Revision of the New World and south-east Asian

Transcription

Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The
nean Society of London, 2005? 2005
144?
415510
Original Article
Lin-
NEW WORLD AND ASIAN VATELLINIK. B. MILLER
Zoological Journal of the Linnean Society, 2005, 144, 415–510. With 321 figures
Revision of the New World and south-east Asian Vatellini
(Coleoptera: Dytiscidae: Hydroporinae) and phylogenetic
analysis of the tribe
KELLY B. MILLER*
Department of Integrative Biology, 401WIDB, Brigham Young University, Provo, Utah 84602, USA
Received August 2004; accepted for publication March 2005
Members of the tribe Vatellini occurring in the New World and south-east Asia are revised. A cladistic analysis of the
tribe is presented including all New World and south-east Asian species and several species of African Derovatellus
Sharp. The results indicate that Mesovatellus Trémouilles is nested within Derovatellus, that Vatellus Aubé is nested
within Macrovatellus Sharp, and that D. (Varodetellus) Biström and Derovatellus are not mutually monophyletic.
Therefore, the following genus–group synonymies are formally recognized; Mesovatellus and
D. (Varodetellus) = Derovatellus (syn. nov.) and Macrovatellus = Vatellus (syn. nov.). Derovatellus floridanus is elevated from status as a synonym or subspecies of D. lentus (Wehncke) to species status (stat. nov.). The following
eight new species are described: Derovatellus roosevelti, D. spangleri, Vatellus drymetes, V. amae, V. annae,
V. pilacaudus, V. wheeleri and V. maculosus. The following species–group synonymies are formally recognized:
Derovatellus ibarrai Spangler 1966 = D. floridanus Fall, 1932 (syn. nov.); and Macrovatellus rudis Sharp, 1882 and
M. marginalis Sharp, 1882 = Vatellus lateralis (Sharp, 1882) (syn. nov.). A detailed revision of the morphology of
members of the tribe is presented along with the description of an abdominal gland system (the ‘speleum’) that is a
synapomorphy for the Vatellini. Keys to species and type information, diagnoses, descriptions, distributions and phylogenetic information are provided for each extant New World and south-east Asian species in the tribe. © 2005 The
Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 415–510.
ADDITIONAL KEYWORDS: cladistics – monograph – morphology – water beetle.
INTRODUCTION
Members of the tribe Vatellini are among the most
morphologically distinctive of the Dytiscidae. Their
long legs, large eyes and characteristic habitus (e.g.
Fig. 1) make them very different in general appearance from more typical members of the family. Some
members of Derovatellus Sharp can be found in very
large numbers in the tropics in ponds with dense
emergent vegetation. As Sharp (1882b) pointed out,
specimens of Vatellus Aubé (most previously assigned
to the genus Macrovatellus Sharp) are relatively
rarely collected. Members of this genus occur in habitats similar to those of Derovatellus.
African species (all historically placed in Derovatellus) have had a relatively long history of taxonomic
*E-mail: [email protected]
treatment including a modern revision and phylogenetic analysis (Biström, 1979). However, the New
World species have received much less attention, and
the species have not been revised since Sharp’s (1882b)
monograph. Spangler (1966b, 1967) described new species of Derovatellus (including the larva) and Biström
(1980b) figured the genitalia of some non-African Derovatellus and discussed their taxonomy. Species in the
other New World genus, Vatellus, have had much less
treatment. The last species described was 50 years ago
(by Guignot, 1955). Since then, the only taxonomic
treatments include Spangler’s (1966a) transfer of Platydessus perforatus Guignot (Bidessini) to Macrovatellus and his description of the larva of Vatellus
mexicanus (Spangler, 1963). More recently, Trémouilles (1995) placed Derovatellus bruchi Zimmermann in
its own, new genus, Mesovatellus Trémouilles.
A single extinct species, Calicovatellus petrodytes
Miller & Lubkin, was described recently from a par-
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 415–510
415
416
K. B. MILLER
K .B
.M
ill e r
1.0mm
Figure 1. Vatellus perforatus, dorsal habitus.
NEW WORLD AND ASIAN VATELLINI
ticularly well-preserved silicaceous specimen from the
Miocene in California (Miller & Lubkin, 2001). This
species was hypothesized to belong to the tribe based
on the short prosternal process (not reaching the
metasternum, characteristic of vatellines) and several
other less discrete characters. The species was further
hypothesized to be the sister to the known extant species based on its metepisternum reaching the middle
coxal cavity externally (not reaching the coxal cavity
in other vatellines). As with many fossils, the specimen of Calicovatellus has a unique combination of
some characters but physically lacks many of the features needed to help clarify its phylogenetic status.
However, its position in the tribe is analysed here
within the context of a revision.
The goal of this work is to revise the New World
members of Vatellini within a phylogenetic context
and to explore the unique morphology of the group.
Since the African species were treated in a careful
revision and subsequent works by Biström (1979,
1980a, 1981, 1982, 1983, 1984, 1986) and Biström &
Roughley (1982) these species are not revised here.
However, a revision of New World species would be
incomplete without a global context. Thus, numerous
African and the single south-east Asian Derovatellus
were examined, and the phylogenetic analysis presented here includes them to provide a more thorough
representation of the group and to test monophyly of
the genera. As the south-east Asian D. orientalis
Wehncke has not been comprehensively illustrated or
described and a few specimens were available, it is
included in this revision.
METHODS
Material: Specimens were borrowed from the following public and private collections in North and South
America and Europe.
AMNH The American Museum of Natural History,
New York, NY (L. Herman).
BMNH The Natural History Museum, London (M.
Brendell). CMNC The Canadian Museum of
Nature Collection, Ottawa, ON (R. Anderson).
CMNH The Carnegie Museum of Natural History,
Pittsburg, PA (R. Davidson).
FSCA The Florida State Collection of Arthropods,
Gainesville, FL (M. C. Thomas).
JBWM J. B Wallis Museum of Entomology, University
of Manitoba, Winnipeg, MB (R.E. Roughley).
LHIC
L. Hendrich Collection, Berlin.
MBIC M. Balke Collection, London.
MCZC The Museum of Comparative Zoology, Harvard University, Cambridge, MA (P. Perkins).
MNHN Museum National d’Histoire Naturelle, Paris
(M. H. Perrin).
417
NHMW Naturhistorisches Museum Wien, Vienna (M.
A. Jäch).
PMIC P. Mazzoldi Collection, Brescia.
USNM The United States National Museum, Washington, DC (P. J. Spangler, W. S. Steiner).
ZSBS
Zoologische Staatssammlung des Bayerischen Staates, Munich (M. Baehr, A.M.
Kühbandner).
Measurements: Measurements were made using an
ocular micrometer in a Wild M3C dissecting microscope. An attempt was made to measure the smallest
and largest individuals examined of each species.
Measurements and their ratios referred to in this
study include: TL, total length (calculated as
HL + PL + EL); HL, head length, measured medially;
PL, medial length of pronotum; EL, length of elytron
from posteromedial apex of pronotum to apex of
elytron; GW, greatest width; PW, greatest width of
pronotum; HW1, width of head across eyes; HW2,
minimum distance between eyes; ML, length of medial
margin of metacoxae; MW, greatest distance between
metacoxal lines.
Association of sexes: In this group, diagnosis of many
species depends on male and female genitalia with few
diagnostic characters overlapping between the sexes.
In these cases, association of males with females of a
given species may be difficult. Species hypotheses in
this project rely primarily on characters of the male
genitalia, and sexes were associated by identifying
series with both males and females of what appear to
be single species in geographical areas where the species do not appear to overlap. Fortunately, closely
related species in this taxon rarely overlap in distribution, but in some cases association of males and
females remains difficult.
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
418
K. B. MILLER
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
2
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).
3
0.05mm
0.05mm
0.05mm
0.05mm
5
4
7
6
0.5mm
0.5mm
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.
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 syn-
419
apomorphy 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
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K. B. MILLER
10
9
ventromedial setae
apicoventral setae
labrum
0.1mm
12
11
epipharynx
8
lacinia
galea
IV
palpiger I
II III
0.25mm
13
palpus
14
subgalea
0.25mm
stipes
palpus
III
cardo
15
III
II
I
II
16
I
0.25mm
palpiger
ligula
17
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.
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
421
Prpo
Prst
Prel
CxI
0.5mm
Mses
Msst
Prpl
Msem
CxII
18
Mtes
Prpo
CxI
Prst
Prel
0.5mm
Mtst
Prpl
0.5mm
20
19
Prst
FeI
CxI
TrI
Msst
1.0mm
FeII
TrII
21
CxII
Prpr
Mtst
22
Prst
CxI
FeI
TrI
0.5mm
Msst
Prpr
CxII
TrII
FeII
Mtst
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.
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
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K. B. MILLER
(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.
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
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.
1.0mm
0.05mm
23
C
rc
24
Sc
R
r-r
2.0mm
RP
RA
MP
r-m
CuA
wc
r-m
25
AP
MP1
RP1
RP2
MP2 AA1
oc
mf
AA2
0.1mm
26
0.1mm
27
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.
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. Proand 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
423
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,
424
K. B. MILLER
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 Hydropori-
nae (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
31
1.0mm
32
28
33
34
0.25mm
35
36
29
30
37
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.
38
39
0.5mm
40
42
0.5mm
45
425
41
46
0.25mm
0.5mm
43
47
44
0.05mm
0.25mm
50
48
49
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.
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’ Hydro-
porinae [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
426
K. B. MILLER
1.0mm
54
0.5mm
A
B
52
0.5mm
B
A
55
0.5mm
53
51
56
0.5mm
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.
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
427
ID
SP
SP
58
RE
RE
TP
57
FD
TP
0.1mm
FC
61
FD
0.25mm
59
FC
SD
60
BC
GC
GS
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.
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).
428
K. B. MILLER
63
64
65
0.25mm
0.5mm
62
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.
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.
VATELLINI SHARP, 1882
Vatellini Sharp, 1882b: 258 (as group of Dytisci
fragmentati); Régimbart, 1895: 11 (new status);
Zimmermann, 1919: 124, 1920: 29 (as tribe of Hydroporinae); Leng, 1920: 76; Guignot, 1959: 59; Nilsson,
Roughley & Brancucci, 1989: 312; Trémouilles,
1995: 24; Nilsson, 2001: 233; Roughley & Larson,
2001: 169.
Vatellinae, Omer-Cooper, 1958: 249.
Diagnosis: This tribe is defined as those Hydroporinae
with the following combination of characters: (1) prosternal process not reaching metasternum (mesocoxae
contiguous) (Figs 21, 22), (2) abdominal sternum VI
with an invaginated, heavily sclerotized gland
(speleum) (Figs 28–30) and (3) female with apically
expanded and broadly truncated process at apex of
spermatheca (e.g. Figs 60, 186). The extinct species,
Calicovatellus petrodytes, was proposed as the sisterspecies to the remaining members of the tribe as the
metepisternum reaches the mesocoxal cavities (Miller
& Lubkin, 2001). A speleum and a triangular sper-
429
0.1mm
66
0.5mm
67
Figures 66–67. Derovatellus orientalis. 66, speleum and abdominal sterna V, VI, dorsal aspect; 67, female genitalia, ventral aspect.
mathecal process have not been observed in Calicovatellus, but the prosternal process does not reach the
metasternum and the general body shape is characteristic of other members of the tribe. Extant species
all possess a metepisternum with its medial apex covered by a lobe of the mesepimeron (the mesepimeron
and the metasternum meet along the mesocoxal
cavity).
Taxonomic history: This taxon was first erected as a
group in Sharp’s (1882b) ‘Dytisci fragmentati’ based
on the separation of the metepisternum from the
mesocoxal cavities (at least externally). The tribe was
placed in the Hydroporinae by subsequent authors
(Zimmermann, 1920; Balfour-Browne, 1940; Crowson,
1955). Omer-Cooper (1958) recognized the group at
the rank of subfamily, but this has not been generally
followed. The phylogenetic position of the group
nested within the Hydroporinae was supported by
Wolfe’s (1985, 1988) analyses, but relationships of this
tribe to others within the Hydroporinae remains
equivocal. Wolfe (1985, 1988) found M. mexicanus to
be sister to a species of Oreodytes, but regarded this
relationship as doubtful. The group was resolved with
members of Hydroporini in a morphological analysis
(Miller, 2001). Most recently, Biström & Nilsson
(2003) proposed a close relationship between Derovatellus, Peschetius Guignot and Necterosoma McLeay.
Prior to this revision there were 52 species (one
species with a single subspecies) recognized in five
genera (one genus with two subgenera). With this
revision there are 57 species in the tribe in three
genera. Changes include three genus-group synonymies, eight new species, one elevation of a subspecies to species rank, and four new species-group
synonymies.
430
K. B. MILLER
KEY
TO THE GENERA OF VATELLINI
Adults
1. Metepisternum extending to mesocoxal cavity externally . . . . . . . . . . . . . . .Calicovatellus Miller & Lubkin (extinct)
1¢. Metepisternum apparently not extending to mesocoxal cavity externally, separated from it by a lobe of the
mesepimeron (Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Pronotum not cordate, variously rounded, widest at or posterior to middle (e.g. Figs 51, 68); metacoxal lines moderately approximated throughout length, not or only slightly divergent anteriorly (e.g. Figs 52, 97); metatrochanter
with ventral (posterior) margin evenly curved, not emarginate and without prominent basal, circular lobe (e.g.
Figs 53, 98); size smaller, TL = 3.0–5.0 mm; male lateral lobes not basally extended, not fused ventrally (e.g. Figs 59,
104); female spermatheca elongate, tubular, medially angulate (e.g. Figs 60, 105); apical margin of ventral apex of
orifice of speleum not lobed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Derovatellus Sharp
2¢. Pronotum variously cordate, widest anterior to middle (e.g. Fig. 1); metacoxal lines generally closely approximated
posteriorly, strongly divergent anteriorly (e.g. Fig. 176); ventral (posterior) margin of metatrochanter with prominent medial emargination (e.g. Figs 267, 268); metatrochanter with prominent basal, circular lobe that extends
medially over surface of metacoxa with leg in anterior position (Figs 267, 268); size larger, TL = 4.5–7.5 mm; male
lateral lobes in basal portion strongly extended and fused along ventral margin (e.g. Fig. 184); female spermatheca
globular, subspherical (e.g. Fig. 186); apical margin of ventral apex of orifice of speleum distinctly lobed (e.g.
Figs 180, 222) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vatellus Aubé
Larvae (Calicovatellus unknown) (adapted from Spangler (1966b))
1. Antennomeres II and III subequal in length and longer than other antennomeres; antennomere IV smallest, slightly
less than half length of antennomere III. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Derovatellus Sharp
1¢. Antennomeres I and II subequal in length and longer than other antennomeres; antennomere IV smallest, about oneseventh length of antennomere III. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vatellus Aubé
CALICOVATELLUS MILLER & LUBKIN, 2001
Calicovatellus Miller & Lubkin, 2001: 891 (type
species: Calicovatellus petrodytes Miller & Lubkin,
2001: 892, by original designation).
Diagnosis: The only known member of this genus differs from other members of this tribe in having the
metepisternum extending medially to the mesocoxal
cavities externally. Other genera have the metepisternum externally excluded from the mesocoxal cavity.
Distribution: This genus is known only from the
Miocene Barstow Formation in the Calico Mountains
of San Bernardino County, California. No extant members of the genus are known.
CALICOVATELLUS PETRODYTES MILLER &
LUBKIN, 2001
Calicovatellus petrodytes Miller & Lubkin, 2001: 892.
Type information: Holotype, sex not determined, in
USNM (Miller & Lubkin, 2001).
Type locality: Lower nodule layer, Barstow Formation,
Mule Canyon, Calico Mountains, Mojave Desert, San
Bernardino County, California, USA.
Diagnosis: This is the only known member of the
genus and is characterized by the features present in
the genus.
Distribution: Calicovatellus petrodytes is known only
from the Barstow Formation, San Bernardino County,
California, USA.
Phylogenetic relationships: This species was hypothesized to be the sister group to the remaining members
of the tribe (Miller & Lubkin, 2001). In the cladistic
analysis conducted for this project, Calicovatellus is
resolved with other members of the tribe (Fig. 218) as
sister to the other Vatellini or as sister to certain
groups within Derovatellus. These conflicting alternative placements are due primarily to missing data
rather than to character conflict.
Discussion: This species was recently described in
detail (Miller & Lubkin, 2001). It is the only known
extinct species in the tribe. Only a single specimen
was investigated by Miller & Lubkin (2001). However,
since then at least one additional specimen has been
isolated from the type formation (M. Thorpe, University of Central Arkansas, pers. comm.) and deposited
at the University of Central Arkansas (B. Waggoner).
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:
431
70
71
72
0.25mm
1.0mm
68
69
0.5mm
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.
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, slender, 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
432
K. B. MILLER
74
0.1mm
73
Figures 73–74. Derovatellus spangleri, female genitalia. 73, ventral aspect; 74, spermatheca and associated structures,
dorsal aspect.
(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
433
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
434
K. B. MILLER
Table 1. Measurements and ratios of selected features of species of Derovatellus and Vatellus. TL, total length (calculated
as HL + PL + EL); HL, medial length of head length; PL, medial length of pronotum; EL, length of elytron from posteromedial apex of pronotum to apex of elytron; GW, greatest width; PW, greatest width of pronotum; HW1, width of head across
eyes; HW2, minimum distance between eyes; ML, length of medial margin of metacoxae; MW, greatest distance between
metacoxal lines
D. bruchi
D. orientalis
D. spangleri
D. roosevelti
D. floridanus
D. peruanus
D. lentus
V. ventralis
V. sahlbergi
V. mexicanus
V. maculosus
V. lateralis
V. haagi
V. wheeleri
V. perforatus
V. tarsatus
V. annae
V. pilacaudus
V. grandis
V. bifenestratus
V. drymetes
V. amae
TL (mm)
GW (mm)
TL/GW
PW (mm)
PL (mm)
PL/PW
HW1/HW2
ML/MW
4.92–4.97
3.58–3.68
3.17–3.86
3.68–3.73
3.45–3.91
3.27–3.80
3.45–3.70
6.44
6.58–7.02
5.01–6.35
5.38–6.14
5.77–7.27
5.22–5.29
5.44–5.82
4.65–4.72
4.65–5.15
4.88–5.50
5.15–5.75
6.12–6.49
5.00–5.11
5.52–5.70
5.15–5.43
2.39–2.48
1.70–1.78
1.43–1.75
1.68–1.84
1.51–1.79
1.50–1.73
1.52–1.82
3.13
3.13–3.40
2.48–2.99
2.53–2.99
2.81–3.36
2.55–2.67
2.39–2.67
2.21–2.30
2.23–2.39
2.37–2.67
2.53–2.81
2.58–2.85
2.27–2.31
2.44–2.65
2.25–2.39
1.98–2.06
2.06–2.10
2.21–2.23
2.03–2.19
2.18–2.28
2.18–2.20
2.04–2.27
2.06
2.06–2.10
2.02–2.12
2.05–2.13
2.06–2.16
1.96–2.07
2.18–2.28
2.02–2.10
2.08–2.15
2.06–2.12
2.04–2.05
2.27–2.38
2.2–2.21
2.16–2.26
2.27–2.29
1.61–1.79
1.24–1.30
1.10–1.33
1.24–1.33
1.2–1.41
1.13–1.27
1.17–1.27
1.93
2.16–2.39
1.47–1.75
1.70–1.84
1.86–2.21
1.54–1.61
1.57–1.68
1.61–1.66
1.45–1.56
1.52–1.70
1.66–1.79
1.93–2.05
1.65–1.72
1.68–1.79
1.66–1.70
0.78–0.80
0.55–0.56
0.48–0.60
0.53–0.58
0.52–0.62
0.51–0.60
0.53–0.55
1.00
1.01–1.08
0.83–0.97
0.87–0.92
0.92–1.15
0.81–0.86
0.82–0.87
0.83–0.92
0.74–0.81
0.69–0.90
0.64–0.87
1.06–1.15
0.83–0.85
0.87–0.92
0.87–0.92
0.44–0.49
0.42–0.45
0.44–0.45
0.43–0.43
0.43–0.44
0.45–0.47
0.42–0.47
0.52
0.45–0.47
0.55–0.56
0.50–0.51
0.49–0.52
0.50–0.52
0.52–0.52
0.50–0.51
0.51–0.53
0.45–0.53
0.39–0.49
0.55–0.56
0.49–0.50
0.51–0.52
0.53–0.54
1.72–1.88
1.80–1.89
2.27–2.56
2.17–2.22
1.96–2.00
2.22–2.25
2.00–2.11
1.73
1.76–1.80
1.79–1.94
1.96–2.00
1.84–1.97
2.00–2.04
1.85–1.86
1.73–1.79
1.79–2.05
1.93–2.00
2.00–2.08
1.88–2.00
1.84–1.96
2.08–2.42
1.73–1.93
3.00–3.25
4.13–4.40
3.60–3.86
3.25–3.71
3.25–3.64
3.53–3.56
3.43–3.71
1.65
2.00–2.18
1.53–1.58
1.74–1.83
1.93–1.96
1.57–1.58
0.95–1.01
1.33–1.37
2.13–2.25
2.12–2.33
2.18–2.57
1.83–1.86
1.44–1.56
1.67–1.74
1.50–1.57
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
435
77
76
78
0.25mm
1.0mm
75
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.
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).
436
K. B. MILLER
0.5mm
80
79
0.1mm
Figures 79–80. Derovatellus roosevelti. 79, speleum and abdominal sterna V, VI, dorsal aspect; 80, female genitalia, ventral aspect.
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
437
83
82
0.1mm
0.25mm
85
1.0mm
81
84
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.
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 mod-
438
K. B. MILLER
0.5mm
86
87
0.1mm
Figures 86–87. Derovatellus floridanus. 86, speleum and abdominal sterna V, VI, dorsal aspect; 87, female genitalia, ventral aspect.
erately 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
slender, very long, strongly coiled near receptacle;
receptacle moderately large, elongate, curved; intermediate duct relatively slender, long, coiled; sper-
matheca elongate, medially curved, triangular process
relatively small; fertilization duct short, curved, slender (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
439
89
90
0.25mm
92
1.0mm
0.1mm
88
91
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.
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
(FIGS 68–74, 106)
SP. NOV.
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
440
K. B. MILLER
95
0.5mm
93
0.1mm
94
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.
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,
National Xingu.
Mato-Grosso,
Jacare,
Parc
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),
given in Table 1.
Coloration. Head yellow; pronotum yellow with narrow line of dark brown along posterior margin; elytron
441
1.0mm
A
0.25mm
B
99
97
0.25mm
B
A
100
0.25mm
98
96
0.5mm
101
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.
dark brown with indistinct yellow, transverse macula
apicomedially on elytron (Fig. 68); venter yellow, dark
red-yellow on metacoxa; appendages yellow.
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 rela-
442
K. B. MILLER
103
102
0.25mm
0.1mm
104
105
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.
tively 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 nar-
rowly rounded (Fig. 71). Lateral lobe broad basally,
apically narrowed, apex broadened, flattened, apex
pointed (Fig. 72).
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
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
0
0
443
500 Miles
500 KM
Figure 106. Derovatellus species distributions. = D. bruchi, = D. peruanus, = D. roosevelti, = D. spangleri.
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’.
444
K. B. MILLER
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
(FIGS 75–80, 106)
SP. NOV.
‘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.
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).
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.
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.
0
0
445
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
500 Miles
500 KM
Figure 107. Derovatellus floridanus distribution.
446
K. B. MILLER
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:
given in Table 1.
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 set-
ose 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).
macula varies from nearly invisible or only visible
laterally to clearly visible, conspicuously yellow and
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:
examined).
Holotype
in
USNM
(not
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 inter-
447
mediate 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:
given in Table 1.
medially (Fig. 88); lateral bead narrow, distinct along
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 punc-
448
K. B. MILLER
tures 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).
macula varies from nearly invisible or only visible laterally to clearly visible, conspicuously yellow and
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 Anto-
nio, 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:
given in Table 1.
lateral margins broadly curved, pronotum widest near
middle (Fig. 96); lateral bead narrow, distinct along
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,
0
0
449
500 Miles
500 KM
Figure 108. Derovatellus lentus distribution.
