VATELLINI, Sharp, 1880

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 : 417-428

MORPHOLOGY OF VATELLINI

The members of this tribe exhibit morphological apomorphies in adults that are unusual among diving beetles and require some explanation and/or novel terminology. The following account is given to provide a basis for the formal descriptions of taxa, to point out and describe characters of particular taxonomic importance, and to provide some context for future comparative morphological studies that include members of the tribe. General aspects of vatelline morphology in the following treatment are based especially on cleared and disarticulated specimens of V. grandis and D. lentus , although other species were also examined to determine generality of the characters.

Body form: Members of Vatellus are elongate and have the lateral outline somewhat to strongly discontinuous between the elytron and pronotum with the pronotum often cordate (e.g. Fig. 1). Members of Derovatellus are continuous or somewhat discontinuous along the margin from the pronotum to elytron in dorsal aspect (e.g. Fig. 68 ) or have the lateral margin of the pronotum strongly curved and the lateral outline strongly discontinuous (e.g. Fig. 51 ). Some members of Vatellus are distinctly flattened dorsally and relatively strongly dorsoventrally compressed.

Surface sculpture: Vatellines, like many Hydroporinae (e.g. see Wolfe & Zimmerman, 1984), have a variety of surface features in the form of punctures, setae and microreticulations that vary between species. All species have scattered setae on the pronotum and elytra. The setae are set in prominent pores (some Vatellus , e.g. Fig. 2 ) or they arise from small depressions ( Derovatellus and some Vatellus , e.g. Figs 3, 4 ) or flat areas (some Vatellus , e.g. Fig. 5 ). The surface between the punctures may be relatively smooth and shiny (many Vatellus , e.g. Fig. 2 ), marked with transverse, impressed lines forming transversely orientated microcells ( Derovatellus , e.g. Fig. 4 ) or marked with short, curved striae bearing very short, spinous, posteriorly directed setae (some Vatellus , e.g. Fig. 3 ). Several Vatellus have irregular cells with small microcavities or irregularities on the surface of most cells ( Fig. 5 ). The surfaces of the metasternum, metacoxae and abdominal sterna exhibit similar patterns of surface sculpturing. In some species of Vatellus a portion of the pronotum and, in some cases, the elytron (especially apically) are marked with a large area of fine shagrination comprising extremely fine microtubercles. When fully developed this shagrination forms a pattern with smooth, shiny, impunctate transverse areas lacking shagrination on each side of the midline of the pronotum ( Fig. 262 ).

Cranium: The head in vatellines is short, rounded and robust. The eyes are located anterolaterally and are very large and protuberant, especially in some species of Vatellus (e.g. Fig. 131 ). The eyes are slightly emarginate along anterior and posterior margins. A paraocular carina extends from the anterior margin of the eye around the eye dorsally, posteriorly and ventrally. There is a distinct cavity posterad of the eye laterally that appears to receive the anterolateral angles of the pronotum. The frontoclypeal suture is effaced medially and indistinct laterally. The anterior clypeal margin in some Vatellus protrudes anteriorly in a distinct, angulate rim ( Fig. 6 ), whereas in other Vatellus and Derovatellus the anterior margin of the clypeus is evenly rounded ( Fig. 7 ). The tentorial arms are very slender and there are no tentorial bridges. The occipital foramen is very large, about one-half the width of the posterior surface of the head.

Antennae: Vatelline antennae are not modified from a form typical of most Hydroporinae. The scape and pedicel are longer and broader than flagellar antennomeres. The antennae are moderately long and antennomeres vary in relative size between species, but not strongly so.

Labrum and epipharynx ( Fig. 8 ): The labrum is broad and apically broadly emarginate medially. The medial margin bears a dense fringe of long fine setae and a series of short stout setae. Laterally on the aboral surface are numerous long, stout setae. Medially on the aboral surface is a broad V-shaped area of dense, short setae. The epipharynx is bordered laterally on each side by a short, stout sclerite. Medially there are two thickened membranous regions bearing short sensory setae.

Mandibles ( Figs 9–14 ): The mandibles in Vatellini are asymmetrical. Each is broad and broadly curved laterally. The apices of each are dorsoventrally broad. The left mandible is strongly excavated apicomedially ( Fig. 13 ). The pointed apex of the right mandible fits into this excavation. The medial margin of each mandible bears a long series of short setae. Medially, each mandible bears a small tooth-like retinaculum. Many Hydroporinae lack mandibular setae, a proposed synapomorphy of the subfamily ( Wolfe, 1985, 1988). However, all Vatellini examined possess a medial fringe of setae and a medial patch of setae on the ventral margin ( Figs 11, 12 ). Vatellus grandis (at least) also possesses long lateral setae ( Figs 9–14 ).

