A new view on the generic classification of the Agabus-group of genera of the Agabini, aimed at solving the problem with a paraphyletic Agabus (Coleoptera: Dytiscidae)

Full text

Koleopterologische Rundschau7017-36Wien, Juni 2000
A new view on the generic classification of the
Agabus-group of genera of
the
Agabini,
aimed at solving the problem with a paraphyletic Agabus
(Coleoptera: Dytiscidae)
A.N. NlLSSON
Abstract
The generic classification of the Agabus-group of genera of the tribe Agabini (Coleoptera: Dytiscidae)
is revised and presented as a checklist of the world species. In the new classification, the following six
genera are recognized: Agabus LEACH, Hydronebrius JAKOVLEV, Hydrotrupes SHARP, Ilybius
ERICHSON,
Ilybiosoma CROTCH, and Platambus THOMSON. The concepts of Ilybius and Platambus
have been expanded, and both genera now include additional species previously placed in Agabus.
The genus Agabus is divided into the three subgenera: Agabus
s.str.,
Acatodes THOMSON, and
Gaurodytes
THOMSON.
The following new generic synonymies are given: Agabinus
CROTCH,
1873,
and Colymbinectes FALKENSTRÖM, 1936 = Platambus THOMSON, 1859; Carrhydrus FALL, 1922 =
Agabus LEACH, 1817. Ilybius lenensis nom.n. is proposed as replacement name for /. aenescens
POPPIUS, 1905.
Key words: Coleoptera, Dytiscidae, Agabini, Agabus-group of genera, generic classification,
checklist, key.
Introduction
Within the Colymbetinae, the Agabini are characterized by the derived presence of
a
linear group
of short, stout setae near the posterior ventroexternal angle of
the
metafemur (BRINCK 1948). A
few other potential autapomorphies of this tribe have been described from larvae (DE MARZO &
NlLSSON 1988, ALARIE & LARSON 1998). However, these characters have so far only been
verified in relatively few species of the Agabini.
With respect to the delimitation of the tribe Agabini, I largely agree with BRINCK (1948), except
that I find his arguments for treating Hydronebriini as a separate tribe weak. Besides the genus
Hydronebrius
JAKOVLEV, 1897 he included also the Nearctic Agabus
cordatus
group in this tribe.
The species in the latter group have in fact the metafemoral row of setae, and the absence of
these setae in Hydronebrius could be viewed as a reduction connected with the strong punctation
of the metafemur in this genus. This setal row is also absent in a few species of Platambus
THOMSON, 1859(BRANCUCCI 1988).
The Nearctic monobasic genus Hydrotrupes was excluded from the Agabini by BEUTEL (1994)
based on larval characters. Later, ALARIE et al. (1998) analysed a larger set of larval characters
which supported that Hydrotrupes belongs to the Agabini. I have included this genus in Agabini,
admitting that its taxonomic position is uncertain.
The generic classification of Agabini needs to be revised. The chief problem being the large
genus Agabus that so far not has been diagnosed by a single autapomorphy. Hopefully, my hypo-
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

18 Koleopt. Rdsch. 70 (2000)
thesis of relationships and the resulting classification given below will be tested by future
studies, in which the use of molecular and morphological characters, including also larvae, will
be combined. Based on my own experience, the application of parsimony analysis on characters
from adult morphology only, will not lead to firmly based hypothesis of relationships within the
Agabini.
Generic classification of Agabini
The tribe Agabini can be separated into two major groups of genera (NILSSON 1997): (1) those
having a pair of elliptical, sublateral clypeal fovea, and female metatibia and -tarsus with a
ventral fringe of natatorial setae present, viz. Andonectes GuÉORGUiEV, \91\,Agametrus SHARP,
1882,
Leuronectes SHARP, 1882, and
Platynectes
RÉGIMBART, 1878; and (2) those having linear
clypeal fovea near anterolateral angles or along entire anterior margin, and having female
metatibia and -tarsus without the ventral setal fringe, i.e. the rest.
The only contradiction to this pattern known to me is Ilybius discedens SHARP, 1882, in which
the female has the ventral setal fringe (LARSON 1987). As this species has all the advanced
characters of the genus Ilybius, the presence of
the
setal fringe in the female is seemingly due to
a reappearance of this character.
In this paper, I will only discuss the second group of genera to which Agabus belongs. My
baseline is to neglect the generic classification currently in use (e.g. GuÉORGUiEV 1971,
FRANCISCOLO 1979: 242, 243) and start to present characters that can be used for the basal
splittings of the group. The new classification proposed for the Agabus-group of genera is
presented in full below.
Review of some relevant characters
1.
Clypeal bead continuous. LARSON (1975: Figs. 278, 279) introduced as a character within
Agabus the development of
a
fine anterior marginal bead on the clypeus. As a continuous bead is
unknown among other Colymbetinae, its presence is viewed as apomorphic. The apomorphic
state occurs in all species here placed in the genus Agabus, except A. striolatus. The additional
presence of the apomorphic state in the Platambus glabrellus- and optatus-gronps is viewed as
due to convergence.
2.
Pronotum with anterior line continuous. NILSSON & LARSON (1990) introduced another
important character in Agabus, i.e. the development of the fine line that follows close to the
anterior margin of the pronotum. A continuous line is viewed as apomorphic as it is unknown in
the Colymbetinae outside Agabini. The apomorphic state of this character is present in all species
placed in the genus Ilybius. Moreover, it is present in the Platambus semivittatus-grovp, in
Ilybiosoma ilybiiformis, and in the following species-groups of Agabus
s.str.:
antennatus,
bifarius,
labiatus,
punctulatus, and uliginosus.
3.
A third important character was first mentioned in SEMENOV'S (1899) description of Apator.
He noted the tiny hairs, pointing straight downward, which arise from punctures along the face of
the epipleuron, except in the basal third and near the elytral apices. Later, LEECH (1942) observed
these hairs also in other Agabus species than A. bifarius. These hairs are somewhat difficult to
see,
even on clean specimens. The absence of such hairs in other agabine genera suggests that
their presence is apomorphic within
Agabus.
As the distribution of this character among Agabus
species is congruent with other characters defining larger groups of
species,
these setae may have
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON: Generic classification of Agabus-group of genera of Agabini (DYTISCIDAE) 19
evolved only once, i.e. they provide a synapomorphy for the species which have them, i.e. most
Agabus
s.str.
