Vatellus perforatus Miller, 2005

Miller, Kelly B., 2005, Revision of the New World and south-east Asian Vatellini (Coleoptera: Dytiscidae: Hydroporinae) and phylogenetic analysis of the tribe, Zoological Journal of the Linnean Society 144 (4), pp. 415-510 : 475-498

VATELLUS PERFORATUS ( GUIGNOT, 1955) COMB. NOV.

( FIGS 1, 203–215 , 314)

Platydessus perforatus Guignot, 1955: 4 . Macrovatellus perforatus ; Spangler, 1966a: 58 (comb.

nov.); Trémouilles, 1995: 27.

Type information: Holotype ♀ in G. Frey Museum, Munich, Germany (not examined). Although the type was not examined, a large number of specimens identified as this species by P. Spangler ( USNM) were. Based on this, I believe Guignot’s species refers to the one described here.

Type locality: Bolivia.

Diagnosis: This species is distinguishable from other Vatellus by the shorter, stouter habitus and relatively small size ( Figs 1, 203 ), relatively coarse punctation and elytra smooth between punctures (elytral surface sculpture as in Fig. 204 ), broad medial portions of the metacoxae ( Fig. 205 ), strongly offset and apically rounded metatrochanter ( Fig. 206 ), weakly produced anterior clypeal margin and distinctive male and female genitalia. The male median lobe is apically abruptly narrowed to a small, ventrally curved point ( Fig. 212 ). The receptacle is relatively large, about the size of the spermatheca, and is ovate ( Figs 214, 215 ). The gonocoxae are long and apically pointed ( Fig. 214 ).

Description:

Habitus. Body outline moderately discontinuous in dorsal aspect, lateral margins of elytra moderately rounded ( Figs 1, 203 ), dorsoventrally moderately compressed. Measurements are given in Table 1.

Coloration. Head, pronotum and elytron dark redbrown; venter dark red brown, darker on metacoxa and metasternum; appendages dark red-brown.

Sculpture and structure. Head with punctures evenly distributed, moderately coarsely punctate, punctures slightly finer on clypeus; anterior clypeal margin distinctly produced; eyes moderately large, not strongly protruding ( Fig. 203 ). Pronotum with punctation evenly distributed, punctures moderately coarse, interpuncture surface shiny; lateral margin strongly rounded anteriorly, widest point near anterior angles, posterior angles slightly acute; lateral bead obscured posteriorly; transverse sulcus absent. Elytron evenly and coarsely punctate, punctures moderately dense, surface between punctures shiny, smooth ( Fig. 204 ). Prosternum short with dense fringe of yellow setae; prosternal process broadly triangular, medially excavated, apex acutely pointed. Metasternum with punctation coarse and dense, medially with punctures slightly finer, interpuncture surface smooth, shiny; anteromedial margin slightly produced ventrally. Abdominal sternal margins not unusually modified; sterna evenly and finely punctate, interpuncture surface smooth, shiny; sixth abdominal sterna with apical lobe flattened, long, subtriangular ( Fig. 209 ); speleum moderately broad throughout length, relatively abruptly narrowed to narrowly rounded apex ( Fig. 210 ). Metacoxa with lateral portion coarsely and densely punctate over entire surface; metacoxal lines relatively broadly separated basally, apically very broadly divergent ( Fig. 205 ); medial portion densely and conspicuously punctate. Metatrochanter very short and rounded ( Fig. 205 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect evenly curved, apically abruptly narrowed to slen- der, slightly hooked apex ( Fig. 212 ); in ventral aspect with lateral margins slightly but evenly convergent, apically abruptly narrowed with small, apical, narrowly rounded point ( Fig. 213 ). Lateral lobe very long, basally very broad, apical portion elongate, straight, expanded medially, apically rounded, with dense apical and dorsal fringe of fine, elongate setae ( Fig. 211 ).

Female genitalia. Gonocoxosternite very broad and robust, apical angle rounded, anterior margin deeply concave, anterior lobe relatively long, slender, apex curved laterally ( Fig. 214 ). Gonocoxa elongate, slen- der, apex narrowly rounded, apodeme elongate and moderately slender ( Fig. 214 ). Bursa copulatrix short, slender ( Fig. 214 ); spermathecal duct moderately long, slender; receptacle robust, elongate, subrenal shaped, subequal in size to spermatheca ( Fig. 214 ); intermediate duct moderately long, robust, sinuate ( Fig. 215 ); spermatheca subspherical, triangular process very long and broad; fertilization duct short ( Fig. 214 ).

Sexually dimorphic characters. Male with setae on mesotrochanter and mesofemur weakly developed. Male and female pro- and mesobasotarsomeres similar in size and length, both with protarsomere III very long and slender ( Figs 207, 208 ).

Intraspecific variation. Specimens differ in the extent of shagrined sculpture on the pronotum with it nearly absent in some specimens and making an extensive, distinct line subposteriorly on others.

Natural history: This species appears to be streaminhabiting, unlike most species in the genus. Specimens were collected from streams in a rainforest among rocks, leaves and sticks. It has also been collected from black lights.

Distribution: Vatellus perforatus is a northern South American species and has been collected from Brazil and Venezuela (Fig. 314).

Phylogenetic relationships: This species occupies a relatively isolated phylogenetic position within Vatellus , but evidence indicates it is relatively closely related to the V. tarsatus and V. grandis clades ( Fig. 316 ).

Material examined: Brazil: Amazonas, Rio Demiti , nr Little Homestead. 0∞35¢N 66∞41¢W, UV light, GE and KE Ball (1, JBWM) . Venezuela: T.P. Amaz. Cerro de la Neblina 1 km S Basecamp 0∞50¢N 66∞10¢W 250 m, 8 February 1985, along small whitewater stream, in pools of dead le, P.J and P.M. Spangler, R. Faitoute, W. Steiner (13, USNM) ; T.P. Amaz. Cerro de la Neblina 1.5 km S Basecamp 0∞50¢N 66∞10¢W 250 m, 9 February 1985, small stream in rainforest, among rocks and leaf p, W.E. Steiner (1, USNM) .

VATELLUS TARSATUS (LAPORTE, 1835)

( FIGS 3 , 216–229 , 313)

Hydroporus tarsatus Laporte, 1835: 106, 1840: 167 .

Vatellus tarsatus ; Aubé, 1837: 218 (comb. nov.), 1838: 449; Gemminger & Harold, 1868: 428; Sharp, 1882b: 285; Régimbart, 1878: 463; Trémouilles, 1995: 27.

Type information: Lectotype (designated here to clarify assignment of this name with this species) ♂ in MNHN labelled, ‘ Hydroporus tarsatus Lap. ♀ Typemihi D.S. [handwritten]/ Hydroporus Vatellus tarsatus Buquet L. Cayenne D. Buquet [handwritten, green label, folded in half, “ Hydroporus ” crossed out]/ Ex.musaeo Dejean [black line around border]/D. Sharp Monogr./ LECTOTYPE Hydroporus tarsatus Laporte, 1835 . des. by K.B. Miller 2003 [red label with double black line border].’

Type locality: French Guyana [‘Cayenne’].

Diagnosis: This species is distinguishable from other Vatellus by the rather strongly demarcated posterior margins of abdominal sterna 2, 3 and 4 in most specimens. The posterior margins on these sterna end abruptly and rise steeply to the surface of the next sternum. Other distinguishing characters include the relatively fine punctures and elytral surface sculpture as in Figure 3 , nearly immaculate dorsal surface and dark brown to black colour, the very slender speleum ( Fig. 223 ), the median lobe apically slender and sharply upturned. The receptacle is about the size of the spermatheca and the intermediate duct is very broad ( Fig. 229 ).

Description:

Habitus. Lateral outline discontinuous in dorsal aspect, lateral margins of elytra moderately rounded ( Fig. 216 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head, pronotum and elytron dark brown to black; venter black to dark red-brown, appendages dark yellow-brown to dark red-brown.

