Vatellus lateralis 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 : 466-475

VATELLUS LATERALIS (SHARP, 1882) COMB. NOV.

Macrovatellus lateralis Sharp, 1882b: 283 ; Zimmermann, 1920: 30; Trémouilles, 1995: 27.

Macrovatellus rudis Sharp, 1882b: 283 ; Zimmermann, 1920: 30; Trémouilles, 1995: 27, syn. nov.

Macrovatellus marginalis Sharp, 1882b: 284 ; Zimmermann, 1920: 30; Trémouilles, 1995: 27, syn. nov.

Macrovatellus deplanatus Zimmermann, 1919: 125 ; 1920: 30; Trémouilles, 1995: 27, syn. nov.

Type information: Macrovatellus lateralis : Lectotype (designated here to clarify assignment of this name with this species) ♂ in BMNH labelled, ‘Type [circular disc with red border]/ Uruguay. [green line transversely across middle of label]/Sharp Coll 1905–313/Type 438G Macrovatellus lateralis Uruguay 11.5. [handwritten]/ SYN- TYPE [circular disc with blue border]/LECTO- TYPE Macrovatellus lateralis Sharp, 1882 . des. K.B. Miller 2002 [red label with black line border].’ There is a single paralectotype in BMNH labelled, ‘Type 438 E [handwritten]/Para-type [circular disc with yellow bor- der]/ Uruguay. [green line transversely across middle of label]/Sharp Coll 1905–313/ Macrovatellus lateralis, Sharp Paratype. [handwritten]/SYN- TYPE [circular disc with blue border]/ PARALECTOTYPE Macrovatellus lateralis Sharp 1882 . des. K.B. Miller 2002 [blue label with black line border].’

Macrovatellus rudis : Lectotype (designated here to clarify assignment of this name with this species) ♂ in BMNH labelled, ‘Type [circular disc with red border]/ S.America. Colombia. [green line transversely across middle of label]/Sharp Coll 1905–313./Type 430 ♂ Macrovatellus rudis Am.Mer. [handwritten]/SYN- TYPE [circular disc with blue border] / LECTOTYPE Macrovatellus rudis Sharp 1882 . des. K.B. Miller 2002 [red label with black line border].’ Sharp indicated that he had multiple specimens (both males and females) when he described this species. However, only a single male in BMNH was examined that appears to be of the original syntype series. It is selected as the lectotype.

Macrovatellus marginalis : Lectotype (designated here to clarify assignment of this name with this species) ♀ in BMNH labelled, ‘Type [circular disc with red border]/ S. America: Brazil. [green line transversely across middle of label]/Santa Rita Sep Sahlberg 1850 [handwritten]/Type 441 Macrovatellus marginalis Santa Rita [handwritten]/SYN- TYPE [circular disc with blue border].’ Sharp had an additional female specimen. Of the two specimens in BMNH labelled as this species and from Sharp’s collection, only one is a female with label information that would indicate it is one of the two original syntypes. Thus it is selected as the lectotype. The other male specimen is probably not of the original type series.

Macrovatellus deplanatus: The type of this species is apparently not in MNHU (M. Uhlig, pers. comm.) nor in ZSBS (M. Baehr, pers. comm.) (the latter the location of most of Zimmermann’s types). The type locality and Zimmermann’s description (1919) strongly suggest that the species is the same as the one treated here as V. lateralis . Two specimens of V. lateralis in ZSBS labelled as ‘ M.? deplanatus ’ (clearly not the types as they are labelled as having been collected in Venezuela) were identified by Guignot, suggesting that he regarded V. lateralis and M. deplanatus as similar, at least. Based on these lines of evidence, I have synonymized this name with V. lateralis . However, until the types are found, this synonymy should be regarded as provisory.

Type locality: Macrovatellus lateralis : Uruguay. Macrovatellus rudis : South America.

Macrovatellus marginalis : Brazil, Santa Rita. Macrovatellus deplanatus : Brazil, Santos.