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
450
K. B. MILLER
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).
of fine, long setae along mesotrochanter and mesofemur. Male pro- and mesobasotarsomeres I and II
(Fig. 99) moderately expanded compared with female
(Fig. 100).
macula varies from nearly invisible or only visible
laterally to clearly visible, conspicuously yellow and
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 spe-
cies, 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, 1st.
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
451
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).
VATELLUS AUBÉ, 1837
Leucorea Laporte, 1835: 106 (unused name of genus of
Vatellini Sharp, objective synonym of Vatellus Aubé;
suppressed by Opinion 1681 (ICZN, 1992)) (type
species: Vatellus tarsatus Laporte, 1835:106, by
monotypy); Laporte, 1840: 167; Gemminger &
Harold, 1868: 428; Nilsson et al., 1989: 300; Nilsson,
1991: 36 (case for suppression).
Vatellus Aubé, 1837: 218 (type species: Hydroporus
tarsatus Laporte, 1835: 106 by monotypy); Aubé,
1838: 448; Lacordaire, 1854: 414; Gemminger &
Harold, 1868: 428; Régimbart, 1878: 454; Sharp,
1882b: 840; Zimmermann, 1919: 124, 1920: 30; Nilsson et al., 1989: 309; Nilsson, 2001: 235. Conserved
in Opinion 1681 (ICZN, 1992).
Macrovatellus Sharp, 1882b: 282 (type species:
Macrovatellus marginalis Sharp, 1882b: 284 by
subsequent designation of Guignot (1946), not Macrovatellus mexicanus Sharp, 1882b: 284 designated
by Leech (1948)); Sharp, 1882a: 8; Zimmermann,
1919: 125, 1920: 30; Nilsson et al., 1989: 300; Nilsson, 2001: 235; Roughley & Larson, 2001: 169, syn.
nov.
Platydessus Guignot, 1955: 3 (type species: Platydessus perforatus Guignot, 1955: 4 by monotypy); Spangler, 1966a: 57 (synonymized with Macrovatellus);
Nilsson et al., 1989: 305; Nilsson, 2001:235, syn.
nov.
Diagnosis: Members of this genus are differentiable
from the other extant genus of the tribe, Derovatellus,
in part by the combination of: (1) lateral body outline
generally strongly discontinuous, pronotum generally
broadest anterior to middle (e.g. Fig. 1); (2) metacoxal
lines approximated posteriorly, anteriorly moderately
to strongly divergent (e.g. Fig. 176); (3) male lateral
lobes strongly extended in basal portion, elongate,
often with corresponding extension of basal portion of
median lobe (e.g. Figs 182–184); (4) spermatheca subspherical, only slightly elongate in some species (e.g.
Fig. 186); (5) apical margin of ventral apex of orifice
of speleum distinctly, but variably, lobed (e.g.
Fig. 209).
452
K. B. MILLER
Distribution: Members of Vatellus occur in the New
World from southern Texas south throughout Central
and South America to Argentina. Most of its members
occur in lowland, subtropical or tropical wetlands.
apparently a Wehncke specimen since it is labelled
exactly the same as the type of V. haagi Wehncke.
Therefore, I have concluded that it must be the holotype of this species.
Taxonomic history: The oldest name for this genus is
Leucorea Laporte (1834, 1835), but this name was
never used. Instead, Vatellus Aubé became the most
commonly used name for the taxon. Both names
included only the type species Hydroporus tarsatus
Laporte (by monotypy), until now. Eventually, Nilsson
et al. (1989) suggested conservation of the name Vatellus. Nilsson (1991) formally proposed doing so and the
name was conserved (ICZN, 1992). However, evidence
from this project indicates that Vatellus tarsatus is
nested well within a group of species historically
placed in a separate genus, Macrovatellus Sharp,
making the latter genus paraphyletic (Fig. 316). Vatellus was delimited based on the character state of distinctive, abrupt posterior margins on the abdominal
sterna. However, in other respects, V. tarsatus resembles members of Macrovatellus, and together the species are monophyletic sharing several distinctive
synapomorphies (see Cladistic analysis below). Therefore, I have chosen to synonymize the two genera. The
conserved name Vatellus has priority over Macrovatellus. Platydessus was described for the species
V. perforatus (Guignot) and placed in the tribe
Bidessini. However, Spangler (1966b) recognized that
the genus belongs in Vatellini and synonymized the
name with Macrovatellus.
Type locality: Brazil.
Natural history: Most of the members of this group
occur in ponds, marshes and small, slow streams with
considerable dense vegetation.
Discussion: Members of this genus have a number of
external characters useful for distinguishing between
species or groups of species. However, positive identification of species generally requires dissection of male
or female genitalia and examination of the speleum.
VATELLUS
VENTRALIS
(SHARP, 1882)
COMB. NOV.
Macrovatellus ventralis Sharp, 1882b: 285; Zimmermann, 1920: 30; Trémouilles, 1995: 27.
Type information: Holotype (by monotypy) in
MNHN labelled, ‘[small, light purple square]/Haag
[handwritten, black line around label]/Brazilia [handwritten, green label with black line around border]/
HOLOTYPE Macrovatellus ventralis Sharp, 1882 [red
label with black line border].’ Sharp (1882b) had a single specimen that he apparently borrowed from
Wehncke. Wehncke’s specimens are in the MNHN. A
single specimen was found among the MNHN
vatelline material that matches Sharp’s description of
the species (including the absence of tarsi) and is
Diagnosis: This species may be distinguished from
other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces alutaceous, elytra
with punctures and fine microreticulation between
them; (2) eyes moderately large, not strongly protruding (Fig. 109); (3) transverse pronotal crease indistinct; (4) apical lobe on abdominal sternum VI very
large, flat, apically rounded (Fig. 113); (5) anterior clypeal margin produced into distinct anteriorly directed
rim; (6) pronotum prominently cordate, lateral margins strongly sinuate; and (7) ventral surface of dorsal
margin of orifice to speleum with small, but prominent, longitudinal carina, dorsal surface of ventral
margin distinctly bilobed to fit over dorsal carina. This
species is similar in many respects to V. grandis and
similar species because they are elongate, evenly
coloured, dorsoventrally flattened and have the anterior margin of the clypeus distinctly produced. However, in V. ventralis the dorsal surfaces are alutaceous,
the punctation is moderately fine and evenly distributed on the head and pronotum, the anteromedial
margin of the metasternum is not swollen, and the
female receptacle is relatively large. Also, in other species of Vatellus the ventral surface of the dorsal margin of the orifice to the speleum has a cavity that
receives a corresponding prominence on the dorsal
surface of the ventral margin. Vatellus ventralis has a
longitudinal, sharp carina on the ventral margin and a
bilobed prominence on the dorsal margin. The dorsal
lobe is extremely thick and has an apical, posteriorly
directed prominence as well. These characters are
subtle, but they make V. ventralis unique.
Description:
Habitus. Lateral outline strongly discontinuous in
dorsal aspect, lateral elytral margins moderately
rounded (Fig. 109), distinctly dorsoventrally flattened.
Coloration. Head brown; pronotum brown; elytron
brown, immaculate (Fig. 109); venter and appendages
brown.
Sculpture and structure. Head with punctation distinct, small, dense, evenly distributed over entire surface except posterior quarter, interpuncture surfaces
prominently microreticulate, surface posterior to eyes
alutaceous, rough, slightly swollen; anterior clypeal
margin anteriorly produced; eyes moderately large,
somewhat protruding. Pronotum densely punctate,
punctures moderately large, pale setae inconspicuous;
pronotum strongly cordate, lateral pronotal margin
KEY
1.
1¢.
2.
2¢.
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3¢.
4.
4¢.
5.
5¢.
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8.
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11.
11¢.
TO THE SPECIES OF
453
VATELLUS
Anterior clypeal margin anteriorly produced into distinct rim (Fig. 6); anteromedial process of metasternum of
most species ventrally produced into distinct tumidity (Fig. 20) (not produced in V. ventralis); body dorsoventrally
flattened; dorsal surface of pronotum usually with fine shagrination between punctures except for smooth, shiny,
impunctate area medially on each side of midline (e.g. Fig. 262); elytra smooth and shiny between punctures or
with very fine shagrination as on pronotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Anterior clypeal margin not or only weakly produced into rim (e.g. Fig. 7); anteromedial process of metasternum
not ventrally produced; body variable, but many species not dorsoventrally flattened; dorsal surface of pronotum
without distinct pattern of shagrination and punctation; elytra of most species with surface microreticulation in
various forms (Figs 3, 5), or a few species or specimens smooth and shiny between punctures . . . . . . . . . . . . . . .5
Anteromedial process of metasternum not ventrally produced; ventral surface of dorsal margin of orifice of
speleum bilobed with a cavity that receives a corresponding longitudinal, carinate prominence on the dorsal
surface of the ventral margin, dorsal lobe of orifice extremely thick and with an apical, posteriorly directed
prominence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. ventralis (Sharp)
Anteromedial process of metasternum ventrally produced into distinct tumidity (e.g. Fig. 20); ventral surface
of dorsal margin of orifice of speleum relatively thin and not lobed, without ventral prominence . . . . . . . . . . . . .3
Male median lobe of aedeagus very slender, curved and sharply pointed apically (Fig. 293) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. drymetes sp. nov.
Median lobe not slender, only moderately curved and less sharply pointed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Median lobe in lateral aspect with apex narrowed and bent ventrad (Fig. 306) . . . . . . . . . . . . . . V. amae sp. nov.
Median lobe in lateral aspect with apex slender (Fig. 282) or broader (Fig. 269) but not bent ventrad . . . . . . . . .5
Median lobe in lateral aspect with apex slender (Fig. 282); lateral lobe with apex long, acutely pointed and membranous (Fig. 285) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. bifenestratus (Zimmermann)
Median lobe in lateral aspect with apex relatively broad (Fig. 269); lateral lobe with apex narrowly rounded, but
not produced into pointed structure (Fig. 271) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. grandis Buquet
Size very large, TL = 6.5–7.1 mm; median lobe long and nearly straight in lateral aspect (Fig. 125), in ventral
aspect with lateral margins parallel to very near apex (Fig. 126); lateral lobe broad, apex densely covered with fine,
long setae (Fig. 127) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. sahlbergi (Sharp)
Size generally smaller, TL < 6.5 mm (some specimens of V. lateralis with TL > 6.5 mm); median lobe various, but
not straight in lateral aspect, in ventral aspect various; lateral lobe with apical setae, but generally only along
margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Prosternal process with apex truncate, with fringe of fine setae; median lobe in lateral aspect slightly bent dorsally
at apex (Fig. 182), in ventral aspect with apex broad and truncate (Figs 183, 185); female bursa very large, flattened
(Figs 186, 187), dorsoventrally sinuate (Fig. 187) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. haagi Wehncke
Prosternal process with apex pointed; median lobe in lateral aspect various, but not characteristically bent dorsally,
in ventral aspect; female bursa smaller, not dorsoventrally sinuate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
Size relatively small (TL = 4.6–4.8 mm); punctures on pronotum, elytron and metacoxae large and coarse (Figs 204,
205); dorsal surfaces smooth and shiny between punctures . . . . . . . . . . . . . . . . . . . . . . . . . V. perforatus (Guignot)
Size generally larger (TL = 4.6–7.3 mm); punctures on pronotum, elytron and metacoxa smaller and obscured; dorsal surfaces of most specimens with fine alutaceous microreticulation between punctures . . . . . . . . . . . . . . . . . . .9
Apex of median lobe in lateral aspect distinctly pointed and curved ventrally (Figs 225, 239, 242), in ventral aspect,
apex acuminate (Figs 224, 226, 240, 241) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
Apex of median lobe not pointed and curved ventrally, not acuminate in ventral aspect . . . . . . . . . . . . . . . . . . . .11
Posterior margins of abdominal sterna 2, 3 and 4 strongly demarcated in most specimens, these margins ending
abruptly and rising steeply to the surface of the next segment; apical slender portion of median lobe relatively
shorter (Fig. 225); intermediate duct between receptacle and spermatheca not tightly coiled, broad (Fig. 229);
speleum very slender (Fig. 223) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. tarsatus (Laporte)
Posterior margins of abdominal sterna not strongly demarcated; apical slender portion of median lobe relatively
longer (Fig. 239); intermediate duct between receptacle and spermatheca long, tightly coiled (Fig. 244); speleum
broad (Fig. 273) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. annae sp. nov.
Dorsal surfaces nearly concolorous, brown; apical lobe at ventral opening of speleum on abdominal segment six
long, with apex nearly spherical (Fig. 251); intermediate duct between receptacle and spermatheca very long,
tightly coiled (Fig. 258) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. pilacaudus sp. nov.
Elytra and, generally, pronotum brown with distinctive pattern of pale maculae; apical lobe at ventral opening of
speleum on abdominal segment six less developed, generally triangular and flattened . . . . . . . . . . . . . . . . . . . . .12
454
K. B. MILLER
12. Speleum with apex large, expanded, strongly hooked dorsally (Fig. 196); austral South America (Fig. 314). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. wheeleri sp. nov.
12¢. Speleum with apex smaller, much less expanded, not strongly hooked. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Median lobe with apex broad, subapically slightly constricted, apex broadly rounded (Fig. 155); female bursa broad
with apex irregularly grooved and with numerous pores (Fig. 159) . . . . . . . . . . . . . . . . . . . V. maculosus sp. nov.
13¢. Median lobe with apex more slender, not subapically distinctly constricted, apex more narrowly rounded; female
bursa not so broad and with apex not irregularly grooved and without numerous pores . . . . . . . . . . . . . . . . . . . 14
14. Median lobe with apex slender, evenly convergent to narrow point (Fig. 170), lateral lobes not extending to apex of
median lobe (Fig. 139); female intermediate duct between receptacle and spermatheca sinuate (Fig. 173); central
South America (Fig. 311) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lateralis (Sharp)
14¢. Median lobe with apex moderately broad, slightly expanded subapically, apex slightly curved dorsad, moderately
broadly rounded (Figs 141, 144); lateral lobes extending distinctly beyond apex of median lobe (Fig. 140); female
intermediate duct between receptacle and spermatheca not sinuate (Fig. 146); extreme southern Texas south
throughout Mexico to Costa Rica (Fig. 312) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .V. mexicanus (Sharp)
rounded anteriorly, broadest near anterior angles, posterior angles acute (Fig. 109); lateral bead obscured;
transverse sulcus slightly impressed; surface alutaceous. Elytron with punctation moderately dense,
punctures moderately large, interpuncture surfaces
prominently alutaceous, pale setae inconspicuous,
surface similar to pronotum (Fig. 111). Prosternum
short, roughly sculptured, glabrous; prosternal process broad with distinctly rounded medial carina, lateral margins rounded, apex moderately triangularly
produced, glabrous, apex obtusely pointed. Metasternum with punctation fine and sparse, obscured by
alutaceous surface sculpture; anteromedial margin
not produced ventrally. Abdominal sterna sutures not
unusually modified; sterna with punctation sparse
and fine, obscured by surface sculpture; sixth abdominal sternum with apical lobe very large, broad, flattened, apically rounded (Fig. 113); speleum broad
basally, narrowed and parallel sided, apically narrowly rounded (Fig. 114). Metacoxae with lateral portion moderately punctate over entire surface,
punctures shallow, somewhat obscured by alutaceous
sculpturing; metacoxal lines approximate posteriorly,
strongly divergent anteriorly (Fig. 110); medial
portion moderately punctate, punctures finely setose.
Metatrochanter rounded, strongly offset (Fig. 112);
metatarsal claws dentate.
Female genitalia. Gonocoxosternite broad, robust,
medial margin broad, apical angle about 90∞; anterior
margin concave; anterior lobe moderately long, narrow (Fig. 115). Gonocoxa moderately broad, subtriangular, apodeme moderately narrow, longer than main
portion, numerous setae along apicoventral margin
and on dorsal surface (Fig. 115). Bursa copulatrix
moderately small, with two anterior lobes, one on each
side; spermathecal duct relatively long, slender; receptacle spherical, slightly smaller than spermatheca;
intermediate duct short, sinuate (Fig. 116); spermath-
eca nearly spherical, triangular process broad; fertilization duct short, curved (Fig. 115).
Distribution: This species is known only from the type
locality, ‘Brazil’.
Phylogenetic relationships: This species occupies an
unresolved, basal position within Vatellus (Fig. 316).
The lack of information about male character states
undoubtedly contributes to its alternative placements
in the most parsimonious trees, but it also lacks some
of the more distinctive characters used to group taxa
within Vatellus.
Material examined: Only the holotype was examined
of this species.
VATELLUS
SAHLBERGI
(SHARP, 1882)
COMB. NOV.
Macrovatellus sahlbergi Sharp, 1882b: 283; Zimmermann (1920): 30; Guignot, 1957: 3; Trémouilles,
1995: 27.
Type information: Lectotype (designated here to clarify assignment of this name with this species) in
BMNH labelled, ‘Type [circular disc with red border]/
440 [handwritten]/S.America: Brazil. [with green line
transversely across middle of label]/Petropolis, March
Sahlberg. 1850 [handwritten]/Petropolis March 1850
Sahlberg. [handwritten]/Sharp Coll 1905–313./Type
440 Macrovatellus Sahlbergi Petropolis. [handwritten]/SYN- TYPE [circular disc with blue border]/LECTOTYPE Macrovatellus sahlbergi Sharp 1882. des.
K.B. Miller 2002 [red label with black line border].’
Sharp had two specimens upon which he based his
description, and a single paralectotype with similar
label information accompanies the lectotype in
BMNH.
Type locality: Brazil, Petropolis.
455
1.0mm
0.5mm
0.5mm
110
112
109
0.5mm
111
0.5mm
113
114
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.
Diagnosis: This species is very large and has the outline of the body strongly discontinuous between the
pronotum and elytra in dorsal aspect. In lateral aspect
the median lobe is long, slender, straight and apically
abruptly narrowed with a minute, apical point
(Fig. 125). In ventral aspect the median lobe is long
and slender with the lateral margins parallel-sided
and with the apex nearly truncate and with a small,
medial process (Figs 126, 128). The lateral lobe is
broad and bears a large field of dense setae apically
(Fig. 127).
Description:
Habitus. Body outline strongly discontinuous in dorsal aspect, lateral margins of elytra strongly rounded
(Fig. 117), dorsoventrally not compressed. Measurements are given in Table 1.
Coloration. Head dark red; pronotum dark red,
nearly black; elytron dark red with apical, lateral and
humeral small, pale maculae (Fig. 117); venter and
appendages dark red.
Sculpture and structure. Head finely and evenly
punctate, punctures slightly finer anteriorly, surface
456
K. B. MILLER
116
115
0.1mm
Figures 115–116. Vatellus ventralis. 115, female genitalia, ventral aspect; 116, spermatheca and associated structures,
dorsal aspect.
shiny and smooth between punctures; anterior clypeal
margin swollen and rounded, not anteriorly produced;
eyes moderately sized and somewhat protuberant
(Fig. 117). Pronotum medially densely punctate, punctures moderately sized, setae short, inconspicuous,
surface between punctures alutaceous, conspicuously
rough, partially obscuring punctures, laterally punctures nearly entirely obscured; pronotum somewhat
cordate, very broad, broadest slightly anterad of
middle, margins strongly rounded, posterior angles
obtusely angled (Fig. 117); lateral bead obscured;
transverse sulcus present, distinct, pronotum slightly
swollen medially. Elytron densely punctate, punctures
moderately coarse, covering entire surface, strongly
obscured apically, setae short, inconspicuous, surface
between punctures alutaceous similar to pronotum
(Fig. 118); with indistinct, longitudinal raised line
medially on elytron. Prosternum short, rough, glabrous; prosternal process relatively narrow, medially
tectiform, apex narrowly triangular, pointed, flattened. Metasternum rugulose and alutaceous, puncta-
tion entirely obscured and not evident; anteromedial
margin laterally somewhat compressed, but not ventrally expanded. Abdominal sterna with surface irregularly rugulose and alutaceous, punctation obscured;
sixth sternum with apical lobe broad, apex broadly
rounded; speleum long, relatively slender, apex narrowly rounded. Metacoxa with medial portion with
surface conspicuously rugose and alutaceous, punctation fine; metacoxal lines approximated basally,
broadly divergent anteriorly (Fig. 119); medial portion
rugulose, finely punctate. Metatrochanter moderately
elongate and distinctly offset, apex rounded (Fig. 120);
metatarsal claws finely pectinate.
Male genitalia. Median lobe in lateral aspect relatively straight throughout length, lateral margins parallel, basal portion short, narrow; apically abruptly
narrowed to sharply curved apex, without setae
(Fig. 125); in ventral aspect with lateral margins parallel, apex truncate with narrow medial spinous or
truncate process (Figs 126, 128). Lateral lobe with apical portion very broad, apex broadly rounded, with
457
121
0.5mm
1.0mm
A
B
118
0.5mm
0.5mm
119
A B
122
120
117
0.5mm
123
124
0.5mm
Figures 117–124. Vatellus sahlbergi. 117, dorsal habitus; 118, elytral sculpture in area indicated on habitus by dotted
square; 119, medial portion of metacoxae; 120, metatrochanter and metafemur, anterior aspect; 121–122, tarsomeres, dorsal aspect; 121, male; 122, female, A, protarsus; B, mesotarsus; 123, sternum VI, ventral aspect; 124, speleum abdominal
sterna V, VI, dorsal aspect.
moderately dense region of fine setae apically and
along apicodorsal margin; long, stout, subapical spine
absent (Fig. 127).
broad, apical angle rounded, anterior margin curved,
anterior lobe long, relatively slender, posterior margin
with many stout setae (Fig. 129). Gonocoxa long, apex
rounded, apodeme long and relatively slender
(Fig. 129). Bursa copulatrix broad, relatively large,
with anterior thickened region around orifice of spermathecal duct (Fig. 129); spermathecal duct long,
slender; receptacle large, subequal in size to spermatheca, subspherical (Fig. 129); intermediate duct
moderately long, sinuate near spermatheca (Fig. 130);
spermatheca subspherical, large; fertilization duct
moderately long, slender (Fig. 129).
Sexually dimorphic characters. With brushes of
setae on mesotrochanter and base of mesofemur well
458
K. B. MILLER
0.5mm
126
127
0.1mm
125
128
130
129
0.1mm
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.
developed, brush on mesofemur forming an elongate
cup-shape. Male with pro- and mesobasotarsomeres
(Fig. 121) broader and more expanded than in female
(Fig. 122).
Distribution: This relatively rare species is known
from southern Brazil and Paraguay (Fig. 311). This
species was recorded from Bolivia by Guignot (1957),
although the identification may be in error.
Phylogenetic relationships: As with V. ventralis, this
species occupies an unresolved, ambiguous position
basally within the genus (Fig. 316). Unlike that species, both males and females are known, but the spe-
cies lacks some of the more distinctive characters used
to group taxa within Vatellus. In the most parsimonious trees, the species is generally associated with the
V. mexicanus clade either as sister to it in a clade or
with it as a paraphyletic grade with it as sister to the
remaining Vatellus (Figs 320–324).
Material examined: Brazil: Country only (1, USNM);
Petropolis, March 1850 (1, BMNH). Paraguay:
Hohenau. Alto-Parana, 28 August, H Jacob (3,
BMNH).
VATELLUS
MEXICANUS
(SHARP, 1882)
COMB. NOV.
Macrovatellus mexicanus Sharp, 1882b: 284; 1882a: 8;
Zimmermann, 1920: 30; Leng, 1920: 76.
Type information: Holotype (by monotypy) in
BMNH labelled, ‘ Mexico [handwritten on card with
specimen]/Type [circular disc with red border]/Sharp
Coll. 1905.-313./Type 442 mexicanus Mexico [handwritten]/B.C.A. Col. I. 2. Macrovatellus mexicanus,
Sharp. [black line along top and bottom of label]/Holotype [circular disc with red border]/HOLOTYPE Macrovatellus mexicanus Sharp 1882 [red label with black
line border].’ Two additional Sharp specimens accompany the holotype, but are not of the original type
series as Sharp (1882b) indicated he had only the single male specimen. He included these other two specimens in his treatment of Central America (Sharp,
1882a).
