian part of the Kalahari Desert. The biology of these species is virtually unknown, some specimens were collected in baited pitfall traps. Their morphology, however, suggests that they belong to the group of specialized psammophilous histerids. MATERIAL AND METHODS The beetles, after being removed from the original cards were side-mounted on triangle cards and observed under a binocular microscope NIKON 102 with diffuse light. Male genitalia were first macerated in 10% KOH solution for about 15 minutes, cleared in 80% alcohol, macerated in lactic acid with fuchsine heated to 60 ºC for another two hours. Then they were treated with aceto-salycilate heated to 60 ºC for 15 minutes and cleared in xylene. They were subsequently observed in α-terpineol in a small dish. Digital photographs were taken by a Nikon 4500 Coolpix camera and edited in Adobe Photoshop CS3. Based on the photographs, observing the actual genitalia, pencil art was drawn; pen art followed, retracing the pencil art and making minor corrections. For the dissection of mouthparts and observations of antennae, the entire head capsule has been removed from a relaxed specimen and treated in the same sequence of chemical processes as the male genitalia. Scanning Electron Microscopy (SEM) photographs were taken at the Faculty of Agriculture, Hokkaido University, Sapporo, Japan on a JSM 6301F scanning electron microscope. Codens of museum collections: SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany (Wolfgang Schawaller); TMSA Transvaal Museum of Natural History, Pretoria, South Africa (Ruth Müller); SANC South African National Collection of Insects, Pretoria, South Africa (Riaan Stals); CVIEN Private collection of Pierpaolo Vienna, Venezia, Italy. Abbreviations used in the measurements are as follows: PEL length between anterior angles of pronotum and apices of elytra; APW width between anterior angles of pronotum; PPW width between posterior angles of pronotum; EL length of elytron along sutural line; EW maximal width between outer margins of elytra. Separated lines of the same label are marked by slash (/). -44-

3 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN RESULTS Genus Terametopon Vienna, 1987 Terametopon Vienna, 1987: 217. GOMY & VIENNA 1996: 331 (key to species); MAZUR 1997: 266. Type species: Terametopon levissimestriatus Vienna, 1987 (by monotypy). Diagnosis. The genus Terametopon stands out as the only Subsaharan genus of the subfamily Saprininae that exhibits true psammophilous morphological adaptations: strongly convex body, ventral side of the body with dense vestiture, setose elytral epipleuron, outer margin of meso- and metatibiae with long dense denticles, long protarsal claws. Species of the nominotypical subgenus possess a curious frontoclypeal projection (Figs. 4, 20, 28, 44), not found in any other genus of Saprininae. Species of this genus resemble somewhat superficially several specialized Palaearctic genera, e.g. Xenonychus Wollaston, 1864, Exaesiopus Reichardt, 1926 and Xenophilotis Kryzhanovskij, 1987, but differ from all of them by various combinations of characters. Since the phylogeny of the African genera of Saprininae has as yet not been provided, it seems inappropriate to discuss taxon relationships at this point. Description. Maximum length (PEL) of beetles is 3.0 mm. Body strongly convex, oval to rectangular oval, elytral humeri prominent, dorsal surface punctate; cuticle ochre, reddishbrown to brown. Pronotal sides moderately to strongly narrowing apically, anterior angles usually acute; pronotal marginal stria reduced to anterior pronotal angles in species of the subgenus Psammoprinus, otherwise complete. Elytral striae thin, occasionally erased by coarse punctation. Palpal organs (sensu KOVARIK & TISHECHKIN 2004) of labial and maxillary palpi well developed, visible; mentum without notch on anterior margin; distal of maxillary palpomere three to five times longer than penultimate palpomere. Penultimate labial palopomere with single long seta. Antennomere before antennal club not saucer-like, resembling other antennomeres of flagellum. Antennal club depressed dorsoventrally. Underside of body with dense vestiture, elytral epipleuron setose. Preapical foveae of prosternum present or absent. Prosternal striae (carinal and lateral) present or absent; prosternal process setose or glabrous. Meso-metaventral sutural stria (sensu KANAAR 1997) lacking. Species of the nominotypical subgenus with curious prominent frontoclypeal projection, unique for the subfamily. Protibia with two large distal teeth bearing a short denticle followed by another three to four low teeth topped with short denticle; protibial spur well developed, bent, originating from near tarsal base. Meso-and metatibiae with long, dense denticles on outer margin, tarsal claws thin, usually long. Coxites of 8 th abdominal segment of male genitalia on apical part with short setae, laterally or dorsolaterally, as well as with apical inflatable membrane. Parameres of aedeagus fused on basal half to basal two-thirds. -45-

4 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) Subgenus Terametopon Vienna, 1987 Terametopon Vienna, 1987: 217. GOMY & VIENNA 1996: 331 (key to species); MAZUR 1997: 266. Type species: Terametopon levissimestriatus Vienna, 1987 (by monotypy). Terametopon (Terametopon) levissimestriatus Vienna, 1987 (Figs. 1-9, 10-16) Terametopon levissimestriatus Vienna, 1987: 218., Type locality: Northern Namibia, Awahuab dunes. Terametopon (Terametopon) levissimestriatus: GOMY & VIENNA 1996: 331 (keyed), figs. 5, 11, 14; MAZUR 1997: 266. Type specimen examined. Paratype (TMSA), mounted on its side, right mesotibia missing, with genitalia mounted in Canada Balsam on a separate slide, with printed label: S.W.Afr., Kaokoveld / Kunene R. W. Hartnog / S E, with consecutive printed label: ; E-Y: 2066 / dunes & humm., day / leg. Perinth, Müller, with consecutive printed label: Terametopon levissimestriatus / Vienna / P. Vienna det. 1985, followed by another red printed label: Paratype, and another yellow pencil label: , added by myself. Redescription. Body length. PEL mm, APW mm, PPW mm, EL mm, EW mm. Body roundly oval, head moderately sized, elytral humeri prominent, moderately convex, cuticle light reddish-brown, elytra lighter than head and pronotum; legs, antennae and mouthparts reddish-brown. Antennal scape dilated, strongly thickened, upper surface densely, coarsely punctate with numerous dense long setae; club round, depressed dorsoventrally, without visible articulation, surface imbricate, almost completely glabrous apart from one deep, almost circular slit-like sensory area on ventral surface (Fig. 5); surface proximal to it fringed with very short sparse setae; Reichardt s organ not examined. Mouthparts. Labrum (Fig. 4) very curiously shaped for the subfamily, about as long as wide, semicircular with anteromedian projection, basally strongly elevated, labral disc flat, with microscopic granulose punctation, labral pits absent; mandibles coarsely and densely punctate, mandibular apex very acutely pointed, subapical tooth present only on left mandible, rather small; maxillary cardo with few short setae on lateral margin; stipes triangular, with three long well sclerotized setae; distal maxillary palpomere strongly thickened, its width almost equals its length, approximately five times as long as penultimate palpomere, palpal organ visible; distal labial palpomere thickened, width almost equal length, apex with sensory area, palpal organ visible, distal labial palpomere with one very long seta; mentum rectangular, anterior margin straight, without median notch, with deep punctures fringed with long setae, lateral margins glabrous, disc with regular circular wrinkles, almost glabrous, only with two microscopic setae; other parts of the mouth not examined. Clypeus very short, almost vertical, hidden under massive median frontoclypeal projection (Fig. 4) in a form of irregular truncated tetrahedron with rounded anterior margin and constricted base, frontal stria rather thick, present only on frontal projection, supraorbital stria swollen on apical half, carinate, curved; disc of frons rugulose-punctate; eyes flattened, almost invisible from above. -46-