Sculpture and structure. Head finely punctate, punctures dense and confused in places, somewhat finer anteriorly, surface slightly shagrined between punctures; anterior clypeal margin rounded; eyes large, protruding. Pronotum evenly punctate, punctures fined and moderately dense, surface alutaceous between punctures, more commonly so in females, as in Figure 3 , each puncture bearing a fine, relatively long seta, with long, distinct subbasal transverse line of shagrination; pronotum cordate, broadest near apical angles, lateral margins moderately strongly rounded anteriorly, posterior angles slightly acute and pointed ( Fig. 216 ); transverse sulcus absent to slightly impressed. Elytron evenly covered with fine punctures, punctures moderately dense, each bearing a relatively long, fine, pale seta ( Fig. 217 ), interpuncture surface shiny, smooth in most males and most females, some with microsculpture as in Figure 3 , making surface alutaceous. Prosternum short, slightly roughened, with numerous setae; prosternal process broad, lateral margins evenly curved, medially excavated, apically triangular, flattened, apex acutely pointed. Metasternum medially finely punctate, laterally with punctures more coarse and shallow, confused with surface slightly shagrined; anteromedial margin not expanded, apex slightly deflexed and pointed. Abdominal sterna with sutural margins abruptly margined, particularly between visible sterna 2, 3 and 4; sterna finely and irregularly punctate, surface between punctures shiny, smooth; sixth sternum with apical lobe very long, narrow, apically narrowly rounded; speleum broad basally, strongly narrowed subbasally, apex long and extremely slender to very narrowly rounded apex ( Fig. 223 ). Metacoxa with lateral portion irregularly and shallowly punctate, punctures confused, surface between punctures slightly shagrined; metacoxal lines very closely approximated posteriorly, anteriorly moderately divergent ( Fig. 218 ); medial portion finely and sparsely punctate. Metatrochanters short, rounded, strongly offset ( Fig. 219 ); metatarsal claws weakly dentate.

Male genitalia. Median lobe in lateral aspect evenly curved, slightly broadened apicomedially, basal portion subtriangular, not strongly elongate; apex distinctly narrowed, apically sharply pointed and distinctly curved ventrally, with numerous small spines on apicoventral surface ( Fig. 225 ); in ventral aspect narrow, evenly narrowed to acutely pointed apex ( Figs 224, 226 ). Lateral lobe with apical portion strongly curved basally and relatively straight in apical portion, moderately broad, apex rounded, with large field of fine, elongate setae along apical and lateral margins, subapical stout, elongate, curved setae absent ( Fig. 227 ).

Female genitalia. Gonocoxosternite with apex narrowly rounded, medial, anterior and posterior margins arcuate, anterior lobe moderately long, narrow ( Fig. 228 ). Gonocoxa elongate-triangular, apex narrowly rounded, ventral surface with numerous long, spinous setae, apodeme relatively short, slender ( Fig. 228 ). Bursa copulatrix small, inconspicuous; spermathecal duct very long, coiled, slender; receptacle spherical, subequal in size to spermatheca ( Fig. 228 ); intermediate duct relatively short, slightly twisted ( Fig. 229 ); spermatheca spherical, triangular process narrow basally, very broadly expanded apically; fertilization duct relatively short ( Fig. 228 ).

Sexually dimorphic characters. Male with very weakly developed mesotrochanteric and mesofemoral brushes of setae. Male with pro- and mesobasotarsomeres ( Fig. 220 ) slightly broader than in female ( Fig. 221 ), male protarsomere III ( Fig. 220A ) distinctly shorter than female protarsomere III ( Fig. 221A ).

Intraspecific variation. Historically the defining apomorphy for this taxon was the strongly demarcated sutures between the abdominal sterna. This feature is somewhat variable, but all specimens exhibit this character state to a greater or lesser degree. Although most specimens are nearly black overall, some specimens are dark brown.

Natural history: Vatellus tarsatus has been collected from a pond and a whitewater stream.

Distribution: This is a northern South American species with specimens collected from Brazil, French Guyana, Suriname and Venezuela (Fig. 313).

Phylogenetic relationships: This species and the following two together form a clade ( Fig. 316 ). They are similar in several characters and in overall similarity in less discrete characters. This clade is relatively closely related to V. perforatus and the V. grandis clade ( Fig. 316 ).

Material examined: Brazil: Para, Rio Xingu Camp , 52∞22¢W 3∞39¢ S Altamira, c. 60 km S, 11 October 1986, pond at 2nd palm grove on trail, P. Spangler and R. Crombie (4, USNM) ; Para, Rio Xingu Camp , 52∞22¢W 3∞39¢ S Altamira, c. 60 km S, 12 October 1986, pond at 2nd palm grove on trail, P. Spangler and O. Flint (2, USNM) ; Para, Rio Xingu Camp , 52∞22¢W 3∞39¢ S Altamira, c. 60 km S, 15 October 1986, pond at 2nd palm grove on trail, P. Spangler and O. Flint (2, USNM) ; Rio Marauia , 24 January 1963, E Fittkau (2, ZSBS) . French Guyana: Country only (1, MNHN) . Suriname: 3rd trib of Colakreek , 9 October 1989, N Nieser (1, NHMW) ; Brokopondo, Coesewijnaproject , 9 April 1970, N Nieser (1, NHMW) ; Marowijne, Mooi Wannakreek , 54∞05¢W 05∞30¢N, 1 December 1989, N Nieser (1, NHMW) ; nr. Sabakoekreek , 55∞11¢W 05∞26¢N, 28 July 1989, N Nieser (1, NHMW) ; Zanderijsavanne, 1st trib. of Colskrrek , 55∞14¢W 05∞27¢N, 15 September 1989, N Nieser (2, NHMW) . Venezuela: T.F. Amaz. Cerro de la Neblina , 1 km S Basecamp 0∞50¢N 66∞10¢W, 11 February 1985, 140 m (6, USNM) ; T.P. Amaz. Cerro de la Neblina 1.5 km S Basecamp 0∞50¢N 66∞10¢W 140 m, 15 February 1985, along small whitewater stream, in pools of dead leaves, P.J and P.M Spangler, R. Faitoute, W. Steiner (2, USNM) ; T.P. Amaz. Cerro de la Neblina 1.5 km S Basecamp 0∞50¢N 66∞10¢W 250 m, 11 February 1985, along small whitewater stream, in pools of dead leaves, P.J and P.M Spangler, R. Faitoute, W. Steiner (3, USNM) .

VATELLUS ANNAE MILLER SP. NOV.

( FIGS 216–229 , 313)

Type information: Holotype ♂ in USNM labelled, ‘ BRAZIL: Matto Grosso Jacaré P.N.Xingu xi.1961 Alvarenga e Werner leg. / HOLOTYPE Vatellus annae Miller 2004 [red label with double black line border].’

Paratypes: 1 in MCZC labelled same as holotype. Bolivia: Beni, 3.4 km W San Borja. 14∞52¢57≤S 66∞46¢03≤W, 14 July 1998, KB Miller (1, KBMC) . Brazil: Mato Grosso, Jacare-Parque Nat Xingu , November 1965, BLT, M Alvarengo, WCA Bokermann (1, FSCA) ; Matto Grosso, Tapirape , 15 November 1961, light, B Malkin (1, FSCA) ; Pres Matto Grosso, 1886, P Germain (4, MNHN) ; Rio Maravia , 24 January 1933, E. Fittkau (2, MBIC) . Peru: Loreto, Yacumana Lodge, Jct Rio Maranon, Rio Ucayall 73∞6¢W 4∞8¢S, 20 August 1994, light in woods, Skelley (4, LHIC) . Suriname: 55∞14¢W 05∞27¢N Brokopondo, 1st trib of Kolakreek , 15 September 1969, N. Nieser (2, MBIC) . Trinidad: Cumuto 1929, Darlington (68, MCZC) ; La Brea, Darlington (1, MCZC) ; St Augustine , April 1929, Darlington (1, MCZC) ; St Augustine , April 1929, Darlington (3, MCZC) .

