Derovatellus, SHARP, 1882

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 : 430-450

DEROVATELLUS SHARP, 1882

Derovatellus Sharp, 1882b: 282 (type species: Vatellus lentus Wehncke, 1876: 356 , by monotypy); Zimmermann, 1919: 124, 1920: 30; Omer-Cooper 1958: 249; Guignot, 1959: 59; Young, 1954: 50; Biström, 1979: 2; Nilsson et al., 1989: 293; Trémouilles, 1995: 26; Nilsson, 2001: 233.

Varodetellus Biström, 1979: 4 (as subgenus of Derovatellus ) (type species: Derovatellus africanus Régimbart, 1889: 55 , by original designation); Nilsson et al., 1989: 309; Nilsson, 2001: 234, syn. nov.

Mesovatellus Trémouilles, 1995: 26 (Type species: Derovatellus bruchi Zimmermann, 1919: 125 by original designation); Nilsson, 2001: 235, syn. nov.

Diagnosis: Members of this genus are differentiable from the other extant genus of the tribe, Vatellus , in part by the combination of: (1) lateral body outline generally not as strongly discontinuous, pronotum generally broadest at middle or posterior to middle (e.g. Figs 51 , 68 ), (2) metacoxal lines closely approximated (e.g. Figs 52 , 97 ), (3) male lateral lobes and median lobe without strongly extended basal portions (e.g. Figs 57, 59 , 103, 104 ), (4) spermatheca elongate, slen- der, medially bent (e.g. Fig. 60 ), (5) apical margin of ventral apex of orifice of speleum not distinctly lobed.

Distribution: Members of Derovatellus occur in Africa, North and South America and south-east Asia. In Africa the genus occurs throughout the continent. The single disjunct south-east Asian species, D. orientalis Wehncke , occurs in Borneo and Malaysia. In the New World, Derovatellus species occur in southern Florida south throughout the Caribbean Islands and lowland South America to Argentina.

Taxonomic history: This genus was first described by Sharp (1882b) to include the single species D. lentus (Wehncke) , which was previously placed in Hydroporus Clairville. Derovatellus was divided into two subgenera by Biström (1979) based partly on the nature of the female genitalia. Derovatellus (Varodetellus) was defined as those species with a large, spherical receptacle (e.g. Fig. 60 ) whereas Derovatellus s.s. have a small receptacle and large, coiled and twisted intermediate duct ( Fig. 105 ). In addition, members of D. (Varodetellus) are darker in colour dorsally. However, based on evidence presented here (see under ‘Cladistic analysis’ below) recognition of D. (Varodetellus) results in a paraphyletic Derovatellus s.s. ( Fig. 316 ). A large, subspherical receptacle is plesiomorphic ( Fig. 320). Varodetellus is hereby synonymized with Derovatellus .

Mesovatellus Trémouilles was described for the Neotropical species D. bruchi Zimmermann. Within the Neotropical fauna the morphology of this species is unique. However, within the context of Vatellini as a whole, Mesovatellus is nested within Derovatellus ( Fig. 320; see under ‘Cladistic analysis’ below) and is hereby synonymized with that genus.

Natural history: Large numbers of individuals of Neotropical species can often be collected in ponds and slow streams with considerable emergent vegetation.

They also come to black lights, often in very large numbers.

Discussion: New World members of this genus are only differentiable by dissection (either male or female) at this time, though geographical distribution can also be useful in some cases. Characters used previously, such as the degree or type of dorsal punctation, sculpturing of the head or pronotum and extent of the maculae on the elytra are not useful for distinguishing among species.

DEROVATELLUS BRUCHI ZIMMERMANN, 1919

( FIGS 51–69 , 106 )

Derovatellus bruchi Zimmermann, 1919: 125, 1920: 30 ; Blackwelder, 1944: 75; Biström, 1980b: 78.

Mesovatellus bruchi ; Trémouilles, 1995: 26 (comb. nov.).

Type information: Lectotype ♂, in ZSBS, labelled ‘ Argentina [handwritten]/Type [handwritten]/ Samml.A. Zimmermann/ Holotypus Derovatellus bruchi A.Zimm. Staatssamml.München [taxon name and author handwritten, pink label]/ Derovatellus bruchi / Zool. Staatssig. München [blue label].’ Biström KEY TO THE NEW WORLD AND SOUTH- EAST ASIAN SPECIES OF DEROVATELLUS

1. Lateral margin of pronotum strongly curved, lateral outline distinctly discontinuous between pronotum and elytron ( Figs 51 , 62 ); speleum relatively short, apex extending anteriorly slightly beyond anterior margin of sternum V ( Figs 56 , 66 ); south-east Asia and Argentina..........................................................2

1¢. Lateral margin of pronotum slightly to moderately curved, lateral outline slightly to moderately discontinuous between pronotum and elytron (e.g. Figs 68 , 75 , 96 ); speleum extremely long, nearly 2¥ length of ventral length of sternum V and VI together (e.g. Figs 69 , 93 , 101 ); North and South America...............................3

2. Male median lobe in lateral aspect relatively broad ( Fig. 63 ), in dorsal aspect with lateral margins broadly convex, not abruptly expanded subapically ( Fig. 64 ); speleum subapically very strongly narrowed, apex expanded into small bulb; south-east Asia........................................................... D. orientalis Wehncke

2¢. Male median lobe in lateral aspect relatively slender ( Fig. 57 ), in dorsal aspect with lateral margins broadly concave, subapically somewhat expanded ( Fig. 58 ); speleum with apex relatively broadly rounded; southern South America ( Fig. 106 ).................................................................... D. bruchi Zimmermann

3. Apex of male median lobe with two very long, slender, sharply pointed processes which are closely approximated in dorsal aspect and evenly curved ventrad in lateral aspect ( Figs 82, 83, 84 ); female with fertilization duct extremely long, much longer than spermatheca ( Fig. 87 ); receptacle very large and spherical, directed posterad ( Fig. 87 ).......................................................................... D. floridanus Fall

3¢. Apex of male median lobe without two very long, slender processes, or with processes not closely approximated and evenly curved dorsad in lateral aspect; female with fertilization duct much shorter than spermatheca or with receptacle not spherical and/or not directed posterad.......................................................4

4. Male median lobe very robust, broadly expanded medially in lateral aspect, with two long, lateral, slender processes which are evenly curved dorsad in lateral aspect ( Fig. 70 ); speleum with apex not expanded as a dorsally directed bulb ( Fig. 69 ).................................................................. D. spangleri sp. nov.

4¢. Male median lobe very slender or only moderately expanded subapically in lateral aspect, without lateral processes; speleum with apex expanded into a dorsally directed bulb (e.g. Figs 79 , 93 , 101 )............................5

5. Apex of male median lobe in ventral aspect broadly truncate with small medial emargination ( Fig. 90 ), in lateral aspect with apex narrowly rounded ( Fig. 89 ); female with duct between spermatheca and receptacle long, robust and strongly coiled ( Figs 94, 95 )...................................................... D. peruanus Spangler

5¢. Apex of male median lobe in ventral aspect abruptly narrowed to pair of short, very sharp, closely approximated spines, in lateral aspect with apex narrowly to sharply pointed and curved ventrad; female with duct between spermatheca and receptacle either very short ( Fig. 80 ) or more slender and tightly coiled ( Fig. 105 )................6

6. Median lobe in lateral aspect broadly expanded subapically ( Fig. 76 ); female with duct between spermatheca and receptacle very short, receptacle very large, directed anterad, ‘triangular’ spermathecal process actually apically rounded with lateral angulate lobes ( Fig. 80 )........................................ D. roosevelti sp. nov.

6¢. Median lobe in lateral aspect slender throughout ( Fig. 103 ); female with duct between spermatheca and receptacle extremely long, strongly coiled, receptacle small, inconspicuous, spermathecal process triangular, apex broad with margin linear ( Fig. 105 )........................................................... D. lentus (Wehncke)

(1980b) designated the lectotype, figured the female genitalia (from a paralectotype) and commented on the unusual morphology of the species.