5 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Figs. 1-9: Terametopon (Terametopon) levissimestriatus Vienna. 1 habitus, dorsal view; 2 same, ventral view; 3 mesotibia, dorsal view; 4 head, labrum, dorsal view; 5 antennal club, ventral view; 6 prosternum; 7 mesoventrite; 8 propygidium and pygidium; 9 protibia, ventral view. -47-

6 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) Pronotal sides (Fig. 1) strongly convergent anteriorly; apical angles blunt, anterior emargination for head shallow; pronotal epipleuron with dense yellow setae; marginal stria fine, complete and feebly carinate on outer side; disc entirely covered with very coarse, dense punctation, punctures separated by less than half their own diameter forming elongate wrinkles laterally; scutellum very small, almost invisible. Elytral epipleura with dense thin yellow setae; marginal epipleural stria strongly impressed, complete; marginal elytral stria well impressed, carinate on outer side, continued as weakened, almost complete apical elytral stria; oblique humeral stria weakly impressed on basal fourth; internal subhumeral stria weakly impressed, present as a very short median fragment; disc with four weakly impressed dorsal elytral striae 1-4, first stria abbreviated basally, second stria with a basal hook; first and second striae about the same length, reaching about elytral half apically, third stria a little longer, apically little surpassing elytral half, fourth stria basally vaguely connected with complete thin sutural elytra stria, reaching elytral length apically. Elytral disc (Fig. 1) (except for smooth elytral humeri) wholly covered with very coarse and dense round punctation, punctures separated by less than their own diameter, becoming sparser laterally; punctation becomes confluent before elytral apex. Propygidium (Fig. 8) almost completely exposed, with very dense shallow confluent punctures separated by less than half their own diameter; pygidium with similar confluent punctation, punctures getting finer apically. Anterior margin of median portion of prosternum (Fig. 6) almost straight; marginal stria strongly impressed, interrupted laterally; prosternal process flattened, somewhat concave, dorsolaterally with fine punctures separated several times their diameter, interspaces striolate; preapical foveae absent; carinal striae well impressed, approximate, apically united in a rounded loop; descending lateral striae strongly carinate on outer side, convergent anteriorly, their apices not attaining united carinal striae. Anterior margin of mesoventrite (Fig. 7) deeply emarginate medially; marginal stria laterally well impressed, anteriorly widely absent; disc with double punctation, large, sparse shallow punctures with microscopic setae intermingled with sparse tiny punctures; mesometaventral sutural stria absent, meso-metaventral suture thin, somewhat curved outwardly; intercoxal disc of metaventrite with shallow longitudinal median depression, punctation similar to that of mesoventrite. Lateral metaventral stria well impressed, carinate on outer side, straight; lateral disc of metaventrite excavate, with shallow large setiferous punctures; metepisternum evenly covered with much coarser and denser punctation, punctures almost unrecognizable under long dense amber setae. Intercoxal disc of first abdominal sternite with almost complete striae laterally; disc on basal third with punctation similar to that of metaventrite, punctation weakens medially; lateral sides of all visible abdominal sternites setose. Protibia dilated, outer margin with two large distal triangular teeth topped with short denticle, followed by three low teeth topped with short denticle diminishing in size in proximal direction; outer part of ventral surface of protibia (Fig. 9) with shallow sparse punctures and wrinkles, median and inner parts of ventral surface smooth; inner margin of protibia with a sparse row of short setae; apical margin of protibia without denticles; tarsi thin, claws of apical tarsomere about equal its length; protibial spur long, bent, growing out from near tarsal base. -48-

7 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Mesotibia (Fig. 3) slender, outer margin with a sparse row of long denticles growing in size in proximal direction, inner-lateral surface next to outer margin with one row of dense long well sclerotized setae; outer-lateral margin with one row of short stout denticles; outerlateral surface smooth; apical margin with four strongly sclerotized denticles, inner most one the longest, several times as long as other three; claws of apical tarsomere almost straight, Figs : Terametopon (Terametopon) levissimestriatus Vienna, male genitalia. 10 coxites of 8 th segment, ventral view; 11 same, dorsal view; 12 same, lateral view; 13 9 th, 10 th tergite, dorsal view, spiculum gastrale, ventral view; 14 same, lateral view; 15 aedeagus, dorsal view; 16 same, lateral view. -49-

8 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) longer than half its length; metatibia in all aspects similar to mesotibia, but denticles on outer margin sparser. Male genitalia. Coxites of 8 th segment (Figs ) longitudinally separated medially, apically with inflatable membrane with dense minute setae; ventrally with sparse pores; coxites on apical fourth ventrally and laterally with numerous short setae. Tergite of 8 th segment and coxites fused laterally (Fig. 12). Morphology of 9 th segment (Figs ) typical for the subfamily; spiculum gastrale (ventral sclerite of 9 th segment) expanded on both ends. Basal piece of aedeagus (Fig. 15) (phallobase) well developed, ratio of its length: length of parameres approximately 1 : 1.5; parameres fused almost along basal half; aedeagus curved ventrad (Fig. 16). Biology. Unknown. Remarks. VIENNA s (1987) description mentions the presence of preapical foveae in this species; however, the borrowed male paratype does not possess them. P. Vienna (pers. comm.) confirmed that during the study of the type material he mistakenly mentioned the presence of preapical foveae in this species. Distribution. Namibia. Terametopon (Terametopon) foveatus sp. nov. (Figs ) Type locality: Namibia, Kalahari Desert, Chulon Farm, Narib Ost, S E. Type specimen examined. Holotype (CVIEN), mounted on its side on a triangular point, left antennal flagellum missing, apical tarsomere of left protarsus missing, left metatarsus missing, right metatarsus broken off, glued to the same triangular point as specimen, female genitalia and pygidium glued to the same triangular point as the specimen; with handwritten label: S. Afr. n o 1934 / leg. Endrödy-Younga ; with consecutive printed label: Terametopon levissimestriatus / Vienna / P. Vienna det. 1987, with consecutive red label, printed: HOLOTYPUS / Terametopon (Terametopon) / foveatus sp. nov. / Des. T. Lackner 2009, with a following pencil yellow label: D08-089, added by myself. According to P. Vienna and Ruth Müller (TMSA) Endrödy-Younga s locality n o 1934 stands for: Namibia / Kalahari, fm Chulon / Narib Ost, S E / groundtraps / 14.viii.1982 / leg. Penrith. Description. Body length. PEL mm, APW mm, PPW mm, EL mm, EW mm. Body rectangular oval, head moderately sized, elytral humeri prominent, moderately convex, cuticle light reddish-brown, elytra lighter than head and pronotum; legs, antennae and mouthparts reddish-brown. Antennal scape dilated, strongly thickened, upper surface densely, coarsely punctate with numerous dense long setae; club (Fig. 21) round, depressed dorsoventrally, without visible articulation, surface imbricate, almost completely glabrous apart from one deep, almost circular slit-like sensory area on ventral surface (Fig. 22); surface proximal to it fringed with very short sparse setae; Reichardt s organ not examined. Mouthparts. Labrum very curiously shaped for the subfamily, about as long as wide, semicircular with anteromedian projection, basally strongly elevated, labral disc flat, with microscopic granulose punctation, labral pits absent; mandibles coarsely and densely punctate, mandibular apex very acutely pointed, subapical tooth present only on left mandible, rather small; cardo of maxilla with few short setae on lateral margin; stipes triangular, with three long well sclerotized setae; distal maxillary palpomere strongly thickened, its width almost -50-