Type locality: Brazil, Matto Grosso, Jacaré, P.N. Xingu [presumably the Indigenous Park of the Xingu].

Diagnosis: This species is most similar to V. tarsatus and V. pilacaudus in having overall fine punctation and elytral sculpture as in Figure 3 , relatively concolorous dorsal surfaces and similar habitus. Also, these species have the apical lobe on abdominal sternum VI long and apically nearly spherical (e.g. Fig. 222 ). Vatellus annae differs from V. tarsatus in lacking a strong, abrupt demarcation between abdominal sterna and having the slender apex of the median lobe more elongate and more sharply pointed apically ( Fig. 239 ). Also, the intermediate duct in V. annae is very long and very coiled on the dorsal surface of the spermatheca ( Fig. 244 ) whereas the intermediate duct in V. tarsatus is shorter and broad ( Fig. 229 ). The speleum is relatively broad in V. annae ( Fig. 273 ) whereas in V. tarsatus it is very slender ( Fig. 223 ). Vatellus pilacaudus shares the presence of the long, coiled intermediate duct, but in V. pilacaudus the apex of the median lobe is not long, slender and pointed ( Fig. 255 ).

Description:

Habitus. Lateral outline moderately discontinuous in dorsal aspect, lateral margins of elytra moderately strongly rounded ( Fig. 230 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head, pronotum and elytron brown-red; venter and appendages brown-red.

Sculpture and structure. Head finely punctate, punctures sparse, very fine to absent on clypeus, surface between punctures shiny; anterior clypeal margin slightly protruded; eyes large, protruding ( Fig. 230 ). Pronotum evenly punctate, punctures fine and moderately dense, surface generally alutaceous between punctures, each puncture bearing a fine, relatively long seta; pronotum cordate, broadest near apical angles, lateral margins moderately strongly rounded anteriorly, posterior angles slightly acute and pointed ( Fig. 230 ); transverse sulcus slightly impressed. Elytron evenly covered with fine punctures, punctures moderately dense ( Fig. 231 ), each bearing a relatively long, fine, pale seta, interpuncture surface shiny, smooth, or with distinctive microsculpture making surface alutaceous similar to pronotum. Prosternum short, slightly roughened, glabrous; prosternal process broad, lateral margins evenly curved, medially excavated, apically triangular, flattened, apex acutely pointed. Metasternum medially finely punctate, laterally with punctures more coarse and shallow, confused with surface between punctures shiny; anteromedial margin not expanded, not deflexed. Abdominal sternal sutures not usually modified; sterna finely and irregularly punctate, surface between punctures shiny, smooth; sixth sternum with apical lobe well-developed, apically rounded ( Fig. 236 ); speleum broad, slightly narrowed medially, apex broadly rounded ( Fig. 237 ). Metacoxa with lateral portion irregularly and shallowly punctate, punctures confused, surface between punctures slightly shagrined; metacoxal lines closely approximated posteriorly, anteriorly strongly divergent ( Fig. 232 ); medial portion finely and sparsely punctate. Metatrochanters short, rounded, strongly offset ( Fig. 233 ); metatarsal claws weakly dentate.

Male genitalia. Median lobe in lateral aspect narrow, medially somewhat broadened, basal portion broad, moderately elongate; apex very slender, apically sharply pointed and curved ventrally, with distinctive region of small subapical spines on ventral surface ( Figs 239, 242 ); in ventral aspect narrow, abruptly narrowed subapically, apex very slender, very narrowly rounded, nearly pointed ( Figs 240, 241 ). Lateral lobe with apical portion evenly curved, narrow, with very few setae apically; long, stout, subapical setae absent ( Fig. 238 ).

Female genitalia. Gonocoxosternite robust, medial margin very broad, apex nearly right-angled; posterior margin concave; anterior lobe relatively robust, short ( Fig. 243 ). Gonocoxa moderately broad, main portion approximately triangular, apex pointed, apodeme long and slender ( Fig. 243 ). Bursa copulatrix small and slender, undifferentiated; spermathecal duct moderately long, slender; receptacle spherical, less than 1/2 size of spermatheca ( Fig. 243 ); intermediate duct long, strongly and tightly coiled between receptacle and spermatheca, adpressed to dorsal surface of spermatheca ( Fig. 244 ); spermatheca spherical, triangular process relatively short, very broad; fertilization duct short, slightly curved ( Fig. 243 ).

Sexually dimorphic characters. Male with moderately developed mesotrochanteric and mesofemoral brushes of setae. Male pro- and mesobasotarsomeres ( Fig. 234 ) broadened in male compared with female ( Fig. 235 ), protarsomere III longer than mesotarsomere III in both sexes ( Figs 234A, 235A ).

Natural history: Vatellus annae has been collected at black lights.

Distribution: This species is relatively widespread in northern South America and has been collected in Bolivia, Brazil, Peru, Suriname and Trinidad (Fig. 313).

Phylogenetic relationships: This species is a member of the V. tarsatus clade along with V. pilacaudus ( Fig. 316 ). Within this clade, V. annae groups with V. pilacaudus based on the common presence of relatively closely approximated metacoxal lines ( Figs 231 , 247 ) (compared with other Vatellus ), the presence of a long intermediate duct coiled on the dorsal surface of the female spermatheca ( Figs 244 , 255 ), and the relatively small female receptacle ( Figs 243 , 257 ).

Etymology: This species is named in honour of my daughter, Annette (Annie) Miller.

VATELLUS PILACAUDUS MILLER SP. NOV.

( FIGS 245–258 , 313)

Type information: Holotype ♂ in USNM labelled, ‘ BOLIVIA: Beni, 40 km E San Borja, Estacion Biologica Beni, Estancia El Porvenir ; 6–8 September 1987; W. E. Steiner / At black light; open grass savanna and marsh / HOLOTYPE Vatellus pilacaudus Miller 2004 [red label with double black line border].’

Paratypes: 38 in USNM labelled same as holotype. Bolivia: Beni, 2.0 km E San Borja . 14∞52¢02≤S 66∞43¢41≤W, 15 July 1988, shaded pool in streambed, KB Miller (2, KBMC) ; 6.7 km NW Trinidad. 14∞47¢27≤S 64∞56¢47≤W, 18 July 1998, blacklight, KB Miller (1, KBMC) ; Prov. Cercado, 9.5 km N Trinidad. 14∞46¢24¢S 64∞58¢00¢W, 28 June 1999, KB Miller (1, KBMC) .

Type locality: Bolivia, Departmento Beni, 40 km E San Borja, Estacion Biologica Beni, Estancia El Porvenir .

Diagnosis: This species is most similar to V. tarsatus and V. annae in having small punctures, surface microsculpture (when present) as in Figure 3 , relatively concolorous dorsal surfaces and similar habitus. Also, these species have the apical lobe on abdominal sternum VI long and very well developed. Vatellus pilacaudus differs from V. tarsatus in lacking strong, abrupt demarcations between abdominal sterna. Also, the apex of the median lobe is broader and more narrowly rounded in lateral aspect ( Fig. 255 ), not slender and upturned as in V. tarsatus and V. annae . Also, the intermediate duct in this species is very long and tightly coiled on the dorsal surface of the spermatheca ( Fig. 258 ), unlike in V. tarsatus . In this species, the metacoxal lines are not as strongly divergent apically as in most species (ML/MW = 2.18–2.57) ( Fig. 247 ).

Description:

Habitus. Lateral outline discontinuous in dorsal aspect, lateral margins of elytra broadly rounded ( Fig. 245 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head and pronotum yellow-brown; elytron yellow-brown, slightly lighter along lateral and anterior margins, often with small, indistinct marginal macula apicomedially ( Fig. 245 ); venter yellow-red; appendages yellow-brown.