Diagnosis: This species may be distinguished from other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces distinctly alutaceous, elytral surface sculpture as in Figure 5 ; (2) eyes large, protruding ( Fig. 161 ); (3) transverse pronotal crease distinctive ( Fig. 161 ); (4) male pro- and mesobasotarsomeres ( Fig. 165 ) much broader than in female ( Fig. 166 ); (5) apical lobe on abdominal sternum VI present but not strongly expressed ( Fig. 167 ); (6) male and female genitalia distinctive ( Figs 169– 173 ). This species is very similar to V. mexicanus from which is distinguishable by the median lobe more slender and apically slightly deflexed and the intermediate duct between the spermatheca and receptacle longer and more strongly sinuate. The apices of the lateral lobes extend to or beyond the apex of the median lobe in V. mexicanus ( Fig. 140 ) whereas in V. lateralis the apices of the lateral lobes end well short of the apex of the median lobe ( Fig. 139 ).

Description:

Habitus. Lateral outline distinctly discontinuous in dorsal aspect, lateral elytral margins evenly rounded ( Fig. 161 ), body not dorsoventrally flattened. Measurements are given in Table 1.

Coloration. Head yellow; pronotum brown, irregularly yellow along anterior and lateral margins; elytron dark brown to yellow-brown with yellow maculae subapically and laterally along margins, basally with few yellow maculae ( Fig. 161 ); venter and appendages red brown to light brown, infuscate, especially laterally on metacoxae.

Sculpture and structure. Head with punctation distinct, variable in size, coarse medially between eyes, becoming finer anteriorly, interpuncture surfaces smooth, shiny to slightly microreticulate, surface posterior to eyes alutaceous, rough; anterior clypeal margin rounded; eyes large, strongly protruding ( Fig. 161 ). Pronotum densely punctate, but punctures strongly obscured by rough, alutaceous sculpturing similar to Figure 5 , pale setae moderately long; pronotum cordate, lateral pronotal margin rounded anteriorly, broadest near anterior angles, posterior angles acute ( Fig. 161 ); lateral bead obscured; transverse sulcus distinctly impressed. Elytron with punctation dense but edges of punctures strongly obscured by dense alutaceous microsculpturing similar to Figure 5 , light setae moderately dense and conspicuous. Prosternum short, roughly sculptured, glabrous; prosternal process broad with distinctly rounded medial carina, lateral margins rounded, apex triangularly produced, glabrous, apex pointed. Metasternum with punctation dense laterally, punctures large and shallow, somewhat obscured by alutaceous sculpturing; anteromedial margin slightly produced ventrally. Abdominal sternal sutures not unusually modified; sterna with punctation dense over entire lateral surface, finer medially, obscured by alutaceous sculpturing; sixth abdominal sternum with apical lobe inconspicuously produced, triangular, not strongly produced ( Fig. 167 ); speleum broad basally, sides strongly convergent, apex moderately rounded, not strongly curved ( Fig. 168 ). Metacoxae with lateral portion densely punctate over entire surface, punctures shallow, somewhat obscured by alutaceous sculpturing; metacoxal lines strongly approximate posteriorly, strongly divergent anteriorly ( Fig. 163 ); medial portion moderately punctate, punctures finely setose. Metatrochanter rounded, offset ( Fig. 164 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect evenly curved, apically gradually narrowed, becoming slen- der, apex narrowly rounded, distinctly but not sharply curved dorsally, with few subapical setae, basal portion elongate and moderately broad ( Fig. 170 ); in ventral aspect moderately broad, margins slightly convergent apically, somewhat sinuate basomedially, apex narrowed relatively abruptly, apex acutely rounded ( Fig. 171 ). Lateral lobe relatively broad, apical portion evenly and shallowly curved, broad to broadly rounded apex, apical an apicomedial margin with dense series of fine setae; subapical stout, curved setae absent ( Fig. 169 ).

Female genitalia. Gonocoxosternite moderately robust, medial margin moderately broad, very irregular; anterior margin strongly concave; anterior lobe relatively slender and elongate ( Fig. 172 ). Gonocoxa broad, medial margin broadly rounded, apex broadly rounded; apodeme short and slender ( Fig. 172 ). Bursa copulatrix moderate in size, not modified; spermathecal duct moderately long; receptacle ovoid, about half the size of spermatheca; intermediate duct moderately long, strongly sinuate ( Fig. 173 ); spermatheca subspherical, triangular process moderately long, broad; fertilization duct short ( Fig. 172 ).