9 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN equals its length, approximately five times as long as penultimate, palpal organ visible; distal labial palpomere thickened, its width almost equals its length, apex with sensory area, palpal organ visible, penultimate labial palpomere with one very long seta; mentum rectangular, anterior margin straight, without median notch, with deep punctures fringed with long setae, Figs : Terametopon (Terametopon) foveatus sp. nov. 17 habitus, dorsal view; 18 same, ventral view; 19 metatibia, dorsal view; 20 head, dorsal view; 21 antennal club, dorsal view; 22 same, ventral view; 23 pygidium; 24 prosternum. -51-

10 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) lateral margins with two or three short setae, disc regular with circular wrinkles, almost glabrous, only with two microscopic setae; other parts of the mouth not examined. Clypeus very short, almost vertical, hidden under massive cube-like frontoclypeal projection (Fig. 20), frontal stria fine, shortened, present only on frontoclypeal projection, supraorbital stria well impressed, little carinate, curved; disc of frons rugulose-punctate; eyes very flattened, invisible from above. Pronotal sides (Fig. 17) strongly convergent anteriorly; apical angles acute, anterior emargination for head deep, semicircular; pronotal epipleuron with dense yellow setae; marginal stria fine, complete and little carinate on outer side; disc entirely covered with very coarse and dense punctation, punctures separated by less than half their own diameter forming deep elongate wrinkles laterally; scutellum very small, almost invisible. Elytral epipleura rugulose-punctate, punctures with sparse thin yellow setae; marginal epipleural stria well impressed, complete; marginal elytral stria well impressed, carinate on outer side, continued as weakened but complete apical elytral stria; oblique humeral stria doubled, faintly impressed on basal fourth; internal subhumeral stria weakly impressed, present as a very short median fragment; disc with four thin dorsal elytral striae 1-4, about the same length, little surpassing elytral length apically, fourth stria basally connected with complete thin sutural elytral stria, reaching elytral apex. Elytral disc wholly covered with regular round punctation, punctures separated by two to several times their diameter, becoming sparser laterally; before elytral apex punctation becomes denser, interspaces striolate. Propygidium almost completely exposed, with very dense shallow punctures separated by less than half their own diameter; pygidium (Fig. 23) with similar but somewhat sparser punctation, punctures getting finer apically. Anterior margin of median portion of prosternum (Fig. 24) almost straight; marginal stria strongly impressed, interrupted laterally; prosternal process flattened, somewhat concave, dorsolaterally with coarse punctures separated several times their diameter, interspaces striolate; preapical foveae large and deep; carinal striae well impressed, convergent on prosternal apophysis, thence slightly divergent apically and united in a rounded loop; descending lateral striae strongly carinate on outer side, slightly convergent anteriorly, their apices not attaining united carinal striae. Anterior margin of mesoventrite deeply emarginate medially; marginal stria laterally well impressed, anteriorly widely absent; disc with sparse wide punctures of different sizes; mesometaventral sutural stria absent, meso-metaventral suture thin, somewhat curved outwardly; intercoxal disc of metaventrite with punctation similar to that of mesoventrite, punctures becoming sparser and finer laterally. Lateral metaventral stria well impressed, carinate on outer side, obliquely arcuate, stopping short of metacoxa; lateral disc of metaventrite excavate, with shallow large setiferous punctures; metepisternum evenly covered with much coarser and denser punctation, punctures separated by less than half their own diameter, fringed with long amber setae. Intercoxal disc of first abdominal sternite with almost complete striae laterally; disc with punctation similar to that of metaventrite; lateral sides of all visible abdominal sternites setose. Protibia dilated, outer margin with two large distal triangular teeth topped with short denticle, followed by three low teeth topped with short denticle diminishing in size in -52-

11 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN proximal direction; outer part of ventral surface of protibia with sparse punctures and shallow wrinkles, median and inner parts of ventral surface smooth; inner margin of protibia with a sparse row of short setae; apical margin of protibia without denticles; tarsi thin, claws of apical tarsomere about equal its length; protibial spur long, bent, growing out from near tarsal base. Mesotibia slender, outer margin with a sparse row of long denticles growing in size in proximal direction, inner-lateral surface next to outer margin with one row of dense long well sclerotized setae; outer-lateral margin with one row of short stout denticles; outer-lateral surface smooth; apical margin with four strongly sclerotized denticles, the inner most one the longest, several times as long as other three; claws of apical tarsomere almost straight, longer than half its length; metatibia (Fig. 19) somewhat thicker than mesotibia, medially with one long denticle, followed by three much longer denticles near tarsal insertion; innerlateral surface with a single row of long, well sclerotized setae; outer-lateral margin and surface similar to mesotibia, but denticles on outer-lateral margin much sparser; claws of apical tarsomere straight, about half its length; metafemora not thickened. Biology. Unknown. Etymology. Specific epithet of this species refers to well developed foveae of the prosternal process. Distribution. Namibia, Kalahari Desert. Differential diagnosis. Most similar to Terametopon (T.) endroedyi, differing from it chiefly by differently shaped frontoclypeal projection, well impressed all four dorsal elytral striae (only first and second striae are well impressed with T. (T.) endroedyi), and punctures of mesoventrite not fringed with microscopic setae. Terametopon (Terametopon) endroedyi Vienna, 1987 (Figs , 34-40) Terametopon endroedyi Vienna, 1987: 220., Type locality: Namibia, dunes west of Felseneck. Terametopon (Terametopon) endroedyi: GOMY & VIENNA 1996: 331 (keyed), figs. 7, 8; MAZUR 1997: 266. Type specimen examined. Holotype (TMSA), mounted on its side on a triangular point, right metatarsus missing, with genitalia mounted in Canada Balsam on a separate slide, with printed label: S.W.Afr. Namib / dunes W. Felseneck / 24.12S 15.50E, with another printed label: ; E-Y: 863A / groundtraps / leg. Endrödy- Younga, with another printed label: groundtraps with / faeces bait, with another black- ink written label with male symbol, followed by another red label, printed: HOLOTYPUS / Terametopon endroedyi / sp. nov. / det. P. Vienna 1984, and another red-margined label, printed: HOLOTYPUS / Terametopon / endroedyi / P. Vienna, followed by another yellow pencil label: , added by myself. Additional specimen examined. NAMIBIA: SE Tsondab Plains / SE CC S4 / 15.vii.1976 / Dept. of Entomology / University of Pretoria (1 SANC). Redescription. Body length. PEL mm, APW mm, PPW mm, EL mm, EW mm. Body roundly oval, moderately convex, head moderately sized, elytral humeri prominent, cuticle ochre-colored, elytra lighter than head and pronotum; legs, antennae and mouthparts yellowish-brown. -53-