Sculpture and structure. Head with fine and dense punctation, slightly finer anteriorly, surface finely microreticulate between punctures, shiny; anterior clypeal margin rounded; eyes large, strongly protruding ( Fig. 245 ). Pronotum evenly covered with dense, moderately coarse punctation, surface finely microreticulate on most specimens; lateral margins moderately curved, widest point submedially, posterior angles slightly acute to slightly obtuse ( Fig. 245 ); lateral bead present and distinct along entire length; transverse sulcus moderately distinct laterally. Elytron densely punctate over entire surface, interpuncture surface similar to pronotum, setae moderately long and conspicuous. Prosternum short, alutaceous, glabrous; prosternal process moderately broad, slightly excavated, with only slight tumidity medially, apex produced into elongate, flattened triangle, apex acutely pointed. Metasternum with punctation laterally dense, punctures large, medially more finely punctate, interpuncture surface shiny, smooth; anteromedial margin not produced, slightly pointed. Abdominal sternal sutures not unusually modified; finely punctate medially, more coarsely punctate laterally, surface smooth, shiny between punctures; sixth sternum with apical lobe strongly produced, distinctly rounded and subspherical apically; speleum moderately broad, apically narrowly rounded ( Fig. 252 ). Metacoxa with lateral portion densely punctate over entire surface, punctures moderate in size, surface shiny between punctures; metacoxal lines approximate posteriorly, distinctly but only moderately divergent anteriorly ( Fig. 247 ); medial portion finely punctate. Metatrochanter strongly rounded and offset ( Fig. 248 ); metacoxal claws slightly dentate.

Male genitalia. Median lobe in lateral aspect evenly curved throughout length, moderately broad, slightly broader subapically, apex moderately narrowly rounded, basal portion broad ( Fig. 255 ); in ventral aspect with lateral margins subparallel, moderately convergent to narrowly rounded apex ( Figs 254, 256 ). Lateral lobe very long, apical portion moderately broad, evenly curved, slightly expanded medially ( Fig. 253 ).

Female genitalia. Gonocoxosternite broad, robust, apical angle narrowly rounded, somewhat acute, anterior margin irregularly concave, anterior lobe relatively short, robust ( Fig. 257 ). Gonocoxa moderately elongate, medial margin broadly curved, apex rounded, apodeme long, slender ( Fig. 257 ). Bursa copulatrix broad, short, relatively undifferentiated ( Fig. 257 ); spermathecal duct moderately long, slen- der; receptacle small, less than half size of spermatheca, spherical ( Fig. 257 ); intermediate duct very long, very tightly coiled on dorsal surface of spermatheca ( Fig. 258 ); spermatheca spherical, large; fertilization duct short, slightly curved ( Fig. 257 ).

Sexually dimorphic characters. Male with very welldeveloped setal region on mesotrochanter, setae on medial portion of mesofemur developed into elongate, cup-shaped form. Male pro- and mesobasotarsomeres ( Fig. 249 ) distinctly expanded compared with female ( Fig. 250 ), but not exceptionally broad, female protarsomeres ( Fig. 250A ) very long and slender.

Intraspecific variation. Most specimens are uniformly light brown, but a few have more extensive lateral, yellow margins on the elytron. Also, some specimens, both male and female, exhibit fine microsculpturing on the pronotum and elytron and a few female specimens are very strongly microsculptured dorsally. Other specimens are smooth between the punctures on the pronotum and elytron.

Natural history: Specimens have been collected at a black light in open savannah and marshland and from a shaded, stagnant pool in a streambed.

Distribution: Specimens have been collected only from areas in Bolivia (Fig. 313).

Phylogenetic relationships: This species is a member of the V. tarsatus clade ( Fig. 316 ). Within this clade, V. pilacaudus groups with V. annae ( Fig. 316 ) (see comments under that species).

Etymology: This species is named from the Latin words pila, meaning ‘ball’, and caudus, meaning ‘tail’, in reference to the semispherical lobe of the sixth abdominal sternum.

VATELLUS GRANDIS BUQUET, 1840

( FIGS 2, 6 , 8–17 , 24–27 , 38, 39, 42–44, 48, 49 , 259–273 , 314)

Vatellus grandis Buquet, 1840: 394 ; Gemminger & Harold, 1868: 428.

Macrovatellus grandis ; van den Branden, 1885: 19 (comb. nov.); Sharp, 1882b: 282, 826; Zimmermann, 1920: 30; Régimbart, 1878: 463; Trémouilles, 1995: 27.

Type information: Lectotype (designated here to clarify assignment of this name with this species) ♂ in MNHN labelled, ‘Cayenne [green label]/Coll.C.Felsche Geschenk 1907 [green label]/ Vatellus grandis Buquet. [handwritten]/1 29/ Macrovatellus grandis [handwritten]/ LECTOTYPE Vatellus grandis Buquet, 1840 . des. by K.B. Miller 2003 [red label with double black line border].’ Buquet apparently had more than one specimen on which he based this Description: I found only a single specimen in MNHN that appears to be one of Buquet’s specimens. It was selected as the lectotype.

Type locality: French Guyana [‘Cayenne’], upper part of Oyapok River.

Diagnosis: This species can be distinguished from other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced ( Fig. 6 ); (2) with surfaces relatively coarsely punctate and with surfaces between the punctures generally shiny, elytral surface sculpture as in Figures 2 and 260 ; (3) with a transverse impunctate area on each side of the midline on the pronotum ( Fig. 262 ); (4) eyes relatively small ( Fig. 259 ); (5) body relatively elongate and dorsoventrally depressed ( Fig. 259 ); (6) lateral pronotal margin relatively weakly curved ( Fig. 259 ); (7) anterior portion of the metasternum expanded ventrally; and (8) male and female genitalia distinctive ( Figs 269–273 ). The median lobe is long and slender with the apex bluntly rounded ( Fig. 269 ). The lateral lobe has the apical portion relatively straight with the apex relatively lacking in setae, but with two or three long, stiff setae on the subapical, ventral surface ( Fig. 271 ). The female receptacle is relatively small and elongate ( Fig. 272 ). This species and the following three are very similar in having a distinctive, flattened habitus, similar punctation, a distinct impunctate area on each side of the pronotum ( Fig. 262 ), the ventrally expanded anterior portion of the metasternum and the anteriorly produced anterior clypeal margin ( Fig. 6 ). Vatellus grandis , V. amae and V. drymetes are nearly indistinguishable externally, but have distinctive genitalia. Vatellus bifenestratus is larger and has the pronotum more prominently cordate ( Fig. 274 ).

Description:

Habitus. Body outline somewhat discontinuous in dorsal aspect, lateral margins of elytra not strongly rounded ( Fig. 259 ), dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head, pronotum, and elytron brown-red; ventral surfaces black except venter of head, pronotal epipleuron, propleuron, prosternum and small portions of posterior margins of abdominal sterna brown-red.

Sculpture and structure. Head finely and moderately densely punctate, slightly more dense anteriorly on frons and clypeus; anterior clypeal margin strongly anteriorly produced; eyes moderate in size, not strongly protruding. Pronotum with punctures relatively large laterally, denser medially and along posterior margin, punctures with fine pale, moder-

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ately elongate setae, area between punctures generally with fine shagrination formed by dense field of very fine tubercles except for transverse impunctate areas medially on each side of midline ( Fig. 262 ); pronotum broadest anteromedially, lateral margins relatively parallel-sided, only curved at extreme anterior end ( Fig. 259 ); lateral bead obscured except at extreme anterior end; transverse groove absent. Elytron strongly and densely punctate over entire surface, punctures finer and slightly more dense apically, each puncture with fine seta, surface between punctures smooth and shiny ( Fig. 260 ). Prosternum short, setose; prosternal process broad, flattened with short, indistinct, rounded medial carina, lateral margins rounded, convergent to narrowly rounded apex. Metasternum laterally with moderately large punctures, relatively sparsely distributed, medially with more dense, finer punctures, surface smooth, shiny; anteromedial margin strongly expanded ventrally into distinct tumidity. Abdominal sternal sutures not unusually modified; sterna laterally with moderately large, moderately dense punctures, interpuncture surface smooth, shiny; apical lobe on sixth abdominal sternum strongly produced, long, apically narrowly rounded, flattened ( Fig. 265 ); speleum elongate, moderately narrow, apically pointed ( Fig. 266 ). Metacoxa with lateral portion with punctures moderately large, relatively dense, interpuncture surface smooth, shiny; metacoxal lines strongly approximate posteriorly, anteriorly conspicuously divergent ( Fig. 261 ); medial portion with surface shiny, finely punctate. Metatrochanter strongly rounded, strongly offset ( Figs 267, 268 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect relatively straight medially, apically slightly narrowed to rounded apex, without subapical setae, basal portion elongate and narrow, broadened apically, slightly curved ( Fig. 269 ); in ventral aspect with lateral margins approximately parallel, subapically abruptly narrowed to narrowly rounded apex ( Fig. 270 ). Lateral lobe long, basally broad, with apical portion straight and narrow to acute, narrow apex; with 2–3 subapical, stout, elongate, curved setae on ventral margin ( Fig. 271 ).