Type locality: Argentina, Buenos Aires.

Diagnosis: This species is larger than any other New World Derovatellus (TL = 4.9–5.0 mm). The lateral margins of the pronotum are very strongly curved making the lateral outline strongly discontinuous between the pronotum and elytron in dorsal aspect ( Fig. 51 ). The male median lobe in lateral aspect is slender with the apex narrow and narrowly rounded ( Fig. 57 ). In ventral aspect the lateral margins are evenly concave, and the lobe is distinctly expanded subapically ( Fig. 58 ). The male lateral lobe is very slender in the apical half ( Fig. 59 ). The speleum is relatively short extending slightly beyond the anterior margin of the abdominal sternum V ( Fig. 56 ).

Description:

Habitus. Body outline strongly discontinuous in dorsal aspect, lateral elytral margins rounded ( Fig. 51 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head red-brown, darker brown along posterior margin, some specimens with a medial pale area; pronotum yellow, with brown medially along posterior and anterior margins; elytron dark brown with transverse, irregular pale macula apicomedially ( Fig. 51 ); venter red-brown; appendages red-brown.

Sculpture and structure. Head with punctation relatively fine, surfaces very finely microreticulate and shiny between punctures; anterior clypeal margin rounded; eyes moderately large, not protuberant ( Fig. 51 ). Pronotum finely, sparsely and evenly punctate, surface between punctures alutaceous, setae relatively long and fine; pronotum broadest slightly anterior of middle, lateral margins broadly curved, more strongly so anteriorly ( Fig. 51 ); lateral bead distinct anteriorly, obscured posteriorly; transverse sulcus absent. Elytron with fine, moderately dense punctures obscured somewhat by alutaceous surface texture, setae fine, pale and relatively long. Prosternum short, transversely carinate; prosternal process strongly tectiform medially, lateral margins rounded to narrowly rounded apex. Metasternum with punctation coarse along metacoxal margins, medially finely, shallowly punctate, surface between punctures microreticulate; anteromedial margin unmodified. Abdominal sterna finely punctate, surface alutaceous; sixth sternum apically broadly rounded, with marginal fringe of fine setae; speleum moderately long, distinctly constricted medially, apex broad, broadly rounded, basal half with surface scabrous ( Fig. 56 ). Metacoxa with lateral portion with punctures large and dense laterally, medially finer, surface between punctures alutaceous; metacoxal lines very closely approximated, only slightly divergent anteriorly ( Fig. 52 ). Metatrochanter moderately rounded anteriorly, somewhat offset ( Fig. 53 ); metatarsal claws minutely serrate.

Male genitalia. Median lobe in lateral aspect curved basally to about half of length, thereafter relatively straight, expanded along ventral margin about threequarters distance from base, apex very slender to very narrowly rounded apex, slightly curved ventrally ( Fig. 57 ); in ventral aspect moderately broad, broadly expanded subapically, apex acutely rounded, with elongate, longitudinal, area of thin chitin from apex medially along lobe ( Fig. 58 ). Lateral lobe very slender except medially broad, apex slightly expanded and irregularly curved, bearing numerous pores and short setae ( Fig. 59 ).

Female genitalia. Gonocoxosternite moderately broad with apex acutely rounded, medial margin relatively straight, anterior lobe long, broader basally, somewhat narrowed apically, apex relatively narrowly truncate ( Fig. 60 ). Gonocoxa subtriangular, apex acutely and sharply rounded, apodeme elongate, slender, slightly curved laterally ( Fig. 60 ). Bursa copulatrix small, inconspicuous; spermathecal duct moderately long, slender; receptacle elongate, rounded, large; intermediate duct moderately long, moderately broad, tightly twisted; spermatheca elongate, medially curved, robust, triangular process relatively short, broad; fertilization duct very long, distinctly twisted and looped ( Figs 60, 61 ).

Sexually dimorphic characters. Male with moderately well-developed setae on mesotrochanter and medial portion of mesofemur, setae long and fine. Male pro- and mesotarsomeres ( Fig. 54 ) very broadly expanded laterally compared with female ( Fig. 55 ).

Distribution: This species is known from Argentina ( Fig. 106 ).

Phylogenetic relationships: Derovatellus bruchi is sister to a clade containing African species of Derovatellus and the only Oriental species of Vatellini , D. orientalis . This relationship is based mainly on the relatively rounded lateral pronotal margin broadest medially (Character 6).

Discussion: This species differs considerably from the other New World Derovatellus in several ways including coloration, habitus ( Fig. 51 ), the shape of the speleum ( Fig. 56 ) and a few other minor features. No doubt this is the reason this taxon was given its own genus by Trémouilles (1995). As the other Neotropical species are together monophyletic relative to D. bruchi ( Fig. 316 ), placing it in its own genus was not unreasonable. However, given a broader perspective and examination of species of Derovatellus from elsewhere in the world, D. bruchi is nested within the genus Derovatellus ( Fig. 316 ) and should not be placed alone in a separate genus.

Material examined: Argentina: Buenos Aires, November 1922 (1, MNHN) ; Buenos Aires, 20 December 1904, C Bruch (1, MNHN) ; Prov . Buenos Aires, Ing. R. Otamendi, 6 January 1988, Archangelsky (2, LHIC) .

DEROVATELLUS ORIENTALIS WEHNCKE, 1883

( FIGS 62–66 )

Derovatellus orientalis Wehncke, 1883: 149 ; Régimbart, 1899: 193; Zimmermann, 1920: 30; Biström, 1980b: 78 (lectotype designation).

Type information: Lectotype ♂, in BMNH, designated by Biström (1980b) (not examined). There is a single paralectotype ( Biström, 1980b).

Type locality: Borneo.

Diagnosis: This species has distinctive male genitalia with a median lobe that is relatively robust in both dorsal and lateral aspects and with the apex narrowly pointed ( Figs 63, 64 ). The speleum is relatively short, extending slightly beyond the anterior margin of abdominal sternum V ( Fig. 66 ). It is parallel-sided to near the apex where it is very abruptly constricted and slender to an apex that is slightly expanded and rounded ( Fig. 66 ). The lateral margins of the pronotum are very strongly curved making the lateral outline strongly discontinuous between the pronotum and elytron in dorsal aspect ( Fig. 62 ). This is the only species known from south-east Asia.

Description:

Habitus. Body outline moderately discontinuous in dorsal aspect, lateral elytral margins moderately rounded ( Fig. 62 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow; pronotum yellow with two brown maculae at posterior margin, one on each side of midline; elytron light brown ( Fig. 62 ); venter yellow, dark red-yellow on metacoxa; appendages yellow.

Sculpture and structure. Head finely punctate, punctures sparse, even more so on clypeus, surface between punctures finely microreticulate, shiny; anterior clypeal margin rounded; eyes large, moderately protuberant ( Fig. 62 ). Pronotum punctate, punctures moderately dense and large, finer and less dense laterally, surface smooth and shiny between punctures, with very fine microreticulation between punctures in some areas, setae very fine, moderately long; pronotum with lateral margins broadly curved, pronotum widest medially ( Fig. 62 ); lateral bead narrow, distinct along entire margin, laterally slightly dentate; posterior angles obtusely rounded; transverse sulcus absent. Elytron with punctures similar to medial surface of pronotum, surface microreticulate between punctures, somewhat alutaceous. Prosternum very short, transversely carinate; prosternal process with lateral margins rounded, laterally moderately beaded, apex rounded, medially broadly tectiform. Metasternum impunctate except medially with fine, finely setose punctures, surface microreticulate, with mesh of small cells over most of surface, shiny; anteromedial margin broadly and flatly beaded, medially slightly produced ventrally. Abdominal sterna very finely punctate laterally, visible sternum I with coarse punctures along anterior and posterior margins, surfaces of sterna shiny; sixth sternum with apex rounded, apex of opening of speleum rounded, without conspicuous setae. Speleum long, with lateral margins parallel, apically narrowed to slender, short duct, apex a small, rounded bulb ( Fig. 66 ). Metacoxa with lateral portion densely and coarsely punctate over most of surface, microreticulate and shiny between punctures, microreticulation consisting of mesh of small cells; metacoxal lines moderately approximated posteriorly, anteriorly slightly divergent; medial portion finely punctate, shiny. Metatrochanter broad, apically slightly obliquely truncate, somewhat offset; metatarsal claws finely serrate.