Type locality: Mexico.
other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces distinctly alutaceous (Fig. 5); (2) eyes large, protruding (Fig. 131); (3)
transverse pronotal crease distinctive (Fig. 131); (4)
male pro- and mesobasotarsomeres (Fig. 135) much
broader than in female (Fig. 136); (5) apical lobe on
abdominal sternum VI present but not strongly
expressed (Fig. 137); (6) male and female genitalia distinct (Figs 140–146). This species is very similar to
V. lateralis from which it is distinguishable by the
median lobe broader and apically not deflexed or as
strongly pointed. In addition, the intermediate duct
between the spermatheca and receptacle is shorter
and not strongly sinuate. The lateral lobes end well
short of the apex of the median lobe in V. lateralis
(Fig. 139) whereas in V. mexicanus the apices of the
lateral lobes extend to or beyond the apex of the
median lobe (Fig. 140).
Description:
Habitus. Lateral outline strongly discontinuous in
dorsal aspect, lateral elytral margins strongly
rounded (Fig. 131), not dorsoventrally flattened. Measurements are given in Table 1.
459
Coloration. Head yellow; pronotum brown, irregularly yellow along anterior and lateral margins;
elytron dark brown to yellow-brown with yellow maculae subapically and laterally along margins, basally
with few yellow maculae (Fig. 131); venter and
appendages red brown to light brown, infuscate, especially laterally on metacoxae.
Sculpture and structure. Head with punctation distinct, variable in size, coarse medially between eyes,
becoming finer anteriorly, interpuncture surfaces
smooth, shiny to slightly microreticulate, surface posterior to eyes alutaceous, rough; anterior clypeal
margin rounded; eyes large, strongly protruding.
Pronotum densely punctate, but punctures strongly
obscured by rough, alutaceous sculpturing, surface as
in Figure 5, pale setae moderately long; pronotum cordate, lateral pronotal margin rounded anteriorly,
broadest near anterior angles, posterior angles acute;
lateral bead obscured; transverse sulcus distinctly
impressed. Elytron with punctation dense but edges
of punctures strongly obscured by dense alutaceous
sculpturing as on pronotum (Fig. 132), light setae
moderately dense and conspicuous. Prosternum short,
roughly sculptured, glabrous; prosternal process
broad with distinctly rounded medial carina, lateral
margins rounded, apex triangularly produced, glabrous, apex pointed. Metasternum with punctation
dense laterally, punctures large and shallow, somewhat obscured by alutaceous sculpturing; anteromedial margin slightly produced ventrally. Abdominal
sternal sutures not unusually modified; sterna with
punctation dense over entire lateral surface, finer
medially, obscured by alutaceous sculpturing; sixth
abdominal sternum with apical lobe inconspicuously
produced, triangular, not strongly produced; speleum
broad basally, sides strongly convergent, apex moderately rounded, not strongly curved. Metacoxae with
lateral portion densely punctate over entire surface,
sculpturing; metacoxal lines strongly approximate
posteriorly, strongly divergent anteriorly; medial portion moderately punctate, punctures finely setose.
Metatrochanter rounded, offset; metatarsal claws
dentate.
Male genitalia. Median lobe in lateral aspect evenly
but not strongly curved, relatively narrow, narrowed
subapically, apex slightly curved dorsally and narrowly rounded, basal portion elongate, slender, without subapical spines (Figs 141, 144); in ventral aspect
narrow apically, broader basally, lateral margins convergent, apex narrowly rounded (Fig. 142). Lateral
lobe broad to broadly rounded apex, with large region
of fine, dense, elongate setae; stout subapical setae
absent (Fig. 143).
medial margin very broad, apical angle about 90∞;
460
K. B. MILLER
A
1.0mm
0.5mm
B
135
132
0.5mm
0.5mm
133
B
A
136
134
131
0.5mm
0.5mm
137
138
Figures 131–138. Vatellus mexicanus. 131, dorsal habitus; 132, elytral sculpture in area indicated on habitus by dotted
anterior margin concave; anterior lobe moderately
long, moderately broad (Fig. 145). Gonocoxa very
broad, about as long as wide, apex obtusely rounded,
apodeme moderately narrow, about as long as main
portion, numerous setae along apicoventral margin
and on dorsal surface, ventral surface with large
region of semicircular striae with very short, spinous,
adpressed setae (Fig. 145). Bursa copulatrix moderate
in size, with four distinct, finger-like lobes; spermathecal duct relatively short, relatively narrow; receptacle spherical, nearly equal in size to spermatheca
(Fig. 145); intermediate duct moderately long, relatively straight (Fig. 146); spermatheca nearly spheri-
cal, triangular process very broad; fertilization duct
short, curved (Fig. 145).
Sexually dimorphic characters. Male with poorly
developed
mesotrochanteric
and
mesofemoral
brushes of setae, broad with lateral portions longer
forming an elongate cup-shaped structure. Male with
protarsomeres 1–3 (Fig. 135A) substantially broader
than in female (Fig. 136A), protarsomere III distinctly shorter than in female; mesotarsomeres of
male (Fig. 135B) broader than in female (Fig. 136B),
but not conspicuously so; male mesotarsomere III
(Fig. 135B) not much longer than mesotarsomere III
(Fig. 136B).
461
142
141
143
140
139
0.5mm
0.1mm
146
144
145
0.1mm
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.
Intraspecific variation. This species varies significantly in size. A series from Belize (USNM) is
nearly entirely black with the maculae nearly
absent.
Natural history: Vatellus mexicanus has been collected from black lights. It has also been collected from
a variety of habitats including dry tropical forest, clear
pools in a stream bed, pools in a temporary stream, a
462
K. B. MILLER
shallow pool in a gravel pit and a stock pond. In Texas
the species occurs in permanent or temporary ponds
with grassy margins (Jasper & Challet, 2002).
Distribution: This species is widespread along the
western and eastern low areas of Mexico and is also
known from Belize and Costa Rica (Fig. 312).
Although Leech & Chandler (1956) speculated the
species may occur in southern California, it has not
been collected there though it is known from the
northern Baja Peninsula and northern Sonora.
Recently, V. mexicanus was reported from Dimmit,
LaSalle and Zapata Counties, Texas, USA (Jasper &
Challet, 2002).
Phylogenetic relationships: This species is sister to a
clade of several very similar species (Fig. 316). These
species are similar in overall shape, the presence of
distinct maculations, similar cuticular microsculpturing (Fig. 5) and shape of the male median lobe.
Material examined: Belize: Cayo Dist. San Ignacio
(8 km S), 23 May 1986, stock pond, P.J. Spangler, R.A.
Faitoute (30, USNM). Costa Rica: 16 mi S La Cruz, 13
July 1965, PJ Spangler (6, USNM); Finca Jenny,
30 km N de Liberia, Prov Guana, 240 m, 14 April
1993, E Araya (30, JBWM); Guanac. Est. Sta. Rosa,
300 m LN 313000, 359300, black light in dry tropical
forest, D.A. Pollock (2, LHIC); Guanacaste, Palo Verde
Nat Park, 10∞20¢43≤N 85∞16¢24≤W, 8 June 2000, shallow pool, gravel pit, RE Roughley (3, JBWM; Playa
Naranjo, Sta Rosa, PN Guanacaste Prov Guan, March
1991, E Alcazar (13, JBWM); Santa Rosa, 1 March
1993, K.G. Bernhardt (5, LHIC); Santa Rosa N P,
Guanacaste, 23 May 1980, DH Janzen and W Hallwachs (1, JBWM); Sector Cerro Cocori, Fca. De E Rojas,
150 m, Prov. Lamon, March 1993, E Rojas (1, JBWM).
Mexico: Baja Calif. Sur, San Bartolo Dam, 1 May
1947, I LaRivers (1, USNM); Baja California, Gulf of
California, Isla del Carmen, 19–20 July 1984, SE
Miller (1, USNM); Baja California Sur, 3 mi W Agua
Caliente, 22 August 1997, WD Shepard (1, LHIC);
Campeche 21 mi E Campeche, 27 July 1964, P.J.
Spangler (2, USNM); Chis. 4 mi W Cuahtemoe, 30
August 1963, JR Zimmermann (1, USNM); Chis. 5 mi
S Las Cruces, 23 August 1965, PJ Spangler (1,
USNM); Chis. Chipa de Corzo, 4 mi E, 1 November
1963, JR Zimmermann (1, USNM); E. of Chiapas, 5
August 1957, D. Lauck (16, USNM); Jalapa, 10 mi E,
27 August 1962, JR Zimmerman (1, USNM); Jalsico,
La Huerta, 3.5 mi N, 22 March 1971, JR Zimmerman
(1, USNM); Lower California, San Felipe, 1926, JD
Sherman (1, USNM); Morelus, nr Tigolna, 22 June
1963, R Woodruff (5, FSCA); Nayarit San Blas, 26
July 1963, PJ Spangler (48, USNM); Nayarit, San
Blas, 5 mi E, 31 July 1962, JR Zimmerman (1,
USNM); Nuevo Leo, Puente Critsolinas, 17 mi S
Monterey, 19 July 1969, FN Young (1, FSCA); Oaxaca,
Juchitan, 3 mi E, 7 September 1964, JR Zimmerman
(1, JBWM); Oaxaca, Juchitan, 3 mi E, 7 September
1964, JR Zimmerman (1, USNM); Oaxaca, Salina
Cruz, 6 September 1964, JR Zimmerman (1, USNM);
Oaxaca, Tehuantepec, 2 September 1963, JR Zimmerman (4, USNM); Oaxaca, Tehuantepec, 11 June 1964,
JC and D Pallister (2, AMNH); Oaxaca, Zanatepec, 22
July 1964, P.J. Spangler (14, USNM); Oaxaca, Zanatepec 8 km E, 23 May 1981, pools in temp stream, PJ
Spangler (1, USNM); Puebla, nr Maria Andres, 10
September 1964, JR Zimmerman (1, USNM); San
Luis Potosi, 12 mi S Cd. Mante, 21 June 1975, CA Triplehorn (1, FSCA); San Luis Potosi, 15 mi E C del
Maiz, 19 November 1948, HB Leech (1, FSCA); San
Luis Potosi, 29 mi N Valley, 19 August 1954, FN
Young (1, FSCA); San Luis Potosi, Antiguo Morelos,
23 March 1963, JR Zimmerman (1, USNM); San Luis
Potosi, Ciudad Valles, 22 August 1964, PJ Spangler
(18, USNM); San Luis Potosi, Comcoa, Rio Axtla, 23
March 1963, JR Zimmerman (1, USNM); San Luis
Potosi, El Salto Falls, 14 June 1963, black light, RE
Woodruff (1, FSCA); Sinaloa, 54.0 mi S Culiacan, 23
April 1969, 540¢, ME Irwin (1, USNM); Sinaloa, 7 mi
N Rosario, 24 July 1963, PJ Spangler (5, USNM);
Sinaloa, Concordia, 8 mi E R15, 12 December 1962,
JR Zimmerman (1, USNM); Sinaloa, Culiacan, 21
July 1959, at light, 250¢, HE Evans (1, FSCA);
Sinaloa, Elota, 1 mi S, 01 August 1962, JR Zimmerman (1, USNM); Sinaloa, pools in stream, 6 mi w El
Limon, 21 June 1965, FN Young (1, FSCA); Sonora,
13 m SE Alamos, 30 October 1972, K Stephens (3,
FSCA); Sonora, 20 mi W Moctequma, 28 September
1980, RD Gordon (1, USNM); Sonora, Alamos, 15 July
1963, PJ Spangler (2, USNM); Sonora, Alamos, 17
September 1980, R. Gordon (1, USNM); Sonora,
Moctezuma 14 mi W, 27 September 1974, clear pools
in stream bed, JR Zimmerman (1, USNM); Sonora,
Rio Moccsorito Arroyo Huerta, 24 June 1971, R Vijenhoes (1, FSCA); Tamaulipas, 4 mi N Ciudad Mante, 3
September 1964, DR Whitehead (1, USNM); Tamaulipas, Limon Rio Frio on Nat. Hgw., 18 December 1940,
FN Young (20, FSCA); Tamaulipas, Limon, Rio Frio on
Nat Hgw, 18 December 1940, FN Young (1, USNM);
Tamaulipas, Llera, 20 mi S, 24 March 1963, JR Zimmerman (1, USNM); Tamaulipas, Nuevo Morelos, 28
August 1965, PJ Spangler (1, USNM); Tamaulipas,
Rio Guayalejo nr Magiscatzin, 11 June 1960, FN
Young (6, FSCA); Tamaulipas, San Jose, April 1910,
JD Sherman (1, USNM); Tehuantepec, A Boucard (4,
MNHN); Vera Cruz, Puente Nacional, Flint and Ortiz
(2, USNM); Veracruz, 15 mi SE Tantoyuca, 28 August
1965, PJ Spangler (6, USNM); Veracruz, Cuitlahuac,
10–12 August 1964, PJ Spangler (75, USNM). United
States: (all from Jasper & Challet, 2002), Texas, Dimmit Co., Chaparral WMA, pond in SW Plot 3 W, Rat-
tlesnake Hill, 25 March 2000, SK Jasper (9, TAMU);
Chaparral WMA, Seco Pond, 26 March 2000, SK Jasper (3, TAMU); LaSalle Co., Chaparral WMA, pon in
SW Plot 14, Rosindo Laguna, 25 March 2000, SK Jasper (1, TAMU); LaSalle Co., Chaparral WMA, Scotty
Pond, 26 March 2000, SK Jasper (1, TAMU); Zapata
Co., Stream at US83, 5.4 mi N TX16, 18 June 1999,
SK Jasper (6, TAMU).
VATELLUS MACULOSUS MILLER
(FIGS 147–160, 311)
SP. NOV.
‘VENEZUELA Guar.,15Km.S. Calabozo II-9-13-1969
P. & P.Spangler/Collected in Lago deLosPatos/HOLOTYPE Vatellus maculosus Miller 2004 [red label with
double black line border].’
Colombia: Meta, 10 km S Villavicencio, P and P. Spangler (3, USNM). Guyana: Rupununi R. Wichabai, 23
April 1961, shady pool, cut off from river in dry season,
stony bottom with much leaf debris; < 1 ft deep, 10 yds
diam, RH McConnel (2, BMNH). Venezuela: Guarico,
15 km S Calabozo, Lago de los Patos, P and P. Spangler (2, USNM).
Type locality: Venezuela, Guarico, 15 km S Calabozo,
Lago de los Patos.
A
0.5mm
463
1.0mm
B
148
151
0.5mm
152
0.5mm
149
B
A
150
147
0.5mm
0.5mm
153
154
Figures 147–154. Vatellus maculosus. 147, dorsal habitus; 148, elytral sculpture in area indicated on habitus by dotted
464
K. B. MILLER
other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces distinctly alutaceous, punctures obscured, elytral surface sculpture
as in Figure 5; (2) eyes large, protruding (Fig. 147); (3)
transverse pronotal crease distinctive (Fig. 147); (4)
male pro- and mesobasotarsomeres (Fig. 151) much
broader than in female (Fig. 152); (5) apical lobe on
abdominal sternum VI present but not strongly
expressed (Fig. 153); (6) male and female genitalia distinct (Figs 155–160). This species is mainly separable
from similar species, V. mexicanus, V. lateralis and
other similarly alutaceous species, by the shape of the
apex of the male median lobe, which is rounded and
distinctly constricted subapically (Fig. 155).
Description:
(Fig. 147), not dorsoventrally compressed. Measurements are given in Table 1.
Coloration. Head yellow; pronotum brown medially
and along anterior and posterior margins, yellow to yellow-brown laterally; elytron dark brown with lateral
and apical yellow maculae, basally with broad, irregular yellow band on most specimens, some with basal
band reduced to few, small maculae (Fig. 147); venter
yellow to yellow-brown, darker on mesocoxae and
metasternum; appendages yellow to yellow-brown.
Sculpture and structure. Head densely and finely
punctate medially, nearly impunctate anteriorly on
frons and clypeus, surface between punctures very
finely microreticulate, shiny; anterior clypeal margin
rounded; eyes moderately large, protruding. Pronotum
densely punctate, punctures shallow and obscured by
microreticulation and alutaceous, rough surface
microsculpture, short, fine, pale setae present; pronotum broadest submedially, not strongly cordate, lateral margin only shallowly rounded along anterior
half, posterior angles slightly obtuse to right-angled
(Fig. 147); lateral bead obscured; transverse sulcus
distinct, mainly laterally (Fig. 147). Elytron finely and
densely punctate (Fig. 148), punctures shallow and
obscured by alutaceous, rough surface as on pronotum, punctures even more obscured apically, each
puncture with short, fine, pale seta. Prosternum
rough, glabrous; prosternal process broad, flat, lateral
margins rounded, apex broadly pointed, glabrous.
Metasternum finely punctate, punctures obscured by
alutaceous surface texture; anteromedial margin not
swollen, pointed medially. Abdominal sterna finely
punctate over entire surface, punctures nearly obliterated by alutaceous surface texture; sixth sternum
with apical lobe well developed, flattened, apically
rounded (Fig. 153); speleum moderately broad, basally
narrowed to narrowly rounded apex, slightly curved
dorsally at apex (Fig. 154). Metacoxa with lateral portion very finely punctate, punctures nearly obliterated
by alutaceous surface texture; metacoxal lines only
moderately approximated posteriorly, broadly divergent anteriorly (Fig. 149); medial portion alutaceous.
Metatrochanter moderately rounded, somewhat
elongate and offset (Fig. 150); metatarsal claws finely
dentate.
slightly curved throughout length, medially expanded
with dorsal margin less strongly curved than ventral
margin, basal portion very long, narrow; apex constricted slightly subapically (Fig. 155), apex rounded,
with numerous minute spinous setae subapically
(Fig. 158); in ventral aspect broader in basal half, narrowed medially and relatively evenly narrowed to
pointed apex (Figs 156, 158). Lateral lobe broad, apex
relatively straight, apex slightly expanded, apically
very broadly rounded, with very dense region of setae
at apex (Fig. 157).
broad, apical angle acutely rounded, anterior margin
somewhat concave, anterior lobe long, broad, apically
broadly rounded (Fig. 159). Gonocoxa elongate, slender, apex moderately narrowly rounded, apodeme
long and slender (Fig. 159). Bursa copulatrix very
broad, irregularly punctate and striate anteriorly
(Fig. 159); spermathecal duct long, slender; receptacle
spherical, approximately half size of spermatheca;
intermediate duct short, relatively straight (Fig. 160);
spermatheca moderately elongate, curved, anterior
portion subspherical and large, posterior portion
narrowed posteriorly; fertilization duct very short
(Fig. 159).
Sexually dimorphic characters. Brushes of setae on
mesotrochanter and mesofemur of male well-developed, that of mesofemur elongate and cup-shaped in
form. Male with pro- and mesobasotarsomeres
(Fig. 151) very distinctly broader than in female
(Fig. 152); female protarsomere III (Fig. 152A) distinctly longer than in male (Fig. 151A).
Intraspecific variation. This species varies in the
extent of pale maculation on the elytron and pronotum. In most specimens the pronotum is nearly
entirely pale and the elytron has a broad transverse
subbasal band of yellow, but in at least one of the specimens studied the pronotum is nearly entirely dark
brown and the basal elytral macula is restricted to the
lateral margin.
Natural history: Vatellus maculosus has been collected from a shady pool with a mineral substrate and
much leaf debris.
Distribution: This is a northern South American species with specimens known from Colombia, Guyana
and Venezuela (Fig. 311).
465
0.5mm
156
157
155
0.1mm
160
158
0.1mm
159
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.
466
K. B. MILLER
Phylogenetic relationships: This species is part of the
clade containing species similar to V. mexicanus
(Fig. 316). However, within this group this species
does not have clear affinities with any other particular
species.
Etymology: This species is named maculosus, Latin
for ‘spotted’, in reference to the maculate elytral
coloration.
VATELLUS
LATERALIS
(SHARP, 1882)
COMB. NOV.
Macrovatellus lateralis Sharp, 1882b: 283; Zimmermann, 1920: 30; Trémouilles, 1995: 27.
Macrovatellus rudis Sharp, 1882b: 283; Zimmermann,
1920: 30; Trémouilles, 1995: 27, syn. nov.
Macrovatellus marginalis Sharp, 1882b: 284; Zimmermann, 1920: 30; Trémouilles, 1995: 27, syn. nov.
Macrovatellus deplanatus Zimmermann, 1919: 125;
1920: 30; Trémouilles, 1995: 27, syn. nov.
Type information: Macrovatellus lateralis: Lectotype
(designated here to clarify assignment of this name
with this species) in BMNH labelled, ‘Type [circular
disc with red border]/Uruguay. [green line transversely
across middle of label]/Sharp Coll 1905–313/Type 438G
Macrovatellus lateralis Uruguay 11.5. [handwritten]/
SYN- TYPE [circular disc with blue border]/LECTOTYPE Macrovatellus lateralis Sharp, 1882. des. K.B.
Miller 2002 [red label with black line border].’ There is
a single paralectotype in BMNH labelled, ‘Type 438 E
[handwritten]/Para-type [circular disc with yellow border]/Uruguay. [green line transversely across middle of
label]/Sharp Coll 1905–313/Macrovatellus lateralis,
Sharp Paratype. [handwritten]/SYN- TYPE [circular
disc with blue border]/PARALECTOTYPE Macrovatellus lateralis Sharp 1882. des. K.B. Miller 2002 [blue
label with black line border].’
Macrovatellus rudis: Lectotype (designated here to
clarify assignment of this name with this species) in
BMNH labelled, ‘Type [circular disc with red border]/
S.America. Colombia. [green line transversely across
middle of label]/Sharp Coll 1905–313./Type 430 Macrovatellus rudis Am.Mer. [handwritten]/SYNTYPE [circular disc with blue border]/LECTOTYPE
Macrovatellus rudis Sharp 1882. des. K.B. Miller 2002
[red label with black line border].’ Sharp indicated
that he had multiple specimens (both males and
females) when he described this species. However,
only a single male in BMNH was examined that
appears to be of the original syntype series. It is
selected as the lectotype.
Macrovatellus marginalis: Lectotype (designated
here to clarify assignment of this name with this species) in BMNH labelled, ‘Type [circular disc with red
border]/S. America: Brazil. [green line transversely
across middle of label]/Santa Rita Sep Sahlberg 1850
[handwritten]/Type 441 Macrovatellus marginalis
Santa Rita [handwritten]/SYN- TYPE [circular disc
with blue border].’ Sharp had an additional female
specimen. Of the two specimens in BMNH labelled as
this species and from Sharp’s collection, only one is a
female with label information that would indicate it is
one of the two original syntypes. Thus it is selected as
the lectotype. The other male specimen is probably not
of the original type series.
Macrovatellus deplanatus: The type of this species is
apparently not in MNHU (M. Uhlig, pers. comm.) nor
in ZSBS (M. Baehr, pers. comm.) (the latter the location of most of Zimmermann’s types). The type locality
and Zimmermann’s description (1919) strongly suggest that the species is the same as the one treated
here as V. lateralis. Two specimens of V. lateralis in
ZSBS labelled as ‘M. ?deplanatus’ (clearly not the
types as they are labelled as having been collected in
Venezuela) were identified by Guignot, suggesting
that he regarded V. lateralis and M. deplanatus as
similar, at least. Based on these lines of evidence, I
have synonymized this name with V. lateralis. However, until the types are found, this synonymy should
be regarded as provisory.
Type locality: Macrovatellus lateralis: Uruguay.
Macrovatellus rudis: South America.
Macrovatellus marginalis: Brazil, Santa Rita.