Female genitalia. Gonocoxosternite very broad and robust, apical angle broadly rounded, anterior margin broadly concave, anterior lobe relatively short, slender ( Fig. 272 ). Gonocoxa elongate and slender, apex relatively narrowly rounded, apodeme elongate and slen- der ( Fig. 272 ). Bursa copulatrix moderately long, somewhat expanded apically ( Fig. 272 ); spermathecal duct moderately long, narrow; receptacle small, much smaller than spermatheca, elongate and slender, apically expanded ( Fig. 272 ); intermediate duct moderately long, with several S-curves ( Fig. 273 ); spermatheca subspherical, triangular process large and broadly expanded; fertilization duct short and curved ( Fig. 272 ).

Sexually dimorphic characters. Male with very weakly developed mesotrochanteric and mesofemoral brushes of setae. Male with pro- and mesobasotarsomeres ( Fig. 263 ) distinctly expanded compared with female ( Fig. 264 ); female protarsomere III ( Fig. 264A ) much longer and more slender than in male ( Fig. 263A ), female ( Fig. 264B ) and male ( Fig. 263B ) mesotarsomeres I–III more similar in size, but female mesotarsomeres narrower.

Intraspecific variation. This species is relatively consistent in characters. The most conspicuous variation is in the extent of shagrination on the pronotum and elytra. In a few specimens there is little or no shagrination. In others the entire pronotum and elytron is shagrinated. However, in most there is a characteristic pattern of shagrination on the pronotum ( Fig. 262 ) and some or none on the elytra.

Natural history: Vatellus grandis has been collected from subtropical marshes and moist forest and small forest pools.

Distribution: This is a moderately commonly collected species known from Brazil, Bolivia, French Guyana, Peru, Suriname and Trinidad (Fig. 314).

Phylogenetic relationships: This species groups with several others based on the common presence of similar cuticular sculpture (smooth and shiny between punctures ( Fig. 2 ) and with dense shagrination on the pronotum except for small lateral patches (e.g. Fig. 262 )) in most specimens, the anteromedial portion of the metasternum distinctly swollen ventrad (e.g. Fig. 20 ), a distinct pencil of stiff setae on the ventroapical surface of the male lateral lobe (e.g. Fig. 271 ), and very small female receptacles (e.g. Fig. 272 ).

Material examined: Bolivia: Beni, 2.0 km E San Borja , shaded pool in streambed. 14∞52¢02≤S 66∞43¢46≤W, 15 July 1998, KB Miller (1, KBMC) ; 6.7 km NW Trinidad. 14∞47¢27≤S 64∞56¢47≤W, 18 June 1999, blacklight, KB Miller (3, KBMC) . Brazil: Para stream near Aldeia Coraci , 3 December 1965, B Malkin (2, FSCA) ; Para, nr Aldeia Coraci , stream 11 km W Caninde, 3 December 1964, B Malkin (1, FSCA) . French Guyana: 1907, Geschenk (1, MNHN) ; Lawa River shore opposite Anapaike village , 25 November 1963, Borys, Malkin (14, FSCA) . Peru: Madre de Dios; Rio Tambopata Res. 30 air km SW Pto. Maldonado, 290 m, subtropical moist forest, J.B. Heppner (15, USNM) . Suriname: Carolina Ck 10 km from Zanderij , 18 November 1962, waterhole in forest, B Malkin (3, FSCA) ; Krekka-Phedra Road, Suriname dist 25 Tiny , 18 November 1962, tiny forest pool, Borys, Malkin (2, FSCA) ; Zanderijsavanne, Parakreek , 55∞10¢W 05∞25¢N, 20 November 1989, N Neiser (1, NHMW) . Trinidad: Camuto 1929, Darlington (69, MCZC) ; Cumuto, 1929, Darlington (8, MCZC) ; Sangre Grande, Cacoa ravines, April 1913, R Thaxter (12, MCZC) ; St Augustine , April 1929, Darlington (28, MCZC) .

VATELLUS BIFENESTRATUS ( ZIMMERMANN, 1921) COMB. NOV.

( FIGS 274–287 , 314)

Macrovatellus bifenestratus Zimmermann, 1921: 191 ; Trémouilles, 1995: 27.

Type information: Lectotype (designated here to clarify assignment of this name with this species), ♀ in ZSBS labelled, ‘Brazilien/Mato-Grosso Corumba/Type [handwritten on blue, circular disc]/Samml.A. Zimmermann/Zool. Staatssig. München [blue label]/ Holotypus Macrovatellus bifenestratus A.Zimm. Zool. Staatsammlg.München [taxon name and author handwritten, red label].’ A single male paralectotype accompanies the specimen labelled, ‘Brazilien/ Mato-Grosso Corumba/Type [handwritten on blue, circular disc]/Samml.A. Zimmermann/Zool. Staatssig. München [blue label]/Paratypoid Macrovatellus bifenestratus A.Zimm. Zool. Staatsammlg.München [taxon name and author handwritten].’

Type locality: Brazil, Mato-Grosso, Corumba.

Diagnosis: This species can be distinguished from other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced (as in Fig. 6 ); (2) elytra relatively coarsely punctate and with surfaces between the punctures generally shiny ( Fig. 2 ); (3) with a transverse impunctate area on each side of the midline on the pronotum; (4) eyes relatively small ( Fig. 274 ); (5) body relatively elongate and dorsoventrally depressed ( Fig. 274 ); (6) lateral pronotal margin distinctly curved ( Fig. 274 ); and (7) male genitalia distinctive ( Figs 282–285 ). The median lobe in lateral aspect is slender with the apex long, straight and apically narrowly rounded ( Figs 282, 284 ). The lateral lobe in lateral aspect has the apical portion long and straight with an apical, curved, apically sharply pointed flange ( Fig. 285 ).

Description:

Habitus. Body outline moderately discontinuous in dorsal aspect, lateral margins of elytra not strongly rounded ( Fig. 274 ), dorsoventrally moderately compressed. Measurements are given in Table 1.

Coloration. Head, pronotum and elytron dark red; venter red to testaceous in areas; appendages yellowred to dark red.