12 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) Antennal scape dilated and strongly thickened, upper surface densely, coarsely punctate with numerous dense long setae; club round, depressed dorsoventrally, without visible articulation, surface imbricate, almost completely glabrous apart from one deep, almost circular slit-like sensory area on ventral surface topped with thick, tooth-paste-like structure (Fig. 27); surface proximal to it fringed with very short, sparse setae; Reichardt s organ not examined. Mouthparts. Labrum very curiously shaped for the subfamily, about as long as wide, semicircular with anteromedian projection, basally strongly elevated, labral disc flat, with microscopic granulose punctation, labral pits absent; mandibles coarsely and densely punctate, mandibular apex very acutely pointed, subapical tooth present only on left mandible, rather small; cardo of maxilla with few short setae on lateral margin; stipes triangular, with three long well sclerotized setae; distal maxillary palpomere strongly thickened, its width about half its length, approximately five times as long as penultimate palpomere, palpal organ visible, with a tiny sensory area on apex; distal labial palpomere thickened, width about half length, apex with tiny sensory area, palpal organ visible, penultimate labial palpomere with one very long seta; mentum rectangular, anterior margin straight, without median notch, with several long setae, lateral margins glabrous, disc with regular circular wrinkles, almost glabrous, only with two microscopic setae; other parts of the mouth not examined. Clypeus very short, almost vertical, hidden under massive triangular frontoclypeal projection with constricted base (Fig. 28), frontal stria unrecognizable, supraorbital stria swollen on apical half, carinate, curved; disc of frons coarsely rugulose-punctate; eyes flattened, almost invisible from above. Pronotal sides (Fig. 25) moderately convergent anteriorly; apical angles acute, anterior emargination for head shallow; pronotal epipleuron with dense yellow setae; marginal stria fine, complete and little carinate on outer side; disc entirely covered with very coarse and dense punctation, punctures separated by less than half their own diameter forming elongate wrinkles laterally; scutellum very small, almost invisible. Elytral epipleura with sparse thin yellow setae; marginal epipleural stria well impressed, complete; marginal elytral stria well impressed, carinate on outer side, continued as complete apical elytral stria; oblique humeral stria faintly impressed on basal fourth; only firsth and partly second dorsal striae recognizable, shortened, present only on basal third, other elytral striae erased by coriarious punctation; sutural elytral stria thin, complete, apically connected with apical stria. Elytral disc (except for smooth elytral humeri) wholly covered with coriarious punctation, punctures separated their own to twice their diameter; before elytral apex punctation disappears. Propygidium (Fig. 31) almost completely exposed, with very dense shallow confluent punctures separated by less than half their own diameter; pygidium with similar confluent punctation, punctures getting finer apically. Anterior margin of median portion of prosternum (Fig. 29) almost straight; marginal stria well-impressed, interrupted laterally; prosternal process flattened, somewhat concave, dorsolaterally with fine punctures separated several times their diameter, interspaces striolate; pre-apical foveae small, rather deep; carinal striae well impressed, divergent on apophysis, thence approximate, apically united in a rounded loop; descending lateral striae strongly carinate on outer side, convergent anteriorly, their apices not attaining united carinal striae. -54-

13 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Figs : Terametopon (Terametopon) endroedyi Vienna. 25 habitus, dorsal view; 26 same, ventral view; 27 antennal club, ventral view; 28 head dorsal, view; 29 prosternum; 30 mesoventrite; 31 propygidium and pygidium; 32 protibia, ventral view; 33 lateral disc of metaventrite and metepisternum. -55-

14 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) Anterior margin of mesoventrite (Fig. 30) deeply emarginate medially; marginal stria laterally well impressed, anteriorly widely absent; disc with double punctation: large, sparse shallow punctures with microscopic setae intermingled with sparse tiny punctures; mesometaventral sutural stria absent, meso-metaventral suture thin, almost straight; intercoxal disc of metaventrite almost flat, punctation similar to that of mesoventrite, becoming sparser laterally. Lateral metaventral stria shortened, obliquely arcuate, carinate on outer side; Figs : Terametopon (Terametopon) endroedyi Vienna, male genitalia. 34 coxites of 8 th segment, ventral view; 35 same, dorsal view; 36 same, lateral view; 37 9 th, 10 th tergite, dorsal view, spiculum gastrale, ventral view; 38 same, lateral view; 39 aedeagus, dorsal view; 40 same, lateral view. -56-

15 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN lateral disc of metaventrite (Fig. 33) excavate, with shallow large setiferous punctures; metepisternum evenly covered with much coarser and denser punctation, punctures almost unrecognizable under long dense amber setae. Intercoxal disc of first abdominal sternite with almost complete striae laterally; disc basally little depressed, on basal third with punctation similar to that of metaventrite, punctation weakens medially; lateral sides of all visible abdominal sternites setose. Protibia (Fig. 32) dilated, outer margin with two large distal triangular teeth topped with short denticle, followed by three low teeth topped with short denticle diminishing in size in proximal direction; outer part of ventral surface of protibia with sparse punctures and shallow wrinkles, median and inner parts of ventral surface smooth; inner margin of protibia with a sparse row of short setae; apical margin of protibia without denticles; tarsi thin, claws of apical tarsomere little longer than its length; protibial spur short, bent, growing out from near tarsal base. Mesotibia slender, outer margin with a sparse row of long denticles growing in size in proximal direction, inner-lateral surface next to outer margin with one row of dense long well sclerotized setae; outer-lateral margin with one row of short stout denticles; outer-lateral surface smooth; apical margin with four strongly sclerotized denticles, inner-most one the longest, several times as long as other three; claws of apical tarsomere almost straight, almost equal its length; metatibia in all aspects similar to mesotibia, but denticles on outer margin sparser; metafemori not thickened. Male genitalia. Coxites of 8 th segment (Figs ) longitudinally separated medially, apically with inflatable membrane with sparse minute setae; coxites on apical tenth ventrally and laterally with few short setae. Tergite of 8 th segment and coxites fused laterally (Fig. 36). Morphology of 9 th segment (Figs ) typical for the subfamily; spiculum gastrale (ventral sclerite of 9 th segment) expanded on both ends. Basal piece of aedeagus (Fig. 39) (phallobase) well developed, ratio of its length : length of parameres approximately 1 : 5; parameres fused almost along basal half; aedeagus slightly curved ventrad (Fig. 40). Biology. Unknown. Collected in sandy dunes by pitfall traps baited with feces. Remarks. VIENNA (1987) in the original description of this species mentions the presence of the pre-apical foveae, a fact that has been confirmed by the study of the holotype. In the paper by GOMY & VIENNA (1996), where a drawing of the prosternal region of this species is given, the pre-apical foveae are not depicted. P. Vienna (pers. comm.) confirmed that this is a mistake, as the specimen that has been used for the drawing in fact possesses pre-apical foveae. Distribution. Namibia. Terametopon (Terametopon) prominentinotus Gomy &Vienna, 1999 (Figs , 48-56) Terametopon (Terametopon) prominentinotus Gomy & Vienna, 1999: 5, figs. 2-5, 7, 8. Type locality: Southern Namibia, Gobabeb. Type specimen examined. Holotype (TMSA), mounted on its side on a triangular point, left mesotibia broken off, glued to the same triangular point as specimen (tarsus missing), left metatibia missing, tarsomeres 3-5 of the right -57-