Macrovatellus deplanatus: Brazil, Santos.
other Vatellus by the following combination of characters: (1) dorsal and ventral surfaces distinctly alutaceous, elytral surface sculpture as in Figure 5; (2) eyes
large, protruding (Fig. 161); (3) transverse pronotal
crease distinctive (Fig. 161); (4) male pro- and mesobasotarsomeres (Fig. 165) much broader than in
female (Fig. 166); (5) apical lobe on abdominal sternum VI present but not strongly expressed (Fig. 167);
(6) male and female genitalia distinctive (Figs 169–
173). This species is very similar to V. mexicanus from
which is distinguishable by the median lobe more
slender and apically slightly deflexed and the intermediate duct between the spermatheca and receptacle
longer and more strongly sinuate. The apices of the
lateral lobes extend to or beyond the apex of the
median lobe in V. mexicanus (Fig. 140) whereas in
V. lateralis the apices of the lateral lobes end well
short of the apex of the median lobe (Fig. 139).
Description:
Habitus. Lateral outline distinctly discontinuous in
dorsal aspect, lateral elytral margins evenly rounded
(Fig. 161), body not dorsoventrally flattened. Measurements are given in Table 1.
Coloration. Head yellow; pronotum brown, irregularly yellow along anterior and lateral margins;
0.5mm
165
B
0.5mm
162
467
A
0.5mm
0.5mm
163
A
B
166
0.5mm
161
164
167
168
0.5mm
Figures 161–168. Vatellus lateralis. 161, dorsal habitus; 162, elytral sculpture in area indicated on habitus by dotted
elytron dark brown to yellow-brown with yellow maculae subapically and laterally along margins, basally
with few yellow maculae (Fig. 161); venter and
appendages red brown to light brown, infuscate, especially laterally on metacoxae.
smooth, shiny to slightly microreticulate, surface posterior to eyes alutaceous, rough; anterior clypeal mar-
gin rounded; eyes large, strongly protruding
(Fig. 161). Pronotum densely punctate, but punctures
strongly obscured by rough, alutaceous sculpturing
similar to Figure 5, pale setae moderately long; pronotum cordate, lateral pronotal margin rounded anteriorly, broadest near anterior angles, posterior angles
acute (Fig. 161); lateral bead obscured; transverse sulcus distinctly impressed. Elytron with punctation
dense but edges of punctures strongly obscured by
dense alutaceous microsculpturing similar to
468
K. B. MILLER
171
0.5mm
169
170
173
172
0.25mm
Figures 169–173. Vatellus lateralis. 169–171, male genitalia; 169, right lateral lobe, lateral aspect; 170, median lobe, right
lateral aspect; 171, median lobe, ventral aspect; 172–173, female genitalia; 172, ventral aspect; 173, spermatheca and associated structures, dorsal aspect.
Figure 5, light setae moderately dense and conspicuous. Prosternum short, roughly sculptured, glabrous;
prosternal process broad with distinctly rounded
medial carina, lateral margins rounded, apex triangularly produced, glabrous, apex pointed. Metasternum
with punctation dense laterally, punctures large and
shallow, somewhat obscured by alutaceous sculpturing; anteromedial margin slightly produced ventrally.
Abdominal sternal sutures not unusually modified;
sterna with punctation dense over entire lateral surface, finer medially, obscured by alutaceous sculpturing; sixth abdominal sternum with apical lobe
inconspicuously produced, triangular, not strongly
produced (Fig. 167); speleum broad basally, sides
strongly convergent, apex moderately rounded, not
strongly curved (Fig. 168). Metacoxae with lateral portion densely punctate over entire surface, punctures
shallow, somewhat obscured by alutaceous sculpturing; metacoxal lines strongly approximate posteriorly,
strongly divergent anteriorly (Fig. 163); medial portion moderately punctate, punctures finely setose.
Metatrochanter rounded, offset (Fig. 164); metatarsal
claws minutely dentate.
curved, apically gradually narrowed, becoming slender, apex narrowly rounded, distinctly but not sharply
curved dorsally, with few subapical setae, basal portion elongate and moderately broad (Fig. 170); in ventral aspect moderately broad, margins slightly
convergent apically, somewhat sinuate basomedially,
apex narrowed relatively abruptly, apex acutely
rounded (Fig. 171). Lateral lobe relatively broad, apical portion evenly and shallowly curved, broad to
broadly rounded apex, apical an apicomedial margin
with dense series of fine setae; subapical stout, curved
setae absent (Fig. 169).
robust, medial margin moderately broad, very irregular; anterior margin strongly concave; anterior lobe
relatively slender and elongate (Fig. 172). Gonocoxa
broad, medial margin broadly rounded, apex broadly
rounded; apodeme short and slender (Fig. 172). Bursa
copulatrix moderate in size, not modified; spermathecal duct moderately long; receptacle ovoid, about half
the size of spermatheca; intermediate duct moderately
long, strongly sinuate (Fig. 173); spermatheca subspherical, triangular process moderately long, broad;
fertilization duct short (Fig. 172).
Sexually dimorphic characters. Mesotrochanter and
mesofemur of male with setal brushes well developed,
that of mesofemur moderately elongate. Male with
pro- and mesobasotarsomeres (Fig. 165) considerably
broader than in female (Fig. 166).
Intraspecific variation. This species varies substantially in size and shape. There is also considerable
variation in degree of expression of the cuticular
microsculpture with females exhibiting it to a greater
degree. The extent of the maculations is highly variable, but maculae are generally distinct though in
some cases they may be difficult to distinguish from
the background coloration.
Natural history: This species is known from black
lights, lowland marshes and subtropical moist
forest.
Distribution: Vatellus lateralis has been collected from
Bolivia, Brazil, Paraguay, Peru and Uruguay
(Fig. 311).
469
(Fig. 316). However, within this group this species does
not have clear affinities with any other particular species. It is very similar to V. mexicanus in many features.
Discussion: The species is widespread and relatively
commonly collected throughout tropical South America. It has been described several times. Examination
of the types of V. lateralis, V. rudis and V. marginalis
indicate that these names refer to the same species.
Vatellus lateralis comes first in Sharp’s (1882b)
monograph, so I selected it as the senior name. The
strong similarity between this species and
V. mexicanus may indicate that the two should be
classified as a single species. However, consistent differences in male and female genitalia and allopatric
distributions suggest that the two are distinctive.
Specimens distributed between the southern range of
V. mexicanus and the northern range of V. lateralis
should be examined in the future for intermediate
character states.
Material examined: Bolivia: Beni, Beni Stn. Palm
Camp, NE San Borja, at light, R.W. Brooks (1, USNM);
Santa Ana, 14 August 1937, HE Hinton (16, BMNH);
Trinidad, July–August 1937, HE Hinton (6, BMNH);
Trinidad, 29–30 July 1937, HE Hinton (4, BMNH);
Beni, 7.0 km SW Trinidad. 14∞52¢12≤S 64∞57¢32≤W, 18
July 1998, weedy pond, KB Miller (6, KBMC); Beni,
Prov. Cercado 9.5 km N Trinidad. 14∞46¢34≤S
64∞50¢00≤S, 17 June 1999, shallow marsh, KB Miller
(2, KBMC). Brazil: Alto da Serra be Sao Paulo 1929 (2,
MNHN); Bahia Encruzilhada, 960 m, 7 November
1972, Moacir Alvarenga (1, FSCA); Caraca, Minas
Gerais, December 1885, E Gouveille (4, MNHN); Caraqa, 1884, P Germain (17, MNHN); Encruzilhada,
980 m, Bahia, November 1974, M Alvarenga (3,
MNHN); Minas Gerias Aguas Vermlehas, December
1983, BLT, Moacir Alvarenga (6, FSCA); Minas Gerias,
Aguas Vermelhas, December 1983, BLT, M Alvarenga
(7, FSCA); Par. Curitiba, 29 June 1969, P and P. Spangler (7, USNM); Prov. Minas Gerais Aquas Vermelhas,
December 1998, FZ Vaz-de-Mello (33, LHIC); Rio
Grande del Norte, Natal, March 1952, M Alvarenga (3,
FSCA). Paraguay: Central Department, Cacupe Road,
Arroyo Yagua Resa y Ypacai, 10 April 1980, PJ Spangler, MM Culzoni, D Wood (1, USNM). Peru: Madre de
Dios; Rio Tambopata Res., 30 air km SW Pto Maldonado, 26–30 November 1979, subtropical moist forest, JP Heppner (1, USNM). Uruguay: (1, MNHN);
Artigma Supulturez, Rio Charreis Pionda de Negro
Muezto, 10 December 1966b, CS Carbonell, MA Mono,
FR San Martin (1, FSCA); Montevideo, Al Cassabo, 26
March 1932 (1, FSCA); Payendui Rio Uruguay Barra
Arroyo Guavigo, 9–11 February 1970, MS Moratorio,
MA Mono, GJ Wibmer (1, FSCA); Rivera Arriera, 3–7
470
K. B. MILLER
B
A
0.5mm
0.1mm
175
178
0.5mm
0.5mm
176
B
A
179
177
174
0.5mm
0.5mm
180
181
Figures 174–181. Vatellus haagi. 174, dorsal habitus; 175, elytral sculpture in area indicated on habitus by dotted square;
176, medial portion of metacoxae; 177, metatrochanter and metafemur, anterior aspect; 178–179, tarsomeres, dorsal aspect;
178, male; 179, female, A, protarsus; B, mesotarsus; 180, sternum VI, ventral aspect; 181, speleum abdominal sterna V, VI,
dorsal aspect.
February 1933, CS Carbonell (1, FSCA); Rocha Ruta
10, Aguas Dulces, 09–11 October 1970, MS Moratorio,
MA Mono, GJ Wibmer (2, FSCA).
VATELLUS HAAGI WEHNCKE, 1876
(FIGS 174–188, 313)
Vatellus haagi Wehncke, 1876: 357.
Macrovatellus haagi; Sharp, 1882b: 284 (comb. nov.);
Régimbart, 1899b:1, 1903: 46; Zimmermann, 1919:
124, 1920: 30; Trémouilles, 1995: 27.
Type information: Lectotype (designated here to clarify assignment of this name with this species), in
MNHN labelled, ‘[small, light purple square]/Haag
[handwritten, black line around label]/Brazilia [handwritten, green label with black line around border]/
Haagi [handwritten, black line around border, additional illegible word below “Haagi”/LECTOTYPE
Vatellus haagi Wehncke, 1876. des. by K.B. Miller
2003 [red label with double black line border].’
Wehncke (1876) did not indicate the number of specimens on which he based his description, although
471
184
0.5mm
0.1mm
182
183
185
ID
SP
ID
RE
SP
188
TP
0.1mm
RE
FD
GS
SD
TP
FC
186
BC
187
GC
RA
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.
Nilsson (2001) concluded that there are syntypes.
Wehncke’s specimens went to Oberthur and thence to
the Museum National d’Histoire Naturelle, Paris
(MNHN). A single specimen in MNHN was examined
that matches his description, and the label informa-
tion currently on the specimen suggests that it, at
least, is a specimen on which he based the Description:
It is selected as the lectotype.
Type locality: Brazil, Corrientes.
472
K. B. MILLER
crease distinctive (Fig. 174); (4) male pro- and
mesobasotarsomeres (Fig. 178) much broader than in
female (Fig. 179); (5) apical lobe on abdominal
sternum VI present but not strongly expressed
(Fig. 180); (6) male and female genitalia distinct
(Figs 182–188). In this species the male median lobe
is apically broadly truncate in ventral aspect
(Figs 183, 185) and distinctly deflexed in lateral
aspect (Fig. 182). The apex is strongly beset with
short, proximally directed spines (Fig. 185). The
female bursa is the largest of any Vatellus and is
broad, strongly dorsoventrally compressed and
strongly sinuate (Figs 186, 187). The prosternal process is apically relatively blunt, not flattened and apically pointed as in most Vatellus.
Description:
Coloration. Head yellow; pronotum yellow, yellowbrown medially, narrowly brown along anterior and
broadly along posterior margins; elytron dark brown
to yellow-brown with yellow maculae subapically and
laterally along margins, basal margin irregularly and
broadly yellow (Fig. 174); venter and appendages redbrown to light brown, infuscate, especially laterally on
metacoxae.
smooth, shiny to slightly microreticulate, surface posterior to eyes alutaceous, rough; anterior clypeal
margin rounded; eyes large, strongly protruding.
Pronotum densely punctate, but punctures strongly
obscured by rough, alutaceous sculpturing similar to
Figure 5, pale setae moderately long; pronotum cordate, lateral pronotal margin rounded anteriorly,
broadest near anterior angles, posterior angles
slightly obtuse; lateral bead obscured; transverse sulcus distinctly impressed. Elytron with punctation
dense but edges of punctures strongly obscured by
dense alutaceous sculpturing similar to pronotum,
light setae moderately dense and conspicuous. Prosternum short, roughly sculptured, glabrous; prosternal process broad, flat, medially with conspicuous
protrusion to rounded carina, lateral margins parallelsided, apex broad, truncate with fringe of fine setae.
Metasternum with punctation dense laterally, punctures large and shallow, somewhat obscured by aluta-
ceous sculpturing; anteromedial margin narrowly
produced ventrally, apex acute. Abdominal sternal
sutures not unusually modified; sterna with punctation dense over entire lateral surface, finer medially,
obscured by alutaceous sculpturing; sixth abdominal
sternum with apical lobe narrow, triangular, not
strongly produced; speleum broad basally, sides
strongly convergent, apex narrowly rounded, not
strongly curved. Metacoxae with lateral portion
densely punctate over entire surface, punctures shallow, somewhat obscured by alutaceous sculpturing;
metacoxal lines strongly approximate posteriorly,
strongly divergent anteriorly (Fig. 176); medial portion moderately punctate, punctures finely setose.
Metatrochanter rounded, offset (Fig. 177); metatarsal
claws minutely dentate.
Male genitalia. Median lobe in lateral aspect moderately narrow, slightly expanded medially, apex narrowly rounded, distinctly curved dorsally, with large
field of short spines over ventrolateral surfaces, basal
portion long, narrow (Fig. 182); in ventral aspect with
lateral margins approximately parallel, apex abruptly
truncate, broad (Figs 183, 185). Lateral lobe very long,
slender, with apical portion evenly curved, apically
distinctly expanded and broadly rounded, densely covered with long, fine setae along lateral and apical
margins; with long, stout, subapical setae absent
(Fig. 184).
apical angle rounded, medial margin membranous,
anterior margin distinctly concave, anterior lobe very
long, robust, apically expanded and truncate
(Fig. 186). Gonocoxa robust, main portion broadest
medially, apex obtusely rounded, with numerous
spinous setae along apicoventral margins and over
dorsal surface, apodeme narrow, as long as main portion (Fig. 186). Bursa copulatrix very long, robust,
broad and dorsoventrally flattened, apex folded dorsally (Figs 186, 187); spermathecal duct moderately
long, narrow; receptacle spherical, about two-thirds
size of spermatheca (Fig. 186); intermediate duct relatively short, curved but not strongly coiled or twisted
(Fig. 188); spermatheca subspherical, produced posteriorly, triangular process relatively narrow basally,
broadly expanded and rounded apically; fertilization
duct short, curved (Fig. 186).
Sexually dimorphic characters. Male with welldeveloped mesotrochanteric brush of setae and
mesofemoral brush well-developed in elongate cupshaped form. Male pro- and mesobasotarsomeres
(Fig. 178) much more broadly expanded than in
female (Fig. 179), similar in length (Figs 178, 179).
Intraspecific variation. This species is somewhat
variable in coloration and extent of maculations on the
elytra. However, the species is, in general, relatively
uniform in characters.
Natural history: Vatellus haagi has been collected at
lights and from roadside pools.
Distribution: Vatellus haagi is found is southern South
America and has been collected from Argentina,
Bolivia, Paraguay and Uruguay (Fig. 313).
species.
Material examined: Argentina: Ao. P Verne, 4 km N
Va San Jose, 15 November 1973, OS Flint (1, USNM);
Bs. As. Laguna Chascomus at Ao. San Felton, 20 February 1974, OS Flint (1, USNM); Buenos Aires, 17
November 1912 (3, BMNH); Buenos Aires, v Belgrano,
10–15 October 1905, R Thaxter (1, MCZC); Chaco,
Charata, March 1991 (1, LHIC); Concordia, Prov Etre
Rios (1, MNHN); Cord, Carlos Paz, 26–27 May 1969,
PJ Spangler (1, USNM); Eldorado Misiones, 8 December 1964 (1, MNHN); Entre Rios, Colon, 1 February
1988 (1, LHIC); Estancia la Noria, Rio San Javier,
Santa Fe, 19 December 1911, GE Bryant (2, BMNH);
Hohenau. Alto-Parana, 6 September 1937, H Jacob (1,
BMNH); Misiones, Jardin America, Salto Tabay, 22
December 1990, riverside forest UV light, S & J Peck
(1, CMNC); Prov. Buenos Aires, Colon, 29 December
1991, at light, Archangelsky (3, LHIC); Prov. Chaco
San Bernardo, 1 February 1982, at light, Dilorio (3,
LHIC); Prov. Entre Rios, Department Colon/Liebig,
October 1989, R Foerster (9, LHIC); Rio Paraguay,
Formosa, 4 December 1973, OS Flint (1, USNM);
Santa Fe Reconquista, 13 February 1979, RE Woodruff, H Cordo (1, FSCA); Villa Ana, FCSFe, 25
December, at light, KJ Hayward (1, BMNH); Villa
Ana, P.C.S. Fe, January 1926, KJ Hayward (1,
USNM). Bolivia: San Antonio de Parapeti Rio Parapeti Santa Cruz, 15–19 July 1964, B Malkin (1,
FSCA); Santa Cruz, 11–12 May 1969, P and P Spangler (2, USNM); Santa Cruz, 60 mi N Santa Cruz,
Saavedra Exp. Sta., 3–5 January 1960, R Cumming
(1, USNM); Santa Cruz, Ayacucho, 13–14 May 1969, P
and P Spangler (12, USNM); Sta. Cruz, Prov. Chiquitos, 1.3 km WSW San Jose, 28 June 1999, stream pool,
KB Miller (16, KBMC). Paraguay: December 1936 (1,
MNHN); (1, MNHN); Asuncion, 7 October 1980, DC
Lowrie (3, USNM); Asuncion, 8 January 1991, Dreshsel (1, LHIC); Cen. 15 km NE Asuncion, 21 June 1969,
P and P Spangler (11, USNM); Cen. 2 km N Luque,
23–24 June 1969, P and P Spangler (32, USNM); Cen.,
Ypacarai, 22 June 1969, P and P Spangler (1, USNM);
Cord. San Bernardino, 22 June 1969, P and P Spangler (18, USNM); Department Central, Lambare,
1991, Drechsel (1, LHIC); Department Central, Asuncion, 5 October 1991, U. Drechsel (1, LHIC); Hohenau,
473
Alto-Parana, 5 September 1937, H Jacob (1, BMNH);
Paraguari Department Ybycui 20 km SE, ruta La
Rosada, Ybycui, 13 April 1980, roadside ponds, PJ
Spangler (2, USNM). Uruguay: Colonia, Carmelo, 11
January 1962, C Morey, M Monne, CS Carbonell (3,
FSCA); Montevideo, Pocitos, 01 November 1932, CS
Carbonell (1, FSCA); Payeandu, Rio Uruguay, Barra
Arroyo Guaviyu, 9–11 February 1970 (1, FSCA).
VATELLUS WHEELERI MILLER
(FIGS 189–202, 314)
SP. NOV.
‘ARGENTINA, Chaco Rcho.Barrangueras Puerto Vilelas 5 December 1973 O. S. Flint, Jr./HOLOTYPE
Vatellus wheeleri Miller 2004 [red label with double
black line border].’
Argentina: Corr. Ituazingo, 20 December 1990, wet
grassland UV, S & J Peck (12, CMNC). Paraguay:
Department Central, Asuncion, 2 October 1992,
Dreschsel (3, LHIC); Department Guaira, Calle Florida, LHIC (2, LHIC); Department Guaira, Garay, 10
August 1992, U. Dreschsel (2, LHIC); Department
Paraguari, Sapucay, 9 April, Dreschsel (1, LHIC).
Type locality: Argentina, Chaco, Barranqueras.
crease distinctive; (4) male pro- and mesobasotarsomeres (Fig. 193) much broader than in female (Fig. 194);
(5) apical lobe on abdominal sternum VI present but
weakly expressed (Fig. 195); (6) speleum long, anteriorly expanded, strongly curved dorsally (Fig. 196); and
(7) male and female genitalia distinct (Figs 197–202).
The median lobe is apically relatively broadly rounded
in ventral aspect (Figs 197, 199), but not as truncated
as in V. haagi. The gonocoxae in this species are more
robust and strongly rounded than most species
(Fig. 201). However, this species is very similar to
V. mexicanus, V. lateralis and V. haagi and it appears
that the shape of the speleum (Fig. 196) is its strongest diagnostic feature.
Description:
Habitus. Body outline moderately discontinuous in
dorsal aspect, lateral margins of elytra moderately
strongly rounded (Fig. 189), not dorsoventrally compressed. Measurements are given in Table 1.
Coloration. Head yellow-brown; pronotum dark
brown, irregularly yellow anterolaterally; elytron
dark brown to yellow-brown with yellow maculae sub-
474
K. B. MILLER
A
B
1.0mm
0.5mm
193
190
0.5mm
0.5mm
B
A
191
194
192
189
0.5mm
0.5mm
195
196
Figures 189–196. Vatellus wheeleri. 189, dorsal habitus; 190, elytral sculpture in area indicated on habitus by dotted
apically and laterally along margins (Fig. 189); venter
and appendages dark red-brown.
Sculpture and structure. Head with punctation distinct, moderately coarse, absent to extremely fine and
spares in large, triangular area on frons and clypeus,
interpuncture surfaces smooth, shiny to slightly
microreticulate, surface posterior to eyes alutaceous,
rough; anterior clypeal margin rounded; eyes large,
prominently protruding (Fig. 189). Pronotum densely
punctate, but punctures obscured by alutaceous sculpturing similar to that shown in Figure 5, pale setae
moderately long; pronotum cordate, lateral pronotal
margin rounded anteriorly, broadest near anterior
angles, posterior angles slightly obtuse (Fig. 189); lateral bead obscured; transverse sulcus moderately
impressed, visible mainly laterally. Elytron with punctation dense but edges of punctures obscured by
alutaceous sculpturing as on pronotum, light setae
moderately dense and conspicuous. Prosternum short,
shagrined, lightly setose; prosternal process broad, lateral margins rounded, basomedially slightly tectiform,
medially excavated, apex flattened, sharply triangular,
acuminate to slightly obtusely pointed. Metasternum
with punctation dense laterally, punctures large and
shallow, somewhat obscured by alutaceous sculpturing; anteromedial margin narrowly produced ven-
trally, apex acute. Abdominal sternal sutures not
unusually modified; sterna with punctation dense over
entire lateral surface, finer medially, obscured by
alutaceous sculpturing; sixth abdominal sternum with
apical lobe moderately produced, rounded, somewhat
flattened; speleum broad, distinctly expanded, apex
dorsally hooked dorsally (Fig. 196). Metacoxae with
lateral portion densely punctate over entire surface,
sculpturing; metacoxal lines strongly approximate
posteriorly, strongly divergent anteriorly (Fig. 191);
medial portion moderately punctate, punctures finely
setose. Metatrochanter strongly rounded, offset
(Fig. 192); metatarsal claws minutely dentate.
slightly and evenly curved throughout length, slender,
basal portion elongate, relatively straight and narrow,
slightly narrowed subapically (Fig. 198); apex narrowly rounded, with numerous short spines subapically on ventral margin (Fig. 197); in ventral aspect
moderately broad basally, narrower in apical twothirds, apically broadly pointed (Figs 197, 199). Lateral lobe with apical portion slightly and evenly
curved, apex broadly rounded, with large apical and
lateral region of long, fine setae; long, stout, subapical
setae absent (Fig. 200).
broad, apical angle rounded, anterior margin deeply
curved, anterior lobe elongate, moderately broad, sinuate (Fig. 201). Gonocoxa short, broad and robust,
apex broadly rounded, apodeme short and slender
(Fig. 201). Bursa copulatrix broad, irregular (Fig. 201);
spermathecal duct long, narrow; receptacle large, subequal in size to spermatheca, spherical (Fig. 201);
intermediate duct broad, curved, strongly curved
(Fig. 202); spermatheca spherical, large; fertilization
duct large, abruptly and broadly expanded (Fig. 201).