4.
Most species of Agabini have a simple penis with a tapering apex. A ventral subapical spine
is,
however, present in many Agabus species, and I have used this character as a synapomorphy
of the subgenus Acatodes. Its absence in
A.
audeni WALLIS,
A.
anthracinus (MANNERHEIM), A.
coxalis SHARP, and possibly also A. inexspectatus NlLSSON, is viewed as secondary loss (cf.
LARSON 1989,1991).
5.
Saw-like female gonocoxae (ovipositor) provide an autapomorphy of
Ilybius
s.str.
Moreover,
FERY
& NlLSSON (1993) suggested a transformation series in the evolution of the ovipositor in
which the row of denticles have evolved along the external ridge found in some species
traditionally placed with Agabus. Externally ridged gonocoxae is here used as a unique
synapomorphy of the species included in a more broadly delimited genus Ilybius.
6. In his key to Nearctic species-groups, LARSON (1989) characterized some groups of Agabus on
the prosternai process having its lateral beads broadly inflated posterior of procoxae (cf. LARSON
& WOLFE 1998: Figs. 21-26). Later, NILSSON (1997) used the same feature to characterize a
redelimited
A.
optatus-group.
I have found this character present also in the two species formerly
assigned to the genus Agabinus, as well as in most species of Platambus as revised by
BRANCUCCI (1988). The character is here used as a synapomorphy of the species included in a
more broadly delimited genus Platambus.
7.
In the same key, LARSON (1989) noted the broad separation of the mesocoxae present in his
Agabus obtusatus-group. This character was given as a ratio of 1.5 or more between the
narrowest distance between the mesocoxae and the narrowest distance between the procoxae.
The character is here used as a synapomorphy of
a
more broadly delimited genus
Platambus.
It is
most weakly developed in the
P.
americanus-
and
P.
confusus-gvowps.
New limits for
Agabus,
Ilybius and Platambus
From my unpublished studies of the available morphological characters of the adults, it seems
that no solution without much conflict is possible to find. A central problem is the conflict
between the two characters represented by a continuous clypeal bead and a prosternai process
with lateral bead inflated posteriorad of procoxae (NlLSSON 1997). I now prefer to give priority
to the last character, supported also by the wide mesocoxal separation of the species which have
it. This character-complex is used to diagnose an expanded genus Platambus, adding to the
species included by BRANCUCCI (1988) the dibasic Nearctic genus Agabinus CROTCH,
1873,
plus
the following species-groups traditionally placed with Agabus: americanus, confusus, optatus,
semivittatus and spinipes. Besides the chiefly East Palearctic and Oriental Platambus
s.str.,
and
the optatus-group in eastern Asia and eastern North America, the resulting expanded genus
Platambus includes also elements from western North America.
I suggest that the genus Agabus is restricted to the species not included in Platambus s.l. that
have the clypeal bead continuous. Consequently, Carrhydrus FALL, 1922, and Metronectes
SHARP, 1882, will join Agabus (cf. BALKE et
al.
1997), whereas many species currently placed in
Agabus will have to be transferred to other genera.
Moreover, I think that Ilybius should be expanded to include all species of Agabini with a conti-
nuous anterior line on pronotum and the clypeal bead broken (FERY & NILSSON 1993). The
widened concept of this genus is also supported by a female ovipositor with a lateral ridge, and
metacoxal lines reduced anteriorly. Consequently, the Agabus
chalconatus-,
A. erichsoni- (sensu
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

20 Koleopt. Rdsch. 70 (2000)
FERY & NILSSON 1993), and
A.
opacus-groups
(sensu LARSON 1996) will have to be transferred
to Ilybius.
Some species currently placed in Agabus, i.e. the bifarius-, labiatus-, punctulatus-, and
uliginosus-groups, have both the clypeal and anterior pronotal beads continuous. Other
characters will have to answer the question whether they belong to Agabus or Ilybius. For the
moment they should better stay in Agabus
s.str.
The remaining species
The species that fall outside these new concepts of
Platambus,
Agabus and Ilybius are those in
Hydronebrius, and the Agabus (in traditional sense) groups cordatus (sensu NiLSSON 1992),
seriatus (sensu LARSON 1997), striolatus (monobasic), and
kermanensis
(monobasic). No unique
synapomorphy has been found that could unite all the remaining species. Consequently, they
have to be separated into two or more groups.
Besides the enigmatic SE Palearctic genus Hydronebrius, the remaining groups include 15
species, 12 of which are
Nearctic.
The three other species are the Ethiopian
A.
discicollis ANCEY,
1882,
plus the Palearctic A. striolatus (GYLLENHAL, 1808) and A. kermanensis J.BALFOUR-
BROWNE, 1939.
BALFOUR-BROWNE (1939) described Ranagabus as a subgenus of Agabus, with the single
species A. kermanensis. The combination of characters found in this species was such that they
"separate the species so sharply from all the groups of SHARP that I consider the only course to
follow is the creation of a separate subgenus" (BALFOUR-BROWNE 1939: 106). The only
subsequent reference to this species is GUEORGUIEV (1965) who added to the original description
and illustrated the penis.
My interest for A. kermanensis was triggered by the study of Afrotropical Agabus (NiLSSON
1992),
especially the search for close relatives of
A.
discicollis,
and the other, Nearctic, species of
the A.
cordatus-group.
I now believe that A. kermanensis belongs to the A. cordatus-group and
that it is the sister-species of
A.
discicollis.
I have earlier (NiLSSON 1992) placed
A.
discicollis together with the four Nearctic species in the
A.
cordatus-group (LARSON 1989). After the examination of
A.
kermanensis I am inclined to
believe that it is the sister species of A. discicollis. The following characters separate A.
kermanensis from A. discicollis and the other species of the A. cordatus-group: (1) head
orientated anteriorly, (2) pronotum broad, (3) prosternai process long and recieved into well-
defined impression on metasternum, and (4) legs short with punctation weak.
The corresponding character states displayed by A. discicollis and the A. cordatus-group most
probably represent adaptations to high altitudes that have evolved independently in
A.
discicollis
and the Nearctic stock (NILSSON 1992). Character
no.
3 is especially interesting as the reduction
of the prosternai process and the metasternal groove is probably connected with the more
ventrally orientated head of the high-altitude specialists. A similar reduction has probably
occurred in the species of the Ethiopian A
ambulator-group
(cf. NiLSSON 1992), that I now think
should be included in the
A.
confinis-group.