Sculpture and structure. Head densely punctate, punctures moderate in size medially, finer anteriorly on frons and clypeus, surface shiny and smooth; anteriorly clypeal margin distinctly protruding; eyes relatively small, slightly protuberant ( Fig. 274 ). Pronotum moderately punctate, covered with sculpturing consisting of extremely fine tubercles making surface shagrined except for two transverse areas on each side of midline which are smooth, shiny and impunctate similar to Figure 2 , each puncture with pale, short, fine seta; pronotum cordate, broadest near anterior angles, lateral margins broadly curved anteriorly, posterior angles subacute ( Fig. 274 ); lateral bead somewhat obscured, especially posteriorly; transverse sulcus absent. Elytron densely punctate, punctures moderately coarse, finer posteriorly, surface between punctures shiny, smooth ( Fig. 275 ). Prosternum moderately short; setose; prosternal process broad, flattened, lateral margins rounded, apex narrowly pointed. Metasternum coarsely punctate laterally, punctures relatively sparse, finer medially, surface between punctures shiny, smooth; anteromedial margin conspicuously swollen medially, ventrally produced. Abdominal sterna finely punctate laterally, surface smooth, shiny; sixth sternum with apical lobe produced, flattened, apically narrowly rounded ( Fig. 280 ); speleum broad, relatively short, apically narrowly rounded ( Fig. 281 ). Metacoxa with lateral portion relatively coarsely punctate over entire surface; lines approximate basally, strongly divergent anteriorly ( Fig. 276 ); medial portion finely and densely punctate. Metatrochanter short, rounded, ball-shaped, offset ( Fig. 277 ); metatarsal claws finely dentate.

Male genitalia. Median lobe in lateral aspect evenly curved, narrowed subapically, apex narrowly rounded, slightly curved ventrally, without subapical setae but with numerous distinct pores distributed randomly over surfaces, basal portion broad, robust ( Figs 282, 284 ); in ventral aspect narrow, apically with lateral margins evenly convergent to relatively sharp apex ( Fig. 283 ). Lateral lobe with apical portion narrow, margins parallel to acute apex, apex with prolonged, apically sharply pointed and curved flange, apicomedial margin with area of dense, fine, elongate setae; subapical stout, curved setae absent ( Fig. 285 ).

Female genitalia. Gonocoxosternite very broad and robust, apical angle moderately rounded, anterior margin concave, anterior lobe relatively short, slender ( Fig. 286 ). Gonocoxa elongate and slender, irregularly curved, apex relatively narrowly rounded, apodeme elongate and slender ( Fig. 286 ). Bursa copulatrix short and slender ( Fig. 286 ); spermathecal duct moderately long, slender; receptacle small, much smaller than spermatheca, spherical ( Fig. 286 ); intermediate duct moderately long, with several tight curves, particularly near spermatheca ( Fig. 287 ); spermatheca subspherical, triangular process large and very broadly expanded; fertilization duct short and curved ( Fig. 286 ).

Sexually dimorphic characters. Male with weakly developed mesofemoral brush of setae and moderately developed mesotrochanteric setal brush. Male pro- and mesobasotarsomeres ( Fig. 278 ) not strongly expanded, only slightly more expanded than female tarsomeres ( Fig. 279 ), male protarsomeres relatively slender, protarsomere III ( Fig. 278A ) much longer than mesotarsomere III ( Fig. 278B ).

Distribution: This species is known from the type locality, Corumba, Mato Grosso, Brazil and central Bolivia (Fig. 314).

Phylogenetic relationships: This species is a member of the V. grandis clade ( Fig. 316 ). Within this group, the species are extremely similar except for particular species-level autapomorphies, and this species does not group with any other particular member of the group.

Material examined: Bolivia: Beni, Prov. Cercado, 9.5 km N Trinidad 14∞46¢34≤S 64∞58¢00≤W, 18 June 1999, KB Miller (1, KBMC) .

VATELLUS DRYMETES MILLER SP. NOV.

( FIGS 288–297 , 313)

Type information: Holotype ♂ in USNM labelled, ‘ PERU: Madre de Dios; Rio Tambopata Res ; 30 air km. SW Pto.Maldonado, 290 m 11-15 XI 1979 JB. Heppner subtropical moist forest / HOLOTYPE Vatellus drymetes Miller 2004 [red label with double black line border].’

Paratypes: 1 ♂ in USNM labelled same as paratype.

Type locality: Peru, Madre de Dios, Rio Tambopata Res, 30 air km SW Pto. Maldonado.

Diagnosis: This species can be distinguished from other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced as in Figure 6 ; (2) with pronotum and elytra relatively coarsely punctate and with surfaces between the punctures generally shiny, surface sculpture as in Figure 2 ; (3) with a transverse impunctate area on each side of the midline on the pronotum (as in Fig. 262 ); (4) eyes moderate in size, not conspicuously protuberant ( Fig. 288 ); (5) body relatively elongate and dorsoventrally depressed ( Fig. 288 ); (6) lateral pronotal margin moderately curved ( Figs 288 ); and (7) male genitalia distinctive ( Figs 293–295 ). The median lobe in lateral aspect is very long, curved and slender with the apex extremely slender, and apically sharply pointed ( Fig. 293 ). The lateral lobe is irregular in shape, very broad basally with the apical portion long and slender with two or three long, stiff setae subapically on the ventral surface ( Fig. 295 ).

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Description:

Habitus. Body outline moderately discontinuous in dorsal aspect, lateral margins of elytra not strongly rounded ( Fig. 288 ), body dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head, pronotum, and elytron brown-red; ventral surfaces black except venter of head, pronotal epipleuron, propleuron, prosternum and small portions of posterior margins of abdominal sterna brown-red.

Sculpture and structure. Head finely and moderately densely punctate, slightly more dense anteriorly on frons and clypeus; anterior clypeal margin strongly anteriorly produced; eyes moderate in size, not strongly protruding ( Fig. 288 ). Pronotum with punctures relatively large laterally, denser medially and along posterior margin, punctures with fine pale, moderately elongate setae, area between punctures smooth, with dense shagrination consisting of dense field of very fine tubercles except for medial transverse impunctate areas on each side of midline similar to Figure 262 ; pronotum broadest anteromedially, lateral margins curved at extreme anterior end, pronotum cordate, posterior angles acute ( Fig. 288 ); lateral bead obscured except at extreme anterior end; transverse groove absent. Elytron strongly and densely punctate over entire surface ( Fig. 290 ), punctures finer and slightly more dense apically, each puncture with fine seta, surface similar to pronotum. Prosternum short, setose; prosternal process broad, flattened with short, indistinct, rounded medial carina, lateral margins rounded, convergent to triangular, pointed apex. Metasternum laterally with moderately large punctures, relatively sparsely distributed, medially with more dense, finer punctures, surface smooth, shiny; anteromedial margin strongly expanded ventrally into distinct tumidity. Abdominal sternal sutures not unusually modified; sterna laterally with moderately large, moderately dense punctures, interpuncture surface smooth, shiny; apical lobe on sixth abdominal sternum strongly produced, long, apically round, flattened ( Fig. 296 ); speleum elongate, moderately narrow, apically narrowly rounded ( Fig. 297 ). Metacoxa with lateral portion with punctures moderately large, relatively dense, interpuncture surface smooth, shiny; metacoxal lines strongly approximate posteriorly, anteriorly strongly divergent ( Fig. 291 ); medial portion with surface shiny, finely punctate. Metatrochanter strongly rounded, strongly offset ( Fig. 289 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect extremely slender and narrowed to sharply pointed apex, basal portion broad, broadly rounded; without setae at apex ( Fig. 293 ); in ventral aspect relatively narrow, lateral margins parallel through most of length, apically with margins evenly narrowed to pointed apex ( Fig. 294 ). Lateral lobe broad basally, apex straight, elongate and slender, with distinct ventral furrow, apex narrowly rounded, with numerous setae apicomedially and apicodorsally ( Fig. 295 ).

Female genitalia. Female not observed.

Sexually dimorphic characters. Male with very weakly developed mesotrochanteric and mesofemoral brushes of setae. Female not observed, but male pro- and mesobasotarsomeres not strongly expanded, protarsomere III very long and slender ( Fig. 292 ).

Intraspecific variation. No variation was noted in the few specimens examined.

Natural history: This species has been collected from 290 m in subtropical moist forest.

Distribution: Vatellus drymetes is known only from the type locality in Peru (Fig. 313).

Phylogenetic relationships: This species is a member of the V. grandis clade ( Fig. 316 ). Within this group this species does not group with any other particular member of the group.