Sexually dimorphic characters. Mesotrochanter and mesofemur of male with setal brushes well developed, that of mesofemur moderately elongate. Male with pro- and mesobasotarsomeres ( Fig. 165 ) considerably broader than in female ( Fig. 166 ).

Intraspecific variation. This species varies substantially in size and shape. There is also considerable variation in degree of expression of the cuticular microsculpture with females exhibiting it to a greater degree. The extent of the maculations is highly variable, but maculae are generally distinct though in some cases they may be difficult to distinguish from the background coloration.

Natural history: This species is known from black lights, lowland marshes and subtropical moist forest.

Distribution: Vatellus lateralis has been collected from Bolivia, Brazil, Paraguay, Peru and Uruguay ( Fig. 311 ).

Phylogenetic relationships: This species is part of the clade containing species similar to V. mexicanus ( Fig. 316 ). However, within this group this species does not have clear affinities with any other particular species. It is very similar to V. mexicanus in many features.

Discussion: The species is widespread and relatively commonly collected throughout tropical South America. It has been described several times. Examination of the types of V. lateralis , V. rudis and V. marginalis indicate that these names refer to the same species. Vatellus lateralis comes first in Sharp’s (1882b) monograph, so I selected it as the senior name. The strong similarity between this species and V. mexicanus may indicate that the two should be classified as a single species. However, consistent differences in male and female genitalia and allopatric distributions suggest that the two are distinctive. Specimens distributed between the southern range of V. mexicanus and the northern range of V. lateralis should be examined in the future for intermediate character states.

Material examined: Bolivia: Beni, Beni Stn. Palm Camp , NE San Borja, at light, R.W. Brooks (1, USNM) ; Santa Ana , 14 August 1937, HE Hinton (16, BMNH) ; Trinidad, July–August 1937, HE Hinton (6, BMNH) ; Trinidad, 29–30 July 1937, HE Hinton (4, BMNH) ; Beni , 7.0 km SW Trinidad. 14∞52¢12≤S 64∞57¢32≤W, 18 July 1998, weedy pond, KB Miller (6, KBMC) ; Beni , Prov. Cercado 9.5 km N Trinidad. 14∞46¢34≤S 64∞50¢00≤S, 17 June 1999, shallow marsh, KB Miller (2, KBMC) . Brazil: Alto da Serra be Sao Paulo 1929 (2, MNHN) ; Bahia Encruzilhada , 960 m, 7 November 1972, Moacir Alvarenga (1, FSCA) ; Caraca , Minas Gerais, December 1885, E Gouveille (4, MNHN) ; Caraqa , 1884, P Germain (17, MNHN) ; Encruzilhada , 980 m, Bahia, November 1974, M Alvarenga (3, MNHN) ; Minas Gerias Aguas Vermlehas , December 1983, BLT, Moacir Alvarenga (6, FSCA) ; Minas Gerias, Aguas Vermelhas , December 1983, BLT, M Alvarenga (7, FSCA) ; Par. Curitiba , 29 June 1969, P and P. Spangler (7, USNM) ; Prov. Minas Gerais Aquas Vermelhas , December 1998, FZ Vaz-de-Mello (33, LHIC) ; Rio Grande del Norte, Natal , March 1952, M Alvarenga (3, FSCA) . Paraguay: Central Department, Cacupe Road, Arroyo Yagua Resa y Ypacai , 10 April 1980, PJ Spangler, MM Culzoni, D Wood (1, USNM) . Peru: Madre de Dios; Rio Tambopata Res. , 30 air km SW Pto Maldonado, 26–30 November 1979, subtropical moist forest, JP Heppner (1, USNM) . Uruguay: (1, MNHN) ; Artigma Supulturez, Rio Charreis Pionda de Negro Muezto , 10 December 1966 b, CS Carbonell, MA Mono, FR San Martin (1, FSCA) ; Montevideo, Al Cassabo , 26 March 1932 (1, FSCA) ; Payendui Rio Uruguay Barra Arroyo Guavigo , 9–11 February 1970, MS Moratorio, MA Mono, GJ Wibmer (1, FSCA) ; Rivera Arriera , 3–7 February 1933, CS Carbonell (1, FSCA) ; Rocha Ruta 10, Aguas Dulces , 09–11 October 1970, MS Moratorio, MA Mono, GJ Wibmer (2, FSCA) .