Sexually dimorphic characters. Male with welldeveloped, elongate cup-shaped brush of setae on
mesofemora and mesotrochanter with dense region of
setae; male pro- and mesobasotarsomeres (Fig. 193)
much broader than in female (Fig. 194), female protarsomere III (Fig. 194A) longer than male protarsomere III (Fig. 193A).
Natural history: This species has been collected from a
UV light in wet grassland.
Distribution: Vatellus wheeleri is known from southern South America and has been collected from Argentina and Paraguay (Fig. 314).
species.
475
Etymology: This species is named in honour of Quentin D. Wheeler (The Natural History Museum, London) for his considerable contribution to taxonomic
theory and knowledge of Coleoptera, for his extensive
support of me during various research projects and for
his highly valued friendship.
VATELLUS PERFORATUS (GUIGNOT, 1955) COMB. NOV.
(FIGS 1, 203–215, 314)
Platydessus perforatus Guignot, 1955: 4.
Macrovatellus perforatus; Spangler, 1966a: 58 (comb.
nov.); Trémouilles, 1995: 27.
Type information: Holotype in G. Frey Museum,
Munich, Germany (not examined). Although the type
was not examined, a large number of specimens identified as this species by P. Spangler (USNM) were.
Based on this, I believe Guignot’s species refers to the
one described here.
Type locality: Bolivia.
Diagnosis: This species is distinguishable from other
Vatellus by the shorter, stouter habitus and relatively
small size (Figs 1, 203), relatively coarse punctation
and elytra smooth between punctures (elytral surface
sculpture as in Fig. 204), broad medial portions of the
metacoxae (Fig. 205), strongly offset and apically
rounded metatrochanter (Fig. 206), weakly produced
anterior clypeal margin and distinctive male and
female genitalia. The male median lobe is apically
abruptly narrowed to a small, ventrally curved point
(Fig. 212). The receptacle is relatively large, about the
size of the spermatheca, and is ovate (Figs 214, 215).
The gonocoxae are long and apically pointed
(Fig. 214).
Description:
dorsal aspect, lateral margins of elytra moderately
rounded (Figs 1, 203), dorsoventrally moderately compressed. Measurements are given in Table 1.
Coloration. Head, pronotum and elytron dark redbrown; venter dark red brown, darker on metacoxa
and metasternum; appendages dark red-brown.
Sculpture and structure. Head with punctures
evenly distributed, moderately coarsely punctate,
punctures slightly finer on clypeus; anterior clypeal
margin distinctly produced; eyes moderately large, not
strongly protruding (Fig. 203). Pronotum with punctation evenly distributed, punctures moderately coarse,
interpuncture surface shiny; lateral margin strongly
rounded anteriorly, widest point near anterior angles,
posterior angles slightly acute; lateral bead obscured
posteriorly; transverse sulcus absent. Elytron evenly
and coarsely punctate, punctures moderately dense,
surface between punctures shiny, smooth (Fig. 204).
476
K. B. MILLER
0.25mm
0.1mm
199
198
200
197
0.1mm
201
202
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,
477
A
1.0mm
0.5mm
B
207
204
0.5mm
0.5mm
B
205
A
208
206
0.5mm
203
0.5mm
209
210
Figures 203–210. Vatellus perforatus. 203, dorsal habitus; 204, elytral sculpture in area indicated on habitus by dotted
Prosternum short with dense fringe of yellow setae;
prosternal process broadly triangular, medially excavated, apex acutely pointed. Metasternum with punctation coarse and dense, medially with punctures
slightly finer, interpuncture surface smooth, shiny;
anteromedial margin slightly produced ventrally.
Abdominal sternal margins not unusually modified;
sterna evenly and finely punctate, interpuncture surface smooth, shiny; sixth abdominal sterna with apical
lobe flattened, long, subtriangular (Fig. 209); speleum
moderately broad throughout length, relatively
abruptly narrowed to narrowly rounded apex
(Fig. 210). Metacoxa with lateral portion coarsely and
densely punctate over entire surface; metacoxal lines
relatively broadly separated basally, apically very
broadly divergent (Fig. 205); medial portion densely
and conspicuously punctate. Metatrochanter very
short and rounded (Fig. 205); metatarsal claws
minutely dentate.
evenly curved, apically abruptly narrowed to slender, slightly hooked apex (Fig. 212); in ventral aspect
with lateral margins slightly but evenly convergent,
apically abruptly narrowed with small, apical, nar-
478
K. B. MILLER
0.5mm
212
213
211
215
0.1mm
214
Figures 211–215. Vatellus perforatus. 211–213, male genitalia; 211, right lateral lobe, lateral aspect; 212, median lobe,
right lateral aspect; 213, median lobe, ventral aspect; 214–215, female genitalia; 214, ventral aspect; 215, spermatheca and
associated structures, dorsal aspect.
rowly rounded point (Fig. 213). Lateral lobe very
long, basally very broad, apical portion elongate,
straight, expanded medially, apically rounded, with
dense apical and dorsal fringe of fine, elongate setae
(Fig. 211).
Female genitalia. Gonocoxosternite very broad and
robust, apical angle rounded, anterior margin deeply
concave, anterior lobe relatively long, slender, apex
curved laterally (Fig. 214). Gonocoxa elongate, slender, apex narrowly rounded, apodeme elongate and
moderately slender (Fig. 214). Bursa copulatrix short,
slender (Fig. 214); spermathecal duct moderately long,
slender; receptacle robust, elongate, subrenal shaped,
subequal in size to spermatheca (Fig. 214); intermediate duct moderately long, robust, sinuate (Fig. 215);
spermatheca subspherical, triangular process very
long and broad; fertilization duct short (Fig. 214).
Sexually dimorphic characters. Male with setae on
mesotrochanter and mesofemur weakly developed.
Male and female pro- and mesobasotarsomeres similar in size and length, both with protarsomere III very
long and slender (Figs 207, 208).
Intraspecific variation. Specimens differ in the
extent of shagrined sculpture on the pronotum with it
nearly absent in some specimens and making an
extensive, distinct line subposteriorly on others.
Natural history: This species appears to be streaminhabiting, unlike most species in the genus. Specimens were collected from streams in a rainforest
among rocks, leaves and sticks. It has also been collected from black lights.
Distribution: Vatellus perforatus is a northern South
American species and has been collected from Brazil
and Venezuela (Fig. 314).
Phylogenetic relationships: This species occupies a relatively isolated phylogenetic position within Vatellus,
but evidence indicates it is relatively closely related to
the V. tarsatus and V. grandis clades (Fig. 316).
Material examined: Brazil: Amazonas, Rio Demiti, nr
Little Homestead. 0∞35¢N 66∞41¢W, UV light, GE and
KE Ball (1, JBWM). Venezuela: T.P. Amaz. Cerro de la
Neblina 1 km S Basecamp 0∞50¢N 66∞10¢W 250 m, 8
February 1985, along small whitewater stream, in
pools of dead le, P.J and P.M. Spangler, R. Faitoute, W.
Steiner (13, USNM); T.P. Amaz. Cerro de la Neblina
1.5 km S Basecamp 0∞50¢N 66∞10¢W 250 m, 9 February 1985, small stream in rainforest, among rocks and
leaf p, W.E. Steiner (1, USNM).
VATELLUS TARSATUS (LAPORTE, 1835)
(FIGS 3, 216–229, 313)
Hydroporus tarsatus Laporte, 1835: 106, 1840: 167.
Vatellus tarsatus; Aubé, 1837: 218 (comb. nov.), 1838:
449; Gemminger & Harold, 1868: 428; Sharp, 1882b:
285; Régimbart, 1878: 463; Trémouilles, 1995: 27.
MNHN labelled, ‘Hydroporus tarsatus Lap. Typemihi D.S. [handwritten]/Hydroporus Vatellus tarsatus Buquet L. Cayenne D. Buquet [handwritten,
green label, folded in half, “Hydroporus” crossed out]/
Ex.musæo Dejean [black line around border]/D. Sharp
Monogr./LECTOTYPE Hydroporus tarsatus Laporte,
479
1835. des. by K.B. Miller 2003 [red label with double
black line border].’
Type locality: French Guyana [‘Cayenne’].
Diagnosis: This species is distinguishable from other
Vatellus by the rather strongly demarcated posterior
margins of abdominal sterna 2, 3 and 4 in most specimens. The posterior margins on these sterna end
abruptly and rise steeply to the surface of the next
sternum. Other distinguishing characters include the
relatively fine punctures and elytral surface sculpture
as in Figure 3, nearly immaculate dorsal surface and
dark brown to black colour, the very slender speleum
(Fig. 223), the median lobe apically slender and
sharply upturned. The receptacle is about the size of
the spermatheca and the intermediate duct is very
broad (Fig. 229).
Description:
Habitus. Lateral outline discontinuous in dorsal
aspect, lateral margins of elytra moderately rounded
Coloration. Head, pronotum and elytron dark brown
to black; venter black to dark red-brown, appendages
dark yellow-brown to dark red-brown.
punctures dense and confused in places, somewhat
finer anteriorly, surface slightly shagrined between
punctures; anterior clypeal margin rounded; eyes
large, protruding. Pronotum evenly punctate, punctures fined and moderately dense, surface alutaceous
between punctures, more commonly so in females, as
in Figure 3, each puncture bearing a fine, relatively
long seta, with long, distinct subbasal transverse line
of shagrination; pronotum cordate, broadest near apical angles, lateral margins moderately strongly
rounded anteriorly, posterior angles slightly acute and
pointed (Fig. 216); transverse sulcus absent to slightly
impressed. Elytron evenly covered with fine punctures, punctures moderately dense, each bearing a relatively long, fine, pale seta (Fig. 217), interpuncture
surface shiny, smooth in most males and most females,
some with microsculpture as in Figure 3, making surface alutaceous. Prosternum short, slightly roughened, with numerous setae; prosternal process broad,
lateral margins evenly curved, medially excavated,
apically triangular, flattened, apex acutely pointed.
Metasternum medially finely punctate, laterally with
punctures more coarse and shallow, confused with surface slightly shagrined; anteromedial margin not
expanded, apex slightly deflexed and pointed. Abdominal sterna with sutural margins abruptly margined,
particularly between visible sterna 2, 3 and 4; sterna
finely and irregularly punctate, surface between punctures shiny, smooth; sixth sternum with apical lobe
480
K. B. MILLER
0.5mm
1.0mm
220
217
B
A
0.5mm
0.5mm
218
A
B
221
219
216
0.5mm
0.5mm
222
223
Figures 216–223. Vatellus tarsatus. 216, dorsal habitus; 217, elytral sculpture in area indicated on habitus by dotted
very long, narrow, apically narrowly rounded; speleum
broad basally, strongly narrowed subbasally, apex long
and extremely slender to very narrowly rounded apex
(Fig. 223). Metacoxa with lateral portion irregularly
and shallowly punctate, punctures confused, surface
between punctures slightly shagrined; metacoxal lines
very closely approximated posteriorly, anteriorly moderately divergent (Fig. 218); medial portion finely and
sparsely punctate. Metatrochanters short, rounded,
strongly offset (Fig. 219); metatarsal claws weakly
dentate.
curved, slightly broadened apicomedially, basal portion
subtriangular, not strongly elongate; apex distinctly
narrowed, apically sharply pointed and distinctly
curved ventrally, with numerous small spines on apicoventral surface (Fig. 225); in ventral aspect narrow,
evenly narrowed to acutely pointed apex (Figs 224,
226). Lateral lobe with apical portion strongly curved
basally and relatively straight in apical portion, moderately broad, apex rounded, with large field of fine,
elongate setae along apical and lateral margins, subapical stout, elongate, curved setae absent (Fig. 227).
Female genitalia. Gonocoxosternite with apex narrowly rounded, medial, anterior and posterior margins
arcuate, anterior lobe moderately long, narrow
0.1mm
481
226
227
224
225
0.5mm
229
228
0.1mm
Figures 224–229. Vatellus tarsatus. 224–227, male genitalia; 224, apex of median lobe, ventral aspect; 225, median lobe,
right lateral aspect; 226, median lobe, ventral aspect; 227, right lateral lobe, lateral aspect; 228–229, female genitalia; 228,
(Fig. 228). Gonocoxa elongate-triangular, apex narrowly rounded, ventral surface with numerous long,
spinous setae, apodeme relatively short, slender
(Fig. 228). Bursa copulatrix small, inconspicuous;
spermathecal duct very long, coiled, slender; receptacle spherical, subequal in size to spermatheca
(Fig. 228); intermediate duct relatively short, slightly
twisted (Fig. 229); spermatheca spherical, triangular
482
K. B. MILLER
process narrow basally, very broadly expanded apically; fertilization duct relatively short (Fig. 228).
Sexually dimorphic characters. Male with very
weakly developed mesotrochanteric and mesofemoral
brushes of setae. Male with pro- and mesobasotarsomeres (Fig. 220) slightly broader than in female
(Fig. 221), male protarsomere III (Fig. 220A) distinctly
shorter than female protarsomere III (Fig. 221A).
Intraspecific variation. Historically the defining
apomorphy for this taxon was the strongly demarcated
sutures between the abdominal sterna. This feature is
somewhat variable, but all specimens exhibit this
character state to a greater or lesser degree. Although
most specimens are nearly black overall, some specimens are dark brown.
Natural history: Vatellus tarsatus has been collected
from a pond and a whitewater stream.
Distribution: This is a northern South American species with specimens collected from Brazil, French
Guyana, Suriname and Venezuela (Fig. 313).
Phylogenetic relationships: This species and the following two together form a clade (Fig. 316). They are
similar in several characters and in overall similarity
in less discrete characters. This clade is relatively
closely related to V. perforatus and the V. grandis
clade (Fig. 316).
Material examined: Brazil: Para, Rio Xingu Camp,
52∞22¢W 3∞39¢S Altamira, c. 60 km S, 11 October 1986,
pond at 2nd palm grove on trail, P. Spangler and R.
Crombie (4, USNM); Para, Rio Xingu Camp, 52∞22¢W
3∞39¢S Altamira, c. 60 km S, 12 October 1986, pond at
2nd palm grove on trail, P. Spangler and O. Flint (2,
USNM); Para, Rio Xingu Camp, 52∞22¢W 3∞39¢S Altamira, c. 60 km S, 15 October 1986, pond at 2nd palm
grove on trail, P. Spangler and O. Flint (2, USNM); Rio
Marauia, 24 January 1963, E Fittkau (2, ZSBS).
French Guyana: Country only (1, MNHN). Suriname:
3rd trib of Colakreek, 9 October 1989, N Nieser (1,
NHMW); Brokopondo, Coesewijnaproject, 9 April
1970, N Nieser (1, NHMW); Marowijne, Mooi Wannakreek, 54∞05¢W 05∞30¢N, 1 December 1989, N Nieser
(1, NHMW); nr. Sabakoekreek, 55∞11¢W 05∞26¢N, 28
July 1989, N Nieser (1, NHMW); Zanderijsavanne, 1st
trib. of Colskrrek, 55∞14¢W 05∞27¢N, 15 September
1989, N Nieser (2, NHMW). Venezuela: T.F. Amaz.
Cerro de la Neblina, 1 km S Basecamp 0∞50¢N
66∞10¢W, 11 February 1985, 140 m (6, USNM); T.P.
Amaz. Cerro de la Neblina 1.5 km S Basecamp 0∞50¢N
66∞10¢W 140 m, 15 February 1985, along small whitewater stream, in pools of dead leaves, P.J and P.M
Spangler, R. Faitoute, W. Steiner (2, USNM); T.P.
Amaz. Cerro de la Neblina 1.5 km S Basecamp 0∞50¢N
66∞10¢W 250 m, 11 February 1985, along small white-
water stream, in pools of dead leaves, P.J and P.M
Spangler, R. Faitoute, W. Steiner (3, USNM).
VATELLUS ANNAE MILLER SP.
(FIGS 216–229, 313)
NOV.
‘BRAZIL: Matto Grosso Jacaré P.N.Xingu xi.1961
Alvarenga e Werner leg./HOLOTYPE Vatellus annae
Miller 2004 [red label with double black line border].’
Paratypes: 1 in MCZC labelled same as holotype.
Bolivia: Beni, 3.4 km W San Borja. 14∞52¢57≤S
66∞46¢03≤W, 14 July 1998, KB Miller (1, KBMC). Brazil: Mato Grosso, Jacare-Parque Nat Xingu, November
1965, BLT, M Alvarengo, WCA Bokermann (1, FSCA);
Matto Grosso, Tapirape, 15 November 1961, light, B
Malkin (1, FSCA); Pres Matto Grosso, 1886, P Germain (4, MNHN); Rio Maravia, 24 January 1933, E.
Fittkau (2, MBIC). Peru: Loreto, Yacumana Lodge, Jct
Rio Maranon, Rio Ucayall 73∞6¢W 4∞8¢S, 20 August
1994, light in woods, Skelley (4, LHIC). Suriname:
55∞14¢W 05∞27¢N Brokopondo, 1st trib of Kolakreek,
15 September 1969, N. Nieser (2, MBIC). Trinidad:
Cumuto 1929, Darlington (68, MCZC); La Brea, Darlington (1, MCZC); St Augustine, April 1929, Darlington (1, MCZC); St Augustine, April 1929, Darlington
(3, MCZC).
Type locality: Brazil, Matto Grosso, Jacaré, P.N.
Xingu [presumably the Indigenous Park of the Xingu].
Diagnosis: This species is most similar to V. tarsatus
and V. pilacaudus in having overall fine punctation
and elytral sculpture as in Figure 3, relatively concolorous dorsal surfaces and similar habitus. Also, these
species have the apical lobe on abdominal sternum VI
long and apically nearly spherical (e.g. Fig. 222).
Vatellus annae differs from V. tarsatus in lacking a
strong, abrupt demarcation between abdominal sterna
and having the slender apex of the median lobe more
elongate and more sharply pointed apically (Fig. 239).
Also, the intermediate duct in V. annae is very long
and very coiled on the dorsal surface of the spermatheca (Fig. 244) whereas the intermediate duct in
V. tarsatus is shorter and broad (Fig. 229). The
speleum is relatively broad in V. annae (Fig. 273)
whereas in V. tarsatus it is very slender (Fig. 223).
Vatellus pilacaudus shares the presence of the long,
coiled intermediate duct, but in V. pilacaudus the
apex of the median lobe is not long, slender and
pointed (Fig. 255).
Description:
Habitus. Lateral outline moderately discontinuous
in dorsal aspect, lateral margins of elytra moderately
strongly rounded (Fig. 230), not dorsoventrally compressed. Measurements are given in Table 1.
483
234
0.5mm
1.0mm
231
B
A
0.5mm
0.5mm
A
B
232
235
233
230
0.5mm
0.5mm
236
237
Figures 230–237. Vatellus annae. 230, dorsal habitus; 231, elytral sculpture in area indicated on habitus by dotted square;
dorsal aspect.
Coloration. Head, pronotum and elytron brown-red;
venter and appendages brown-red.
Sculpture and structure. Head finely punctate, punctures sparse, very fine to absent on clypeus, surface
between punctures shiny; anterior clypeal margin
slightly protruded; eyes large, protruding (Fig. 230).
Pronotum evenly punctate, punctures fine and moderately dense, surface generally alutaceous between
punctures, each puncture bearing a fine, relatively
long seta; pronotum cordate, broadest near apical
angles, lateral margins moderately strongly rounded
anteriorly, posterior angles slightly acute and pointed
(Fig. 230); transverse sulcus slightly impressed.
Elytron evenly covered with fine punctures, punctures
moderately dense (Fig. 231), each bearing a relatively
long, fine, pale seta, interpuncture surface shiny,
smooth, or with distinctive microsculpture making
surface alutaceous similar to pronotum. Prosternum
short, slightly roughened, glabrous; prosternal process
broad, lateral margins evenly curved, medially excavated, apically triangular, flattened, apex acutely
pointed. Metasternum medially finely punctate, later-
484
K. B. MILLER
ally with punctures more coarse and shallow, confused
with surface between punctures shiny; anteromedial
margin not expanded, not deflexed. Abdominal sternal
sutures not usually modified; sterna finely and irregularly punctate, surface between punctures shiny,
smooth; sixth sternum with apical lobe well-developed,
apically rounded (Fig. 236); speleum broad, slightly
narrowed medially, apex broadly rounded (Fig. 237).
Metacoxa with lateral portion irregularly and shallowly punctate, punctures confused, surface between
punctures slightly shagrined; metacoxal lines closely
approximated posteriorly, anteriorly strongly divergent (Fig. 232); medial portion finely and sparsely
punctate. Metatrochanters short, rounded, strongly
offset (Fig. 233); metatarsal claws weakly dentate.
Male genitalia. Median lobe in lateral aspect narrow, medially somewhat broadened, basal portion
broad, moderately elongate; apex very slender, apically sharply pointed and curved ventrally, with distinctive region of small subapical spines on ventral
surface (Figs 239, 242); in ventral aspect narrow,
abruptly narrowed subapically, apex very slender,
very narrowly rounded, nearly pointed (Figs 240, 241).
Lateral lobe with apical portion evenly curved, narrow, with very few setae apically; long, stout, subapical setae absent (Fig. 238).
Female genitalia. Gonocoxosternite robust, medial
margin very broad, apex nearly right-angled; posterior
margin concave; anterior lobe relatively robust, short
(Fig. 243). Gonocoxa moderately broad, main portion
approximately triangular, apex pointed, apodeme long
and slender (Fig. 243). Bursa copulatrix small and
slender, undifferentiated; spermathecal duct moderately long, slender; receptacle spherical, less than 1/2
size of spermatheca (Fig. 243); intermediate duct long,
strongly and tightly coiled between receptacle and
spermatheca, adpressed to dorsal surface of spermatheca (Fig. 244); spermatheca spherical, triangular process relatively short, very broad; fertilization duct
short, slightly curved (Fig. 243).
Sexually dimorphic characters. Male with moderately developed mesotrochanteric and mesofemoral
brushes of setae. Male pro- and mesobasotarsomeres
(Fig. 234) broadened in male compared with female
(Fig. 235), protarsomere III longer than mesotarsomere III in both sexes (Figs 234A, 235A).
Natural history: Vatellus annae has been collected at
black lights.
Distribution: This species is relatively widespread in
northern South America and has been collected
in Bolivia, Brazil, Peru, Suriname and Trinidad
(Fig. 313).
Phylogenetic relationships: This species is a member
of the V. tarsatus clade along with V. pilacaudus
(Fig. 316). Within this clade, V. annae groups with
V. pilacaudus based on the common presence of relatively closely approximated metacoxal lines (Figs 231,
247) (compared with other Vatellus), the presence of a
long intermediate duct coiled on the dorsal surface of
the female spermatheca (Figs 244, 255), and the relatively small female receptacle (Figs 243, 257).
Etymology: This species is named in honour of my
daughter, Annette (Annie) Miller.
VATELLUS
PILACAUDUS MILLER
(FIGS 245–258, 313)
SP. NOV.
‘BOLIVIA: Beni, 40 km E San Borja, Estacion Biologica Beni, Estancia El Porvenir; 6–8 September 1987;
W. E. Steiner/At black light; open grass savanna and
marsh/HOLOTYPE Vatellus pilacaudus Miller 2004
[red label with double black line border].’
Bolivia: Beni, 2.0 km E San Borja. 14∞52¢02≤S
66∞43¢41≤W, 15 July 1988, shaded pool in streambed,
KB Miller (2, KBMC); 6.7 km NW Trinidad.
14∞47¢27≤S 64∞56¢47≤W, 18 July 1998, blacklight, KB
Miller (1, KBMC); Prov. Cercado, 9.5 km N Trinidad.
14∞46¢24¢S 64∞58¢00¢W, 28 June 1999, KB Miller (1,
KBMC).