16 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) metatarsus missing, left mesotarsus missing, both protarsi, both antennae and left mandible missing, with genitalia mounted in Canada Balsam on a separate slide, with printed label: GOBABEB / 70m. S.E. of / Walvis Bay, with another printed label: Southern / Namib / Desert, with a consecutive printed label: SOUTH WEST / AFRICA, followed by another printed label: sandy dunes, and another printed label: October, followed by another blackink written label with male symbol and another red label, printed: HOLOTYPUS/ Terametopon (Terametopon) / prominentinotus sp. nov. / Y. Gomy et P. Vienna des., 1998, followed by another yellow, pencil label: D09-039, added by myself. Redescription. Body length. PEL mm, APW mm, PPW mm, EL mm, EW mm. Body ovoid, convex, head comparatively small, elytral humeri prominent, cuticle reddish-brown; legs, antennae and mouthparts reddish-brown. Antennal scape dilated and strongly thickened, upper surface with numerous dense long setae; club broken off. Mouthparts. Labrum missing; mandibles coarsely and densely punctate, mandibular apex broken off, left mandible missing; cardo of maxilla with few short setae on lateral margin; stipes triangular, with three long well sclerotized setae; rest of maxillae broken off, missing; labial palpi missing; mentum rectangular, anterior margin straight, without median notch, with several long setae, lateral margins and disc with several setae. Clypeus smooth, transverse, about twice as broad as long, with massive prominent frontoclypeal projection (Fig. 44) in a form of inverted pyramid, base of this projection growing out from anterior frontal margin; frontal, supraorbital striae absent; disc of frons uneven, surface (including the frontoclypeal projection) very coarsely rugulose-lacunose; eyes very flattened, invisible from above. Pronotal sides (Fig. 41) strongly convergent anteriorly; apical angles very acute, anterior emargination for head very deep, circular; pronotal epipleuron with dense yellow setae; marginal stria almost entirely absent, present only on anterior angles as a thin rudiment; disc (apart from finely punctate apical angles and a short band of fine punctation behind head) entirely covered with very coarse and dense round-elongate punctures forming coarse elongate wrinkles laterally; scutellum very small, almost invisible. Elytral epipleura with sparse thin yellow setae; marginal epipleural stria well impressed, complete; marginal elytral stria well impressed, carinate on outer side, bisinuate, reaching elytral apex, apical elytral stria absent; all other elytral striae absent. Elytral disc wholly covered with very coarse and dense rugulose-lacunose hollow punctation, punctures separated less than their own diameter. Propygidium (Fig. 45) completely exposed, about 1.5 times as broad as long, laterally with sparse setae, evenly covered with very dense shallow confluent punctures separated by less than half their own diameter; pygidium long, about 2.5 times as long as broad, with confluent punctation similar to that of propygidium, but punctures sparse, getting finer apically, laterally with short yellow setae. Anterior margin of median portion of prosternum (Fig. 47) almost straight, bisinuate; marginal stria shortly present laterally, next obliterated; prosternal process constricted, very narrow, somewhat concave, with deep punctures fringed with long amber setae; pre-apical foveae absent, both sets of prosternal striae absent. Anterior margin of mesoventrite (Fig. 46) deeply emarginate medially; marginal stria absent; disc with large, oval hollow punctures with modified petal-like microscopic seta; meso-metaventral sutural stria absent, meso-metaventral suture very faintly impressed, almost unrecognizable beneath coarse punctures; intercoxal disc of metaventrite almost flat, -58-

17 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Figs : Terametopon (Terametopon) prominentinotus Gomy & Vienna. 41 habitus, dorsal view; 42 same, ventral view; 43 mesotibia, dorsal view; 44 head, 45 propydigium and pygidium; 46 mesoventrite; 47 prosternum. -59-

18 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) punctation similar to that of mesoventrite; lateral metaventral stria very shortened, obliquely arcuate; lateral disc of metaventrite excavate, punctures similar to those of intercoxal disc of metaventrite, fringed with long setae; metepisternum with identical punctation, punctures with long amber setae, denser than those of lateral disc of metaventrite. Figs : Terametopon (Terametopon) prominentinotus Gomy & Vienna, male genitalia. 48 coxites of 8 th segment, ventral view; 49 same, dorsal view; 50 same, lateral view; 51 9 th and 10 th tergite, dorsal view; 52 same, lateral view; 53 spiculum gastrale, ventral view; 54 same, lateral view; 55 aedeagus, dorsal view; 56 same, lateral view. -60-

19 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Intercoxal disc of first abdominal sternite with dense punctation, punctures finer and sparser to those of metaventrite, lateral stria vaguely impressed, thin, shortened apically; lateral sides of all visible abdominal sternites setose. Protibia dilated, outer margin with two large distal triangular teeth topped with short denticle, followed by three low teeth topped with short denticle diminishing in size in proximal direction; outer part of ventral surface of protibia with sparse punctures and shallow wrinkles, median and inner parts of ventral surface smooth; inner margin of protibia with a dense row of long setae; apical margin of protibia without denticles; tarsi broken off; protibial spur short, bent, growing out from near tarsal base. Mesotibia (Fig. 43) slender, denticles of outer margin all broken off, inner-lateral surface next to outer margin with one row of dense long well sclerotized setae, another row of much shorter setae present medially; denticles of outer-lateral margin all broken off; outer-lateral surface smooth; apical margin with four strongly sclerotized denticles, inner-most one the longest, several times as long as other three; claws of apical tarsomere broken off; somewhat more thickened and dilated than metatibia, otherwise in all aspects similar to it, denticles on outer margin and outer-lateral margins all broken off; tarsi broken off; metafemori thickened. Male genitalia. Coxites of 8 th segment (Figs ) longitudinally fused medially, apically with inflatable membrane; coxites ventrally with dense minute spores; on apical tenth laterally with dense short setae. Tergite of 8 th segment and coxites fused laterally (Fig. 50). Morphology of 9 th segment (Figs ) typical for the subfamily; spiculum gastrale (Figs ) (ventral sclerite of 9 th segment) expanded on both ends. Basal piece of aedeagus (Fig. 55) (phallobase) well developed, ratio of its length : length of parameres approximately 1 : 2; parameres fused almost along basal two-thirds; aedeagus slightly curved ventrad (Fig. 56). Biology. Unknown. Distribution. Namibia. Remarks. This species, as already mentioned by GOMY & VIENNA (1999) possesses characters that are typical for the species of the subgenus Psammoprinus: comparatively small head, strongly convergent pronotal sides with very acute anterior angles, rudimentary marginal pronotal stria (present only on anterior pronotal angles), reduction of elytral striae, strongly constricted, keel-like prosternal process devoid of both sets of striae. Aedeagus of this species is most similar to Terametopon (P.) namibiensis. Furthermore, the frontoclypeal projection of this species is somewhat different as well, its base is growing out from anterior margin of frons and apically it continues on well defined quadrate clypeus. Clypeus of the species included in the nominotypical subgenus is almost vertical and hidden under the frontoclypeal projection. However, since there is only a single, somewhat damaged specimen available (both antennal clubs, that would be helpful in subgeneric placement are missing, as is labrum and one mandible), it is tentatively left in the nominotypical subgenus, until further material is discovered and the phylogeny of species is studied from the cladistical viewpoint. -61-