VATELLUS HAAGI WEHNCKE, 1876

( FIGS 174–188 , 313)

Vatellus haagi Wehncke, 1876: 357 .

Macrovatellus haagi ; Sharp, 1882b: 284 (comb. nov.); Régimbart, 1899b:1, 1903: 46; Zimmermann, 1919: 124, 1920: 30; Trémouilles, 1995: 27.

Type information: Lectotype (designated here to clarify assignment of this name with this species), ♂ in MNHN labelled, ‘[small, light purple square]/Haag [handwritten, black line around label]/Brazilia [handwritten, green label with black line around border]/ Haagi [handwritten, black line around border, additional illegible word below “Haagi”/ LECTOTYPE Vatellus haagi Wehncke, 1876 . des. by K.B. Miller 2003 [red label with double black line border].’ Wehncke (1876) did not indicate the number of specimens on which he based his description, although Nilsson (2001) concluded that there are syntypes. Wehncke’s specimens went to Oberthur and thence to the Museum National d’Histoire Naturelle, Paris ( MNHN). A single specimen in MNHN was examined that matches his description, and the label information currently on the specimen suggests that it, at least, is a specimen on which he based the Description: It is selected as the lectotype.

Type locality: Brazil, Corrientes. Diagnosis: This species may be distinguished from other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces distinctly alutaceous, elytral surface sculpture as in Figure 5 ; (2) eyes large, protruding ( Fig. 174 ); (3) transverse pronotal crease distinctive ( Fig. 174 ); (4) male pro- and mesobasotarsomeres ( Fig. 178 ) much broader than in female ( Fig. 179 ); (5) apical lobe on abdominal sternum VI present but not strongly expressed ( Fig. 180 ); (6) male and female genitalia distinct ( Figs 182–188 ). In this species the male median lobe is apically broadly truncate in ventral aspect ( Figs 183, 185 ) and distinctly deflexed in lateral aspect ( Fig. 182 ). The apex is strongly beset with short, proximally directed spines ( Fig. 185 ). The female bursa is the largest of any Vatellus and is broad, strongly dorsoventrally compressed and strongly sinuate ( Figs 186, 187 ). The prosternal process is apically relatively blunt, not flattened and apically pointed as in most Vatellus .

Description:

Habitus. Body outline strongly discontinuous in dorsal aspect, lateral margins of elytra strongly rounded ( Fig. 174 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow; pronotum yellow, yellowbrown medially, narrowly brown along anterior and broadly along posterior margins; elytron dark brown to yellow-brown with yellow maculae subapically and laterally along margins, basal margin irregularly and broadly yellow ( Fig. 174 ); venter and appendages redbrown to light brown, infuscate, especially laterally on metacoxae.