Type locality: Bolivia, Departmento Beni, 40 km E San
Borja, Estacion Biologica Beni, Estancia El Porvenir.
Diagnosis: This species is most similar to V. tarsatus
and V. annae in having small punctures, surface
microsculpture (when present) as in Figure 3, relatively concolorous dorsal surfaces and similar habitus. Also, these species have the apical lobe on
abdominal sternum VI long and very well developed.
Vatellus pilacaudus differs from V. tarsatus in lacking
strong, abrupt demarcations between abdominal
sterna. Also, the apex of the median lobe is broader
and more narrowly rounded in lateral aspect
(Fig. 255), not slender and upturned as in V. tarsatus
and V. annae. Also, the intermediate duct in this species is very long and tightly coiled on the dorsal
surface of the spermatheca (Fig. 258), unlike in
V. tarsatus. In this species, the metacoxal lines are
not as strongly divergent apically as in most species
(ML/MW = 2.18–2.57) (Fig. 247).
Description:
Habitus. Lateral outline discontinuous in dorsal
aspect, lateral margins of elytra broadly rounded
Coloration. Head and pronotum yellow-brown;
elytron yellow-brown, slightly lighter along lateral
and anterior margins, often with small, indistinct
485
0.1mm
0.5mm
239
241
240
242
238
244
243
1.0mm
Figures 238–244. Vatellus annae. 238–242, male genitalia; 238, right lateral lobe, lateral aspect; 239, median lobe,
right lateral aspect; 240, median lobe, ventral aspect; 241, apex of median lobe, ventral aspect; 242, apex of median
lobe, right lateral aspect; 243–244, female genitalia; 243, ventral aspect; 244, spermatheca and associated structures,
dorsal aspect.
marginal macula apicomedially (Fig. 245); venter yellow-red; appendages yellow-brown.
Sculpture and structure. Head with fine and dense
punctation, slightly finer anteriorly, surface finely
microreticulate between punctures, shiny; anterior
clypeal margin rounded; eyes large, strongly protruding (Fig. 245). Pronotum evenly covered with dense,
moderately coarse punctation, surface finely microreticulate on most specimens; lateral margins moderately curved, widest point submedially, posterior
486
K. B. MILLER
A
0.5mm
B
1.0mm
249
246
0.5mm
0.5mm
247
B
A
250
248
245
0.5mm
0.5mm
251
252
Figures 245–252. Vatellus pilacaudus. 245, dorsal habitus; 246, elytral sculpture in area indicated on habitus by dotted
angles slightly acute to slightly obtuse (Fig. 245); lateral bead present and distinct along entire length;
transverse sulcus moderately distinct laterally.
Elytron densely punctate over entire surface, interpuncture surface similar to pronotum, setae moderately long and conspicuous. Prosternum short,
alutaceous, glabrous; prosternal process moderately
broad, slightly excavated, with only slight tumidity
medially, apex produced into elongate, flattened triangle, apex acutely pointed. Metasternum with punctation laterally dense, punctures large, medially more
finely punctate, interpuncture surface shiny, smooth;
anteromedial margin not produced, slightly pointed.
finely punctate medially, more coarsely punctate laterally, surface smooth, shiny between punctures;
sixth sternum with apical lobe strongly produced, distinctly rounded and subspherical apically; speleum
moderately broad, apically narrowly rounded
(Fig. 252). Metacoxa with lateral portion densely
punctate over entire surface, punctures moderate in
size, surface shiny between punctures; metacoxal
lines approximate posteriorly, distinctly but only
moderately divergent anteriorly (Fig. 247); medial
portion finely punctate. Metatrochanter strongly
rounded and offset (Fig. 248); metacoxal claws
slightly dentate.
curved throughout length, moderately broad, slightly
broader subapically, apex moderately narrowly
rounded, basal portion broad (Fig. 255); in ventral
aspect with lateral margins subparallel, moderately
convergent to narrowly rounded apex (Figs 254, 256).
Lateral lobe very long, apical portion moderately
broad, evenly curved, slightly expanded medially
(Fig. 253).
apical angle narrowly rounded, somewhat acute, anterior margin irregularly concave, anterior lobe relatively short, robust (Fig. 257). Gonocoxa moderately
elongate, medial margin broadly curved, apex
rounded, apodeme long, slender (Fig. 257). Bursa copulatrix broad, short, relatively undifferentiated
(Fig. 257); spermathecal duct moderately long, slender; receptacle small, less than half size of spermatheca, spherical (Fig. 257); intermediate duct very long,
very tightly coiled on dorsal surface of spermatheca
(Fig. 258); spermatheca spherical, large; fertilization
duct short, slightly curved (Fig. 257).
Sexually dimorphic characters. Male with very welldeveloped setal region on mesotrochanter, setae on
medial portion of mesofemur developed into elongate,
cup-shaped form. Male pro- and mesobasotarsomeres
(Fig. 249) distinctly expanded compared with female
(Fig. 250), but not exceptionally broad, female protarsomeres (Fig. 250A) very long and slender.
Intraspecific variation. Most specimens are uniformly light brown, but a few have more extensive
lateral, yellow margins on the elytron. Also, some
specimens, both male and female, exhibit fine microsculpturing on the pronotum and elytron and a few
female specimens are very strongly microsculptured
dorsally. Other specimens are smooth between the
punctures on the pronotum and elytron.
Natural history: Specimens have been collected at a
black light in open savannah and marshland and from
a shaded, stagnant pool in a streambed.
Distribution: Specimens have been collected only from
areas in Bolivia (Fig. 313).
of the V. tarsatus clade (Fig. 316). Within this clade,
V. pilacaudus groups with V. annae (Fig. 316) (see
comments under that species).
Etymology: This species is named from the Latin
words pila, meaning ‘ball’, and caudus, meaning ‘tail’,
in reference to the semispherical lobe of the sixth
abdominal sternum.
487
VATELLUS GRANDIS BUQUET, 1840
(FIGS 2, 6, 8–17, 24–27, 38, 39, 42–44, 48,
49, 259–273, 314)
Vatellus grandis Buquet, 1840: 394; Gemminger &
Harold, 1868: 428.
Macrovatellus grandis; van den Branden, 1885: 19
(comb. nov.); Sharp, 1882b: 282, 826; Zimmermann,
1920: 30; Régimbart, 1878: 463; Trémouilles, 1995: 27.
MNHN labelled, ‘Cayenne [green label]/Coll.C.Felsche
Geschenk 1907 [green label]/Vatellus grandis Buquet.
[handwritten]/1 29/Macrovatellus grandis [handwritten]/LECTOTYPE Vatellus grandis Buquet, 1840. des.
by K.B. Miller 2003 [red label with double black line
border].’ Buquet apparently had more than one specimen on which he based this Description: I found only
a single specimen in MNHN that appears to be one of
Buquet’s specimens. It was selected as the lectotype.
Type locality: French Guyana [‘Cayenne’], upper part
of Oyapok River.
Diagnosis: This species can be distinguished from
other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced (Fig. 6); (2)
with surfaces relatively coarsely punctate and with
surfaces between the punctures generally shiny, elytral surface sculpture as in Figures 2 and 260; (3) with
a transverse impunctate area on each side of the midline on the pronotum (Fig. 262); (4) eyes relatively
small (Fig. 259); (5) body relatively elongate and dorsoventrally depressed (Fig. 259); (6) lateral pronotal
margin relatively weakly curved (Fig. 259); (7) anterior portion of the metasternum expanded ventrally;
and (8) male and female genitalia distinctive
(Figs 269–273). The median lobe is long and slender
with the apex bluntly rounded (Fig. 269). The lateral
lobe has the apical portion relatively straight with the
apex relatively lacking in setae, but with two or three
long, stiff setae on the subapical, ventral surface
(Fig. 271). The female receptacle is relatively small
and elongate (Fig. 272). This species and the following
three are very similar in having a distinctive, flattened
habitus, similar punctation, a distinct impunctate area
on each side of the pronotum (Fig. 262), the ventrally
expanded anterior portion of the metasternum and the
anteriorly produced anterior clypeal margin (Fig. 6).
Vatellus grandis, V. amae and V. drymetes are nearly
indistinguishable externally, but have distinctive genitalia. Vatellus bifenestratus is larger and has the
pronotum more prominently cordate (Fig. 274).
Description:
Habitus. Body outline somewhat discontinuous in
dorsal aspect, lateral margins of elytra not strongly
488
K. B. MILLER
0.1mm
0.5mm
255
254
0.5mm
256
253
258
257
0.1mm
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,
rounded (Fig. 259), dorsoventrally compressed. Measurements are given in Table 1.
Coloration. Head, pronotum, and elytron brown-red;
ventral surfaces black except venter of head, pronotal epipleuron, propleuron, prosternum and small
portions of posterior margins of abdominal sterna
brown-red.
Sculpture and structure. Head finely and moderately densely punctate, slightly more dense anteriorly on frons and clypeus; anterior clypeal margin
strongly anteriorly produced; eyes moderate in size,
not strongly protruding. Pronotum with punctures
relatively large laterally, denser medially and along
posterior margin, punctures with fine pale, moder-
A
489
B
0.5mm
1.0mm
260
263
A
B
0.5mm
0.5mm
261
0.5mm
264
259
262
0.5mm
265
266
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.
ately elongate setae, area between punctures generally with fine shagrination formed by dense field of
very fine tubercles except for transverse impunctate
areas medially on each side of midline (Fig. 262);
pronotum broadest anteromedially, lateral margins
relatively parallel-sided, only curved at extreme
anterior end (Fig. 259); lateral bead obscured except
at extreme anterior end; transverse groove absent.
Elytron strongly and densely punctate over entire
surface, punctures finer and slightly more dense apically, each puncture with fine seta, surface between
punctures smooth and shiny (Fig. 260). Prosternum
short, setose; prosternal process broad, flattened
with short, indistinct, rounded medial carina, lateral
margins rounded, convergent to narrowly rounded
apex. Metasternum laterally with moderately large
punctures, relatively sparsely distributed, medially
with more dense, finer punctures, surface smooth,
shiny; anteromedial margin strongly expanded ventrally into distinct tumidity. Abdominal sternal
sutures not unusually modified; sterna laterally with
moderately large, moderately dense punctures, interpuncture surface smooth, shiny; apical lobe on sixth
abdominal sternum strongly produced, long, apically
narrowly rounded, flattened (Fig. 265); speleum elongate, moderately narrow, apically pointed (Fig. 266).
Metacoxa with lateral portion with punctures moderately large, relatively dense, interpuncture surface
490
K. B. MILLER
smooth, shiny; metacoxal lines strongly approximate
posteriorly, anteriorly conspicuously divergent
(Fig. 261); medial portion with surface shiny, finely
punctate. Metatrochanter strongly rounded, strongly
offset (Figs 267, 268); metatarsal claws minutely
dentate.
Male genitalia. Median lobe in lateral aspect relatively straight medially, apically slightly narrowed to
rounded apex, without subapical setae, basal portion
elongate and narrow, broadened apically, slightly
curved (Fig. 269); in ventral aspect with lateral margins approximately parallel, subapically abruptly narrowed to narrowly rounded apex (Fig. 270). Lateral
lobe long, basally broad, with apical portion straight
and narrow to acute, narrow apex; with 2–3 subapical,
stout, elongate, curved setae on ventral margin
(Fig. 271).
robust, apical angle broadly rounded, anterior margin
broadly concave, anterior lobe relatively short, slender
(Fig. 272). Gonocoxa elongate and slender, apex relatively narrowly rounded, apodeme elongate and slender (Fig. 272). Bursa copulatrix moderately long,
somewhat expanded apically (Fig. 272); spermathecal
duct moderately long, narrow; receptacle small, much
smaller than spermatheca, elongate and slender,
apically expanded (Fig. 272); intermediate duct
moderately long, with several S-curves (Fig. 273);
spermatheca subspherical, triangular process large
and broadly expanded; fertilization duct short and
curved (Fig. 272).
brushes of setae. Male with pro- and mesobasotarsomeres (Fig. 263) distinctly expanded compared with
female (Fig. 264); female protarsomere III (Fig. 264A)
much longer and more slender than in male
(Fig. 263A), female (Fig. 264B) and male (Fig. 263B)
mesotarsomeres I–III more similar in size, but female
mesotarsomeres narrower.
Intraspecific variation. This species is relatively consistent in characters. The most conspicuous variation
is in the extent of shagrination on the pronotum and
elytra. In a few specimens there is little or no shagrination. In others the entire pronotum and elytron is
shagrinated. However, in most there is a characteristic pattern of shagrination on the pronotum (Fig. 262)
and some or none on the elytra.
Natural history: Vatellus grandis has been collected
from subtropical marshes and moist forest and small
forest pools.
Distribution: This is a moderately commonly collected
species known from Brazil, Bolivia, French Guyana,
Peru, Suriname and Trinidad (Fig. 314).
Phylogenetic relationships: This species groups with
several others based on the common presence of similar cuticular sculpture (smooth and shiny between
punctures (Fig. 2) and with dense shagrination on the
pronotum except for small lateral patches (e.g.
Fig. 262)) in most specimens, the anteromedial portion
of the metasternum distinctly swollen ventrad (e.g.
Fig. 20), a distinct pencil of stiff setae on the ventroapical surface of the male lateral lobe (e.g. Fig. 271), and
very small female receptacles (e.g. Fig. 272).
Material examined: Bolivia: Beni, 2.0 km E San
Borja, shaded pool in streambed. 14∞52¢02≤S
66∞43¢46≤W, 15 July 1998, KB Miller (1, KBMC);
6.7 km NW Trinidad. 14∞47¢27≤S 64∞56¢47≤W, 18 June
1999, blacklight, KB Miller (3, KBMC). Brazil: Para
stream near Aldeia Coraci, 3 December 1965, B
Malkin (2, FSCA); Para, nr Aldeia Coraci, stream
11 km W Caninde, 3 December 1964, B Malkin (1,
FSCA). French Guyana: 1907, Geschenk (1, MNHN);
Lawa River shore opposite Anapaike village, 25
November 1963, Borys, Malkin (14, FSCA). Peru:
Madre de Dios; Rio Tambopata Res. 30 air km SW Pto.
Maldonado, 290 m, subtropical moist forest, J.B. Heppner (15, USNM). Suriname: Carolina Ck 10 km from
Zanderij, 18 November 1962, waterhole in forest, B
Malkin (3, FSCA); Krekka-Phedra Road, Suriname
dist 25 Tiny, 18 November 1962, tiny forest pool,
Borys, Malkin (2, FSCA); Zanderijsavanne, Parakreek, 55∞10¢W 05∞25¢N, 20 November 1989, N
Neiser (1, NHMW). Trinidad: Camuto 1929, Darlington (69, MCZC); Cumuto, 1929, Darlington (8, MCZC);
Sangre Grande, Cacoa ravines, April 1913, R Thaxter
(12, MCZC); St Augustine, April 1929, Darlington (28,
MCZC).
VATELLUS
BIFENESTRATUS
(ZIMMERMANN, 1921)
COMB. NOV.
(FIGS 274–287, 314)
Macrovatellus bifenestratus Zimmermann, 1921: 191;
Trémouilles, 1995: 27.
Type information: Lectotype (designated here to clarify assignment of this name with this species), in
ZSBS labelled, ‘Brazilien/Mato-Grosso Corumba/Type
[handwritten on blue, circular disc]/Samml.A. Zimmermann/Zool. Staatssig. München [blue label]/Holotypus Macrovatellus bifenestratus A.Zimm. Zool.
Staatsammlg.München [taxon name and author handwritten, red label].’ A single male paralectotype
accompanies the specimen labelled, ‘Brazilien/
Mato-Grosso Corumba/Type [handwritten on blue,
circular disc]/Samml.A. Zimmermann/Zool. Staatssig.
München [blue label]/Paratypoid Macrovatellus
bifenestratus A.Zimm. Zool. Staatsammlg.München
[taxon name and author handwritten].’
491
271
269
0.5mm
270
267
1.0mm
268
273
272
0.25mm
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.
492
K. B. MILLER
278
1.0mm
0.5mm
B
A
275
0.5mm
279
0.5mm
276
A
277
0.5mm
0.5mm
B
274
0.5mm
280
281
Figures 274–281. Vatellus bifenestratus. 274, dorsal habitus; 275, elytral sculpture in area indicated on habitus by dotted
Type locality: Brazil, Mato-Grosso, Corumba.
other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced (as in
Fig. 6); (2) elytra relatively coarsely punctate and with
surfaces between the punctures generally shiny
(Fig. 2); (3) with a transverse impunctate area on each
side of the midline on the pronotum; (4) eyes relatively
small (Fig. 274); (5) body relatively elongate and dorsoventrally depressed (Fig. 274); (6) lateral pronotal
margin distinctly curved (Fig. 274); and (7) male genitalia distinctive (Figs 282–285). The median lobe in
lateral aspect is slender with the apex long, straight
and apically narrowly rounded (Figs 282, 284). The
lateral lobe in lateral aspect has the apical portion
long and straight with an apical, curved, apically
sharply pointed flange (Fig. 285).
Description:
rounded (Fig. 274), dorsoventrally moderately compressed. Measurements are given in Table 1.
Coloration. Head, pronotum and elytron dark red;
venter red to testaceous in areas; appendages yellowred to dark red.
493
284
0.5mm
283
0.1mm
285
282
287
286
0.25mm
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.
494
K. B. MILLER
Sculpture and structure. Head densely punctate,
punctures moderate in size medially, finer anteriorly
on frons and clypeus, surface shiny and smooth; anteriorly clypeal margin distinctly protruding; eyes relatively small, slightly protuberant (Fig. 274). Pronotum
moderately punctate, covered with sculpturing consisting of extremely fine tubercles making surface shagrined except for two transverse areas on each side of
midline which are smooth, shiny and impunctate similar to Figure 2, each puncture with pale, short, fine
seta; pronotum cordate, broadest near anterior angles,
lateral margins broadly curved anteriorly, posterior
angles subacute (Fig. 274); lateral bead somewhat
obscured, especially posteriorly; transverse sulcus
absent. Elytron densely punctate, punctures moderately coarse, finer posteriorly, surface between punctures shiny, smooth (Fig. 275). Prosternum moderately
short; setose; prosternal process broad, flattened, lateral margins rounded, apex narrowly pointed. Metasternum coarsely punctate laterally, punctures
relatively sparse, finer medially, surface between
punctures shiny, smooth; anteromedial margin conspicuously swollen medially, ventrally produced.
Abdominal sterna finely punctate laterally, surface
smooth, shiny; sixth sternum with apical lobe produced, flattened, apically narrowly rounded (Fig. 280);
speleum broad, relatively short, apically narrowly
rounded (Fig. 281). Metacoxa with lateral portion relatively coarsely punctate over entire surface; lines
approximate basally, strongly divergent anteriorly
(Fig. 276); medial portion finely and densely punctate.
Metatrochanter short, rounded, ball-shaped, offset
(Fig. 277); metatarsal claws finely dentate.
curved, narrowed subapically, apex narrowly rounded,
slightly curved ventrally, without subapical setae but
with numerous distinct pores distributed randomly
over surfaces, basal portion broad, robust (Figs 282,
284); in ventral aspect narrow, apically with lateral
margins evenly convergent to relatively sharp apex
(Fig. 283). Lateral lobe with apical portion narrow,
margins parallel to acute apex, apex with prolonged,
apically sharply pointed and curved flange, apicomedial margin with area of dense, fine, elongate setae;
subapical stout, curved setae absent (Fig. 285).
robust, apical angle moderately rounded, anterior
margin concave, anterior lobe relatively short, slender
(Fig. 286). Gonocoxa elongate and slender, irregularly
curved, apex relatively narrowly rounded, apodeme
elongate and slender (Fig. 286). Bursa copulatrix
short and slender (Fig. 286); spermathecal duct moderately long, slender; receptacle small, much smaller
than spermatheca, spherical (Fig. 286); intermediate
duct moderately long, with several tight curves, particularly near spermatheca (Fig. 287); spermatheca
subspherical, triangular process large and very
broadly expanded; fertilization duct short and curved
(Fig. 286).
Sexually dimorphic characters. Male with weakly
developed mesofemoral brush of setae and moderately
developed mesotrochanteric setal brush. Male proand mesobasotarsomeres (Fig. 278) not strongly
expanded, only slightly more expanded than female
tarsomeres (Fig. 279), male protarsomeres relatively
slender, protarsomere III (Fig. 278A) much longer
than mesotarsomere III (Fig. 278B).
Distribution: This species is known from the type
locality, Corumba, Mato Grosso, Brazil and central
Bolivia (Fig. 314).
Phylogenetic relationships: This species is a member of
the V. grandis clade (Fig. 316). Within this group, the
species are extremely similar except for particular species-level autapomorphies, and this species does not
group with any other particular member of the group.
Material examined: Bolivia: Beni, Prov. Cercado,
9.5 km N Trinidad 14∞46¢34≤S 64∞58¢00≤W, 18 June
1999, KB Miller (1, KBMC).
VATELLUS DRYMETES MILLER
(FIGS 288–297, 313)
SP. NOV.
‘PERU: Madre de Dios; Rio Tambopata Res; 30 air
km.SW Pto.Maldonado,290 m 11-15 XI 1979 JB.
Heppner subtropical moist forest/HOLOTYPE Vatellus drymetes Miller 2004 [red label with double black
line border].’
Paratypes: 1 in USNM labelled same as paratype.
Type locality: Peru, Madre de Dios, Rio Tambopata
Res, 30 air km SW Pto. Maldonado.
other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced as in
Figure 6; (2) with pronotum and elytra relatively
coarsely punctate and with surfaces between the
punctures generally shiny, surface sculpture as in
Figure 2; (3) with a transverse impunctate area on
each side of the midline on the pronotum (as in
Fig. 262); (4) eyes moderate in size, not conspicuously
protuberant (Fig. 288); (5) body relatively elongate
and dorsoventrally depressed (Fig. 288); (6) lateral
pronotal margin moderately curved (Figs 288); and (7)
male genitalia distinctive (Figs 293–295). The median
lobe in lateral aspect is very long, curved and slender
with the apex extremely slender, and apically sharply
pointed (Fig. 293). The lateral lobe is irregular in
shape, very broad basally with the apical portion long
and slender with two or three long, stiff setae subapically on the ventral surface (Fig. 295).
495
0.5mm
289
0.5mm
293
294
0.5mm
290
1.0mm
0.5mm
295
291
292
0.5mm
B
288
A
0.5mm
296
297
Figures 288–297. Vatellus drymetes. 288, dorsal habitus; 289, metatrochanter and metafemur, anterior aspect; 290, elytral
sculpture in area indicated on habitus by dotted square; 291, medial portion of metacoxae; 292, male tarsomeres, dorsal aspect,
A, protarsus; B, mesotarsus; 293–295, male genitalia; 293, median lobe, right lateral aspect; 294, median lobe, ventral aspect;
295, right lateral lobe, lateral aspect; 296, sternum VI, ventral aspect; 297, speleum abdominal sterna V, VI, dorsal aspect.
Description:
rounded (Fig. 288), body dorsoventrally compressed.
ventral surfaces black except venter of head, pronotal
epipleuron, propleuron, prosternum and small portions
of posterior margins of abdominal sterna brown-red.
Sculpture and structure. Head finely and moderately densely punctate, slightly more dense anteriorly
on frons and clypeus; anterior clypeal margin strongly
anteriorly produced; eyes moderate in size, not
strongly protruding (Fig. 288). Pronotum with punctures relatively large laterally, denser medially and
along posterior margin, punctures with fine pale, moderately elongate setae, area between punctures
smooth, with dense shagrination consisting of dense
496
K. B. MILLER
field of very fine tubercles except for medial transverse
impunctate areas on each side of midline similar to
Figure 262; pronotum broadest anteromedially, lateral margins curved at extreme anterior end, pronotum cordate, posterior angles acute (Fig. 288); lateral
bead obscured except at extreme anterior end; transverse groove absent. Elytron strongly and densely
punctate over entire surface (Fig. 290), punctures
finer and slightly more dense apically, each puncture
with fine seta, surface similar to pronotum. Prosternum short, setose; prosternal process broad, flattened
margins rounded, convergent to triangular, pointed
apex. Metasternum laterally with moderately large
punctures, relatively sparsely distributed, medially
with more dense, finer punctures, surface smooth,
shiny; anteromedial margin strongly expanded ventrally into distinct tumidity. Abdominal sternal
sutures not unusually modified; sterna laterally with
moderately large, moderately dense punctures, interpuncture surface smooth, shiny; apical lobe on sixth
abdominal sternum strongly produced, long, apically
round, flattened (Fig. 296); speleum elongate, moderately narrow, apically narrowly rounded (Fig. 297).