Male genitalia. Median lobe in lateral aspect relatively slender, evenly curved, apex pointed and very finely curved ventrad ( Fig. 63 ); in ventral aspect moderately broad, lateral margins evenly and broadly curved, apex narrowly rounded ( Fig. 64 ). Lateral lobe broad basally, apical portion evenly narrowed, sinuate, apically narrowly rounded, ventral margin with many setae ( Fig. 65 ).

Female genitalia. Gonocoxosternite elongate, apex acutely rounded, medial margin broad, slightly concave, anterior lobe long, slender ( Fig. 67 ). Gonocoxa elongate, subtriangular, apex narrowly rounded, apodeme very long, slender, straight ( Fig. 67 ). Bursa copulatrix small, inconspicuous; spermathecal duct slender, very long, strongly coiled near receptacle; receptacle moderately large, elongate, curved; intermediate duct relatively slender, long, coiled; spermatheca elongate, medially curved, triangular process relatively small; fertilization duct short, curved, slen- der ( Fig. 67 ).

Sexually dimorphic characters. Male pro- and mesobasotarsomeres moderately expanded compared with female.

Distribution: This species is restricted to Borneo and Malaysia.

Phylogenetic relationships: This species is sister to a clade containing members of Derovatellus from Africa ( Fig. 316 ). These African species were part of the nominal subgenus as defined by Biström (1979). The only close affinities of D. orientalis with the Neotropical fauna are with the unusual D. bruchi , which is more basal in the clade containing D. orientalis .

Discussion: The specimens examined and illustrated here differ slightly from published illustrations of D. orientalis ( Biström, 1980b) . The male median lobe differs from the published illustrations of the genitalia of the type by being somewhat broader apically in both ventral and lateral aspects. However, the lateral lobe is similarly sinuate in each species (fig. 3c in Biström, 1980b; Fig. 63 ). In addition, the female genitalia are extremely similar (fig. 3d in Biström, 1980b; Fig. 67 ). Specimens of Derovatellus from south-east Asia are very rare. A more thorough survey of the Derovatellus fauna of this region will undoubtedly improve knowledge of the species limits and variation in these characters.

Material examined: Malaysia: Lake Chini , 31 December 1995, Mazzoldi (4, PMIC) .

DEROVATELLUS SPANGLERI MILLER SP. NOV.

( FIGS 68–74 , 106 )

Type information: Holotype ♂ in USNM labelled, ‘ BRAZIL, M.G. Jacare, P.N.Xingu XI-1965, at lite M. Alvarenga / HOLOTYPE Derovatellus spangleri Miller, 2004 [red label with double black line border].

Paratypes: Argentina: BA Zelaya , 18 February 1968, OS Flint (1, USNM) . Brazil: Para, Rio Xingu Camp 52∞22¢W 3∞39¢S, Altamira c. 60 km S, 15 October 1986, P Spangler, O Flint (2, USNM) . Colombia: Amaz. Leticia , 12–15 March 1969, P and P Spangler (2, USNM) . Peru: Loreta, Yacumama Lodge, nr jct Rio Maranon , Rio Uceyali , 73.6∞ W 4.8 ∞S, 6–20 August 1994, small light in woods, LHIC (16, LHIC) ; Loreto San Antonio , August 1965, light trap, JC Hitchcock (1, USNM) ; Loreto, Yacumama Lodge 73.6∞ W 4.8 ∞S, Maronon, Rio Ucayall , 20 August 1994, small light in woods, Skelley (26, LHIC) ; Madre de Dios; Rio Tambopata Res., 30 air km SW Pto Maldonado , 26–20 November 1979, subtropical moist forest, JP Heppner (3, USNM) ; San Antonio Loreto, August 1965, light trap, JC Hitchcock (1, USNM) .

Type locality: Brazil, Mato-Grosso, Jacare, Parc National Xingu.

Diagnosis: In this species the male median lobe is very robust and bears a pair of long dorsolateral spines that are slightly curved dorsad in lateral aspect ( Fig. 70 ). The speleum is extremely long, extending nearly to the metathorax. The apex is narrowed, not bulbous and not curved dorsad ( Fig. 69 ). In the female, the receptacle is relatively large and prominent with a long, thick, twisted intermediate duct between the spermatheca and receptacle ( Figs 73, 74 ).

Description:

Habitus. Body outline continuous in dorsal aspect, lateral elytral margins moderately rounded ( Fig. 68 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow; pronotum yellow with narrow line of dark brown along posterior margin; elytron dark brown with indistinct yellow, transverse macula apicomedially on elytron ( Fig. 68 ); venter yellow, dark red-yellow on metacoxa; appendages yellow.

Sculpture and structure. Head finely punctate, punctures sparse, even more so on clypeus, surface between punctures finely microreticulate, shiny; anterior clypeal margin rounded; eyes large, not protuberant ( Fig. 68 ). Pronotum finely punctate, punctures moderately dense, finer and less dense laterally, surface smooth and shiny between punctures, with very fine microreticulation between punctures in some areas, setae very fine, moderately long; pronotum with lateral margins broadly curved, pronotum widest slightly posterad of middle ( Fig. 68 ); lateral bead narrow, distinct along entire margin; posterior angles obtusely rounded; transverse sulcus absent. Elytron with punctures moderately large, fairly dense, setae elongate and fine, surface very lightly microreticulate between punctures. Prosternum very short, transversely carinate; prosternal process with lateral margins rounded, broadly beaded, apex rounded, medially broadly tectiform. Metasternum impunctate except medially with fine, finely setose punctures, surface transversely microreticulate, shiny; anteromedial margin broadly and flatly beaded, medially narrowly carinate posterad of bead. Abdominal sterna moderately punctate, surface shiny; sixth sternum with apex truncate, with apical marginal fringe of fine setae. Speleum exceptionally long, broad, lateral margins approximately parallel for most of length, apex relatively abruptly narrowed to narrowly rounded apex, not dorsally curved ( Fig. 69 ). Metacoxa with lateral portion densely punctate medially and anteriorly, surface microreticulate and shiny between punctures; with metacoxal lines narrowly approximated posteriorly, anteriorly slightly divergent; medial portion finely punctate, shiny. Metatrochanter elongate, apically narrowly rounded, not strongly offset; metatarsal claws minutely serrate.

Male genitalia. Median lobe in lateral aspect robust, broad, apically with long, slender spine, apex of spine slightly bent dorsad, surface of spine bearing numerous punctures ( Fig. 70 ); in dorsal aspect robust, broad, with two long, slender apicolateral spines, apices of spines slightly expanded, medial portion apically narrowly rounded ( Fig. 71 ). Lateral lobe broad basally, apically narrowed, apex broadened, flattened, apex pointed ( Fig. 72 ).

Female genitalia. Gonocoxosternite moderately elongate, apex rounded, medial margin convex, anterior lobe long, relatively broad, apex narrowly rounded ( Fig. 73 ). Gonocoxa elongate-triangular, apically narrowed, apex narrowly rounded, apodeme approximately straight, narrow, elongate ( Fig. 73 ). Bursa copulatrix small, inconspicuous; spermathecal duct moderately long, very slender, prominently expanded and tightly twisted near insertion of duct into receptacle; receptacle relatively large, subspherical, intermediate duct broad, broadly twisted; spermatheca very long, slender, strongly curved, slightly constricted in two places medially, triangular process moderately expanded apically; fertilization duct relatively long ( Figs 73, 74 ).

Sexually dimorphic characters. Male with dense row of fine, long setae along mesotrochanter and mesofemur. Male pro- and mesobasotarsomeres moderately expanded compared with female.