20 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) Subgenus Psammoprinus Gomy & Vienna, 1996 Psammoprinus Gomy & Vienna, 1996: 330. Type species: Terametopon namibiensis Mazur, 1993 (original designation). Diagnosis. Differing from the species of the nominotypical subgenus by the absence of frontoclypeal projection; larger body size; flattened frons with a vague curved stria. Furthermore, dorsal upper two-thirds (roughly) of antennal club are covered with dense short sensilla intermingled with scattered longer sensilla; lower third (roughly) glabrous, ventrally sensilla surrounds entire club leaving out large median circular glabrous patch. Species of the nominotypical subgenus have dorsal surface of their antennal club imbricate, almost completely glabrous apart from one deep, almost circular slit-like sensory area that is surrounded by sparse short setae. Terametopon (Psammoprinus) namibiensis Mazur, 1993 (Figs , 67-73) Terametopon namibiensis Mazur, 1993: 4, figs Type locality: Namibia, Kromhoek Farm. Terametopon (Psammoprinus) namibiensis: GOMY & VIENNA 1996: 331 (keyed), figs. 1, 2, 4, 9, 12, 15; MAZUR 1997: 266. Type specimen examined. Holotype (SMNS), mounted on its side on a triangular point, right antennal flagellum broken off, glued to the same point as specimen, right protarsus missing, with printed label: (19)90 Schülle / Namibia Farm / Kromhoek, followed by printed label: Terametopon / namibiensis / sp. n. / det. S. Mazur 92, followed by red label (printed): HOLOTYPE, with a following yellow pencil label: D08-091, added by me. Additional specimens examined. BOTSWANA: Drotsky s Cave / Ngamiland / Apr. 65 W.D.Haacke (1, TMSA, P. Vienna det.). NAMIBIA: South West Africa / Kuiseb Namib Dunes / (19)59 / H. Dick Brown (1, TMSA, P. Vienna det.); South West Africa / Namib/Naukluft Park / Dunes near Gobabeb / 23.34S 15.03E. / 1983 R. Oberpreiler (1, SANC, det. by myself); SE 25, 15 Ac4 / Jan 1977 / Namib Expedition / Holm, Kirsten & Scholz (1, SANC, det. by myself). Redescription. Body length. PEL mm, APW mm, PPW mm, EL mm, EW mm. Body ovoid, head comparatively very small, elytral humeri prominent, dorsally moderately convex, ventrally very convex, cuticle dark brown; legs, antennae and mouthparts light brown. Antennal scape (Fig. 65) strongly thickened and dilated, impunctate, with one ring of numerous strongly sclerotized longish setae; club round, flattened, without visible articulation, dorsally (Fig. 60) upper two-thirds (roughly) covered with dense short sensilla intermingled with scattered longer sensilla, lower third (roughly) glabrous, ventrally (Fig. 61) sensilla surrounds entire club leaving out large median circular glabrous patch; Reichardt s organ in a form of two small, circular vesicles situated on inner and outer lateral margins, respectively; two other small circular sensory areas present in middle of antennal club. Mouthparts. Labrum semicircular, finely punctate, with a large shallow depression; labral pits absent; mandibles finely punctate, stout, mandibular apex very acutely pointed, subapical tooth very large, globular, present on both mandibles; cardo of maxilla with two short setae on lateral margin; stipes triangular, with three long well sclerotized setae; distal maxillary -62-

21 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Figs : Terametopon (Psammoprinus) namibiensis (Mazur). 57 habitus, dorsal view; 58 same, ventral view; 59 protibia, ventral view; 60 antennal club, dorsal view; 61 same, ventral view; 62 mesotibia, dorsal view; 63 mesoventrite; 64 prosternum; 65 head, dorsal view; 66 same, ventral view. -63-

22 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) palpomere thickened, its width more than half length, apex with nipple-like projection bearing sensory area, approximately five times as long as penultimate, palpal organ present, well developed; lacinial tooth absent; distal labial palpomere (Fig. 66) thickened, width almost equal length, apex with nipple-like projection bearing sensory area, palpal organ well-visible, distal labial palpomere with one very long seta; mentum subtrapezoid, anterior margin without notch, slightly projecting anteriorly, with several longish well sclerotized setae, lateral margins and disc of mentum glabrous. Clypeus (Fig. 65) large, flattened, with dense fine punctation; frons densely punctate, broadly, but shallowly depressed medially, this depression faintly anteriorly marked by a curved vague stria, frontal and supraorbital striae absent; anterolateral corners of frons conspicuously projected; eyes very flattened, invisible from above. Pronotal sides (Fig. 57) strongly convergent anteriorly; apical angles very acute, anterior pronotal margin with a deep semicircular incision for head; marginal stria weakly impressed, present only behind head and on anterior angles, next obliterated; disc with dense and coarse deep punctures, separated by half to their own diameter, in anterolateral angles and behind head punctation becoming sparser and finer; epipleura with long amber setae; scutellum very small, almost invisible. Elytral humeri prominent, epipleura with short thin setae; marginal epipleural stria thin, complete, shortly continued along elytral apex; marginal elytral stria well impressed, carinate on outer margin, almost reaching elytral apex. Oblique humeral stria vaguely impressed on basal fifth; internal subhumeral stria absent; dorsal elytral striae 1-3 vaguely impressed, almost unrecognizable beneath coarse punctation, 1 st and 3 rd about the same length, reaching about elytral half apically, 2 nd s bit longer, reaching about three-fourths of elytral length apically; 4 th dorsal and sutural striae absent. Elytral disc wholly covered with very dense hollow elongate deep punctation; punctures separated by about half their diameter reaching elytral apex. Propygidium almost completely exposed, with very dense and coarse punctures separated by less than half their diameter; pygidium long, convex medially, basally covered with punctation similar to that of propygidium, punctures becoming sparser and finer apically. Anterior margin of median portion of prosternum (Fig. 64) rounded; marginal stria anteriorly absent; preapical foveae absent; prosternal process very compressed, concave, densely setose, with scattered deep punctures; lateral striae divergent basally, subparallel, obliterated on apical third; carinal striae shortly impressed on coarsely punctate and setose prosternal apophysis. Anterior marginal stria of mesoventrite (Fig. 63) emarginate medially; laterally stria well impressed, carinate; disc with very deep round hollow punctures of various sizes; mesometaventral sutural stria absent, meso-metaventral suture thin, curved outwardly; intercoxal disc of metaventrite flat, wholly covered with punctation similar to that of mesoventrite; lateral metaventral stria shortened, obliquely arcuate. Lateral disc of metaventrite excavate, with very dense and coarse elongate punctures fringed with long setae; metepisternum covered with even coarser and denser punctures fringed with long amber setae. Intercoxal disc of the first visible abdominal sternite convex, lateral striae rudimentary, very shortened and vaguely impressed; disc with deep round punctures separated by about their to twice their diameter; lateral disc of all visible abdominal sternites setose laterally. Protibia dilated, outer margin with two large distal triangular teeth topped with short -64-