The Afrotropical species of
the
group are seemingly
closely related to
A.
turcmenus GuiGNOT, 1957.
The parameres of
A.
discicollis and A. kermanensis are very similar, and more strongly setose
than in the Nearctic species. The median lobes are also quite similar, although the basal apodeme
is much larger in A. kermanensis. The broad metafemur with its laminar posteroexternal angle
and the medially truncate or slightly excavate last abdominal segment are unique to A.
kermanensis.
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON:
Generic classification
of
Agabus-gwup
of
genera
of
Agabini
(DYTISCIDAE) 21
The morphological differences between A. kermanensis and A. discicollis are in the Nearctic
paralled by those between the A. cordatus- and A. sma/ws-groups (sensu
LARSON
1989), with
the high-altitude specialists found in the A. cordatus-group. Phylogenetically, it would probably
make sense to assign the Old and New World species to one species-group each, viz. the
Ilybiosoma discicollis- and seriatum-gxoups, respectively. This solution has been used in my
classification (see below).
The only apomorphic character (within the Agabus-gxoup of genera) found in all species of the
redelimited seriatus-, discicollis-, and striolatus-groups is the presence of anteroventral
spiniferous punctures along the entire length of the metatibia. Consequently, this character is
used to diagnose a genus with the available name Ilybiosoma
CROTCH,
1873, with the type
species Ilybius regularis
CROTCH,
1852. These punctures are fewer and more sparsely set in A.
striolatus than in the other species, in which they form a more or less continuous row. As these
punctures also occur in several other Agabus species, like those in the tristis-gxoup, the
possibility to place A. striolatus in Agabus {Gaurodytes) should also be considered further. Until
more evidence will become available, I prefer a conservative view, and have consequently kept
A.
striolatus in this position in my classification (see below).
Revised key to Agabus-group of genera and subgenera
1.
Female metatibia and -tarsus with ventral setal fringe present. Clypeus with pair of sublateral,
elliptical fovea
Platynectes-group
of
genera
Female metatibia
and
-tarsus with ventral setal fringe reduced (except
in
Ilybius discedens).
Clypeus with medially broken
or
continuous linear fovea along anterior margin {Agabus-
group
of
genera)
2
2.
Prosternai process
in
most species with lateral bead broadly inflated posterior
of
procoxae.
Mesocoxae widely separated. Epipleuron broad
in
many species, also
in
posterior
half...
Platambus
Prosternai process with lateral bead
not
broadly inflated posterior
of
procoxae. Mesocoxae
more narrowly separated. Epipleuron narrow
in
posterior half
3
3.
Metafemur with linear group
of
setae
at
posteroexternal angle reduced Hydronebrius
Metafemur with linear group
of
setae
at
posteroexternal angle present
4
4.
Clypeus with linear fovea more
or
less continuous {Agabus)
5
Clypeus with linear fovea broadly interrupted medially
7
5.
Penis with subapical ventral spine subgenus Acatodes
Penis without subapical ventral spine
6
6. Paramere stylate. Pronotum with anterior bead broadly interrupted medially.. subgenus Gaurodytes
Paramere strap-like. Many species with pronotum having anterior bead continuous, and/or
metasternal wing narrow subgenus Agabus
s.str.
7.
Pronotum with continuous fine line along anterior margin Ilybius
Pronotum with anterior bead broadly interrupted medially Ilybiosoma
Classification of Agabus-group of genera
Distributions coded as: (E) Ethiopian, (EP) East Palearctic (Asian part), (H) Holarctic, (NA)
Nearctic (including Mexico), (O) Oriental, (P) Palearctic, and (WP) West Palearctic (European
plus North African parts). As in the forthcoming new Palearctic Coleoptera catalogue (Eds. Lobi
& Smetana), all of China and the Himalayan provinces of India are included in the Palearetic
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

22
Koleopt.
Rdsch.
70 (2000)
Region. The type region is given for each species. Synonyms are given in alphabetical order.
Names are categorized as: [HN] preoccupied homonym, [IN] indication to, i.e. new name for
misidentification, [NO] nomen oblitum, [RN] replacement name. Only available names based on
extant species are listed. When possible, names of species groups are based on species with
Holarctic distributions.
Genus
Agabus LEACH, 1817 (E, H, O)
Autapomorphy: clypeus with fine marginal bead more or less continuous.
Subgenus Acatodes
THOMSON,
1859 (E, H, O)
Autapomorphy: penis with subapical spine.