Intraspecific variation. The extent of the elytral macula varies from nearly invisible or only visible laterally to clearly visible, conspicuously yellow and extending across the entire width of the elytron.

Natural history: This species has been collected at light and in ‘subtropical moist forest’.

Distribution: Derovatellus spangleri is widespread in South America and is known from Argentina, Brazil, Colombia and Peru ( Fig. 106 ). It is apparently relatively rarely collected.

Phylogenetic relationships: This species is sister to the remaining Neotropical species of Derovatellus excluding D. bruchi ( Fig. 316 ). Other Neotropical species have the speleum exceptionally long and with the apex bulbous and upturned whereas that of D. spangleri is shorter and lacks the apical bulb ( Fig. 69 ).

Etymology: This species is named in honour of Paul J. Spangler for his contribution to our knowledge of this genus and for his valuable research efforts in Dytiscidae in general.

DEROVATELLUS ROOSEVELTI MILLER SP. NOV.

( FIGS 75–80 , 106 )

Type information: Holotype ♂ in USNM labelled, ‘ ECUADOR, Los Rios, Bababoyo 21 June 1975 at blacklite Cohen, Langley , Monnig / HOLOTYPE Derovatellus roosevelti Miller 2004 [red label with double black line border].

Paratypes: 12 in USNM labelled same as holotype. Ecuador: Esmer., La Union , 3 February 1979, at black light, Cohen, Langley, Monnig (4, USNM) . Venezuela: T.F. Amaz. Cerro de la Neblina 1 km S basecamp 0∞50¢N 66∞10¢W, 140 m, 11 February 1985, along small whitewater stream, pools of dead leaves, PJ and PM Spangler, R Faitoute, W Steiner (1, USNM) .

Type locality: Ecuador, Los Rios, Bababoyo.

Diagnosis: This species is similar to D. lentus in the shape of the median lobe except it is more robust in both ventral and lateral aspect ( Figs 76, 77 ). In lateral aspect the apex of the lobe is gradually broadened subapically ( Fig. 76 ). Apically it is abruptly narrowed and terminates in a small slender, short, apical spine ( Fig. 76 ). In dorsal aspect the male median lobe is broad subapically and apically terminates in a pair of slender, sharp, closely approximated spines ( Fig. 77 ). The female receptacle is large with the spermathecal duct inserted on its ventral side ( Fig. 80 ). There is a small lobe on the spermathecal duct near this insertion ( Fig. 80 ). The intermediate duct is very short, the spermatheca is long and slender and the ‘triangular process’ is apically rounded and not triangular ( Fig. 80 ).

Description:

Habitus. Body outline somewhat discontinuous in dorsal aspect, lateral elytral margins moderately rounded ( Fig. 75 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head red-brown, darker brown along posterior margin; pronotum yellow, with brown medially along posterior and anterior margins; elytron dark brown with transverse, irregular pale macula apicomedially ( Fig. 75 ); venter red-brown; appendages red-brown.

Sculpture and structure. Head with punctation relatively fine, surfaces very finely microreticulate and shiny between punctures; anterior clypeal margin rounded; eyes moderately large, not protuberant ( Fig. 75 ). Pronotum finely, sparsely and evenly punctate, surface between punctures lightly alutaceous, setae relatively long and fine; pronotum broadest slightly anterior of middle, lateral margins broadly curved, more strongly so anteriorly ( Fig. 75 ); lateral bead distinct anteriorly, obscured posteriorly; transverse sulcus absent. Elytron with fine, moderately dense punctures obscured somewhat by alutaceous surface texture, setae fine, pale and relatively long. Prosternum short, transversely carinate; prosternal process strongly tectiform medially, lateral margins rounded to narrowly rounded apex. Metasternum with punctation coarse along metacoxal margins, medially finely, shallowly punctate, surface between punctures microreticulate; anteromedial margin unmodified. Abdominal sterna finely punctate, surface alutaceous; sixth sternum apically broadly rounded, with marginal fringe of fine setae; speleum very long, apically with prominent dorsally directed bulb ( Fig. 79 ). Metacoxa with lateral portion with punctures large and dense laterally, medially finer, surface between punctures alutaceous; metacoxal lines very closely approximated, only slightly divergent anteriorly. Metatrochanter moderately rounded anteriorly, slightly offset; metatarsal claws minutely serrate.

Male genitalia. Median lobe in lateral aspect relatively broad, evenly curved, ventral surface distinctly expanded and subapically rounded, apex a stout, ventrally curved spine, apex acutely pointed ( Fig. 76 ); in ventral aspect moderately broad, apically broadly rounded with two short, stout, distinct spines narrowly pointed and closely approximate ( Fig. 77 ). Lateral lobe stout, broadly subtriangular, apex flattened and expanded, with few sparse and short setae subapically and numerous punctures and very short setae along dorsal margin ( Fig. 78 ).

Female genitalia. Gonocoxosternite moderately elongate, apex somewhat acutely rounded, medial margin convex, anterior lobe very long, slightly curved, apex broadly rounded ( Fig. 80 ). Gonocoxa triangular, apically narrowed, apex very narrowly rounded to pointed, apodeme curved laterally, narrow, elongate ( Fig. 80 ). Bursa copulatrix small, rounded; base of spermathecal duct robust, narrowed, spermathecal duct very long, very slender, with distinct, small, posteriorly directed lobe near insertion of duct into receptacle; receptacle large, spherical, directed anteriorly, spermathecal duct insertion near base on dorsal side, receptacle broadly attached directly to base of spermatheca; spermatheca very long, slender, strongly curved, triangular process broadest basally where process is expanded into lateral points, apex broadly rounded; fertilization duct relatively long, broad medially ( Fig. 80 ).

Sexually dimorphic characters. Male with moderately dense row of fine setae along mesotrochanter and mesofemur. Male pro- and mesobasotarsomeres moderately expanded compared with female.

Natural history: This species has been collected at black lights and from a ‘small whitewater stream’ in ‘pools of dead leaves’.

Distribution: Derovatellus roosevelti is known from Ecuador and Venezuela ( Fig. 106 ).

Phylogenetic relationships: This species belongs to a clade containing Neotropical members of Derovatellus excluding D. bruchi and D. spangleri ( Fig. 316 ). Within this group it occupies an unresolved position relative to the other members ( Fig. 316 ).

Etymology: This species is named for Theodore Roosevelt: Rough Rider, Nobel Peace Prize winner, conservationist, progressive conservative, hunter, trust buster, historian, cowboy, biologist, and one of the greatest presidents of the United States of America.

DEROVATELLUS FLORIDANUS FALL, 1932

( FIGS 81–87 , 107 )

Derovatellus floridanus Fall, 1932: 146 .

Derovatellus lentus floridanus ; Young, 1954: 51 (stat. nov.); Spangler, 1966b: 14; Biström, 1980b: 78; Epler, 1996:3.40; Nilsson, 2001: 234; Roughley & Larson, 2001: 169.

Derovatellus ibarri Spangler, 1966b: 11 , incorrect original spelling, unavailable.

Derovatellus ibarrai Spangler, 1966b: 11 (including description of larva); Trémouilles, 1995: 26; Nilsson, 2001: 234, syn. nov..

Type information: Derovatellus floridanus : holotype ♂ in MCZC labelled, ‘Royal Palm Park. Fla. VCSB 1-7-30 [date handwritten]/ Derovatellus TYPE floridanus [handwritten, ‘TYPE’ underlined with red line]/M.C.Z Type 23884 [red label, number handwritten]/H. C. FALL COLLECTION.

Derovatellus ibarrai : holotype ♂ in USNM (not examined).

The holotype of D. floridanus was examined and found to be the same as D. ibarrai based on Spangler’s (1966b) illustration. Although the holotype of D. ibarrai was not examined, the male genitalia are so distinct that there can be little doubt as to the identity of this taxon. The taxon was regarded as a subspecies of D. lentus by Young (1954), and this has been followed until now. Previous interpretations of D. floridanus emphasized external characters which are inadequate for reliably defining these species.