Etymology: This species is named from the Greek word drymo, meaning ‘forest’, and the suffix - etes, meaning ‘to dwell’, in reference to the tropical forest area of the type locality (Tambopata Reserve, Peru).

VATELLUS AMAE MILLER SP. NOV.

( FIGS 298–310 , 314)

Type information: Holotype ♂ in USNM labelled, ‘ BRAZIL: Para ; Rio Xingu Camp (52∞22¢W 3∞39¢S) Altamira (c. 60 km S) 15 October 1986 P. Spangler & O. Flint / Colln. #24, pond at 2nd palm grove on trail 1 / HOLOTYPE Vatellus amae Miller 2004 [red label with double black line border].’

Paratypes: 2 in USNM labelled same as holotype. Brazil: Para , Rio Iriri Camp , 52∞40¢W 3∞50¢ S Altamira, c. 100 km S, Lago Sao Joao north side of Iriri River, P. Spangler and O. Flint (1, USNM); Para, Rio Xingu Camp, 52∞22¢W 3∞39¢ S Altamira, c. 60 km S, pond at 2nd palm grove on trail, P. Spangler and O. Flint (3, USNM) .

Type locality: Brazil, Para, Rio Xingu Camp, c. 60 km S Altamira, 52∞22¢W 3∞39¢S.

Diagnosis: This species can be distinguished from other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced (as in Fig. 6 ); (2) with surfaces relatively coarsely punctate and with surfaces between the punctures generally shiny, surface sculpture as in Figure 2 ; (3) with a transverse impunctate area on each side of the midline on the pronotum (as in Fig. 262 ); (4) eyes relatively small ( Fig. 298 ); (5) body relatively elongate and dorsoventrally depressed ( Fig. 298 ); (6) lateral pronotal margin relatively weakly curved ( Figs 298 ); and (7) male and female genitalia distinctive ( Figs 306–310 ). The median lobe in lateral aspect is slender with the apex narrowed to a very narrowly rounded apex that is slightly curved ventrally ( Fig. 306 ). The lateral lobe has the apical portion long and narrow with two or three long, stiff, subapical setae ( Fig. 308 ). The female receptacle is relatively small and spherical ( Fig. 309 ).

Description:

Habitus. Body outline moderately discontinuous in dorsal aspect, lateral margins of elytra not strongly rounded ( Fig. 298 ), dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head, pronotum, and elytron brown-red; ventral surfaces dark red-brown. Appendages redbrown.

Sculpture and structure. Head finely and moderately densely punctate, slightly more dense anteriorly on frons and clypeus; anterior clypeal margin relatively strongly anteriorly produced; eyes moderate in size, not strongly protruding ( Fig. 298 ). Pronotum with punctures relatively large laterally, denser medially and along posterior margin, punctures with fine pale, moderately elongate setae, area between punctures smooth, shiny on anterior and medial portions of pronotum, posterior half, lateral edges and a transverse anterior area with very fine shagrination consisting of dense field of very fine tubercles except for transverse impunctate, smooth areas medially on each side of midline similar to those shown in Figure 262 ; pronotum broadest anteromedially, lateral margins curved mainly in anterior half, moderately cordate, posterior angles acute ( Fig. 298 ); lateral bead obscured except at extreme anterior end; transverse groove absent. Elytron strongly and densely punctate over entire surface, punctures finer and slightly more dense apically ( Fig. 299 ), each puncture with fine seta, surface smooth and shiny between punctures. Prosternum short, setose; prosternal process broad, flattened with short, indistinct, rounded medial carina, lateral margins rounded, convergent to broadly rounded apex, with few long, fine setae apically. Metasternum laterally with moderately large punctures, relatively sparsely distributed, medially with more dense, large punctures, surface smooth, shiny; anteromedial margin strongly expanded ventrally into distinct tumidity. Abdominal sternal sutures not unusually modified; sterna laterally with moderately large, moderately dense punctures, interpuncture surface smooth, shiny; apical lobe on sixth abdominal sternum strongly produced, long, apically round, flattened ( Fig. 304 ); speleum elongate, relatively narrow, apically narrowly rounded ( Fig. 305 ). Metacoxa with lateral portion with punctures moderately large, relatively dense, interpuncture surface smooth, shiny; metacoxal lines strongly approximate posteriorly, anteriorly strongly divergent ( Fig. 300 ); medial portion with surface shiny, finely punctate. Metatrochanter strongly rounded, strongly offset ( Fig. 301 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect moderately straight in apical half, slender, apically gradually narrowed to slender, slightly hooked, narrowly rounded apex ( Fig. 306 ); in ventral aspect with lateral margins evenly and gradually convergent to narrowly rounded to nearly pointed apex ( Fig. 307 ). Lateral lobe long, basally slender, apical portion straight and narrow to rounded apex; with 2–3 subapical, stout, elongate, curved setae on ventral margin ( Fig. 308 ).

Female genitalia. Gonocoxosternite very broad and robust, apical angle moderately narrowly rounded, anterior margin relatively narrowly concave, anterior lobe relatively short, narrow, apex rounded ( Fig. 309 ). Gonocoxa short, robust, rounded, apex broadly rounded, apodeme elongate, slender ( Fig. 309 ). Bursa copulatrix robust, broad, curved dorsally and gradually narrowed to base of spermathecal duct ( Fig. 309 ); spermathecal duct long; receptacle small, about quarter size of spermatheca, spherical ( Fig. 309 ); intermediate duct relatively short, slender, only slightly sinuate ( Fig. 310 ); spermatheca elongate-spherical, large, triangular process moderately long, very broadly expanded; fertilization duct moderately long, relatively straight ( Fig. 309 ).

Sexually dimorphic characters. Male with very weakly developed mesotrochanteric and mesofemoral brushes of setae. Male with pro- and mesobasotarsomeres ( Fig. 302 ) only somewhat expanded compared with female ( Fig. 303 ), both male and female with protarsomere III relatively elongate ( Figs 302A, 303A ), but slightly longer in female ( Fig. 303A ).

Intraspecific variation. This species is relatively constant in characters in the few specimens examined. There is some minor variation in the extent of the microsculpture present on the pronotum.

Natural history: Specimens have been collected from ponds in palm woods.

Distribution: This species is known only from a few close localities in Brazil (Fig. 314).

Phylogenetic relationships: This species is a member of the V. grandis clade ( Fig. 316 ). Within this group this species does not group with any other particular member of the group.

Etymology: This species is named in honour of my wife, Amy Beth M. Miller.

Figures

Figures 203–210. Vatellus perforatus. 203, dorsal habitus; 204, elytral sculpture in area indicated on habitus by dotted square; 205, medial portion of metacoxae; 206, metatrochanter and metafemur, anterior aspect; 207–208, tarsomeres, dorsal aspect; 207, male; 208, female, A, protarsus; B, mesotarsus; 209, sternum VI, ventral aspect; 210, speleum abdominal sterna V, VI, dorsal aspect.

Figures 211–215. Vatellus perforatus. 211–213, male genitalia; 211, right lateral lobe, lateral aspect; 212, median lobe, right lateral aspect; 213, median lobe, ventral aspect; 214–215, female genitalia; 214, ventral aspect; 215, spermatheca and associated structures, dorsal aspect.

Figures 315–316. Vatellini species, consensus cladograms. 315, Consensus cladogram of 72 most parsimonious cladograms (length = 78, CI = 67, RI = 92) resulting from cladistic analysis of Vatellini including Calicovatellus petrodytes. 316, Consensus cladogram of five most parsimonious cladograms (length = 74, CI = 70, RI = 93) resulting from cladistic analysis of Vatellini not including Calicovatellus petrodytes. Numbers above branches are Bremer support values.

Figures 2–7. Vatellini species, morphology. 2–5, elytral sculpture; 2, Vatellus grandis; 3, V. tarsatus; 4, Derovatellus lentus; 5, V. mexicanus. 6–7, Vatellus species, crania, lateral aspect; 6, V. grandis; 7, V. mexicanus.