Maxilla ( Fig. 15 ): The cardo is stout and apically expanded. The stipes is small and narrow medially. The subgalea is elongate and irregular in shape. The palpus comprises four palpomeres and a basal palpiger. The palpiger and palpomeres I–III are relatively short and stout. Palpomere IV is longer than the basal three and bears an oblique, interrupted ring of sensory setae submedially and a small apical patch of sensillae. The galea consists of two long, slender segments. The lacinia is broad, strongly sinuate and sharply pointed apically. Its medial margin bears numerous long, stout setae.

Labium ( Figs 16, 17 ): The mentum is very broad and flattened with large anterolateral lobes, similar to most Dytiscidae . The ligula is stout and broad with a dorsal transverse excavation which bears stout setae on its lateral surface and a large field of dense, fine setae medially. The anterior surface bears a few long setae. The palpiger is broad and stout. The palpus consists of three palpomeres. The second bears a few apicodorsal setae. The apical palpomere bears a small field of sensillae apically.

Prothorax: The pronotum varies from subtrapezoidal to conspicuously cordate. In Derovatellus the lateral margins are evenly curved and the pronotum is widest medially (e.g. Fig. 51 ) or near the posterior angles (e.g. Fig. 68 ). In Vatellus the lateral margins are somewhat ( Fig. 259 ) to extremely ( Fig. 109 ) sinuate and are broadest anterad of the middle. The lateral margins are distinctly dentate ( Figs 19 , 262 ). In some species of Vatellus there is a transverse crease or depression in the surface of the pronotum slightly posterad of the middle (e.g. Fig. 230 ). The prosternal pore ( Wolfe, 1985, 1988) in most Hydroporinae is located at or near the anterolateral apex of the prosternum (where the anterolateral margin of the prosternum meets the pronotal epipleuron) ( Fig. 18 ). In vatellines the anterolateral margin of the prosternum is shifted medially and the prosternal pore is shifted posteromedially ( Fig. 19 ). The pore bears a conspicuous lobe ( Fig. 19 ).

In most Dytiscidae , the prosternal process extends between the pro- and mesocoxae where its apex contacts the anterior margin of the metasternum. In a few taxa the prosternal process is relatively short but still more-or-less attains the metasternum (e.g. the Ilybius opacus -group of species), whereas in others the process does not extend between the mesocoxae and the mesocoxae are contiguous. This last group includes several phylogenetically disparate groups like Stictotarsus minipi Larson , the Bidessini genus Tyndallhydrus Sharp , some species of Hygrotus (Coelambus) Thomson , and all members of Vatellini . In vatellines the prosternal process extends ventrally between the procoxae and is curved dorsally posterad of the procoxae ( Figs 21, 22 ). The mesocoxae are contiguous. The apex of the prosternal process is strongly flexed dorsad. The apex may be rounded or flattened and truncate to sharply pointed.

Mesothorax: Sharp (1882b) differentiated between Derovatellus and Macrovatellus based on the relative visibility of the mesosternum on each side of the prosternal process. However, I have not found this to be a useful character. In each group it is visible to approximately the same degree. The mesothorax is narrow and relatively undifferentiated.

Metathorax: As recognized most notably by Sharp (1882b), in extant vatellines the metepisternum does not appear to extend to the mesocoxal cavity, instead it is separated from it by the mesepimeron. This is a condition putatively shared in Hydradephaga with Gyrinidae , Hygrobiidae , Noteridae and Laccophilini. These other taxa differ from Vatellini in having the mesepimeron separated both externally and internally from the mesocoxal cavity. In vatellines the metepisternum extends to the cavity, but the posteromedial angle of the mesepimeron extends ventrally as a lobe over the medial end of the metepisternum ( Fig. 20 ). The metasternum is medially broadly convex. In some species of Vatellus the anteromedial process is prominently swollen ventrad ( Fig. 20 ). In Calicovatellus the metepisternum extends to the mesocoxal cavity externally.

Elytron: The elytron is relatively undifferentiated in vatellines. Dorsally there are no conspicuous striae, carinae or plicae. Occasionally very slightly raised longitudinal lines can be discerned on some species, particularly V. mexicanus and closely related species. Ventrally the apicolateral lobe (elytral ligula) is singular and simple, though it may be quite large in some species, especially in some Vatellus . The lateral carina is low and relatively short with no medial lobe. There are elongate friction pads along the lateral and medial margins comprising fields of minute tubercles or spines.