Metacoxa with lateral portion with punctures moderately large, relatively dense, interpuncture surface
smooth, shiny; metacoxal lines strongly approximate
posteriorly, anteriorly strongly divergent (Fig. 291);
medial portion with surface shiny, finely punctate.
Metatrochanter strongly rounded, strongly offset
(Fig. 289); metatarsal claws minutely dentate.
extremely slender and narrowed to sharply pointed
apex, basal portion broad, broadly rounded; without
setae at apex (Fig. 293); in ventral aspect relatively
narrow, lateral margins parallel through most of
length, apically with margins evenly narrowed to
pointed apex (Fig. 294). Lateral lobe broad basally,
apex straight, elongate and slender, with distinct ventral furrow, apex narrowly rounded, with numerous
setae apicomedially and apicodorsally (Fig. 295).
Female genitalia. Female not observed.
brushes of setae. Female not observed, but male proand mesobasotarsomeres not strongly expanded, protarsomere III very long and slender (Fig. 292).
Intraspecific variation. No variation was noted in
the few specimens examined.
Natural history: This species has been collected from
290 m in subtropical moist forest.
Distribution: Vatellus drymetes is known only from the
type locality in Peru (Fig. 313).
of the V. grandis clade (Fig. 316). Within this group
this species does not group with any other particular
member of the group.
Etymology: This species is named from the Greek word
drymo, meaning ‘forest’, and the suffix -etes, meaning
‘to dwell’, in reference to the tropical forest area of the
type locality (Tambopata Reserve, Peru).
VATELLUS AMAE MILLER SP.
(FIGS 298–310, 314)
NOV.
‘BRAZIL: Para; Rio Xingu Camp (52∞22¢W 3∞39¢S)
Altamira (c. 60 km S) 15 October 1986 P. Spangler &
O. Flint/Colln.#24, pond at 2nd palm grove on trail 1/
HOLOTYPE Vatellus amae Miller 2004 [red label with
double black line border].’
Brazil: Para, Rio Iriri Camp, 52∞40¢W 3∞50¢S Altamira,
c. 100 km S, Lago Sao Joao north side of Iriri River, P.
Spangler and O. Flint (1, USNM); Para, Rio Xingu
Camp, 52∞22¢W 3∞39¢S Altamira, c. 60 km S, pond at
2nd palm grove on trail, P. Spangler and O. Flint (3,
USNM).
Type locality: Brazil, Para, Rio Xingu Camp, c. 60 km
S Altamira, 52∞22¢W 3∞39¢S.
other members of the genus by the following: (1) anterior margin of clypeus anteriorly produced (as in
Fig. 6); (2) with surfaces relatively coarsely punctate
and with surfaces between the punctures generally
shiny, surface sculpture as in Figure 2; (3) with a
transverse impunctate area on each side of the midline on the pronotum (as in Fig. 262); (4) eyes relatively small (Fig. 298); (5) body relatively elongate and
dorsoventrally depressed (Fig. 298); (6) lateral pronotal margin relatively weakly curved (Figs 298); and (7)
male and female genitalia distinctive (Figs 306–310).
The median lobe in lateral aspect is slender with the
apex narrowed to a very narrowly rounded apex that
is slightly curved ventrally (Fig. 306). The lateral lobe
has the apical portion long and narrow with two or
three long, stiff, subapical setae (Fig. 308). The female
receptacle is relatively small and spherical (Fig. 309).
Description:
rounded (Fig. 298), dorsoventrally compressed. Measurements are given in Table 1.
ventral surfaces dark red-brown. Appendages redbrown.
Sculpture and structure. Head finely and moderately densely punctate, slightly more dense anteriorly
on frons and clypeus; anterior clypeal margin rela-
497
302
0.5mm
0.1mm
A
B
299
0.5mm
303
0.5mm
300
B
A
301
0.5mm
298
0.5mm
304
305
Figures 298–305. Vatellus amae. 298, dorsal habitus; 299, elytral sculpture in area indicated on habitus by dotted square;
dorsal aspect.
tively strongly anteriorly produced; eyes moderate in
size, not strongly protruding (Fig. 298). Pronotum
with punctures relatively large laterally, denser medially and along posterior margin, punctures with fine
pale, moderately elongate setae, area between punctures smooth, shiny on anterior and medial portions of
pronotum, posterior half, lateral edges and a transverse anterior area with very fine shagrination consisting of dense field of very fine tubercles except for
transverse impunctate, smooth areas medially on each
side of midline similar to those shown in Figure 262;
pronotum broadest anteromedially, lateral margins
curved mainly in anterior half, moderately cordate,
posterior angles acute (Fig. 298); lateral bead
obscured except at extreme anterior end; transverse
groove absent. Elytron strongly and densely punctate
over entire surface, punctures finer and slightly more
dense apically (Fig. 299), each puncture with fine seta,
surface smooth and shiny between punctures. Prosternum short, setose; prosternal process broad, flattened
margins rounded, convergent to broadly rounded
apex, with few long, fine setae apically. Metasternum
laterally with moderately large punctures, relatively
sparsely distributed, medially with more dense, large
punctures, surface smooth, shiny; anteromedial margin strongly expanded ventrally into distinct tumidity.
498
K. B. MILLER
sterna laterally with moderately large, moderately
dense punctures, interpuncture surface smooth,
shiny; apical lobe on sixth abdominal sternum
strongly produced, long, apically round, flattened
(Fig. 304); speleum elongate, relatively narrow, apically narrowly rounded (Fig. 305). Metacoxa with lateral portion with punctures moderately large,
relatively dense, interpuncture surface smooth, shiny;
metacoxal lines strongly approximate posteriorly,
anteriorly strongly divergent (Fig. 300); medial portion with surface shiny, finely punctate. Metatrochanter strongly rounded, strongly offset (Fig. 301);
metatarsal claws minutely dentate.
Male genitalia. Median lobe in lateral aspect moderately straight in apical half, slender, apically gradually narrowed to slender, slightly hooked, narrowly
rounded apex (Fig. 306); in ventral aspect with lateral
margins evenly and gradually convergent to narrowly
rounded to nearly pointed apex (Fig. 307). Lateral lobe
long, basally slender, apical portion straight and narrow to rounded apex; with 2–3 subapical, stout, elongate, curved setae on ventral margin (Fig. 308).
robust, apical angle moderately narrowly rounded,
anterior margin relatively narrowly concave, anterior
lobe relatively short, narrow, apex rounded (Fig. 309).
Gonocoxa short, robust, rounded, apex broadly
rounded, apodeme elongate, slender (Fig. 309). Bursa
copulatrix robust, broad, curved dorsally and gradually narrowed to base of spermathecal duct (Fig. 309);
spermathecal duct long; receptacle small, about quarter size of spermatheca, spherical (Fig. 309); intermediate duct relatively short, slender, only slightly
sinuate (Fig. 310); spermatheca elongate-spherical,
large, triangular process moderately long, very
broadly expanded; fertilization duct moderately long,
relatively straight (Fig. 309).
brushes of setae. Male with pro- and mesobasotarsomeres (Fig. 302) only somewhat expanded compared
with female (Fig. 303), both male and female with protarsomere III relatively elongate (Figs 302A, 303A),
but slightly longer in female (Fig. 303A).
Intraspecific variation. This species is relatively constant in characters in the few specimens examined.
There is some minor variation in the extent of the
microsculpture present on the pronotum.
Natural history: Specimens have been collected from
ponds in palm woods.
Distribution: This species is known only from a few
close localities in Brazil (Fig. 314).
of the V. grandis clade (Fig. 316). Within this group
this species does not group with any other particular
member of the group.
Etymology: This species is named in honour of my
wife, Amy Beth M. Miller.
CLADISTIC ANALYSIS
Taxa: All extant species of New World Vatellini were
included in a parsimony analysis of 43 morphological
characters. Also included were several species of African Derovatellus and the south-east Asian species,
D. orientalis. Non-vatelline taxa included were other
Hydroporinae including Stictotarsus striatellus
(LeConte), Hydroporus notabilis LeConte, Hygrotus
impressopunctatus (Schaller), Celina hubbelli Young
and Laccornis difformis (LeConte) (see Table 1). The
cladograms were rooted at L. difformis based on evidence from Wolfe (1985, 1988) and Miller (2001) that
this genus is the sister to a clade containing all other
known genera of Hydroporinae. Coding of Calicovatellus petrodytes resulted in a considerable amount of
missing data for this taxon which severely degraded
resolution due to the taxon’s tendency to group with
other taxa in multiple, disparate places because of
ambiguities in coding. Therefore, this taxon was both
included and excluded from analyses to examine the
effects of doing so.
Character analysis: The data matrix was constructed
and edited using the program WinClada (Nixon, 1999–
2000). Data were analysed using the program NONA
(Goloboff, 1995) as implemented in WinClada heuristics. The commands ‘hold 10000’, ‘hold/20’, ‘mult*50’
and ‘max*’ were used to find the most parsimonious
trees. Character state distributions were examined
under various optimizations using WinClada. Trees
were examined and saved and the strict consensus
was calculated in WinClada. Branch support values
(Bremer, 1994) were calculated using NONA by reading in the consensus of the most parsimonious cladogram and using the commands ‘hold 10000’,
‘suboptimal 20’ and ‘bsupport 20’.
The character matrix is shown in Table 2. The following characters were coded for analysis.
HEAD
1. Anterior clypeal margin: (0) rounded anteriorly
(Fig. 7); (1) produced anteriorly (Fig. 6).
PROTHORAX
2. Lateral crenulations on pronotum: (0) absent; (1)
present (Figs 1, 262).
499
0.5mm
307
308
306
0.2mm
310
309
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.
3. Prosternal process: (0) not attaining metasternum,
mesocoxal cavities contiguous (Figs 21, 22); (1) attaining metasternum, mesocoxal cavities separated.
4. Prosternal process apex: (0) relatively truncate, apically convex (Fig. 22); (1) flattened, apically sharply
angulate (Fig. 21).
5. Pronotum surface sculpture: (0) without shagrination; (1) shagrined except for transverse smooth sections medially on each side of midline (Fig. 262).
Some species with shagrination do not display it to
the same degree in all specimens. In some the shagrination occurs also on the elytra.
500
K. B. MILLER
Table 2. Data matrix for 35 terminals and 43 morphological characters used in cladistic analysis of Vatellini. Additive
characters are marked with ‘+’, inapplicable character states with ‘–’ and unobserved character states with ‘?’
00000 00001 11111 11112 22222 22223 33333 33334 444
12345 67890 12345 67890 12345 67890 12345 67890 123
++
+
+
Laccornis difformis
Celina hubbeli
Hydroporus notabilis
Stictotarsus striatellus
Calicovatellus petrodytes
Derovatellus dagombai
D. nyanzae
D. lugubris
D. bisignatus
D. bruchi
D. orientalis
D. wewalkai
D. fasciatus
D. decellei
D. spangleri
D. roosevelti
D. floridanus
D. peruanus
D. lentus
Vatellus sahlbergi
V. ventralis
V. mexicanus
V. maculosus
V. lateralis
V. haagi
V. wheeleri
V. perforatus
V. tarsatus
V. annae
V. pilacaudus
V. grandis
V. bifenestratus
V. drymetes
V. amae
00110
00110
00110
00110
0?00?
01000
01000
01000
01000
01000
01000
01000
01000
01000
01000
01000
01000
01000
01000
01010
11010
01010
01010
01010
01000
01010
11010
01010
01010
01010
11011
11011
11011
11011
6. Pronotum: (0) broadest at or near posterior margin
(Fig. 68); (1) broadest medially, lateral margins
strongly rounded (Fig. 51); (2) broadest anterad of
middle, pronotum cordate (Fig. 1).
7. Prosternal pore: (0) not located medially, located
along anterior margin near lateral edge of prosternum (Fig. 18); (1) located medially and posterad
(Fig. 19).
MESO/METATHORAX
8. Metepisternum: (0) not reaching mesocoxal cavity
externally (Fig. 20); (1) reaching mesocoxal
cavity.
00100
00100
00100
00103
1?103
01000
01001
01001
01001
11001
11000
11001
11001
11001
01001
01001
01001
01001
01001
21002
21000
21002
21002
21002
21002
21002
21000
21000
21000
21002
21010
21010
21010
21010
-00--00--00--00--????
-3102
03102
03102
03102
13102
-3102
13101
13100
03101
13103
13103
13103
13103
13103
-1122
-1122
-1112
-1112
-1112
-1112
-1112
-0122
-2122
-2122
-2122
-0122
-0122
-0122
-0122
---00
---00
---00
---00
????0
01010
01010
01110
01110
00010
00010
00010
00010
00010
00010
10010
10010
10010
10010
00011
00011
00011
00011
00012
00011
10011
00012
00012
00012
00012
00012
00012
00012
00012
00000
00000
00100
00100
0010?
00101
00101
00101
00101
00101
00101
00101
00101
00101
00101
00101
00101
00101
00101
11121
1112?
11121
11121
11121
11121
11121
11121
11121
11111
11111
11121
11121
11121
11121
00000
00000
00000
00000
?????
00001
00001
00001
00001
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
10000
?????
10000
10100
10100
10100
10100
10010
10110
10110
10100
11000
10000
11000
11010
000-000-00000
00000
?????
00000
00000
00000
00000
00000
00011
00011
00011
00011
01000
01000
01000
00010
01011
00100
???00
10100
10100
10100
10100
10100
00100
00100
00100
00100
10100
00100
001??
00100
00000
00000
00100
00100
?????
11100
11100
11100
11100
11100
11100
11100
11100
11100
11100
11100
11100
11100
11100
10110
10110
10110
10110
10110
10110
10110
10110
10110
10111
10111
10110
10110
?????
10110
0000000
000
???
110
110
110
110
110
110
110
110
110
110
110
110
110
111
010
010
010
010
010
010
010
010
010
011
011
011
011
???
011
9. Anteromedial margin of metasternum: (0) not ventrally swollen; (1) ventrally swollen and deflexed
(Fig. 20).
ELYTRA
10. Maculae on elytron: (0) absent; (1) a light, transverse, subapical fasciae or subapical, lateral macula present (Figs 75, 88; see following character);
(2) several distinct maculae along lateral margin
subapically, medially and at humeral angle
(Fig. 174); (3) longitudinally vittate.
11. Maculae on elytron: (0) a subapical, lateral macula
only present along lateral margin (Fig. 75); (1) a
0
0
501
500 Miles
500 KM
Figure 311. Vatellus species, distribution. = V. lateralis, = V. maculosus, = V. sahlbergi.
subapical pale transverse fascia extending from
lateral margin nearly to elytral suture (Fig. 88).
Those characters with states 0, 2 and 3 in character 10 are coded as inapplicable for this character.
Some species of Derovatellus have a distinct transverse fascia extending nearly across the entire
elytron, whereas others have a distinct lateral
macula at approximately the same position as the
origin of the fascia.
12. Elytral surface sculpture: (0) absent or with very
few, extremely fine punctures, very smooth (Fig. 2)
(some of these specimens bear conspicuous shagrination which is not regarded as the same character because the basic surface sculpture in these
502
K. B. MILLER
0
0
500 Miles
500 KM
Figure 312. Vatellus mexicanus distribution.
specimens is apparently smooth. See character 5.);
(1) with fine irregular cells (Fig. 5); (2) with transverse, curved striae with series of small setae arising from them (Fig. 3); (3) with narrow, transverse
cells (Fig. 4).
22.
23.
ABDOMEN
13. Speleum: (0) absent; (1) present (e.g. Figs 28–37).
14. Apical lobe at apex of sixth abdominal sternum:
(0) absent or weakly developed; (1) present (e.g.
Fig. 209).
15. Length of speleum: (0) very short (Fig. 31); (1)
medium length, not extending beyond anterior
margin of abdominal sternum V (Fig. 32); (2)
long, extending distinctly beyond abdominal sternum V (Fig. 138); (3) extremely long, extending
nearly to thorax (Fig. 101) (additive).
16. Apex of speleum: (0) not upturned or bulbous; (1)
distinctly and characteristically upturned and
bulbous (Figs 28, 30).
17. Divisions of speleum: (0) not divided into two distinct parts; (1) with broad basal part, with apical
part distinctly separated (Figs 34–37).
18. Apex of speleum: (0) not broadly expanded apically; (1) very broadly expanded (Figs 36, 37).
LEGS
19. Legs: (0) relatively short; (1) very long.
20. Metatrochanter apex: (0) not strongly modified;
(1) offset, but elongate, apex rounded (e.g.
Fig. 134); (2) short, apex strongly rounded, laterally offset (Figs 267, 268) (additive).
21. Metatrochanter basal expansion: (0) without
basal modification; (1) with basal circular expansion which extends ventrad under medial portion
24.
25.
of metacoxa when legs anteriorly flexed (Figs 267,
268).
Metatrochanter emargination: (0) not proximally
emarginate along ventral margin; (1) constricted
proximally along ventral margin (Figs 267, 268).
Posterior margin of metacoxal process: (0) without
continuous margin, interrupted medially; (1) with
continuous margin.
Metacoxal lines: (0) relatively parallel-sided or
only slightly diverging anteriorly (e.g. Fig. 97); (1)
approximate posteriorly, somewhat diverging
anteriorly (e.g. Fig. 232); (2) approximate posteriorly, strongly diverging anteriorly (e.g. Fig. 119)
(additive).
Dense brush of setae at base of male mesofemur
and mesotrochanter: (0) absent; (1) present.
MALE
GENITALIA
26. Basal portion of lateral lobes: (0) not unusually
expanded; (1) longitudinally expanded, very long
(Fig. 143).
27. Few, long stiff setae subapically on lateral lobe:
(0) absent; (1) present (Fig. 271).
28. Very short spines on ventral, apical surfaces of
median lobe: (0) absent; (1) present (Fig. 158).
29. Median lobe apex: (0) not slender and curved; (1)
elongate, slender and curved ventrally (Figs 225,
239).
30. Robust, stout spines on apicomedial margin of
lateral lobe: (0) absent; (1) present (Fig. 50).
31. Median lobe: (0) without long basal piece; (1) with
long basal piece (Fig. 141).
32. Two lateral or sublateral processes on male
median lobe: (0) absent; (1) present (Fig. 83).
33. Strut of male genitalia: (0) small, straight
(Fig. 46); (1) long, medially angulate (Fig. 43).
503
" "
0
0
500 Miles
500 KM
Figure 313. Vatellus species, distribution. = V. annae, = V. drymetes, = V. pilacaudus, + = V. haagi, = V. tarsatus.
FEMALE
GENITALIA
34. Intermediate duct of female genitalia: (0) short to
only moderately long; (1) extremely long and
coiled (Fig. 105).
35. Receptacle: (0) large, spherical, globular (Figs 80,
145); (1) small (Figs 105, 272).
36. Apical, flat, truncate process on spermatheca: (0)
absent; (1) present (Fig. 60). The process in
D. roosevelti (Fig. 80) is apically rounded, but the
process seems clearly homologous with the truncate process of other taxa.
37. Spermatheca: (0) subspherical (Fig. 186); (1) long
and curved (Fig. 60).
504
K. B. MILLER
0
0
Figure 314. Vatellus
+ = V. wheeleri.
species,
distribution.
= V. amae,
38. Receptacle: (0) absent; (1) present (Figs 60, 186).
39. Intermediate duct: (0) not invaginated into spermatheca; (1) invaginated. In species of Vatellus
the intermediate duct extends into the spermatheca. This invagination is visible in cleared
specimens.
500 Miles
500 KM
= V. bifenestratus,
= V. grandis,
= V. perforatus,
40. Intermediate duct: (0) not coiled; (1) coiled on dorsal surface of spermatheca (Fig. 258).
41. Spermathecal glands: (0) with small glands over
entire surface of spermatheca (Fig. 186); (1) with
small glands concentrated near posterior apex of
spermatheca (Fig. 105).
2
315
505
Laccornis difformis
Laccornis difformis
Celina hubbeli
Celina hubbeli
C petrodytes
D dagombai
D bruchi
D nyanzae
1
2
D orientalis
D lugubris
1
D wewalkai
D bisignatus
D fasciatus
3
D bruchi
1
D decellei
D orientalis
2
316
D dagombai
D wewalkai
1
1
D nyanzae
D fasciatus
D lugubris
1
D decellei
D bisignatus
D
spangleri
1
2
D spangleri
D roosevelti
1
1
D roosevelti
1
D floridanus
9
D floridanus
D peruanus
1
1
D peruanus
D lentus
D lentus
V ventralis
V ventralis
V sahlbergi
V sahlbergi
V mexicanus
V mexicanus
1
V maculosus
1
1
V maculosus
V
lateralis
1
V lateralis
8
6
V haagi
V haagi
V wheeleri
V wheeleri
V
perforatus
V perforatus
V tarsatus
1
V tarsatus
2
V annae
2
2
V annae
1
1
V pilacaudus
V pilacaudus
V grandis
V grandis
3
V bifenestratus
3
V bifenestratus
V drymetes
V drymetes
V amae
V amae
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.
42. Large field of microspines on ventral surface of
common oviduct: (0) absent; (1) present. Members
of Vatellini possess a large ventral region of
microspines near the base of the common oviduct.
43. Female receptacle: (0) distinctly larger than half
size of spermatheca (Fig. 201); (1) distinctly
smaller than half size of spermatheca (Fig. 286).
RESULTS AND CONCLUSIONS
The analysis with all terminals included resulted in
31 most parsimonious trees (length = 75, CI = 70,
RI = 93), the consensus of which is shown in
Figure 315. Inclusion of Calicovatellus resulted in
decreased resolution within Vatellini, but the tribe,
including C. petrodytes, is monophyletic in this analysis (Fig. 315). Within the tribe a sister-group relationship between Calicovatellus and the remaining
members of the tribe is one of several most parsimonious solutions, whereas the other solutions include
the resolution of Calicovatellus as sister to a variety of
groups within the Derovatellus clade. Calicovatellus
does not group with Vatellus in any of the most parsimonious solutions. The lack of resolution resulting
from inclusion of Calicovatellus appears to be the
result of absence of character state information for a
506
K. B. MILLER
Laccornis difformis
Celina hubbeli
10
3
1730
1 1
4 10123741
2338
0 1 3 1 1
1 1
11
V ventralis
V sahlbergi
1 10
V mexicanus
12
1
14 31
0 2
10
D dagombai
0
1 6 1420212224263339
D nyanzae
18
1 2 2 1 1 1 2 1 1 1
D lugubris
1
D bisignatus
D spangleri
1532
D floridanus
16
3 1
D32roosevelti
1
34
D peruanus
3543
0
1
1
1 1
1 1
20
2
29
318
D lentus
1
D bruchi
6
V lateralis
V haagi
16
0
1
V wheeleri
1
V maculosus
V bifenestratus
28
5 9 43
1 1 1
20
4 2
31
V grandis
V drymetes
29
1
V amae
1
V perforatus
1 1228
V tarsatus
244043
V annae
0 2 1
27
1
1029
10
D orientalis
0
1
D wewalkai
15 15
1 1
D fasciatus
0 11
1
D decellei
1
0
6 1420212224263339
V
ventralis
12
1
2 2 1 1 1 2 1 1 1
V sahlbergi
10
1
V mexicanus
20
1431
2
V lateralis
4 2
1 1
28
V haagi
0 16
1
V wheeleri
1
V maculosus
1228
V tarsatus
244043
V annae
2 1
10 29
2029
1 1 1
V pilacaudus
2 0
2 1
V
perforatus
1
V amae
5 9 2743
2 3 7 8 1319253642
1 0 1 0 1 1 1 1 1
6 1420212224263339
2 2 1 1 1 2 1 1 1
317
V sahlbergi
V mexicanus
2
2
31
1
6 1420212224263339
2 1 1 1 1 2 1 1 1
1 14
0
20
1220
0 2
1
1
10
V ventralis
V sahlbergi
V mexicanus
31
V grandis
27
1
1 1 1
V
drymetes
29
1
V amae
1
V perforatus
29
1 1228
V tarsatus
1
244043
V annae
0 2 1
10 29
1 1 1
2 0
20
1431
2
1 1
1228
2 1
2029
V lateralis
V haagi
0 16
1
V wheeleri
1
V maculosus
V tarsatus
244043
V annae
28
42
10 29
V pilacaudus
2 0
V perforatus
V bifenestratus
5 9 2943
1 1 1
2 1
1
31
1
1 1 0 1
319
27
1
V bifenestratus
V grandis
1
V drymetes
V mexicanus
V lateralis
28
V haagi
0 16
1
V wheeleri
1
6 101420212224263339
V maculosus
2 2 1 1 1 1 2 1 1 1
V ventralis
V bifenestratus
5 9 43
12
31
1
14
2
V pilacaudus
V grandis
V
drymetes
29
V amae
1
1
0 31
4 2
1 2
320
29
1 1 1 1
14
10
2 0
1
27
1
V pilacaudus
1 1 1
3435
20
V lateralis
V haagi
16
0
1
V wheeleri
1
V maculosus
V sahlbergi
V ventralis
V bifenestratus
1 10
5 9 43
4 2
28
31
1 0
1 1 1
1220
0 2
321
29
1
V grandis
V
drymetes
29
1
V amae
1
V perforatus
1 12 28
V tarsatus
244043
V annae
0 2 1
10 29
27
1
1 1 1
2 0
V pilacaudus
507
Figures 317–321. Five equally parsimonious arrangements resulting from cladistic analysis of Vatellini not including Calicovatellus petrodytes (length = 74, CI = 70, RI = 93). 317, one arbitrarily chosen cladogram showing all taxa in analysis.