Figures 216–223. Vatellus tarsatus. 216, dorsal habitus; 217, elytral sculpture in area indicated on habitus by dotted square; 218, medial portion of metacoxae; 219, metatrochanter and metafemur, anterior aspect; 220–221, tarsomeres, dorsal aspect; 220, male; 221, female, A, protarsus; B, mesotarsus; 222, sternum VI, ventral aspect; 223, speleum abdominal sterna V, VI, dorsal aspect.

Figures 224–229. Vatellus tarsatus. 224–227, male genitalia; 224, apex of median lobe, ventral aspect; 225, median lobe, right lateral aspect; 226, median lobe, ventral aspect; 227, right lateral lobe, lateral aspect; 228–229, female genitalia; 228, ventral aspect; 229, spermatheca and associated structures, dorsal aspect.

Figures 238–244. Vatellus annae. 238–242, male genitalia; 238, right lateral lobe, lateral aspect; 239, median lobe, right lateral aspect; 240, median lobe, ventral aspect; 241, apex of median lobe, ventral aspect; 242, apex of median lobe, right lateral aspect; 243–244, female genitalia; 243, ventral aspect; 244, spermatheca and associated structures, dorsal aspect.

Figures 267–273. Vatellus grandis. 267–268, metatrochanter and metafemur, anterior aspect; 267, in flexion; 268, in extension; 269–271, male genitalia; 269, median lobe, right lateral aspect; 270, median lobe, ventral aspect; 271, right lateral lobe, lateral aspect; 272–273, female genitalia; 272, ventral aspect; 273, spermatheca and associated structures, dorsal aspect.

Figures 253–258. Vatellus pilacaudus. 253–256, male genitalia; 253, right lateral lobe, lateral aspect; 254, apex of median lobe, ventral aspect; 255, median lobe, right lateral aspect; 256, median lobe, ventral aspect; 257–258, female genitalia; 257, ventral aspect; 258, spermatheca and associated structures, dorsal aspect.

Figures 230–237. Vatellus annae. 230, dorsal habitus; 231, elytral sculpture in area indicated on habitus by dotted square; 232, medial portion of metacoxae; 233, metatrochanter and metafemur, anterior aspect; 234–235, tarsomeres, dorsal aspect; 234, male; 235, female, A, protarsus; B, mesotarsus; 236, sternum VI, ventral aspect; 237, speleum abdominal sterna V, VI, dorsal aspect.

Figures 245–252. Vatellus pilacaudus. 245, dorsal habitus; 246, elytral sculpture in area indicated on habitus by dotted square; 247, medial portion of metacoxae; 248, metatrochanter and metafemur, anterior aspect; 249–250, tarsomeres, dorsal aspect; 249, male; 250, female, A, protarsus; B, mesotarsus; 251, sternum VI, ventral aspect; 252, speleum abdominal sterna V, VI, dorsal aspect.

Figures 8–17. Vatellus grandis, mouthparts. 8, labrum and epipharynx, ventral aspect. 9–14, mandibles; 9, left mandible, dorsal aspect; 10, right mandible, dorsal aspect; 11, right mandible, ventral aspect; 12, left mandible, ventral aspect; 13, left mandible, medial aspect; 14, right mandible, medial aspect. 15, left maxilla, ventral aspect. 16–17, labium; 16, left lateral aspect; 17, dorsal aspect.

Figures 23–27. Vatellini species, morphology. 23–24, left metathoracic wing; 23, Derovatellus lentus; 24, Vatellus grandis; A, anal; AA, anal anterior; AP, anal posterior; C, costa; Cu, cubitus; CuA, cubitus anal; M, media; mf, submedial binding patch; MP, media posterior; oc, oblongum cell; R, radius; RA, radius anterior; RP, radius posterior; Sc, subcosta; wc, wedge cell. 25, V. grandis, field of protarsal ventral adhesive setae; 26, V. grandis, metafemur, posterior margin; 27, metatarsomere V and metatarsal claws.

Figures 38–50. Vatellini species, morphology. 38–39, Vatellus grandis, metendosternite; 38, dorsal aspect; 39, right lateral aspect. 40–41, Derovatellus lentus, metendosternite; 40, dorsal aspect; 41, right lateral aspect. 42–44, V. grandis, genital capsule; 42, abdominal sternum VIII; 43, spiculum; 44, genital ring. 45–47, D. lentus, genital capsule; 45, abdominal sternum VIII; 46, spiculum; 47, genital ring. 48–49, V. grandis, ejaculatory duct sclerite; 48, lateral aspect; 49, dorsal aspect. 50, D. dagombae, apex of lateral lobe.

Figures 259–266. Vatellus grandis. 259, dorsal habitus; 260, elytral sculpture in area indicated on habitus by dotted square; 261, medial portion of metacoxae; 262, left half of pronotum, dorsal aspect; 263–264, tarsomeres, dorsal aspect; 263, male; 264, female, A, protarsus; B, mesotarsus; 265, sternum VI, ventral aspect; 266, speleum abdominal sterna V, VI, dorsal aspect.

Figures 274–281. Vatellus bifenestratus. 274, dorsal habitus; 275, elytral sculpture in area indicated on habitus by dotted square; 276, medial portion of metacoxae; 277, metatrochanter and metafemur, anterior aspect; 278–279, tarsomeres, dorsal aspect; 278, male; 279, female, A, protarsus; B, mesotarsus; 280, sternum VI, ventral aspect; 281, speleum abdominal sterna V, VI, dorsal aspect.

Figures 18–22. Hydroporinae species, morphology. 18, Hydroporus notabilis, left prothorax, left oblique aspect. 19–21, Vatellus grandis; 19, left prothorax, left oblique aspect; 20, left mesosternum, mesocoxa, metasternum and metacoxa, left oblique aspect; 21, mesosternum, mesolegs, metasternum and metalegs. 22, Derovatellus lentus, mesosternum, mesolegs, metasternum and metalegs. CxI, procoxa; CxII, mesocoxa; FeI, profemur; FeII, mesofemur; Msem, mesepimeron; Mses, mesepisternum; Msst, mesosternum; Mtes, metepisternum; Mtst, metasternum; Prel, pronotal epipleuron; Prpl, propleuron; Prpo, prosternal pore; Prpr, prosternal process; Prst, prosternum; TrI, protrochanter; TrII, mesotrochanter.

Figures 282–287. Vatellus bifenestratus. 282–285, male genitalia; 282, median lobe, right lateral aspect; 283, median lobe, ventral aspect; 284, apex of median lobe, right lateral aspect; 285, right lateral lobe, lateral aspect; 286–287, female genitalia; 286, ventral aspect; 287, spermatheca and associated structures, dorsal aspect.

Figures 288–297. Vatellus drymetes. 288, dorsal habitus; 289, metatrochanter and metafemur, anterior aspect; 290, elytral sculpture in area indicated on habitus by dotted square; 291, medial portion of metacoxae; 292, male tarsomeres, dorsal aspect, A, protarsus; B, mesotarsus; 293–295, male genitalia; 293, median lobe, right lateral aspect; 294, median lobe, ventral aspect; 295, right lateral lobe, lateral aspect; 296, sternum VI, ventral aspect; 297, speleum abdominal sterna V, VI, dorsal aspect.

Figures 298–305. Vatellus amae. 298, dorsal habitus; 299, elytral sculpture in area indicated on habitus by dotted square; 300, medial portion of metacoxae; 301, metatrochanter and metafemur, anterior aspect; 302–303, tarsomeres, dorsal aspect; 302, male; 303, female, A, protarsus; B, mesotarsus; 304, sternum VI, ventral aspect; 305, speleum abdominal sterna V, VI, dorsal aspect.

Figures 306–310. Vatellus amae. 306–308, male genitalia; 306, median lobe, right lateral aspect; 307, median lobe, ventral aspect; 308, right lateral lobe, lateral aspect; 309–310, female genitalia; 309, ventral aspect; 310, spermatheca and associated structures, dorsal aspect.