Metathoracic wing ( Figs 23, 24 ): The flight wings in Vatellini are similar in most respects to other hydroporines. The wings are short, broad and strongly rounded apically. The oblongum cell is transversely oval. The longitudinal triangular space between the radius and radius posterior is very broad. The radial cell is broad and strongly rounded apically. The submedial binding patch (this term based on Lawrence & Britton, 1994) is prominent and consists of two elongate, robust, oblique parallel parts and a pigmented area between them. In Vatellus the wedge cell is relatively large and elongate ( Fig. 24 ) whereas in Derovatellus it is very small ( Fig. 23 ), a trend similar in other small Hydroporinae. Derovatellus has reduced venation ( Fig. 23 ), also similar to other small Hydroporinae. In this genus, the media posterior veins (MP) and radius posterior vein 2 (RP2) are weakly visible to absent ( Fig. 23 ). In Vatellus the RP2 is present and branches from the RP1 distad of the oblongum cell ( Fig. 24 ). In Dytiscidae except Hydroporinae and Laccophilini the RP2 arises from the margin of the oblongum cell ( Goodliffe, 1939; Balfour-Browne, 1944; Wolfe, 1985; Wolfe, 1988).

Legs: The legs of vatellines are unusually long. Pro- and mesotarsomeres I–III on both females and males bear a ventral field of adhesive setae ( Fig. 25 ). The apices of the setae are slightly ovoid ( Fig. 25 ) and often dimorphic with many small setae and fewer large setae interspersed ( Fig. 25 ). These tarsomeres are often more strongly expanded in males than in females, though only slightly more so in some species of Vatellus . All tarsi and tibiae bear long natatory setae along the dorsal and ventral margins. Males of many species, especially in Vatellus , have a large cluster of fine, long setae on the apicoventral surface of the mesotrochanter and the base of the mesofemur. The metacoxae are strongly anteriorly expanded and are relatively long. The metacoxal lines are closely approximated in Derovatellus (e.g. Fig. 97 ). In many species of Vatellus the metacoxal lines are closely approximated posteriorly and are strongly divergent anteriorly (e.g. Fig. 300 ). The lateral portion of the metacoxa is variously punctate from nearly impunctate to very coarsely punctate and in some cases there is a distinctive microsculpture similar to that found on the elytra (see above under Surface sculpture). The anteroventral margins of the metatibia and metatarsomeres of those species examined have a series of very fine dentitions ( Fig. 26 ). The metatarsal claws are unequal in length with the posterior claw slightly longer. The metatarsal claws are denticulate with rounded teeth ( Fig. 27 ). The metatrochanter in Vatellus is often strongly offset (e.g. Fig. 301 ). It bears a prominent basal, circular lobe that extends medially under the surface of the metacoxa when the leg is flexed anteriorly and a distinct, broad emargination in the ventral (posterior) surface (e.g. Figs 267, 268 ). The metatrochanter in Derovatellus is not unusually modified ( Fig. 98 ).

Abdomen: Visible sterna II–V are relatively unmodified compared with other Hydroporinae. However, sternum VI exhibits complex modification in all Vatellini . In vatellines there is an elongate, invaginated, heavily sclerotized structure extending from the apex of the sixth abdominal sternum internally along the dorsal surface of the abdominal sterna ( Figs 28–30 ). Because of the unique nature of this feature, I have coined a new term for it, ‘ speleum ’, Latin for ‘cave’. The speleum is a hollow, sclerotized tube that extends from a broad opening along the apical margin of the sixth visible abdominal sternum. It is generally moreor-less flattened for much of its length and variously dorsoventrally undulate ( Fig. 29 ). The anterior apex is variously expanded and may be dorsally curved ( Fig. 30 ). The apex is typically very darkly pigmented and heavily sclerotized. There appear to be no muscles attached to the structure. The speleum is apparently the reservoir and duct of a large gland system. In fresh specimens of Vatellus and Derovatellus , a large region of white cells is visible surrounding the speleum with numerous tiny ducts emptying into the apex of the speleum or along its length ( Fig. 29 ). Because welldeveloped pygidial glands are present in vatellines, the speleum is apparently not derived from their modification. The orifice forms a groove along the margin of the sixth visible sternum and the ventral margin of the apex may be variously lobed, particularly in Vatellus , or setose. The dorsal surface of the ventral margin of the orifice in Vatellus generally has a swelling that fits into a corresponding cavity on the ventral surface of the dorsal margin. The function of this gland structure is unknown. It occurs in both males and females of all species, though in some Derovatellus (e.g. D. fasciatus Régimbart , Fig. 33 ), it is reduced. The speleum is species-specific in shape in many taxa, particularly in Derovatellus where it exhibits considerable variation, but also in Vatellus . There appears to be very little variation in shape within species. Because of damage to the type specimen, it is not known whether Calicovatellus petrodytes possesses a speleum. No apparently homologous features occur in this position in any of the many other Hydroporinae genera examined.