Sculpture and structure. Head with punctation distinct, variable in size, coarse medially between eyes, becoming finer anteriorly, interpuncture surfaces smooth, shiny to slightly microreticulate, surface posterior to eyes alutaceous, rough; anterior clypeal margin rounded; eyes large, strongly protruding. Pronotum densely punctate, but punctures strongly obscured by rough, alutaceous sculpturing similar to Figure 5 , pale setae moderately long; pronotum cordate, lateral pronotal margin rounded anteriorly, broadest near anterior angles, posterior angles slightly obtuse; lateral bead obscured; transverse sulcus distinctly impressed. Elytron with punctation dense but edges of punctures strongly obscured by dense alutaceous sculpturing similar to pronotum, light setae moderately dense and conspicuous. Prosternum short, roughly sculptured, glabrous; prosternal process broad, flat, medially with conspicuous protrusion to rounded carina, lateral margins parallelsided, apex broad, truncate with fringe of fine setae. Metasternum with punctation dense laterally, punctures large and shallow, somewhat obscured by alutaceous sculpturing; anteromedial margin narrowly produced ventrally, apex acute. Abdominal sternal sutures not unusually modified; sterna with punctation dense over entire lateral surface, finer medially, obscured by alutaceous sculpturing; sixth abdominal sternum with apical lobe narrow, triangular, not strongly produced; speleum broad basally, sides strongly convergent, apex narrowly rounded, not strongly curved. Metacoxae with lateral portion densely punctate over entire surface, punctures shallow, somewhat obscured by alutaceous sculpturing; metacoxal lines strongly approximate posteriorly, strongly divergent anteriorly ( Fig. 176 ); medial portion moderately punctate, punctures finely setose. Metatrochanter rounded, offset ( Fig. 177 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect moderately narrow, slightly expanded medially, apex narrowly rounded, distinctly curved dorsally, with large field of short spines over ventrolateral surfaces, basal portion long, narrow ( Fig. 182 ); in ventral aspect with lateral margins approximately parallel, apex abruptly truncate, broad ( Figs 183, 185 ). Lateral lobe very long, slender, with apical portion evenly curved, apically distinctly expanded and broadly rounded, densely covered with long, fine setae along lateral and apical margins; with long, stout, subapical setae absent ( Fig. 184 ).

Female genitalia. Gonocoxosternite broad, robust, apical angle rounded, medial margin membranous, anterior margin distinctly concave, anterior lobe very long, robust, apically expanded and truncate ( Fig. 186 ). Gonocoxa robust, main portion broadest medially, apex obtusely rounded, with numerous spinous setae along apicoventral margins and over dorsal surface, apodeme narrow, as long as main portion ( Fig. 186 ). Bursa copulatrix very long, robust, broad and dorsoventrally flattened, apex folded dorsally ( Figs 186, 187 ); spermathecal duct moderately long, narrow; receptacle spherical, about two-thirds size of spermatheca ( Fig. 186 ); intermediate duct relatively short, curved but not strongly coiled or twisted ( Fig. 188 ); spermatheca subspherical, produced posteriorly, triangular process relatively narrow basally, broadly expanded and rounded apically; fertilization duct short, curved ( Fig. 186 ).

Sexually dimorphic characters. Male with welldeveloped mesotrochanteric brush of setae and mesofemoral brush well-developed in elongate cupshaped form. Male pro- and mesobasotarsomeres ( Fig. 178 ) much more broadly expanded than in female ( Fig. 179 ), similar in length ( Figs 178, 179 ).

Intraspecific variation. This species is somewhat variable in coloration and extent of maculations on the elytra. However, the species is, in general, relatively uniform in characters.

Natural history: Vatellus haagi has been collected at lights and from roadside pools.

Distribution: Vatellus haagi is found is southern South America and has been collected from Argentina, Bolivia, Paraguay and Uruguay (Fig. 313).

Phylogenetic relationships: This species is part of the clade containing species similar to V. mexicanus ( Fig. 316 ). However, within this group this species does not have clear affinities with any other particular species.