23 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN denticle, followed by four low teeth topped with short denticle diminishing in size in proximal direction; outer part of ventral surface of protibia (Fig. 59) with shallow sparse punctures and wrinkles, inner margin of protibia with a dense row of long setae; apical margin of protibia Figs : Terametopon (Psammoprinus) namibiensis (Mazur) male genitalia. 67 coxites of 8 th segment, ventral view; 68 same, dorsal view; 69 same, lateral view; 70 9 th, 10 th tergite, dorsal view, spiculum gastrale, ventral view; 71 same, lateral view; 72 aedeagus, dorsal view; 73 same, lateral view. -65-

24 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) with two short vague denticles; tarsi thin, claws of apical tarsomere long, several times its length; protibial spur long, bent, growing out from near tarsal base. Mesotibia (Fig. 62) slender, outer margin with long sparse denticles, inner-lateral surface next to outer margin with long well sclerotized setae, with another brush of long setae present medially; outer-lateral margin jagged, with sparse double row of shorter denticles; apical margin with four strongly sclerotized denticles, the inner-most one the longest, several times as long as other three; tarsus slightly thickened, each tarsomere with long, strongly sclerotized amber seta; claws of apical tarsomere slightly bent, very thin, several times its length. Metatibia slightly more dilated and thickened than mesotibia, but otherwise strikingly similar to it; metafemori thickened. Male genitalia. Coxites of 8 th segment (Figs ) longitudinally separated medially, apically with inflatable membrane with dense minute setae; ventrally with dense pores; coxites on apical tenth laterally with several short setae. Tergite of 8 th segment and coxites fused laterally (Fig. 69). Morphology of 9 th segment (Figs ) typical for the subfamily; spiculum gastrale (ventral sclerite of 9 th segment) expanded on both ends. Basal piece of aedeagus (Fig. 72) (phallobase) well developed, ratio of its length : length of parameres approximately 1 : 3; parameres fused almost along entire length; aedeagus slightly curved ventrad (Fig. 73). Since the Holotype is a female, another specimen from Namibia, which was compared to Holotype has been used for the genitalia preparation and description. This specimen bears my pencil label D and is deposited at SANC. Biology. Unknown. Remark. Drotsky s Caves are situated in the Kalahari Desert, northwestern Botswana. It is not known whether the specimen was found inside the cave complex or just near it. Distribution. Botswana, Namibia. New record for Botswana. Terametopon (Psammoprinus) deserticola Gomy &Vienna, 1996 (Figs , 81-85) Terametopon (Psammoprinus) deserticola Gomy & Vienna, 1996: 326, figs. 3, 6, 10, 13, 16. Type locality: Southern Namibia, Gobabeb, Walvis Bay. Type specimen examined. Paratype (TMSA), mounted on its side on a triangular point, left mesotarsus, both antennae, right mid-leg, left hind-leg missing, with genitalia mounted in Canada Balsam on a separate slide, with printed label: GOBABEB / 70m. S.E. of / Walvis Bay, with another printed label: sandy dunes / Trap. No. 45, with another printed label: 16 th / August / 65, followed by another written label with male symbol and another red label, printed: PARATYPE, followed by another printed label: Terametopon / deserticola Gomy et / Vienna / Y. GOMY DET. 1995, with another consecutive red-margined printed label: Paratype / Terametopon / deserticola / sp. n. / Gomy & P. Vienna and another yellow pencil label: D08-038, added by myself. Additional specimens examined. NAMIBIA: SE 2615 Fld4 / 10.vii.1980 / Dep. Entomologie UP. (1, SANC, identified by myself, compared to paratype). Redescription. Body length. PEL mm, APW mm, PPW mm, EL mm, EW mm. Body rectangular oval, head comparatively small, elytral humeri prominent, dorsally moderately convex, ventrally very -66-

25 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN convex, cuticle reddish-brown, pronotum darker than elytra; legs and mouthparts reddish brown. Antennal scape strongly thickened and triangularly dilated, impunctate, with one ring of numerous strongly sclerotized long setae; club round, flattened, without visible articulation, Figs : Terametopon (Psammoprinus) deserticola Gomy & Vienna. 74 habitus, dorsal view; 75 same, ventral view; 76 protibia, dorsal view; 77 prosternum; 78 propygidium and pygidium; 79 mesoventrite; 80 head, dorsal view. -67-

26 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) dorsally upper two-thirds (roughly) covered with dense short sensilla intermingled with scattered longer sensilla, lower third (roughly) glabrous, ventrally sensilla surrounding entire club leaving out large median circular glabrous patch; Reichardts organ not examined. Mouthparts. Labrum semicircular, finely punctate, basally somewhat elevated, gradually sloping down anteriorly; labral pits absent; mandibles (Fig. 80) finely punctate, slender, mandibular apex very acutely pointed, subapical tooth present only on the left mandible, prominent, acute; cardo of maxilla with several short setae on lateral margin; stipes triangular, with three long well sclerotized setae; distal maxillary palpomere thickened, width more than half length, apex with nipple-like projection bearing sensory area, approximately five times as long as penultimate, palpal organ present, well developed; lacinial tooth absent; distal labial palpomere thickened, width about half length, apex with nipple-like projection bearing sensory area, palpal organ well-visible, penultimate labial palpomere with one very long seta; mentum rectangular, anterior margin without notch, anterior, lateral margins and disc with several longish well sclerotized setae. Clypeus (Fig. 80) large, flattened, with dense fine punctation, constricted at base; frons flattened, densely punctate, with curved vague stria present on frontal disc, frontal and supraorbital striae absent; anterolateral corners of frons conspicuously projected; eyes very flattened, invisible from above. Pronotal sides (Fig. 74) strongly convergent anteriorly; apical angles prominent, anterior pronotal margin with a deep semicircular incision for head; marginal stria weakly impressed, present only behind head, on anterior angles and on apical fourth of lateral pronotal margin, next obliterated; median part of disc with dense and coarse deep punctures, separated by half to their own diameter, laterally punctures turn into coarse elongate wrinkles, on anterolateral angles and behind head punctation becoming sparser and finer; epipleura with long amber setae; scutellum very small, almost invisible. Elytral humeri prominent, epipleura with short thin setae; marginal epipleural stria thin, complete; marginal elytral stria well impressed, carinate on outer margin, almost reaching elytral apex; apical elytral stria absent. Oblique humeral stria very vaguely impressed on basal fifth, almost unrecognizable beneath coarse punctation; other elytral striae absent; sutural elytral stria faintly recognizable, complete. Elytral disc wholly covered with very dense and coarse deep punctation, punctures forming elongate rugulose wrinkles; before elytral apex punctation disappears. Propygidium (Fig. 78) completely exposed, almost as long as broad, lateral margins setose, with very dense and coarse shallow punctures separated by less than half their diameter; pygidium long, about twice as long as its basal width, laterally covered with dense brush of thin yellow setae, medially glabrous, punctation of pygidium finer than that of propygidium, apically punctures confluent. Anterior margin of median portion of prosternum (Fig. 77) rounded; marginal stria present anteriorly as very vague rudiment; preapical foveae absent; prosternal process very compressed, concave, impunctate, setose; lateral striae rudimentary, convergent, present only basally; carinal striae absent; prosternal apophysis coarsely and densely punctate. Anterior margin of mesoventrite (Fig. 79) emarginate medially; striae unrecognizable beneath coarse punctation; disc with very deep round hollow punctures separated up to two times their diameter, each puncture with microscopic petal-like modified seta; mesometaventral sutural stria absent, meso-metaventral suture carinate on outer margin, little -68-