Male genitalia: As in other diving beetles, the genitalia of vatellines are rotated 90∞ in repose and 180∞ when extruded ( Balfour-Browne, 1940; Sharp & Muir, 1912). Thus the concave surface of the median lobe is actually the dorsal surface ( Miller & Nilsson, 2003). Members of the tribe have relatively unmodified, bilaterally symmetrical male genitalia. Members of Derovatellus exhibit considerable autapomorphic variation in shape of the median lobe, but do not exhibit many apparent synapomorphies. By contrast, members of Vatellus have the median lobe very long and evenly curved in general with relatively subtle differences in shape. However, they have several prominent features that appear to be synapomorphies within the group. In all species the basal portion of the lateral lobes are dramatically elongated anteriorly (e.g. Fig. 271 ) and are fused along the dorsal margins. The base of the median lobe may also be extremely elongated (e.g. Fig. 269 ) or this portion may be shorter (e.g. Fig. 125 ). The apex of the median lobe varies in shape from species to species, though differences are often subtle. The apices of the lateral lobes are often laterally flattened and expanded (e.g. Fig. 200 ), though they may be narrowed (e.g. Fig. 285 ). Some species have two or three long, stout, curved, subapical setae on the ventral surface (e.g. Fig. 308 ).

Members of Dytiscidae have a ring-like sclerite surrounding the genital capsule considered the joined last dorsal and ventral sclerites by Sharp & Muir (1912) and regarded by Roughley & Pengelly (1981), Franciscolo (1979) and others as the fused basal portion of sternum IX, pleuron IX and tergum IX. In Hydroporinae (and a few other dytiscids such as some Aciliini) the ventral sclerite in this ring is absent. In vatellines this ring is very elongate with the apices of the lateral arms expanded apically ( Figs 44, 47 ). The apices are fused in Vatellus ( Fig. 44 ) whereas in Derovatellus they are separated by clear membrane ( Fig. 47 ). In Vatellus the anteromedial lobes of the fused sterna VIII are greatly prolonged anteriorly ( Fig. 42 ), but

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they are short in Derovatellus ( Fig. 45 ). The spiculum in Vatellus is long, expanded on each end and medially bent ( Fig. 43 ). In Derovatellus it is short, not bent and expanded on one end ( Fig. 46 ). As with other Dytiscidae , there is no differentiated internal sac. There is a sclerotized structure present on the ejaculatory duct ( Figs 48, 49 ). This structure is present throughout Dytiscidae , but it has seemingly not been previously mentioned or described. There has been no published account of the potential function of this ejaculatory sclerite.

Female genitalia: Vatelline female genitalia are, in general configuration, similar to most ‘derived’ Hydroporinae [those taxa with Hydroporine-type female genitalia and receptacle present (Miller, 2001)]. There is considerable variation within the group, however. Members of Vatellus have relatively broad gonocoxosternites with long anterior lobes (e.g. Fig. 186 ) whereas those of Derovatellus are elongate-triangular with the apices relatively acutely rounded and with long anterior lobes (e.g. Fig. 73 ). In both there is generally a dense series of short, fine setae along the entire posterior margin and a series of long, stout, evenly spaced setae along the medial half of the posterior margin and posterior half of the medial margin (e.g. Figs 73 , 186 ). Members of both genera have gonocoxae relatively consistent with the generalized form