Spangler (1966b) introduced two alternative original spellings for a new species, D. ibarri and D. ibarrai . The spelling D. ibarrai was used subsequently by Trémouilles (1995) which, under Article 24.2.3 of the Code, constitutes a fixation of this spelling which is appropriate given that Spangler (1966b) made it clear that the species was named in honour of J. Ibarra. Therefore, the name D. ibarri is an incorrect original spelling and is unavailable.

Type locality: Derovatellus ibarrai : Guatemala, Izabal, 1 mi N Morales, km 239 Atlantic Hwy.

Derovatellus floridanus : USA, Florida, Royal Palm Park.

Diagnosis: This species is distinguishable by the male median lobe of the aedeagus, which is robust with two extremely long spines apically ( Figs 82, 83 ). The spines are very slender, closely approximated and curved ventrally at the apex ( Fig. 82 ). The female receptacle is very large and spherical ( Fig. 87 ). The speleum is extremely long extending nearly to the metathorax ( Fig. 86 ). Its apex is bulbous and curved dorsad ( Fig. 86 ).

Description:

Habitus. Body outline continuous in dorsal aspect, lateral elytral margins moderately rounded ( Fig. 81 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow; pronotum yellow with narrow line of dark brown along posterior margin; elytron dark brown with indistinct yellow, transverse macula apicomedially on elytron ( Fig. 81 ); venter yellow, dark red-yellow on metacoxa; appendages yellow.

Sculpture and structure. Head finely punctate, punctures sparse, even more so on clypeus, surface between punctures finely microreticulate, shiny; anterior clypeal margin rounded; eyes large, not protuberant ( Fig. 81 ). Pronotum finely punctate, punctures moderately dense, finer and less dense laterally, surface smooth and shiny between punctures, with very fine microreticulation between punctures in some areas, setae very fine, moderately long; pronotum with lateral margins broadly curved, pronotum widest medially ( Fig. 81 ); lateral bead narrow, distinct along entire margin; posterior angles obtusely rounded; transverse sulcus absent. Elytron with punctures moderately large, fairly dense, obscured and shallow in areas, setae elongate and fine, surface lightly alutaceous between punctures, especially apically. Prosternum very short, transversely carinate; prosternal process with lateral margins rounded, broadly beaded, apex rounded, medially broadly tectiform. Metasternum impunctate except medially with fine, finely setose punctures, surface transversely microreticulate, shiny; anteromedial margin broadly and flatly beaded, medially narrowly carinate posterad of bead. Abdominal sterna very finely punctate laterally, surface shiny; sixth sternum with apex truncate to broadly emarginate, with apical marginal fringe of fine setae. Speleum exceptionally long, slightly narrowed apically, apex bulbous, rounded, curved dorsally ( Fig. 86 ). Metacoxa with lateral portion densely punctate medially and anteriorly, surface microreticulate and shiny between punctures; with metacoxal lines moderately approximated posteriorly, anteriorly slightly divergent; medial portion finely punctate, shiny. Metatrochanters elongate, apically narrowly rounded, not strongly offset; metatarsal claws minutely serrate.

Male genitalia. Median lobe in lateral aspect broad, becoming slightly broader subapically, apical portion developed into long, slender, curved spine, spine with numerous punctures ( Figs 82, 84 ); in ventral aspect broad, lateral margins rounded, apex bifid, developed into two long, slender, straight spines ( Fig. 83 ). Lateral lobe moderately broad, elongate triangular, apex developed into broad, flattened, leaf-shaped structure, structure with few short setae, apex and dorsal margin with numerous punctures and short, spinous setae ( Fig. 85 ).

Female genitalia. Gonocoxosternite elongate, subtriangular, apex acutely rounded, anterior lobe long, about as long as main portion of gonocoxosternite ( Fig. 87 ). Gonocoxa elongate, subtriangular, apex very narrowly rounded, apodeme long, slightly sinuate ( Fig. 87 ). Bursa copulatrix inconspicuous, small; spermathecal duct slender, moderately long; receptacle large, spherical; intermediate duct relatively broad, long, tightly coiled; spermatheca elongate, medially curved, narrow, triangular process broad; fertilization duct long, evenly curved ( Fig. 87 ).

Sexually dimorphic characters. Male with dense row of fine, long setae along mesotrochanter and mesofemur. Male pro- and mesobasotarsomeres moderately expanded compared with female.

Intraspecific variation. The extent of the elytral macula varies from nearly invisible or only visible laterally to clearly visible, conspicuously yellow and extending across the entire width of the elytron.

Natural history: This species has been collected from black light traps and a shaded pool. In Guatemala the species was collected from small, water-filled depressions and hoof-prints in dense grass near a pond ( Spangler, 1966b), but was very rare along the margins of the pond. In Florida the species has been taken at lights and in a ‘leaf-choked pool in a hammock’ ( Young, 1954).

Distribution: Derovatellus floridanus is widespread around the Caribbean. It is known from Belize, Brazil,

Cuba, Guatemala, Trinidad and Florida in the United States ( Fig. 107 ).

Phylogenetic relationships: This species belongs to a clade containing Neotropical members of Derovatellus excluding D. bruchi and D. spangleri . Within this group it occupies an unresolved position relative to the other members ( Fig. 316 ).

Material examined: Belize: Cayo Dist. E Roaring Cr , 28 April 1984, HL Dozier (2, USNM) ; Stann Cr., Sittee Point, blacklight, on riverbank at Possum Point Biological Station , 23 April 1987, Spangler, Faitoute (2, USNM) . Brazil: Para, Rio Xingu Camp 52∞22¢W 3∞39¢S Altamira c. 60 km S, 8 October 1986, P Spangler, O Flint (1, USNM) . Cuba: Matanzas Prov., Cienaga Zapata, at Playa Larga , 10–11 February 1981, D Davis, P Spangler (3, USNM) ; Matanzas, Buenaventura, 7 km NW, Zapata Swamp , 1 May 1983, PJ Spangler, I Fernandez (24, USNM) ; Matanzas, Palpite 1 km NE Zapata Swamp, 2 May 1983, PJ Spangler (56, USNM) ; Matanzas, Palpite 2 km NE Zapata Swamp, 1 May 1983, PJ Spangler, I Fernandez, WN Mathis (5, USNM) . Guatemala: 1 mi N Morales , 16–18 August 1965, PJ Spangler (27, USNM) ; Peten Tikal , 28 June 1974, WE Steiner (7, USNM) . Trinidad: Cedros , 4 May 1929, Darlington (1, MCZC) ; Cedros , 4 May 1929, Darlington (4, MCZC) ; Cumuto , 1929, Darlington (7, MCZC) ; Cumuto , 1929, Darlington (14, MCZC) . United States: Florida: Collier County, Rt. 41 5 mi E Collier S. S. P., 21 March 2000, mercury vapour light (1, KBMC) ; Collier Seminole St Pk., 4 April 1964, RH Arnett, ER Van Tassell (1, FSCA) ; Highlands Co., Highland Hammock St. Pk. , 22 March 2000, A Kawahara (1, KBMC) ; Dade Co., Canal at Pinecrest , 29 December 1982, FN Young (1, FSCA) ; Dade Co., Pinecrest pool, shaded, 27 December 1980, FN Young (3, FSCA) ; Monroe Co. Big Pine Key, Long Beach , 17 August 1992, UV Light, RE Roughley (20, JBWM) ; Palm Beach Co., W Palm Beach , 23 July 1959, BLT, ML Messec (3, FSCA) .

DEROVATELLUS PERUANUS SPANGLER, 1967

( FIGS 88–94 , 106 )

Derovatellus peruanus Spangler, 1967: 142 ; Trémouilles, 1995: 26.

Type information: Holotype ♂ in USNM (not examined).

Type locality: Peru, Loreto, San Antonio.