[Syn.: Arctodytes THOMSON, 1874; Heteronychus
SEIDLITZ,
1887; Scytodytes
SEIDLITZ,
1887;
Allonychus
ZAITZEV,
1905; Mesogabus
G\1ÉORG\J\EV,
1969]
arcticus-group (H)
anthracinus MANNERHEIM, 1852:304, Alaska (NA)
= scapularis MANNERHEIM, 1852:303
arcticus arcticus (PAYKULL, 1798:201) (Dytiscus), Sweden
(H)
= reticulatus (KlRBY, 1837:71) (Colymbetes)
arcticus alpinus (MOTSCHULSKY, 1860:102) {Colymbetes), Mongolia (EP)
= insignis GUÉORGUIEV, 1969:62
= punctipennis (J.SAHLBERG, 1880:56) {Gaurodytes)
= sibiricus (J.SAHLBERG, 1880:56) {Gaurodytes)
arcticus ochoticus POPPIUS 1908:54, Siberia (EP)
browni
KAMIYA,
1934:181, Manchuria (EP)
= brunneus KAMIYA, 1935:8
[HN]
= orientalis KAMIYA, 1938:36
[RN]
conspicuus
SHARP,
1873:48, Japan (EP)
= deplanatus GUIGNOT, 1952:18
=
procerus (RÉGIMBART, 1883:229) {Platynectes)
granulatus (FALKENSTRÖM, 1936:95)
{Gaurodytes),
China (EP)
sturmii (GYLLENHAL
in
SCHÖNHERR, 1808:18) {Dytiscus), Sweden
(P)
=fallax MUNSTER, 1932:85
= goedelii (VILLA & VILLA, 1833:33) {Colymbetes)
confìnis-group (E, H, O)
ambulator RÉGIMBART, 1895:152, Ethiopia (E)
angusi
NILSSON,
1994:172, Siberia (EP)
approximatus
FALL,
1922:26, Colorado (NA)
audeni
WALLIS,
1933:270, British Columbia (NA)
bergi ZAITZEV, 1913:195, Transcaucasus
(P)
bicolor (KlRBY, 1837:70) {Colymbetes), North America (NA)
= mutus SHARP, 1882:513
canadensis FALL, 1922:27, Manitoba (NA)
clypealis (THOMSON, 1867:107) {Gaurodytes), Sweden
(H)
= scholzi KOLBE, 1916:253
confinis (GYLLENHAL,
1808:511
)
{Dytiscus),
Sweden
(H)
= longulus (LECONTE, 1878:596) {Gaurodytes)
= ovoideus
(CROTCH,
1873:418) {Gaurodytes)
congener
(THUNBERG,
1794:75) {Dytiscus), Sweden
(P)
= congener (PAYKULL, 1798:214) {Dytiscus) [HN]
= daisetsuzanus KAMIYA, 1938:34
=foveolatus MULSANT & Godart, 1860:177
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON:
Generic classification of
Agabus-group
of genera of
Agabini
(DYTISCIDAE) 23
=funkii SEIDLITZ, 1887:92
=fuscotestaceus DALLA
TORRE,
1877:63
= venturii BERTOLINI, 1870:242
costulatus (MOTSCHULSKY, 1859:541) (Colymbetes), Siberia (EP)
= tunkunensis (ZIMMERMANN, 1928:178) (Gaurodytes)
discolor
(HARRIS,
1828:164) (Colymbetes), North America (H)
= levanderi HELLEN, 1929:40
elongatus (GYLLENHAL in C.R.SAHLBERG, 1826:169) (Dytiscus), Scandinavia (H)
= bryanti CARR, 1930:278
immaturus LARSON, 1991:1263, New Brunswick (NA)
inexspectatus
NlLSSON,
1990:157, Siberia (H)
inscriptus (CROTCH, 1873:422) (Gaurodytes), Labrador (NA)
kootenai LARSON, 1991:1279, British Columbia (NA)
lapponicus (THOMSON, 1867:108) (Gaurodytes), Norway (P)
= obovatus (J.SAHLBERG, 1875:176) (Gaurodytes)
= obscuripennis (J.SAHLBERG, 1875:177) (Gaurodytes)
loeffleri
WEWALKA
& NlLSSON, 1990:152, Ethiopia (E)
mackenziensis LARSON, 1991:1267, Northwest Territories (NA)
matsumotoi
SATO
& NlLSSON, 1990:193, Japan (EP)
moestus (CURTIS, 1835:LX) (Colymbetes), Northwest Territories (H)
= borealis SHARP, 1882:513
= nigripalpis J.SAHLBERG, 1880:56
phaeopterus (KlRBY, 1837:70) (Colymbetes), North America (NA)
pseudoclypealis SCHOLZ, 1933:74, Russia (P)
= haraldi Hâk.LlNDBERG, 1933:121
sasquatch LARSON, 1991:1285, California (NA)
setulosus (J.SAHLBERG, 1895:39) (Gaurodytes), Finland (WP)
slovzovi (J.SAHLBERG, 1880:59) (Gaurodytes), Siberia (EP)
smithi BROWN, 1930:88, British Columbia (NA)
subfuscatus SHARP, 1882:514, Massachusetts (NA)
thomsoni (J.SAHLBERG, 1871:407) (Gaurodytes), Finland (H)
= coriaceus (J.SAHLBERG, 1875:174) (Gaurodytes)
tibetanus ZAITZEV, 1908:425, Tibet (EP)
turcmenus GuiGNOT, 1957:93, Turkestan (EP)
zetterstedti
THOMSON,
1856:216, Scandinavia (H)
= browni LEECH, 1938:126 [HN]
fuscipennis-group (H)
ajax FALL, 1922:30, Alberta (NA)
coxalis coxalis SHARP, 1882:535, Siberia (H)
= splichali REITTER, 1899:196
coxalis ermaki (ZAITZEV, 1953:259)
(Gaurodytes),
Siberia (EP)
coxalis schmidti ZAITZEV, 1913:197, Georgia (P)
fuscipennis fuscipennis (PAYKULL, 1798:209) (Dytiscus), Sweden (P)
= eversmanni BALLION, 1855:237
=fossarum (GERMAR, 1824:29) (Dytiscus)
= obscurior (J.SAHLBERG, 1875:199) (Acatodes)
fuscipennis ontarionis FALL, 1922:30, Ontario (NA)
infuscatus AUBÉ, 1838:330, North America (H)
= dubiosus POPPIUS, 1905:20
= gelidus
U.SAHLBERG,
1906:15
kaszabi GUÉORGUIEV, 1972:37, Mongolia (EP)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

24
Koleopt.
Rdsch.