Material examined: Argentina: Ao. P Verne , 4 km N Va San Jose, 15 November 1973, OS Flint (1, USNM) ; Bs. As. Laguna Chascomus at Ao. San Felton , 20 February 1974, OS Flint (1, USNM) ; Buenos Aires, 17 November 1912 (3, BMNH) ; Buenos Aires, v Belgrano , 10–15 October 1905, R Thaxter (1, MCZC) ; Chaco, Charata , March 1991 (1, LHIC) ; Concordia , Prov Etre Rios (1, MNHN) ; Cord, Carlos Paz , 26–27 May 1969, PJ Spangler (1, USNM) ; Eldorado Misiones , 8 December 1964 (1, MNHN) ; Entre Rios, Colon , 1 February 1988 (1, LHIC) ; Estancia la Noria, Rio San Javier , Santa Fe, 19 December 1911, GE Bryant (2, BMNH) ; Hohenau. Alto-Parana , 6 September 1937, H Jacob (1, BMNH) ; Misiones, Jardin America, Salto Tabay , 22 December 1990, riverside forest UV light, S & J Peck (1, CMNC) ; Prov. Buenos Aires, Colon , 29 December 1991, at light, Archangelsky (3, LHIC) ; Prov. Chaco San Bernardo , 1 February 1982, at light, Dilorio (3, LHIC) ; Prov. Entre Rios, Department Colon / Liebig , October 1989, R Foerster (9, LHIC) ; Rio Paraguay, Formosa , 4 December 1973, OS Flint (1, USNM) ; Santa Fe Reconquista, 13 February 1979, RE Woodruff, H Cordo (1, FSCA) ; Villa Ana , FCSFe, 25 December, at light, KJ Hayward (1, BMNH) ; Villa Ana, P.C.S. Fe , January 1926, KJ Hayward (1, USNM) . Bolivia: San Antonio de Parapeti Rio Parapeti Santa Cruz, 15–19 July 1964, B Malkin (1, FSCA) ; Santa Cruz, 11–12 May 1969, P and P Spangler (2, USNM) ; Santa Cruz, 60 mi N Santa Cruz, Saavedra Exp. Sta. , 3–5 January 1960, R Cumming (1, USNM) ; Santa Cruz, Ayacucho , 13–14 May 1969, P and P Spangler (12, USNM) ; Sta. Cruz, Prov. Chiquitos, 1.3 km WSW San Jose , 28 June 1999, stream pool, KB Miller (16, KBMC) . Paraguay: December 1936 (1, MNHN); (1, MNHN) ; Asuncion, 7 October 1980, DC Lowrie (3, USNM) ; Asuncion, 8 January 1991, Dreshsel (1, LHIC) ; Cen. 15 km NE Asuncion, 21 June 1969, P and P Spangler (11, USNM) ; Cen. 2 km N Luque, 23–24 June 1969, P and P Spangler (32, USNM) ; Cen., Ypacarai , 22 June 1969, P and P Spangler (1, USNM) ; Cord. San Bernardino , 22 June 1969, P and P Spangler (18, USNM) ; Department Central, Lambare , 1991, Drechsel (1, LHIC) ; Department Central , Asuncion, 5 October 1991, U. Drechsel (1, LHIC) ; Hohenau, Alto-Parana , 5 September 1937, H Jacob (1, BMNH) ; Paraguari Department Ybycui 20 km SE, ruta La Rosada, Ybycui , 13 April 1980, roadside ponds, PJ Spangler (2, USNM) . Uruguay: Colonia, Carmelo , 11 January 1962, C Morey, M Monne, CS Carbonell (3, FSCA) ; Montevideo, Pocitos , 01 November 1932, CS Carbonell (1, FSCA) ; Payeandu , Rio Uruguay, Barra Arroyo Guaviyu , 9–11 February 1970 (1, FSCA) .

VATELLUS WHEELERI MILLER SP. NOV.

( FIGS 189–202 , 314)

Type information: Holotype ♂ in USNM labelled, ‘ ARGENTINA, Chaco Rcho.Barrangueras Puerto Vilelas 5 December 1973 O. S. Flint, Jr ./ HOLOTYPE Vatellus wheeleri Miller 2004 [red label with double black line border].’

Paratypes: 6 in USNM labelled same as holotype. Argentina: Corr. Ituazingo , 20 December 1990, wet grassland UV, S & J Peck (12, CMNC) . Paraguay: Department Central , Asuncion, 2 October 1992, Dreschsel (3, LHIC) ; Department Guaira, Calle Florida , LHIC (2, LHIC) ; Department Guaira, Garay , 10 August 1992, U. Dreschsel (2, LHIC) ; Department Paraguari, Sapucay , 9 April, Dreschsel (1, LHIC) .

Type locality: Argentina, Chaco, Barranqueras.