Diagnosis: In this species, the male median lobe of the aedeagus is slender and apically truncate, in ventral aspect the apex consists of two truncate lobes with an intermediate emargination ( Fig. 90 ). The receptacle is very small and represented by a small lobe, the intermediate duct is very long and coiled and the spermatheca is dramatically and irregularly swollen at its anterior end ( Figs 94, 95 ). The speleum is exceptionally long with the apex bulbous and curved dorsad ( Fig. 93 ).

Description:

Habitus. Body outline continuous in dorsal aspect, lateral elytral margins moderately rounded ( Fig. 88 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow; pronotum yellow with narrow line of dark brown along posterior margin; elytron dark brown with indistinct yellow, transverse macula apicomedially on elytron ( Fig. 88 ); venter yellow, dark red-yellow on metacoxa; appendages yellow.

Sculpture and structure. Head finely punctate, punctures sparse, even more so on clypeus, surface between punctures finely microreticulate, shiny; anterior clypeal margin rounded; eyes large, not protuberant ( Fig. 88 ). Pronotum finely punctate, punctures moderately dense, finer and less dense laterally, surface smooth and shiny between punctures, with very fine microreticulation between punctures in some areas, setae very fine, moderately long; pronotum with lateral margins broadly curved, pronotum widest medially ( Fig. 88 ); lateral bead narrow, distinct along entire margin; posterior angles obtusely rounded; transverse sulcus absent. Elytron with punctures moderately large, fairly dense, setae elongate and fine, surface very lightly microreticulate between punctures, somewhat alutaceous in areas, especially apically. Prosternum very short, transversely carinate; prosternal process with lateral margins rounded, broadly beaded, apex rounded, medially broadly tectiform. Metasternum impunctate except medially with fine, finely setose punctures, surface transversely microreticulate, shiny; anteromedial margin broadly and flatly beaded, medially narrowly carinate posterad of bead. Abdominal sterna very finely punctate laterally, surface shiny; sixth sternum with apex truncate, with apical marginal fringe of fine setae. Speleum exceptionally long, narrow, with lateral margins approximately parallel, apex bulbous, rounded, curved dorsally. Metacoxa with lateral portion densely punctate medially and anteriorly, surface microreticulate and shiny between punctures; with metacoxal lines moderately approximated posteriorly, anteriorly slightly divergent; medial portion finely punctate, shiny. Metatrochanters elongate, apically narrowly rounded, not strongly offset; metatarsal claws minutely serrate.

Male genitalia. Median lobe in lateral aspect moderately broad, evenly curved, slightly expanded subapically on ventral margin, apex narrowly rounded, slightly dorsally curved, with numerous small punctures around apex ( Figs 89, 91 ); in ventral aspect narrow basally becoming expanded medially, apical half broader, apex broad, truncate, ending in two apical processes, each short, broad and truncate, medially between processes with longitudinal region of very thin chitin ( Fig. 90 ). Lateral lobe broad basally and medially, apical portion narrowed, apically expanded and curved dorsally, apically with few spines, several punctures apically and along dorsal margin ( Fig. 92 ).

Female genitalia. Gonocoxosternite elongate, apex acutely rounded, medial margin broadly curved, anterior lobe moderately long, robust, broad ( Fig. 94 ). Gonocoxa elongate, subtriangular, apex narrowly rounded, apodeme moderately long, slender, straight ( Fig. 94 ). Bursa copulatrix small, inconspicuous; spermathecal duct slender, only moderately long, broadly expanded near receptacle; receptacle very small, consisting of a small lobe on duct; intermediate duct relatively broad, strongly coiled and twisted; spermatheca very elongate, medially curved, very broad and irregularly expanded at anterior end, triangular process long and broadly expanded; fertilization duct moderately long, slender ( Figs 94, 95 ).

Sexually dimorphic characters. Male with dense row of fine, long setae along mesotrochanter and mesofemur. Male pro- and mesobasotarsomeres moderately expanded compared with female.

Intraspecific variation. The extent of the elytral macula varies from nearly invisible or only visible laterally to clearly visible, conspicuously yellow and extending across the entire width of the elytron.

Natural history: This species has been collected at black lights and open grassland and marsh and subtropical forest habitats.

Distribution: This uncommonly collected species is known only from Bolivia and Peru ( Fig. 106 ).

Phylogenetic relationships: This species belongs to the clade of Neotropical Derovatellus where it is sister species to D. lentus ( Fig. 316 ). This is based on the very long, coiled intermediate duct between the receptacle and spermatheca in females (Character 34). This feature is shared with a variety of African species, as well, but the two states appear to be homoplasious ( Fig. 316 ).

Material examined: Bolivia: Beni, 40 km E San Borja, Estacion Biologica Beni, Estancia El Porvenir , 6–8 September 1987, blacklight, open grass savanna and marsh, WE Steiner (4, USNM) ; Santa Cruz, Ichilo Province, Buena Vista , 3 October 1994, 400 m, mercury vapour light, R Ward (1, KBMC) . Peru: Colonia Galleria 15 km E Ecayali, 10–25 September 1981, Borys, Malkin (3, FSCA) ; Loreto 1 km SW Boca del Rio Samiria Vigilante Post no. 1, 04∞40.5¢S 74∞18.9¢W, 15 August 1991, blacklight trap, restringa forest Tr. Norte lot 15, MG Pogue (2, USNM) ; Loreto, San Antonio, August 1965, JC Hitchcock (5, USNM) ; Madre de Dios, Rio Tambopata Res, 30 air km SW Pto Maldonado , 729 m, 16–20 November 1979, subtropical moist forest, JB Heppner (3, USNM) .

DEROVATELLUS LENTUS ( WEHNCKE, 1876)

( FIGS 4 , 22 , 23 , 28–30 , 40, 41, 45–47 , 97–105 , 108)

Vatellus lentus Wehncke, 1876: 356 .

Derovatellus lentus ; Sharp, 1882b: 286 (comb. nov.); Régimbart, 1904: 224; Zimmermann, 1920: 30, Blackwelder, 1944: 75; Spangler, 1966b: 14; Biström, 1980b: 78 (designation of lectotype); Trémouilles, 1995: 26.

Type information: Lectotype ♂, in MNHU, designated by Biström (1980b) (not examined).

Type locality: Puerto Rico.

Diagnosis: This species is identifiable by the median lobe which is relatively slender with the apex abruptly narrowed, slender and distinctly ventrally curved in lateral aspect ( Fig. 103 ). The female receptacle is very small, the intermediate duct is extremely long and irregularly coiled ( Fig. 105 ). The spermathecal duct is exceptionally long and slender ( Fig. 105 ). The speleum is very long, extending nearly to the mesothorax ( Figs 28–30 , 101 ). The apex is bulbous and prominently curved dorsad ( Figs 28–30 , 101 ).

Description:

Habitus. Body outline continuous in dorsal aspect, lateral elytral margins moderately rounded ( Fig. 96 ), not dorsoventrally compressed. Measurements are given in Table 1.

Coloration. Head yellow; pronotum yellow with narrow line of dark brown along posterior margin; elytron dark brown with indistinct yellow, transverse macula apicomedially on elytron ( Fig. 96 ); venter yellow, dark red-yellow on metacoxa; appendages yellow.

Sculpture and structure. Head finely punctate, punctures sparse, even more so on clypeus, surface between punctures finely microreticulate, shiny; anterior clypeal margin rounded; eyes large, not protuberant ( Fig. 96 ). Pronotum finely punctate, punctures moderately dense, finer and less dense laterally, surface smooth and shiny between punctures, with very fine microreticulation between punctures in some areas, setae very fine, moderately long; pronotum with lateral margins broadly curved, pronotum widest near middle ( Fig. 96 ); lateral bead narrow, distinct along entire margin; posterior angles obtusely rounded; transverse sulcus absent. Elytron with punctures moderately large, fairly dense, setae elongate and fine, surface very lightly microreticulate between punctures. Prosternum very short, transversely carinate; prosternal process with lateral margins rounded, broadly beaded, apex rounded, medially broadly tectiform. Metasternum impunctate except medially with fine, finely setose punctures, surface transversely microreticulate, shiny; anteromedial margin broadly and flatly beaded, medially narrowly carinate posterad of bead. Abdominal sterna very finely punctate laterally, surface shiny; sixth sternum with apex narrowly truncate, with apical marginal fringe of fine setae. Speleum exceptionally long, apically narrowed, apex bulbous, rounded, curved dorsally ( Figs 28–30 , 101 ). Metacoxa with lateral portion densely punctate medially and anteriorly, surface microreticulate and shiny between punctures; with metacoxal lines moderately approximated posteriorly, anteriorly slightly divergent; medial portion finely punctate, shiny ( Fig. 97 ). Metatrochanters elongate, apically narrowly rounded, not strongly offset ( Fig. 98 ); metatarsal claws minutely serrate.