70 (2000)
japonicus-group (EP, O)
aequabilis (GSCHWENDTNER, 1923:105) {Gaurodytes), Tien Shan (EP)
aequalis SHARP, 1882:501, Siberia (EP)
amoenus amoenus SOLSKY, 1874:142, Turkestan (EP)
amoenus sinuaticollis RÉGIMBART, 1899:278, India (EP)
bakeri (ZIMMERMANN, 1924:743) {Gaurodytes), Philippines (O)
fulvipennis RÉGIMBART, 1899:277, China (EP)
= chinensis (ZIMMERMANN, 1919:211) {Gaurodytes)
hummeli (FALKENSTRÖM, 1936:2) {Gaurodytes), China (EP)
japonicus japonicus SHARP, 1873:50, Japan (EP)
japonicus ezo
NAKANE,
1989:23, Japan (EP)
japonicus continentalis GUÉORGUIEV, 1970:259 [RN], Primorye (EP)
=falkenstromi
(ZAITZEV,
1953:254) {Gaurodytes) [HN]
japonicus shiroumanus (NAKANE, 1959:98)
{Gaurodytes),
Japan (EP)
kokoosson FENG, 1936:8, China (EP)
mucronatus (FALKENSTRÖM, 1936:89) {Gaurodytes), China (EP)
philippensis (ZIMMERMANN, 1924:743) {Gaurodytes), Philippines (O)
regimbarti ZAITZEV, 1906:174, China (EP)
rufipennis (GSCHWENDTNER, 1933:163) {Gaurodytes), China (EP)
lutosus-group (NA)
griseipennis LECONTE, 1859:5, Wyoming (NA)
lutosus LECONTE, 1853:31 [RN], California (NA)
= discolor LECONTE, 1852:204 [HN]
= leconlei (CROTCH, 1873:417) {Gaurodytes)
[RN]
= mimus LEECH, 1942:132
rumppi
LEECH,
1964:79, California (NA)
obsoletus-group (NA)
ancillus FALL, 1922:24, Oregon (NA)
hoppingi LEECH, 1942:135, California (NA)
morosus LECONTE, 1852:204, California (NA)
=fassiger (MOTSCHULSKY, 1859:170) {Colymbetes)
obliteratus obliteratus LECONTE, 1859:5, Wyoming (NA)
obliteratus nectris LEECH, 1942:133, British Columbia (NA)
obsoletus LECONTE, 1858:15, California (NA)
raffrayi-group (E)
dytiscoides RÉGIMBART, 1908:6, Tanzania (E)
pallidus OMER-COOPER, 1931:786, Ethiopia
(E)
raffrayi SHARP, 1882:501, Ethiopia (E)
=
limbicoUis
RÉGIMBART, 1905:224
ruwenzoricus GUIGNOT, 1936:49, Uganda (E)
sjostedti RÉGIMBART, 1908:5, Tanzania
(E)
Subgenus Agabus s. str. (H)
No autapomorphy found. The following characters occur within the group: male antenna davate;
pronotum with anterior bead continuous; metasternal wing narrow; epipleuron with row of fine
setae;
larval legs provided with secondary swimming hairs.
[Syn.: Eriglenus
THOMSON,
1859;
Apator
SEMENOV,
1899;
Carrhydrus
FALL,
1922; Neonecticus
GUIGNOT, 1951]
aeruginosus-group (NA)
aeruginosus AUBÉ, 1838:298, North America (NA)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON:
Generic classification
of
Agabus-group
of
genera
of
Agabini
(DYTISCIDAE)
25
=
dispositus
GUIGNOT,
1936:189
punctatus
MELSHEIMER,
1844:27,
North
America
(NA)
antennatus-group (NA)
antennatus LEECH, 1939:217 [RN], Nebraska (NA)
= clavatus
LECONTE,
1859:4 [HN]
bifarius-group (H)
bifarius (KlRBY, 1837:71) (Colymbetes), North America (H)
= kessleri HOCHHUTH, 1871:238
clavicornis-group (H)
clavicornis SHARP, 1882:536, Siberia (H)
= verus BROWN, 1931:115
crassipes (FALL, 1922:35) (Carrhydrus) Alberta (NA)
serricornis (PAYKULL, 1799:49) (Dytiscus), Finland (P)
= clavatus (LATREILLE, 1804:166) (Dytiscus)
= minor J.SAHLBERG, 1875:199
=
paykullii
LEACH,
1817:72 [RN]
disintegratus-group (NA)
disintegrates (CROTCH, 1873:416) (Gaurodytes), North America (NA)
taeniolatus (HARRIS, 1828:164) (Colymbetes), North America (NA)
= taeniatus AUBÉ, 1838:311
/a///-group (NA)
falli (ZIMMERMANN, 1934:186) (Gaurodytes) [RN], Manitoba (NA)
= sharpi FALL, 1922:19 [HN]
labiatus-group (H)
fuhaster ZAITZEV, 1906:26, Russia (P)
labiatus (BRAHM, 1791:27) (Dytiscus), Germany (P)
= assimilis (STURM, 1834:112) (Colymbetes)
=femoralis (PAYKULL, 1798:215) (Dytiscus)
= hochhuti ZAITZEV, 1908:CLX [IN; brunneus F. sensu HOCHHUTH 1871]
= impressus (ZOUBKOFF, 1833:317) (Colymbetes)
= transcaucasicus ZAITZEV, 1927:28
mandsuricus (GUIGNOT, 1956:139) (Ilybius), Manchuria (EP)
= charmi (LAFER, 1988:54) (Eriglenus)
pallens POPPIUS, 1905:22, Siberia (H)
= hudsonicus LEECH, 1938:123
= mongolicus GUÉORGUIEV, 1968:27
= zaitzewi
POPPIUS,
1909:12
undulatus (SCHRANK, 1776:70) (Dytiscus), Germany (WP)
= abbreviatus (FABRICIUS, 1787:191) (Dytiscus)
= imperfectus MEIER, 1899:98
= interruptus SCHILSKY, 1888:183
=pictus
MEIER,
1899:98
= ruficeps (MÉNÉTRIES, 1832:141) (Colymbetes)
zimmermanni SCHOLZ, 1920:15, Asia Minor (P)
lineatus-group (EP)
desertorum F.MORAWITZ, 1863:169, Russia (EP)
lineatus GEBLER, 1848:75, Siberia (EP)
punctulatus-group (H)
colymbus LEECH, 1938:125, Manitoba (NA)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

26 Koleopt. Rdsch.
70
(2000)
luteaster
ZAITZEV,
1906:27, Siberia
(EP)
oblongulus FALL, 1922:18, Oregon
(NA)
pisobius LEECH, 1949:246, British Columbia (NA)
punctulatus
AUBÉ,
1838:332, North America (NA)
= aeneolus (CROTCH, 1873:417) (Gaurodytes)
uliginosus-group
(H)
amnicola (J.SAHLBERG, 1880:58) (Gaurodytes), Siberia
(H)
= triton
F'ALL
1922:17
jacobsoni
ZAITZEV,
1905:225, Russia
(P)
margaretae LARSON, 1975:344, Alberta (NA)
uliginosus (LINNAEUS, 1761:216) (Dytiscus), Sweden
(P)
= aeratus STEPHENS, 1828:79 (Colymbetes)
= rf/spar BOLD, 1849:XXIV
= reichei
AUBÉ,
1837:138
vereschaginae ANGUS, 1984:193, Siberia (EP)
Subgenus Gaurodytes
THOMSON,
1859 (E, H, O)
Autapomorphy: paramere stylate.