Diagnosis: This species may be distinguished from other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces distinctly alutaceous, elytral surface sculpture as in Figure 5 ; (2) eyes large, protruding ( Fig. 189 ); (3) transverse pronotal crease distinctive; (4) male pro- and mesobasotarsomeres ( Fig. 193 ) much broader than in female ( Fig. 194 ); (5) apical lobe on abdominal sternum VI present but weakly expressed ( Fig. 195 ); (6) speleum long, anteriorly expanded, strongly curved dorsally ( Fig. 196 ); and (7) male and female genitalia distinct ( Figs 197–202 ). The median lobe is apically relatively broadly rounded in ventral aspect ( Figs 197, 199 ), but not as truncated as in V. haagi . The gonocoxae in this species are more robust and strongly rounded than most species ( Fig. 201 ). However, this species is very similar to V. mexicanus , V. lateralis and V. haagi and it appears that the shape of the speleum ( Fig. 196 ) is its strongest diagnostic feature.

Description:

Habitus. Body outline moderately discontinuous in dorsal aspect, lateral margins of elytra moderately strongly rounded ( Fig. 189 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow-brown; pronotum dark brown, irregularly yellow anterolaterally; elytron dark brown to yellow-brown with yellow maculae subapically and laterally along margins ( Fig. 189 ); venter and appendages dark red-brown.

Sculpture and structure. Head with punctation distinct, moderately coarse, absent to extremely fine and spares in large, triangular area on frons and clypeus, interpuncture surfaces smooth, shiny to slightly microreticulate, surface posterior to eyes alutaceous, rough; anterior clypeal margin rounded; eyes large, prominently protruding ( Fig. 189 ). Pronotum densely punctate, but punctures obscured by alutaceous sculpturing similar to that shown in Figure 5 , pale setae moderately long; pronotum cordate, lateral pronotal margin rounded anteriorly, broadest near anterior angles, posterior angles slightly obtuse ( Fig. 189 ); lateral bead obscured; transverse sulcus moderately impressed, visible mainly laterally. Elytron with punctation dense but edges of punctures obscured by alutaceous sculpturing as on pronotum, light setae moderately dense and conspicuous. Prosternum short, shagrined, lightly setose; prosternal process broad, lateral margins rounded, basomedially slightly tectiform, medially excavated, apex flattened, sharply triangular, acuminate to slightly obtusely pointed. Metasternum with punctation dense laterally, punctures large and shallow, somewhat obscured by alutaceous sculpturing; anteromedial margin narrowly produced ventrally, apex acute. Abdominal sternal sutures not unusually modified; sterna with punctation dense over entire lateral surface, finer medially, obscured by alutaceous sculpturing; sixth abdominal sternum with apical lobe moderately produced, rounded, somewhat flattened; speleum broad, distinctly expanded, apex dorsally hooked dorsally ( Fig. 196 ). Metacoxae with lateral portion densely punctate over entire surface, punctures shallow, somewhat obscured by alutaceous sculpturing; metacoxal lines strongly approximate posteriorly, strongly divergent anteriorly ( Fig. 191 ); medial portion moderately punctate, punctures finely setose. Metatrochanter strongly rounded, offset ( Fig. 192 ); metatarsal claws minutely dentate.

Male genitalia. Median lobe in lateral aspect slightly and evenly curved throughout length, slender, basal portion elongate, relatively straight and narrow, slightly narrowed subapically ( Fig. 198 ); apex narrowly rounded, with numerous short spines subapically on ventral margin ( Fig. 197 ); in ventral aspect moderately broad basally, narrower in apical twothirds, apically broadly pointed ( Figs 197, 199 ). Lateral lobe with apical portion slightly and evenly curved, apex broadly rounded, with large apical and lateral region of long, fine setae; long, stout, subapical setae absent ( Fig. 200 ).

Female genitalia. Gonocoxosternite moderately broad, apical angle rounded, anterior margin deeply curved, anterior lobe elongate, moderately broad, sinuate ( Fig. 201 ). Gonocoxa short, broad and robust, apex broadly rounded, apodeme short and slender ( Fig. 201 ). Bursa copulatrix broad, irregular ( Fig. 201 ); spermathecal duct long, narrow; receptacle large, subequal in size to spermatheca, spherical ( Fig. 201 ); intermediate duct broad, curved, strongly curved ( Fig. 202 ); spermatheca spherical, large; fertilization duct large, abruptly and broadly expanded ( Fig. 201 ).