Male genitalia. Median lobe in lateral aspect slightly and evenly curved, narrow throughout length, slightly expanded subapically, apex narrowed, narrowly rounded and slightly curved dorsally ( Fig. 103 ); in ventral aspect narrow, slightly to distinctly broadened subapically, apex divided into two, short, pointed processes ( Fig. 102 ). Lateral lobe elongate and slender, apex expanded into pointed, flattened, leaf-shaped structure, with few setae apically, numerous setiferous and nonsetiferous punctures around apex and along dorsal margin ( Fig. 104 ).

Female genitalia. Gonocoxosternite elongate, suboval, apex moderately pointed, anterior lobe moderately long, expanded anteriorly ( Fig. 105 ). Gonocoxa elongate, apex narrowly rounded, apodeme long, as long as main portion ( Fig. 105 ). Bursa copulatrix short, inconspicuous; spermathecal duct extremely long, slender and coiled; receptacle reduced; intermediate duct very long, tightly coiled into complicated mass; spermatheca elongate, narrow and curved medially, triangular process moderately broad; fertilization duct moderately long, curved apically ( Fig. 105 ).

Sexually dimorphic characters. Male with dense row of fine, long setae along mesotrochanter and mesofemur. Male pro- and mesobasotarsomeres I and II ( Fig. 99 ) moderately expanded compared with female ( Fig. 100 ).

Intraspecific variation. The extent of the elytral macula varies from nearly invisible or only visible laterally to clearly visible, conspicuously yellow and extending across the entire width of the elytron.

Natural history: Derovatellus lentus has been collected at black lights and from lowland subtropical ponds and pools.

Distribution: This species is very common and widespread in the Neotropical Region. It has been collected in Argentina, Bolivia, Brazil, Colombia, Dominca, the Dominican Republic, Ecuador, French Guyana, Guatemala, Guayana, Haiti ( Young, 1954), Panama, Paraguay, Peru, Puerto Rico, Suriname, Trinidad and Venezuela ( Fig. 108). It appears to be absent from further north in Mexico. Also, specimens from Florida attributed to this species appear to be D. floridanus ; however, the species is known from several areas of the Caribbean, suggesting that it may eventually be found in the US.

Phylogenetic relationships: This species belongs to the clade of Neotropical Derovatellus where it is sister species to D. peruanus ( Fig. 316 ). This is based on the very long, coiled intermediate duct between the receptacle and spermatheca in females ( Fig. 105 , Character 34). This feature also occurs in a variety of African species, but the condition is homoplasious in these two groups ( Fig. 320).

Material examined: Argentina: Corr. Ituzaingo , 20 December 1990, west grassland UV, S & J Peck (3, CMNC) ; Prov. Santa Fe Reconquista , 13 February 1979, blacklight, RE Woodruff (2, USNM) ; Santa Fe Reconquista , 13 February 1979, V Duthic and RE Woodruff (1, FSCA) ; Tuc. 20 km S Tucuman, 23 May 1969, P and P Spangler (4, USNM) . Bolivia: Beni, 40 km E San Borja, Estacion Biologica Beni, Estancia El Porvenir , 6–8 September 1987, black light, open grass savanna and marsh, WE Steiner (4, USNM) ; Beni, 1.8 km E San Borja , 15 July 1998, muddy pool, KB Miller (1, KBMC) ; Beni, 6.7 km NW Trinidad, 14∞47¢27≤S 64∞56¢47≤W, 18 July 1988, KB Miller (9, KBMC) ; Beni , 7.0 km SW Trinidad. 14∞52¢12≤S 64∞57¢32≤W, 18 July 1998, weedy pond, KB Miller (14, KBMC) ; Beni , Prov. Cercado 9.5 km N Trinidad. 14∞46¢34≤S 64∞58¢00≤W, 18 June 1999, KB Miller (4, KBMC) ; Santa Cruz, Santa Cruz , 11–12 May 1969, P and P Spangler (1, USNM) ; Santa Cruz, Ayacucho , 13 May 1969, P & P Spangler (7, USNM) ; Dpto. Sta Cruz, Prov. Chiquitos , 2.7 km S San Jose, 27 June 1999, pool in stream, KB Miller (43, KBMC) . Brazil: Country only (1, USNM) ; Bahia 15 km E Itabuna , 3 July 1969, P and P Spangler (61, USNM) ; Bahia, 15 km E Itabuna , 3 July 1969, P & P Spangler (1, USNM) ; Espirito Santos Linhares , October 1972, BLT, M Alvarenga (20, FSCA) ; Mato Grosso, Cuiaba Agr Exp Sta , 11 April 1972, BLT, WH Whitcomb (2, FSCA) ; Mato Grosso, Jacare Parque Nat. Xingu , November 1965, BLT, M Lavarenga, WCA Bokermann (1, FSCA) ; Matto Grosso (3, MNHN) ; Matto Grosso Jacare, PN Xingu, November 1961, Alvarenga and Werner (1, FSCA) ; Minas Geraes , P Mestreit (1, MNHN) ; Para Stream Nr Aldaia Coraci , 03 December 1964, Broys, Malkin (4, FSCA) ; R.G.N. Ceara-Mirim, 6–7 July 1969, P and P Spangler (1, USNM) ; Sao Paulo, Piracicaba , 24 October 1965, BLT, CA Triplehorn (1, FSCA) . Colombia: Putumayo, Puerto Asis , 4 July 1996, 300 m, R Reitmaier (2, LHIC) ; Rio Frio, Mgd , Darlington (2, MCZC) ; Rio Frio, Mgd , Darlington (17, MCZC) ; Sevilla, Mgd. , Darlington (3, MCZC) . Dominica: Cabrit Swamp , 10–13 May 1965, DR Davis (1, USNM) . Dominican Republic: Distrito Nacional Guerra (4 km N), 15 November 1984, P and P Spangler and RF Faitoute (1, USNM) ; LV Piedra Blanca , 19–20 July 1969, P and P Spangler (1, USNM) ; Sanchez , July 1938, Darlington (4, MCZC) . Ecuador: Nap Limoncocha , 11 June 1977, collected in pools of water hyacinth and water let, PJ Spangler, DR Givens (3, USNM) ; Nap Limoncocha , 13 June 1977, collected in pools of water hyacinth and water let, PJ Spangler, DR Givens (3, USNM) . French Guyana: Lawa R. Shore oppos. Anapaike village , 22–25 November 1963, B Malkin (6, FSCA) ; St Jean du Maroni , le Moult (1, MNHN) . Guatemala: Department Paten, Santa Teresa , 13 April 1935, Hubbs, VanderShalie (1, FSCA) . Guayana: Pirara Ranch 3∞37.3¢N 59∞44.2¢W, 27 April 1995, savanna pond, PJ Spangler, SA Perry (9, USNM) . Panama: Cocle, Penonome (13 km SW), Rio Cocle Auxiliar , 28 May 1983, PJ Spangler, RA Faitoute (1, USNM) ; Cocle, Penonome (13 km SW), Rio Cocle Auxiliar , 6 June 1983, PJ Spangler, RA Faitoute, WF Steiner (6, USNM) . Paraguay: Bolils (1, MNHN) ; Cen. 15 km NE Asuncion, 21 June 1969, P and P Spangler (50, USNM) ; Cen. 2 km N Luque, 23–24 June 1969, P and P Spangler (61, USNM) ; Cen. , 15 km NE Asuncion, 21 June 1969, P and P Spangler (15, USNM) ; Central Department, Aregua , 26–27 April 1980, PJ Spangler (39, USNM) ; Central Department; Aregua , 26–27 April 1980, PJ Spangler et al. (1, USNM) ; Cord. San Bernardino , 22 June 1969, P and P Spangler (54, USNM) ; Cordillera San Bernardino, shore of Lago Ypacarai , 7 November 1987, black light trap, J Kochalka (1, USNM) ; Department Central , Asuncion, 2 October 1992, U Drechsel (14, LHIC) ; Department Guaira, Calle Florida , 5 August, U Drechsel (6, LHIC) ; Department Gualra, Garay , 13 April 1980, L LHIC (55, LHIC) ; Paraguari Department, Ybycui 20 km SE Ruta La Rosada, Ybycui , 13 April 1980, roadside ponds, PJ Spangler (6, USNM) . Peru: Madre de Dios, Parque Manu, Pakitza Cocha Salvador 12∞07¢S 70∞58¢W 250 m, 21 September 1989, RA Faitoute (8, USNM) . Puerto Rico: Hwy 3 km 32.6 nr Palmer , 10 January 1963, PJ Spangler (42, USNM) ; Laguna Cartagena , 20 December 1962, P and P Spangler (1, USNM) ; nr La Cueva del Indio , 14 January 1963, PJ Spangler (1, USNM) . Suriname: Ceropinakreek , 55∞10¢W 05∞31¢N, 22 September 1989, N Nieser (1, NHMW) ; Christian Kondre, Marowije dist, 5–8 October 1963, light, B Malkin (1, FSCA) ; Paramaribo, Guest House of Museum , 1 October 1969 – 1 February 1970, N Nieser (3, NHMW) ; Zanderijsavanne, 1 st. Trib. of Colakreek, 55∞14¢W 05∞27¢N, 19 September 1989, N Nieser (5, NHMW) . Trinidad: La Brea, Darlington (4, MCZC) ; La Brea, Darlington (7, MCZC) ; Mayaro, 28 April 1929, Darlington (1, MCZC) ; Nariva Swamp , 30 July 1963, FN Young (1, FSCA) ; Rio Pan , 21 March 1912, A Busck (2, USNM) ; Riv. Pan , 21 May 1911, August, Busck (1, FSCA) ; Tabogilla Id Pan , 21 February 1912, A Busck (2, USNM) . Venezuela: Bolivar, El Dorado 65 km S, 4 November 1982, JL Hellman (1, USNM) ; Edo. Guar. Corozo Pando 8 km N, 17 August 1984, Eiland and Linares (1, USNM) ; Edo. Miranda Higuerote ( Barlo-vento ), 04 February 1962, C. Bordon (3, FSCA) ; Guar , 32 km SW Calabozo, 11 February 1969, P and P Spangler (3, USNM) ; Guar. Calabozo , 7 February 1969, P and P Spangler (1, USNM) ; Guar. Calabozo 44 km S Hato Masaguaral, 5 March 1986, PJ Spangler (86, USNM) ; Guar. Calabozo 44 km S Hato Masaguatal, 6 March 1986, PJ Spangler (45, USNM) ; Guar. San Fernanado , 12 February 1969, P and P Spangler (33, USNM) ; Guar. San Fernando , 12 February 1969, P and P Spangler (47, USNM) ; Guarico Corozo Pando (8 km N), 20–21 June 1974, blacklight, PW Eiland and V Linares (2, USNM) ; Guarico Corozo Pando 8 km N, 11 June 1984, FW Eiland (61, USNM) ; Guarico Corozo Pando 8 km N, 18 June 1984, blacklight, FW Eiland and V Linares (115, USNM) ; Higuerote n.5 ( Barlovento Edo. Miranda), 04 February 1962, Bordon (2, FSCA) .