[Syn.: Necticus HOPE,
1839
(preocc); Metronectes
SHARP,
1882;
Dichodytes THOMSON,
1886;
Xanthodytes SEIDLITZ,
1887;
Gabinectes GuiGNOT,
1931;
Agabinectes GuiGNOT,
1932;
Dichonectes
GUIGNOT,
1945]
adpressus-group
(H)
adpressus AUBÉ, 1837:169, Dauria
(H)
= angusticollis (J.SAHLBERG, 1871:408) (Gaurodytes)
= haeffnerì
AUBÉ,
1837:170
= sahlbergi SHARP, 1882:517
= solus LEECH, 1949:248
= subquadratus (MOTSCHULSKY, 1860:102) (Colymbetes)
udege NlLSSON, 1994:170, Primorye
(EP)
affìnis-groìip
(H)
qffìnis (PAYKULL, 1798:211) (Dytiscus), Sweden
(P)
= branchiatus (BABINGTON, 1832:329) (Colymbetes)
= guttulus (SCHÖNHERR, 1808:19) (Dytiscus) [IN; guttatus PAYKULL sensu ILLIGER
1801]
biguttulus (THOMSON, 1867:110) (Gaurodytes), Scandinavia
(P)
= boreellus (J.SAHLBERG,
1871:409)
(Gaurodytes)
= ovalis (J.SAHLBERG, 1875:178) (Gaurodytes)
kholini
NlLSSON,
1994:45, Primorye
(EP)
laferi
NlLSSON,
1994:47, Primorye
(EP)
semipunctatus (KlRBY, 1837:69) (Colymbetes), North America (NA)
= stridulator
SHARP,
1882:509
sikhotealinensis (LAFER, 1988:56) (Gaurodytes), Primorye
(EP)
unguicularis (THOMSON, 1867:101) (Eriglenus), Sweden
(P)
velox LEECH, 1939:219, Manitoba (NA)
yakutiae NlLSSON & LARSON, 1990:229, Siberia (EP)
ambiguus-group
(NA)
ambiguus (SAY, 1823:96) (Colymbetes), North America (NA)
=fimbriatus
LECONTE,
1850:214 [RN]
= reticulatus AUBÉ, 1838:355
[HN]
austinii
SHARP,
1882:516, Utah (NA)
erythropterus
(SAY,
1823:95) (Colymbetes), North America (NA)
klamathensis LARSON
&
LEECH, 1989:875, Oregon (NA)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NILSSON:
Generic classification of
Agabus-group
of genera of Agabini (DYTISCIDAE)
27
strigulosus (CROTCH, 1873:422) (Gaurodytes), California (NA)
= nanus
(LECONTE,
1878:452) (Gaurodytes)
aubei-group
(WP)
aubei
PERRIS,
1869:6, Corsica (WP)
=
parallelipennis DESBROCHERS
DES
LOGES,
1871:337
brunneus-group (WP)
brunneus (FABRICIUS, 1798:64) (Dytiscus), Morocco (WP)
= castaneus (GYLLENHAL
in
SCHÖNHERR, 1808:21) (Dytiscus)
=ferrugineus (STEPHENS, 1828:79) (Colymbetes)
= marginicollis FAIRMAIRE, 1860:631
= rotundatus WEHNCKE, 1872:136
= rufulus FAIRMAIRE, 1859:272
didymus (OLIVIER, 1795:26) (Dytiscus), France (WP)
= chalybaeus J. S AHLBERG, 1903:12
= vitreus (PAYKULL, 1798:217) (Dytiscus)
guttatus-group
(P)
africanus PEDERZANI & SCHIZZEROTTO, 1998:88, Tunisia (WP)
balcanicus HLISNIKOVSKY, 1955:101, Bulgaria (WP)
basalis (GEBLER, 1830:65) (Colymbetes), Turkestan (EP)
= abnormicollis BALLION, 1871:329
=
pallidipennis JAKOVLEV, 1897:40
[HN]
= songoricus GEBLER, 1859:450 (Colymbetes)
[RN]
biguttatus (OLIVIER, 1795:26) (Dytiscus), France
(P)
= alligator
NORMAND,
1933:299
= annulatus (ZOUBKOFF, 1833:318) (Colymbetes)
= concii FRANCISCOLO, 1942:137
= consanguineus WOLLASTON, 1864:81
=/ontinalis
(STEPHENS, 1828:66) (Colymbetes)
= indicus RÉGIMBART, 1899:272
= melas AUBÉ, 1837:168
= nigricollis (ZOUBKOFF, 1833:317) (Colymbetes)
= nitidus (FABRICIUS,
1801:265)
(Dytiscus)
= nubiensis RÉGIMBART, 1895:154
= olivieri ZAITZEV, 1908:121
[RN]
=
pauper SCHILSKY, 1888:184
=
picicornis (STEPHENS, 1828:66) (Colymbetes)
= silesiacus LETZNER, 1844:173
= subaquilus GOZIS, 1912:60
binotatus AUBÉ, 1837:161, Sardinia (WP)
blatta
JAKOVLEV,
1897:39, Turkestan (EP)
=picea
JAKOVLEV,
1897:40
brandii HAROLD, 1880:148, China (EP)
=jeholensis
KAMIYA,
1935:8
caraboides SHARP, 1882:494, Syria or Mesopotamia (EP)
= merkli RÉGIMBART, 1885:XIX
cephalotes REICHE, 1861:202, Corsica (WP)
dilatatus (BRULLE, 1832:127) (Colymbetes), France
(P)
= castaneus SHARP, 1882:500
[HN]
= gory
AUBÉ,
1837:162
faldermanni ZAITZEV, 1927:22, Transcaucasia
(P)
= iranicus GUÉORGUIEV, 1965:257
=
palaestinus (ZIMMERMANN, 1934:164) (Gaurodytes)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

28
Koleopt.
Rdsch.