Sexually dimorphic characters. Male with welldeveloped, elongate cup-shaped brush of setae on mesofemora and mesotrochanter with dense region of setae; male pro- and mesobasotarsomeres ( Fig. 193 ) much broader than in female ( Fig. 194 ), female protarsomere III ( Fig. 194A ) longer than male protarsomere III ( Fig. 193A ).

Natural history: This species has been collected from a UV light in wet grassland.

Distribution: Vatellus wheeleri is known from southern South America and has been collected from Argentina and Paraguay (Fig. 314).

Phylogenetic relationships: This species is part of the clade containing species similar to V. mexicanus ( Fig. 316 ). However, within this group this species does not have clear affinities with any other particular species.

Etymology: This species is named in honour of Quentin D. Wheeler (The Natural History Museum, London) for his considerable contribution to taxonomic theory and knowledge of Coleoptera , for his extensive support of me during various research projects and for his highly valued friendship.

Figures

Figure 311. Vatellus species, distribution. - = V. lateralis, Δ = V. maculosus, • = V. sahlbergi.

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 161–168. Vatellus lateralis. 161, dorsal habitus; 162, elytral sculpture in area indicated on habitus by dotted square; 163, medial portion of metacoxae; 164, metatrochanter and metafemur, anterior aspect; 165–166, tarsomeres, dorsal aspect; 165, male; 166, female, A, protarsus; B, mesotarsus; 167, sternum VI, ventral aspect; 168, speleum abdominal sterna V, VI, dorsal aspect.

Figures 169–173. Vatellus lateralis. 169–171, male genitalia; 169, right lateral lobe, lateral aspect; 170, median lobe, right lateral aspect; 171, median lobe, ventral aspect; 172–173, female genitalia; 172, ventral aspect; 173, spermatheca and associated structures, dorsal aspect.

Figures 139–146. Vatellus species. 139, Vatellus lateralis, apices of median and right lateral lobes of aedeagus, right lateral aspect; 140–146, Vatellus mexicanus; 140–144, male genitalia; 140, apices of median and lateral lobes of aedeagus, right lateral aspect; 141, median lobe, right lateral aspect; 142, median lobe, ventral aspect; 143, right lateral lobe, lateral aspect; 144, apex of median lobe, right lateral aspect; 145-146, female genitalia; 145, ventral aspect; 146, 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 174–181. Vatellus haagi. 174, dorsal habitus; 175, elytral sculpture in area indicated on habitus by dotted square; 176, medial portion of metacoxae; 177, metatrochanter and metafemur, anterior aspect; 178–179, tarsomeres, dorsal aspect; 178, male; 179, female, A, protarsus; B, mesotarsus; 180, sternum VI, ventral aspect; 181, speleum abdominal sterna V, VI, dorsal aspect.

Figures 182–188. Vatellus haagi. 182–185, male genitalia; 182, median lobe, right lateral aspect; 183, median lobe, ventral aspect; 184, right lateral lobe, lateral aspect; 185, apex of median lobe, ventral aspect; 186–188, female genitalia; 186, ventral aspect; 187, bursa copulatrix, left lateral aspect; 188, spermatheca and associated structures, dorsal aspect; BC, bursa copulatrix; FC, fertilization sac sclerite; FD, fertilization duct; GC, gonocoxa; GS, gonocoxosternite; ID, intermediate duct; RA, rami; RE, receptacle; SD, spermathecal duct; SP, spermatheca; TP, truncate spermathecal process.

Figures 189–196. Vatellus wheeleri. 189, dorsal habitus; 190, elytral sculpture in area indicated on habitus by dotted square; 191, medial portion of metacoxae; 192, metatrochanter and metafemur, anterior aspect; 193–194, tarsomeres, dorsal aspect; 193, male; 194, female, A, protarsus; B, mesotarsus; 195, sternum VI, ventral aspect; 196, speleum abdominal sterna V, VI, dorsal aspect.

Figures 197–202. Vatellus wheeleri. 197–200, male genitalia; 197, apex of median lobe, ventral aspect; 198, median lobe, right lateral aspect; 199, median lobe, ventral aspect; 200, right lateral lobe, lateral aspect; 201–202, female genitalia; 201, ventral aspect; 202, spermatheca and associated structures, dorsal aspect.