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in Hydroporinae. Members of Vatellus are somewhat variable in the length and degree of roundedness of the gonocoxae whereas those of Derovatellus are generally elongate and apically pointed (e.g. Fig. 73 ). The bursa in most taxa is small and relatively undifferentiated though there may be variation in the presence of lobes (e.g. Fig. 145 ), unusual sculpture near the insertion of the spermathecal duct (e.g. Fig. 159 ), size and shape. In one taxon, V. haagi , the bursa is exceptionally long, flattened and dorsoventrally sinuate ( Figs 186, 187 ). The spermathecal duct varies from moderately long and slender in Vatellus and many Derovatellus to exceptionally long and very slender in a few Derovatellus (e.g. Fig. 80 ). The spermathecal duct near the receptacle, the receptacle and the duct between it and the spermatheca are variable in length, shape and degree of curvature in Vatellus and extremely variable in Derovatellus . Close examination of the spermathecal duct reveals microstructure (e.g. Fig. 186 ). The receptacle in Vatellus is generally relatively large and subspherical (e.g. Fig. 186 ) though it may be reduced (e.g. Fig. 272 ). In Vatellus the intermediate duct between the receptacle and spermatheca is generally relatively short, robust and somewhat sinuate (e.g. Figs 186, 188 ), though in a few species it is very long and strongly coiled on the dorsal surface of the spermatheca (e.g. Fig. 258 ). Members of Derovatellus have the receptacle variable from very large and anteriorly directed (e.g. Fig. 80 ) to reduced or nearly absent (e.g. Fig. 105 ). The intermediate duct may be short or nearly absent (e.g. Fig. 80 ) to long, moderately and strongly twisted, or coiled (e.g. Figs 94, 95 ). A small diverticulum is present on the spermathecal duct in some species (e.g. Fig. 80 ). The spermatheca in vatellines comes in two general forms. In Vatellus it is subspherical anteriorly (e.g. Fig. 186 ) whereas in Derovatellus it is elongate and bent medially (e.g. Fig. 60 ). All vatellines have a broad, subtriangular, truncate process at the posterior apex of the spermatheca which varies in size (e.g. Figs 60 , 186 ). In D. roosevelti it is apically rounded rather than truncate in shape ( Fig. 80 ). Members of both genera have the spermatheca bearing fine gland ducts. In Vatellus the glands cover the surface of the spermatheca ( Fig. 186 ) whereas in Derovatellus the glands occupy a region near the origin of the fertilization duct (e.g. Fig. 105 ). The fertilization duct is generally short but may be longer in some species (e.g. Fig. 80 ). The ventral surface of the common oviduct bears a large, oval region of short spines in all species. Finally, the rami are relatively broad anteriorly and posteriorly convergent to a sharply pointed apex (e.g. Fig. 186 ).

Larva: Third instar larvae of D. ibarrai (= D. floridanus ) and V. mexicanus were described by Spangler (1963, 1966b) and this diagnosis is based on his descriptions. Larvae of Vatellini are characterized by an exceptionally elongate and slender nasale that is apically expanded and rounded. The nasale bears a long, slender, curved process on each side. These rami are apically bifid. The nasale and the lateral rami are prominently spinous. Abdominal segments VII and VIII are entirely sclerotized. Abdominal segment VIII terminates in a very long, slender posterior projection that is subequal in length to the urogomphi. The urogomphi are unsegmented and bear many small setae.

Figures

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

Figures 131–138. Vatellus mexicanus. 131, dorsal habitus; 132, elytral sculpture in area indicated on habitus by dotted square; 133, medial portion of metacoxae; 134, metatrochanter and metafemur, anterior aspect; 135–136, tarsomeres, dorsal aspect; 135, male; 136, female, A, protarsus; B, mesotarsus; 137, sternum VI, ventral aspect; 138, speleum abdominal sterna V, VI, dorsal aspect.

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

Figures 109–114. Vatellus ventralis. 109, dorsal habitus; 110, medial portion of metacoxae; 111, elytral sculpture in area indicated on habitus by dotted square; 112, left metatrochanter and metafemur, anterior aspect; 113, sternum VI, ventral aspect; 114, speleum abdominal sterna V, VI, dorsal aspect.

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

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

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

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

Figures 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 125–130. Vatellus sahlbergi. 125–128, male genitalia; 125, median lobe, right lateral aspect; 126, median lobe, ventral aspect; 127, right lateral lobe, lateral aspect; 128, apex of median lobe, ventral aspect; 129–130, female genitalia; 129, ventral aspect; 130, spermatheca and associated structures, 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.

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

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

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

Figures 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 73–74. Derovatellus spangleri, female genitalia. 73, ventral aspect; 74, 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 155–160. Vatellus maculosus. 155–158, male genitalia; 155, median lobe, right lateral aspect; 156, median lobe, ventral aspect; 157, right lateral lobe, lateral aspect; 158, apex of median lobe, ventral aspect; 159–160, female genitalia; 159, ventral aspect; 160, spermatheca and associated structures, dorsal aspect.

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

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

Figures 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 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 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.