70 (2000)
freudei GUÉORGUIEV, 1975:97, Nepal (EP)
glacialis HOCHHUTH, 1846:218, Caucasus (P)
= armeniacus SHARP, 1882:497
= inguttatus RE1TTER, 1908:223
glazunovi (ZAITZEV, 1953:228) (Gaurodytes), Uzbekistan (EP)
guttatus guttatus (PAYKULL, 1798:211) (Dytiscus), Sweden (P)
= didymoides ROUBAL, 1919:15
= nigripes (COSTA, 1847:97) (Colymbetes)
=
picinus (MARSHAM, 1802:428) (Dytiscus)
= septemseriatus (J.SAHLBERG, 1875:180) (Gaurodytes)
= signatus
(GRIMMER,
1841:32) (Colymbetes)
= styriacus SHARP, 1882:496
= unicolor DALLA
TORRE,
1877:64
guttatus baudii SEIDLITZ, 1887:85, Italy (WP)
heydeni WEHNCKE, 1872:135, Spain (WP)
=
parvulus FRESNEDA & HERNANDO, 1989:14
lobonyx
GUIGNOT,
1952:17, Sikkim (EP)
longissimus RÉGIMBART, 1899:275, Tibet (EP)
maderensis WOLLASTON, 1854:85, Madeira (WP)
ommani ZAITZEV, 1908:424, Tibet (EP)
picotae FOSTER & BlLTON, 1997:113, Portugal (WP)
svenhedini (FALKENSTRÖM, 1932:192) (Gaurodytes), China (EP)
winkleri (GSCHWENDTNER 1923:104) (Gaurodytes), Turkestan (EP)
=
/?a/mncws GUIGNOT, 1955:274
nebulosus-group (P)
conspersus (MARSHAM, 1802:427) (Dytiscus), England (P)
= bulgaricus (CSIKI, 1943:214) (Gaurodytes)
= corsicus GUIGNOT, 1932:571
= gougeletii REICHE, 1863:474
= luniger
KOLENATI,
1845:82
=perlautus
GOZIS,
1912:53
= subnebulosus (STEPHENS, 1828:72) (Colymbetes)
dichrous SHARP, 1878:169, Pamir (EP)
= lederii SEIDLITZ, 1887:94
= luteolus RÉGIMBART, 1899:275
friedrichi (FALKENSTRÖM, 1936:91) (Gaurodytes), China (EP)
godmanni
CROTCH,
1867:385, Azores (WP)
nebulosus (FORSTER, 1771:56) (Dytiscus), Germany (P)
= abdominalis RAGUSA, 1888:7 [HN]
= bipunctatus (FABRICIUS, 1787:190) (Dytiscus) [HN]
= immaculatus (GSCHWENDTNER, 1927:92) (Gaurodytes) [HN]
= mixtus GUIGNOT, 1949:6
= naevius (GMELIN, 1790:1957) (Dytiscus)
= nigromaculatus (GOEZE, 1777:625) (Dytiscus)
= notatus (BERGSTRÄSSER, 1778:31) (Dytiscus)
=
pratensis
SCHAUFUSS,
1881:620
= ragusai
ZAITZEV,
1908:123 [RN]
= tesselatus (GEOFFROY in FOURCROY, 1785:68) (Dytiscus)
safei ABDUL-KARIM & ALI, 1986:277, Iraq (EP)
paludosus-group (P)
alinae (LAFER, 1988:58) (Gaurodytes), Primorye (EP)
paludosus (FABRICIUS, 1801:266) (Dytiscus), Germany (WP)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NILSSON:
Generic classification of
Agabus-group
of genera of Agabini (DYTISCIDAE) 29
= marginalis SHARP, 1882:502
= pallidipennis (LAPORTE, 1834:103) (Colymbetes)
=
politus
(MARSHAM,
1802:419) (Dytiscus)
ragazziï-group (E)
abessinicus (ZIMMERMANN, 1928:177) (Gaurodytes), Ethiopia (E)
crypticoides RÉGIMBART, 1895:152, Ethiopia (E)
galamensis
NILSSON,
1992:174, Ethiopia (E)
perssoni
NlLSSON,
1992:172, Ethiopia (E)
ragazza RÉGIMBART, 1887:638, Ethiopia (E)
=ferrugatus RÉGIMBART, 1905:225
striolatus-group (WP)
striolatus (GYLLENHAL, 1808:508) (Dytiscus), Sweden (WP)
= costatus
GERHARDT,
1910:42
= rectus (BABINGTON, 1841:53) {Colymbetes)
taiwanensis-group (EP)
taiwanensis NlLSSON & WEWALKA, 1994:993, Taiwan (EP)
tristis-group (H)
adustus GUIGNOT, 1954:223, Kashmir (EP)
bipustulatus (LINNAEUS, 1767:667) (Dytiscus), Sweden (E, P)
= abdominalis (COSTA, 1847:134) (Colymbetes)
= acuductus
(MARSHAM,
1802:416) (Dytiscus)
= alpestris HEER, 1837:53
= alpicola
ZAITZEV,
1927:24
= alticola BRUNEAU
DE
MIRÉ
& LEGROS, 1963:875
= callosus (THOMSON, 1884:1031) (Gaurodytes)
= carbonarius (FABRICIUS, 1801:263) (Dytiscus)
= dolomitanus SCHOLZ, 1935:37
=falcozi GUIGNOT, 1931:7
= immaculatus (SCHRANK, 1781:201) (Dytiscus)
= kiesenwetterii SEIDLITZ, 1887:88
= latus (GEBLER, 1841:371) (Colymbetes)
= luctuosus
(GEOFFROY IN FOURCROY,
1785:67) (Dytiscus)
= maurus (ZIMMERMANN, 1919:209) (Gaurodytes)
=
peyerimhojfi BRUNEAU
DE
MIRÉ & LEGROS, 1963:875
=
picipennis
J.SAHLBERG,
1903:5
=
pyrenaeus FRESNEDA & HERNANDO, 1988:35
= regalis PETRI, 1903:49
= remotus J.SAHLBERG, 1913:45
= sexualis
REICHE,
1857:IX
= snowdonius (NEWMAN, 1833:55) (Colymbetes)
= solieri
AUBE,
1837:183
debilipes RÉGIMBART, 1899:273, Simla (EP)
= skarduensis GUIGNOT, 1958:30
= subsericatus RÉGIMBART, 1899:274
leptapsis (LECONTE, 1878:596) (Gaurodytes), Michigan (NA)
melanarius AUBÉ, 1837:180, Russia (WP)
=>/g/<ÌMS SCHIÖDTE, 1841:477
= kotschyi LETZNER, 1849:95
= tarsatus (ZETTERSTEDT, 1837:132) (Dytiscus)
= tatricus ROUBAL, 1938:19
nevadensis Hàk.LlNDBERG, 1939:32, Spain (WP)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

30
Koleopt.
Rdsch.