Figures

Figure 106. Derovatellus species distributions. • = D. bruchi, O = D. peruanus, Δ = D. roosevelti, - = D. spangleri.

Figure 107. Derovatellus floridanus distribution.

Figures 51–56. Derovatellus bruchi. 51, dorsal habitus; 52, medial portion of metacoxae; 53, left metatrochanter and metafemur, anterior aspect; 54–55, tarsomeres, dorsal aspect; 54, male; 55, female, A, protarsus; B, mesotarsus; 56, speleum and abdominal sterna V, VI, dorsal aspect.

Figures 68–72. Derovatellus spangleri. 68, dorsal habitus; 69, speleum and abdominal sterna V, VI, dorsal aspect 70–72, male genitalia; 70, median lobe, right lateral aspect; 71, median lobe, ventral aspect; 72, right lateral lobe, lateral aspect.

Figures 96–101. Derovatellus lentus. 96, dorsal habitus; 97, medial portion of metacoxae; 98, left metatrochanter and metafemur, anterior aspect; 99–100, tarsomeres, dorsal aspect; 99, male; 100, female, A, protarsus; B, mesotarsus; 101, speleum and abdominal sterna V, VI, dorsal aspect.

Figures 57–61. Derovatellus bruchi. 57–59, male genitalia; 57, median lobe, right lateral aspect; 58, median lobe, ventral aspect; 59, right lateral lobe, lateral aspect; 60–61, female genitalia; 60, ventral aspect; 61, 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 102–105. Derovatellus lentus. 102–104, male genitalia; 102, median lobe, ventral aspect; 103, median lobe, right lateral aspect; 104, right lateral lobe, lateral aspect; 105, female genitalia, ventral 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 62–65. Derovatellus orientalis. 62, dorsal habitus; 63-35, male genitalia; 63, median lobe, right lateral aspect; 64, median lobe, ventral aspect; 65, right lateral lobe, lateral aspect.

Figures 66–67. Derovatellus orientalis. 66, speleum and abdominal sterna V, VI, dorsal aspect; 67, female genitalia, ventral aspect.

Figures 75–78. Derovatellus roosevelti. 75, dorsal habitus; 76–78, male genitalia; 76, median lobe, right lateral aspect; 77, median lobe, ventral aspect; 78, right lateral lobe, lateral aspect.

Figures 93–95. Derovatellus peruanus. 93, speleum and abdominal sterna V, VI, dorsal aspect; 94–95, female genitalia; 94, ventral aspect; 95, spermatheca and associated structures, dorsal aspect.

Figures 81–85. Derovatellus floridanus. 81, dorsal habitus; 82–85, male genitalia; 82, median lobe, right lateral aspect; 83, median lobe, ventral aspect; 84, median lobe, apex, right lateral aspect; 85, right lateral lobe, lateral aspect.

Figures 86–87. Derovatellus floridanus. 86, speleum and abdominal sterna V, VI, dorsal aspect; 87, female genitalia, ventral aspect.

Figures 79–80. Derovatellus roosevelti. 79, speleum and abdominal sterna V, VI, dorsal aspect; 80, female genitalia, ventral aspect.

Figures 88–92. Derovatellus peruanus. 88, dorsal habitus; 89–92, male genitalia; 89, median lobe, right lateral aspect; 90, median lobe, ventral aspect; 91, median lobe, apex, right lateral aspect; 92, right lateral lobe, lateral aspect.

Figures 73–74. Derovatellus spangleri, female genitalia. 73, ventral aspect; 74, spermatheca and associated structures, dorsal aspect.

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 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 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 28–37. Vatellini species, morphology. 28–30, Derovatellus lentus, speleum; 28, apical half of speleum, dorsal aspect with gland reservoirs removed; 29, dorsal aspect, dorsal surface of sternum VI, and gland reservoirs present; 30, left lateral aspect. 31–37, Derovatellus species, spelea and dorsal surfaces of sterna V–VI; 31, D. fasciatus; 32, D. decellei; 33, D. wewalkai; 34, D. dagombae; 35, D. nyanzae; 36, D. lugubris; 37, D. bisignatus.

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.