Oblique line indicates clade in which there are multiple alternative equally parsimonious arrangements of taxa; 318–321,
four additional topologies for clade with alternative parsimonious arrangements. Black hashmarks = non-homoplasious
character state changes, white hashmarks = homoplasious character state changes or reversals, numbers above
hashmarks = character numbers, numbers below hashmarks = derived state at character change. Characters mapped using
WinClada ‘fast’ optimization.
large number of characters for Calicovatellus rather
than conflict among characters.
Analysis of these taxa, discluding the Calicovatellus
terminal, resulted in five most parsimonious trees
(length = 73, CI = 72, RI = 93). Monophyly of the tribe
Vatellini (not including Calicovatellus) is supported by
several synapomorphies, including: (1) lateral crenulations present along lateral margin of pronotum
(Character 2, Fig. 262); (2) the prosternal process not
reaching the mesosternum and with the mesocoxae
contiguous (Character 3, Figs 21, 22); (3) the prosternal pore shifted posteromedially (Character 7,
Fig. 19); (3) the metepisternum not reaching the mesocoxal cavity externally (Character 8, Fig. 20); (4)
speleum present (Character 13, Figs 28–30); (5) legs
very long (Character 19); (6) males with a cluster of
setae proximally along the ventral surface of the
mesofemur and mesotrochanter (Character 25); (7) a
prominent truncate process present on the spermatheca (Character 36, Figs 60, 186); and (8) a large region
of small microspines on the ventral surface of the common oviduct (Character 42). Although outgroup sampling of the large and diverse Hydroporinae was not
extensive, Vatellini is resolved nested within the
Hydroporini (Fig. 315).
Within the Vatellini there are two large groups. One
of these includes members of the genus Derovatellus
with Mesovatellus nested within it, requiring placement of the latter genus as a junior synonym of the
former based on the Hennigian principle of naming
only monophyletic groups. In addition, although the
subgenus D. (Varodetellus) is resolved as monophyletic within Derovatellus, the members of Derovatellus
s.s. are not together monophyletic and those characters used by Biström (1979) to define this group are
not synapomorphies, according to this analysis. Therefore, use of these two subgenera should not be
continued, with Varodetellus a junior synonym of
Derovatellus.
Synapomorphies of Derovatellus include the elongate, medially bent spermatheca (Character 37), the
prosternal process apically rounded (Character 4,
homoplasious), and the small spermathecal glands
concentrated near the posterior apex of the spermatheca (Character 41). Within this group there are three
main clades. First, the New World species except
D. bruchi form a monophyletic group that is sister to a
clade containing African species with the austral
South American species D. bruchi at its base. Sister to
the clade containing these species is a clade containing
another group of African species with the Oriental
D. orientalis at its base.
The second group includes Vatellus and Macrovatellus (Fig. 316). The single species previously placed in
Vatellus, V. tarsatus, is nested within Macrovatellus in
all most parsimonious trees. Because there is no clear
justification for dividing these genera into multiple
monophyletic genera (Fig. 316), Macrovatellus is
hereby synonymized with Vatellus.
Synapomorphies of Vatellus include the lateral lobes
strongly elongated and expanded basally and fused
to each other along dorsal margin (Character 26,
Fig. 143), the metatrochanter with a large basal, circular lobe that extends ventrad of the metacoxa when
the leg is anteriorly flexed and with a medial emargination (Characters 21, 22, Figs 267, 268), the metatrochanter moderately to strongly offset from
metafemur (Character 20, Figs 267, 268), the metacoxal lines relatively closely approximated posteriorly
and moderately to strongly divergent anteriorly
(Character 24, Fig. 119), and the intermediate duct
between the receptacle and spermatheca invaginated
into the spermatheca (Character 39). Within Vatellus,
the resolution is not strong mainly because of missing
female characters in some species for which females
are unknown, and some character conflict. However,
within all trees two larger clades are evident. First are
those species centred around V. grandis that are dorsoventrally flattened, possess an anteriorly strongly
protruding clypeal margin and have similar dorsal
sculpture patterns, among other things. The other
large clade includes those species centred around
V. mexicanus that are dorsally maculate and have a
similar type of surface sculpture among other less-discrete similarities. Those taxa in a polytomy at the base
are not especially similar to these other groups. Of
these taxa, V. tarsatus, V. annae and V. pilacaudus
are each similar to each other. Both V. tarsatus and
V. annae have extremely similar male genitalia with
the apex upturned and sharply pointed and the apical
portion of the lateral lobe very long and slender. Vatellus annae and V. pilacaudus have the female intermediate duct tightly coiled on the dorsal surface of the
spermatheca (Figs 244, 258). These three also share a
508
K. B. MILLER
similar dorsal surface sculpturing and general body
shape, suggesting that they may be closely related.
Other species in this polytomy are not so clearly
related to each other or to any in the resolved clades.
ACKNOWLEDGEMENTS
I thank Q. D. Wheeler for his inspiration and support,
especially at the beginning of this project. I thank the
numerous people who provided valuable material,
especially R. E. Roughley, P. Mazzoldi, L. Hendrich
and M. Balke. I thank C. Marshall for advice on several issues regarding morphology. Thanks also to M. F.
Whiting for allowing me liberty to continue my work
on diving beetles. Finally I thank my wife, A. B. M.
Miller, for her continued support of my work. Portions
of this work were funded by NSF grants #DEB0073088 to Q. D. Wheeler and M. Luckow and #DEB0120718 to M. F. Whiting.
REFERENCES
Aubé C. 1836–1838. Hydrocanthares. In: Dejean PF, ed. Iconographie et histoire naturelle des coléoptères d’Europe.
Paris: Méquignon-Marvis, 1–64(1836), 65–224(1837), 225–
416(1838).
Aubé C. 1838. Hydrocanthares et Gyriniens. In: Dejean
PF, ed. Species géneral des coléoptères de la collection de
M. le Comte Dejean. Paris: Méquignon Père et Fils, XVI
+ 804.
Balfour-Browne F. 1940. British water beetles, Vol. 1. London: Ray Society.
Balfour-Browne F. 1944. The wing-venation of the Adephaga
(Coleoptera) with special reference to the Hydradephaga and
some homologies with the Polyphaga. Journal of the Royal
Microscopical Society 63: 55–84.
Biström O. 1979. A revision of the genus Derovatellus Sharp
(Coleoptera, Dytiscidae) in Africa. Acta Entomologica Fennica 35: 1–28.
Biström O. 1980a. Derovatellus nyanzae, new species, with
notes on the genus in Africa (Coleoptera: Dytiscidae). Entomologica Scandinavica 11: 366–368.
Biström O. 1980b. Notes on non-African Derovatellus species
(Coleoptera: Dytiscidae). Entomologica Scandinavica 11:
78–80.
Biström O. 1981. Derovatellus hancocki, new species
(Coleoptera, Dytiscidae) with notes on the genus in Africa.
Annales Entomologici Fennici 47: 69–72.
Biström O. 1982. Derovatellus onorei new species from Congo
(Coleoptera: Dytiscidae). Entomologica Scandinavica 13:
267–268.
Biström O. 1983. New records of Hyphydrus and Derovatellus, with descriptions of 2 new Hyphydrus species
(Coleoptera: Dytiscidae). Acta Entomologica Fennica 49:
117–121.
Biström O. 1984. Description of Hyphydrus holomelas new
species and Hyphydrus gibbosus new species with new
records of Derovatellus, Hyphydrus, Yola, and Yolina
(Coleoptera: Dytiscidae). Annales Entomologici Fennici 50:
89–92.
Biström O. 1986. Derovatellus intermedius new species and
notes on the genus in the Omer-Cooper collection (Coleoptera:
Dytiscidae). Acta Entomologica Fennica 52: 44–45.
Biström O, Nilsson AN. 2003. Taxonomic revision and cladistic analysis of the genus Peschetius Guignot (Coleoptera:
Dytiscidae). Aquatic Insects 25: 125–156.
Biström O, Roughley RE. 1982. Notes on Derovatellus mocquerysi (Coleoptera: Dytiscidae). Entomologica Scandinavica 13: 138–139.
Blackwelder R. 1944. Checklist of the Coleopterous insects of
Mexico, Central America, the West Indies, and South America. Bulletin of the United States National Museum 185: 1–
188.
van den Branden C. 1885. Catalogue des coléoptères carnassiers aquatiques (Haliplidae, Amphizoïdae, Pelobiidae et
Dytiscidae). Annales de la Société Entomologique de Belgique 29: 5–118.
Bremer K. 1994. Branch support and tree stability. Cladistics
10: 295–304.
Buquet L. 1840. Description de plusieurs coléoptères nouveaux appartenant aux genres Lebia, Vatellus, Acmaeodera,
Hammaticherus et Leptura. Annales de la Société Entomologique de France 9: 393–400.
Crowson RA. 1955. The natural classification of the families
of Coleoptera. London: N. Lloyd.
Epler JH. 1996. Identification manual for the water beetles of
Florida (Coleoptera: Dryopidae, Dytiscidae, Elmidae, Gyrinidae, Haliplidae, Hydraenidae, Hydrophilidae, Noteridae,
Psephenidae, Ptilodactylidae, Scirtidae). Tallahassee:
Bureau of Water Resource Protection, Florida Department of
Environmental Protection.
Fall HC. 1932. Random notes and descriptions (Coleoptera).
Bulletin of the Brooklyn Entomological Society 27: 145–148.
Franciscolo ME. 1979. Coleoptera. Haliplidae, Hygrobiidae,
Gyrinidae, Dytiscidae. Bologna: E. Calderini.
Gemminger M, Harold EV. 1868. Catalogus Coleopterorum
hucusque descriptorum synonymicus et Systematicus. Tomus
II. Dytiscidae, Gyrinidae, Hydrophilidae, Staphylinidae, Pselaphidae, Gnostidae, Paussidae, Scydmaenidae, Silphidae,
Trichopterygidae, Scaphidiidae. Monachii: E.H. Gummi,
425–752.
Goloboff P. 1995. NONA, Version 2.0. Fundación e Instituto
Miguel Lillo.
Goodliffe FD. 1939. The taxonomic value of the wing venation
in the larger Dytiscidae (Coleoptera). Transactions of the
Society of British Entomology 6: 23–38.
Guignot F. 1946. Génotypes des Dytiscoidea et des
Gyrinoidea. Revue Française d’Entomologie 13: 112–118.
Guignot F. 1955. Description de nouveaux Dytiscidae principaliemente de l’Amerique de Sud. Bulletin Institut Royal des
Sciences Naturelles de Belgique 31: 1–12.
Guignot F. 1957. Beiträge zur Kenntnis der Insekten fauna
Boliviens. Teil II. Coleoptera. I. Dytiscidae (2. Contribution à
l’etude des dytiscides sudaméricains). Opuscula Zoologica 6:
1–10.
Guignot F. 1959. Revision des Hydrocanthares d’Afrique
(Coleoptera Dytiscoidea), Première partie. Annales du
Musée Royal du Congo Belge Tervuren (Belgique) (ser. 8) Sciences Zoologiques 70: 1–316.
ICZN. 1992. Opinion 1681. Vatellus Aubé, [1837] (Insecta,
Coleoptera): conserved. Bulletin of Zoological Nomenclature
49: 165–166.
Jasper S, Challet GL. 2002. A genus and species new to the
U.S.A., Macrovatellus mexicanus Sharp (Coleoptera: Dytiscidae), in South Texas, with habitat notes. Coleopterists Bulletin 56: 142–143.
Laporte FL. 1834–1835. Études entomologiques ou description d’Insectes nouveaux et observations sur leur synonymie.
Première partie. Paris: Méquignon-Marvis Père et Fils, 1–
94(1834), 95–159(1835).
Laporte FL. 1840. Histoire naturelle des animaux articulés.
1: 1–324.
Lawrence JF, Britton EB. 1994. Australian beetles. Carlton,
Victoria: Melbourne University Press.
Leech HB. 1948. Contributions toward a knowledge of the
insect fauna of Lower California, 11. Coleoptera: Haliplidae,
Dytiscidae, Gyrinidae, Hydrophilidae, Limnebiidae. Proceedings of the California Academy of Science 24: 375–484.
Leech HB, Chandler HP. 1956. Aquatic Coleoptera. In:
Usinger RL, ed. Aquatic insects of California with keys to
North American genera and California species. Berkeley, CA:
University of California Press, 293–371.
Leng CW. 1920. Catalogue of the Coleoptera of America north
of Mexico. Mt. Vernon, NY: J.D. Sherman.
Miller KB. 2001. On the phylogeny of the Dytiscidae
(Coleoptera) with emphasis on the morphology of the female
reproductive tract. Insect Systematics and Evolution 32: 45–
92.
Miller KB, Lubkin SH. 2001. Calicovatellus petrodytes, a
new genus and species of primitive vatelline diving beetle
(Coleoptera: Dytsicidae: Hydroporinae: Vatellini) from the
Miocene Barstow Formation, southern California, USA.
Journal of Paleontology 75: 890–894.
Miller KB, Nilsson AN. 2003. Homology and terminology:
Communicating information about rotated structures in
water beetles. Latissimus 17: 1–4.
Nilsson AN. 1991. Case 2742. Vatellus Aubé, 1837 (Insecta,
Coleoptera): proposed conservation. Bulletin of Zoological
Nomenclature 48: 36–37.
Nilsson AN. 2001. Dytiscidae. World Catalogue of Insects 3: 1–
395.
Nilsson AN, Roughley RE, Brancucci M. 1989. A review of
the genus- and family-group names of the family Dytiscidae
Leach (Coleoptera). Entomologica Scandinavica 20: 287–316.
Nixon KC. 1999–2002. Winclada, Version 1.0000. Published
by the author.
Omer-Cooper J. 1958. Dytiscidae (Coleoptera) from Nyasaland and Southern Rhodesia. IV. Vatellinae, Methlinae and
Hydroporinae, Hyphydrini. Journal of the Entomological
Society of South Africa 21: 249–260.
Régimbart M. 1878. Étude sur la classification des Dytiscidae. Annales de la Société Entomologique de France 5: 8:
447–466+pl, 10: 1–28.
509
Régimbart M. 1889. Dytiscidae et Gyrinidae nouveaux ou
rares de la collection du Musée royal de Leyde. Notes from
the Leyden Museum 11: 51–63.
Régimbart M. 1895. Révision des Dytiscidae et Gyrinidae
d’Afrique, Madagascar et îles voisines. Memoires de la
Société Entomologique de Belgique 4: 1–244.
Régimbart M. 1899a. Revision des Dytiscidae de la région
Indo-Sino-Malaise. Annales de la Société Entomologique de
France 68: 186–367.
Régimbart M. 1899b. Viaggio del Dott. Alfredo Borelli nel
Chaco Boliviano e nella Repubblica Argentino xv. Dytiscidae
de la Bolivie. Bollettino dei Musei di Zoologia ed Anatomia
Comparata della R. Università Di Torino 14: 1–2.
Régimbart M. 1903. Liste des Dytiscidae & Gyrinidae recueillis par le Dr. Philippe Silvestri dans l’Amérique méridionale
de 1898 à 1900. Bollettino della Società Entomologica Italiana 35: 46–74.
Régimbart M. 1904. Dytiscides et gyrinides recuellis au
Vénézuéla et à la Guyarre par M.F. Geay et faisant partie
des collections du Muséum d’Histoire Naturelle. Bulletin du
Muséum National d’Histoire Naturelle 1904 (5): 224–226.
Roughley RE, Larson DJ. 2001. Dytiscidae Leach, 1815. In:
Arnett RH, Thomas MC, eds. American beetles, Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia.
Boca Raton, FL: CRC Press, 156–186.
Roughley RE, Pengelly DH. 1981. Classification, phylogeny,
and zoogeography of Hydaticus Leach (Coleoptera: Dytiscidae) of North America. Quaestiones Entomologicae 17:
249–309.
Sharp D. 1882a. Coleoptera. Tribe Adephaga (continued). In:
Godman FC, Salvin O, eds. Biologia Centrali Americana; or,
contributions to the knowledge of the fauna and flora of Mexico and Central America. Zoology, Botany and Archaeology.
Zoology, Section 15. Insecta, Coleoptera. London: R.H. Porter, 144.
Sharp D. 1882b. On aquatic carnivorous Coleoptera or Dytiscidae. Scientific Transactions of the Royal Dublin Society 2:
179–1003.
Sharp D, Muir F. 1912. The comparative anatomy of the male
genital tube in Coleoptera. Transactions of the Royal Entomological Society of London 60: 477–532.
Spangler PJ. 1963. A description of the larva of Macrovatellus mexicanus Sharp (Coleoptera: Dytiscidae). Coleopterists
Bulletin 17: 97–100.
Spangler PJ. 1966a. Changes in nomenclature and reassignment of Platydessus perforatus (Coleoptera: Dytiscidae).
Coleopterists Bulletin 20: 57–58.
Spangler PJ. 1966b. A new species of Derovatellus from
Guatemala and a description of its larva (Coleoptera: Dytiscidae). Coleopterists Bulletin 20: 11–18.
Spangler PJ. 1967. A new species of Derovatellus from Peru
(Coleoptera: Dytiscidae). Journal of the Kansas Entomological Society 40: 142–145.
Trémouilles ER. 1995. Insecta, Coleoptera, Dytiscidae. Fasciculo 1. Dytiscidae: Methlinae-Hydroporinae. Fauna de
Agua Dulce de la Republica Argentina 37: 1–82.
Wehncke E. 1876. Neue Dytisciden. Stettiner Entomologische
Zeitung 37: 356–360.
510
K. B. MILLER
Wehncke E. 1883. Neue Dytisciden. Deutsche Entomologische
Zeitschrift 27: 146–149.
Wolfe GW. 1985. A phylogenetic analysis of plesiotypic hydroporine lineages with an emphasis on Laccornis des Gozis
(Coleoptera: Dytiscidae). Proceedings of the Academy of Natural Sciences of Philadelphia 137: 132–155.
Wolfe GW. 1988. A phylogenetic investigation of Hydrovatus,
Methlini and other plesiotypic hydroporines (Coleoptera:
Dytiscidae). Psyche 95: 327–344.
Wolfe GW, Zimmerman JR. 1984. Sensilla, punctation, reticulation, and body shape in the Hydroporinae (Coleoptera:
Dytiscidae). International Journal of Insect Morphology and
Embryology 13: 373–387.
Young FN. 1954. The water beetles of Florida. University of
Florida Biology Series 5: 1–238.
Zimmermann A. 1919. Die Schwimmkäfer des Deutschen
Entomologischen Museums zur Berlin-Dahlem. Archiv für
Naturgeschichte 83: 68–249.
Zimmermann A. 1920. Dytiscidae, Haliplidae, Hygrobiidae,
Amphizoidae. In: Schenkling S, ed. Coleopterorum Catalogus. Berlin: Dr W. Junk, 1–326.
Zimmermann A. 1921. Beiträge zur Kenntnis der südamerikanischen Schwimmkäferfauna nebst 41 Neubeschreibungen. Archiv für Naturgeschichte 87: 181–206.
APPENDIX
LIST
OF TAXA IN THE TRIBE
VATELLINI
Vatellini Sharp, 1882
Calicovatellus Miller & Lubkin, 2001
C. petrodytes Miller & Lubkin, 2001 (extinct)
Derovatellus Sharp, 1882
= Varodetellus Biström, 1979 syn. nov.
= Mesovatellus Trémouilles, 1995 syn. nov.
africanus Régimbart, 1889
allaudi Guignot, 1936
assinicus Régimbart, 1889
ater Bilardo & Pederzani, 1978
baloghi Biström, 1979
bisignatus Ahlwarth, 1921
= nyasaenesis Omer-Cooper, 1958
bistroemi Brancucci, 1981
bruchi Zimmermann, 1917
caprai Guignot, 1952
corvus Guignot, 1954
dagombae Biström, 1979
decellei Biström, 1979
dimorphus Guignot, 1936
duplex Guignot, 1956
erratus Guignot, 1979
eupteryx Guignot, 1955
fasciatus Régimbart, 1895
ferrugineus Bilardo & Pederzani, 1978
floridanus Fall, 1932 stat. nov.
= ibarrai Spangler, 1966a syn. nov.
hancocki Biström, 1981
intermedius Biström, 1986
kamerunensis Biström, 1979
lentus (Wehncke, 1876)
lugubris Guignot, 1955
macrocolus Guignot, 1956
marmottani Guignot, 1940
mocquerysi Régimbart, 1895
natalensis Omer-Cooper, 1965
nyanzae Biström, 1980
obscurus Régimbart, 1895
olofi Franciscolo & Sanfilippo, 1991
= bistroemi Franciscolo & Sanfilippo, 1990
(preoccupied)
onorei Biström, 1982
orientalis Wehncke, 1883
peruanus Spangler, 1967
regimbarti Guignot, 1936
roosevelti sp. nov.
ruficollis Régimbart, 1895
spangleri sp. nov.
taeniatus Biström, 1979
wewalkai Biström, 1979
wittei Biström, 1979
Vatellus Aubé, 1837
= Leucorea Laporte, 1835
= Macrovatellus Sharp, 1882 syn. nov.
= Platydessus Guignot, 1955 syn. nov.
amae sp. nov.
annae sp. nov.
drymetes sp. nov.
bifenestratus (Zimmermann, 1921) comb. nov.
pilacaudus sp. nov.
grandis Buquet, 1840
haagi Wehncke, 1876)
lateralis (Sharp, 1882) comb. nov.
= marginalis (Sharp, 1882) syn. nov.
= rudis (Sharp, 1882) syn. nov.
= deplanatus (Zimmermann, 1919) syn. nov.
mexicanus (Sharp, 1882) comb. nov.
perforatus (Guignot, 1955) comb. nov.
maculosus sp. nov.
sahlbergi (Sharp, 1882a) comb. nov.
tarsatus (Laporte, 1835)
ventralis (Sharp, 1882a) comb. nov.
wheeleri sp. nov.

Revision of the New World and south-east Asian

Transcription

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