27 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN Figs : Terametopon (Psammoprinus) deserticola Gomy & Vienna, male genitalia. 81 coxites of 8 th segment, 9 th, 10 th tergite and spiculum gastrale, ventral view; 82 same, dorsal view; 83 coxites of 8 th segment, tergite lateral view; 84 aedeagus, dorsal view; 85 same, lateral view curved outwardly; intercoxal disc of metaventrite flat, wholly covered with very deep coarse oval punctation, punctures separated by their own diameter; lateral metaventral stria rudimentary, obliquely arcuate. Lateral disc of metaventrite excavate, punctures similar to those of metaventrite, laterally punctures fringed with short setae; punctation of metepisternum unrecognizable beneath long amber setae. Intercoxal disc of the first visible abdominal sternite convex, lateral striae thin, shortened, not reaching half of the sternites length apically; punctation finer and sparser than that of metaventrite, punctures separated several times their diameter; lateral disc of all visible abdominal sternites setose laterally. Protibia (Fig. 76) dilated, outer margin with two large triangular distal triangular teeth topped with short denticle, followed by four low teeth topped with short denticle diminishing -69-

28 Lackner T.: Revision of the genus Terametopon, with description of new species (Coleoptera: Histeridae) in size in proximal direction; outer part of ventral surface of protibia with shallow sparse punctures and wrinkles, inner and median part of ventral surface of protibia well separated by curved stria, inner margin with a dense row of short setae; apical margin of protibia without denticles; tarsi thin, claws of apical tarsomere long, several times its length; protibial spur long, bent, growing out from near tarsal base. Mesotibia slender, outer margin with long sparse denticles, inner-lateral surface next to outer margin with long well sclerotized setae, with another brush of long setae present medially; outer-lateral margin with sparse double row of shorter denticles; apical margin with four strongly sclerotized denticles, inner-most one the longest, several times as long as other three; tarsus slightly thickened, each tarsomere with long, strongly sclerotized amber seta; claws of apical tarsomere slightly bent, straight, thin, little longer than its length. Metatibia more dilated than mesotibia, denticles on outer margin shorter, all (except for a single long denticle near tarsal insertion) of equal length; outer-lateral margin with two regular rows of dense short denticles; metafemori thickened. Male genitalia. Coxites of 8 th segment (Figs ) longitudinally separated medially, apically with inflatable membrane; coxites on apical tenth laterally and ventrally with several short setae. Tergite of 8 th segment and coxites fused laterally (Fig. 83). Morphology of 9 th segment typical for the subfamily; spiculum gastrale (ventral sclerite of 9 th segment) expanded on both ends. Basal piece of aedeagus (phallobase) broken off; parameres (Fig. 84) fused on basal half; aedeagus slightly curved ventrad (Fig. 85). Biology. Unknown. Collected in sandy dunes by pitfall traps. Distribution. Namibia. KEY TO THE SPECIES OF THE GENUS TERAMETOPON 1) Head with a distinct frontoclypeal projection (Figs. 4, 20, 28, 44) (subgenus Terametopon Vienna) ) Head without such projection (Figs. 65, 80) (subgenus Psammoprinus Gomy & Vienna) ) Preapical foveae of prosternum absent (Figs. 6, 47, 64, 77) ) Preapical foveae of prosternum present (Figs. 24, 29) ) Prosternal process narrowed, setose, both sets of prosternal striae absent, all elytral striae (except for marginal elytral stria) absent....t. (T.) prominentinotus Gomy & Vienna 6) Prosternal process flattened, not narrowed, glabrous, both sets of prosternal striae present, well developed; elytral striae 1-4 present; marginal pronotal stria well developed, complete.... T. (T.) levissimestriatus Vienna 7) Frontoclypeal projection in form of inverted pyramid; elytral striae 1-2 vaguely visible, striae 3-4 unrecognizable; mesoventrite with punctures fringed with microscopic setae... T. (T.) endroedyi Vienna 8) Frontoclypeal projection in form of a cube; elytral striae 1-4 well impressed, clearly distinguishable; mesoventrite punctate, punctures without microscopic setae.... T. (T.) foveatus sp. nov. 9) Body ovoid, elytral striae 1-3 present; punctation weakens on elytral humeri; subapical tooth globular, present on both mandibles; propygidium and pygidium on lateral sides without long yellow setae; punctation of pronotum does not form coarse elongate wrinkles laterally....t. (P.) namibiensis Mazur 10) Body rectangular-oval, elytral striae 1-4 absent, punctation does not weaken on elytral humeri, subapical tooth present only on left mandible, not globular, acute; propygidium and pygidium laterally with long dense setae; punctation of pronotum forms coarse elongate wrinkles laterally.... T. (P.) deserticola Gomy & Vienna -70-

29 Folia Heyrovskyana, series A, vol. 17(2): 43-72, 2009, ed.: November 20, 2009 ISSN DISCUSSION Most of the species of the genus Terametopon are endemic to the sandy dunes of the Namib Desert, currently known from seven species. Six of those species are revised and redescribed herein, and the seventh, T. (?) truncatus, is not treated here because of the severe alternations of its body and it is left as Terametopon incertae sedis (see Introduction). Species of Terametopon exhibit traits of specialized psammophilous Saprininae known from all greater deserts of the world: underside of body with dense vestiture, swollen femora, strongly convex body, dilated protibia, long and straight protarsal claws. Species of the nominotypical subgenus possess a curious frontoclypeal projection on their head, which is unique within the subfamily Saprininae. The function of this projection is unknown. They are all undoubtedly interrelated, sharing several synapomorphies: well developed and visible palpal organs of labial and maxillary palpi, antennomere before antennal club that is not saucer-like, but shaped as other antennomeres of flagellum, dorsoventrally depressed antennal club, penultimate labial palpomere with one long seta, quadrate mentum without notch on anterior margin, absent meso-metaventral sutural stria, dilated protibia with two large distal teeth topped with short denticle, followed by three short teeth topped by short denticle and coxites of 8 th abdominal segment of male genitalia bearing short setae apically as well as inflatable membrane. It is Fig. 86. Distribution of Terametopon Vienna. -71-

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