70 (2000)
solskii
JAKOVLEV,
1897:40, Turkestan (EP)
tristis AUBÉ, 1838:356, North America (H)
= atratus MANNERHEIM, 1853:157
= crotchi ZAITZEV, 1905:212
= dubius
MANNERHEIM,
1843:221
= kurilensis KAMI YA, 1938:35
=
picea ZAITZEV, 1905:212 [HN]
=piceolus ZAITZEV, 1908:123 [RN]
wollastoni SHARP, 1882:531, Madeira (WP)
= dissimilis (FALKENSTRÖM, 1938:13) (Gaurodytes)
=falkenstromi J.BALFOUR-BROWNE, 1944:352 [RN]
Genus Hydronebrius
JAKOVLEV,
1897 (EP)
Autapomorphy: body punctation very strong; setal row
at
posterolateral angle
of
metafemur
reduced.
amplicollis TOLEDO, 1994:207, China (EP)
cordaticollis (REITTER, 1896:233)
{Agabus),
Uzbekistan (EP)
kashmirensis (VAZIRANI, 1964:145)
(Amphizoa),
Kashmir (EP)
= gvignoti VAZIRANI, 1970:346
mattheyi mattheyi BRANCUCCI, 1980:171, Pakistan (EP)
mattheyi nepalensis BRANCUCCI, 1980:174, Nepal (EP)
Genus Hydrotrupes
SHARP,
1882 (NA)
palpalis SHARP, 1882:492, California (NA)
Genus Ilybiosoma
CROTCH,
1873 (E, H)
Autapomorphy: metatibia with anteroventral spiniferous punctures along entire length.
[Syn.: Nebriogabus GuiGNOT, 1936; Ranagabus
J.BALFOUR-BROWNE,
1939]
discicollis-group (E, EP)
discicollis (ANCEY, 1883:70) {Agabus), Ethiopia (E)
kermanensis (J.BALFOUR-BROWNE, 1939:107)
{Agabus),
Iran (EP)
seriatum-group (NA)
amaroides (SHARP, 1882:33) {Agabus), Mexico (NA)
bjorkmanae (HATCH, 1939:104)
{Agabus),
British Columbia (NA)
= recta (LECONTE, 1869:374) {Anisomera) [HN]
brevicollis (LECONTE, 1857:34) {Agabus), California (NA)
= sobrinus (MOTSCHULSKY, 1859:170) {Colymbetes)
cordatum (LECONTE, 1853:226)
{Anisomera),
New Mexico (NA)
flohrianum (SHARP, 1887:756) {Agabus), Mexico (NA)
=flohri (ZIMMERMANN, 1919:207) {Gaurodytes)
ilybiiformis (ZIMMERMANN, 1928:176) {Gaurodytes), California (NA)
lugens (LECONTE, 1852:203) {Agabus), New Mexico (NA)
= suturalis (CROTCH, 1873:423) {Gaurodytes)
minnesotensis (WALLIS, 1933:268)
{Agabus),
?Minnesota (NA)
pandurum (LEECH, 1942:128) {Agabus), California (NA)
perplexus (SHARP, 1882:498) {Agabus), California (NA)
regularis (LECONTE, 1852:203) {Ilybius), California (NA)
= oblongus (MOTSCHULSKY, 1859:169) {Ilybius)
roguus (LARSON, 1997:123) {Agabus), Oregon (NA)
seriatum (SAY, 1823:97) {Colymbetes), New York (NA)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON:
Generic classification
of
Agabus-group
of
genera
of
Agabini
(DYTISCIDAE)
31
=
arctus
(MELSHEIMER,
1844:27)
{Agabus)
= intersectus (CROTCH, 1873:419) {Gaurodytes)
=parallelus
(LECONTE,
1850:213) {Agabus)
Genus Ilybius
ERICHSON,
1832 (H)
Autapomorphies: female ovipositor with lateral ridge; pronotum with fine anterior bead
continuous; metacoxal lines reduced anteriorad.
[Syn.: Ilyobius
GEMMINGER
&
HAROLD,
1868; Idiolybius Gozis, 1886; Agabidius
SEIDLITZ,
1887;
Asternus
GuiGNOT,
1931;
Parasternus
GuiGNOT,
1936; Ilybidius
GUIGNOT,
1948]
chalconatus-group (H)
albarracinensis (FERY, 1986:345) {Agabus), Spain (WP)
= maestroae (FRESNEDA & HERNANDO, 1987:67) {Agabus)
bedeli (ZAITZEV, 1908:121) {Agabus) [RN], Algeria (WP)
=
politus (REICHE, 1861:369) {Agabus) [HN]
chalconatus (PANZER, 1797:38/17) {Dytiscus), Germany
(P)
= aterrimus (STEPHENS, 1828:79) {Colymbetes)
=fuscoaenescens (RÉGIMBART, 1877:CXLVII) {Agabus)
= nigroaeneus (MARSHAM, 1802:428) {Dytiscus)
dettneri (FERY, 1986:342) {Agabus), Portugal (WP)
gagates (AUBÉ, 1838:306) {Agabus), North America (NA)
hozgargantae (BURMEISTER, 1983:133)
{Agabus),
Spain (WP)
hulae (WEWALKA, 1984:137) {Agabus), Israel (EP)
jaechi (FERY & NlLSSON, 1993:94) {Agabus), Turkey (EP)
lagabrunensis (SCHIZZEROTTO & FERY, 1990:148)
{Agabus),
Italy (WP)
larsoni (FERY & NlLSSON, 1993:98) {Agabus), Québec (NA)
lenensis nom.n. [RN] Siberia (EP)
= aenescens (POPPIUS, 1905:18) {Agabus) [HN]
lenkoranensis (FERY & NlLSSON, 1993:95)
{Agabus),
Azerbaijan (WP)
montanus (STEPHENS, 1828:76) {Colymbetes), England (WP)
= melanocornis (ZIMMERMANN, 1915:223) {Agabus)
neglectus (ERICHSON, 1837:158) {Agabus), Germany (WP)
pederzanii (FERY & NlLSSON, 1993:95) {Agabus), Italy (WP)
pseudoneglectus (FRANCISCOLO, 1972:84) {Agabus), Italy (WP)
= skiathos (HINTERSEHER, 1981:87) {Agabus)
samokovi (FERY & NlLSSON, 1993:91) {Agabus), Bulgaria (WP)
satunini (ZAITZEV, 1913:196) {Agabus), Transcaucasia (WP)
wewalkai (FERY & NlLSSON, 1993:85) {Agabus), Turkey (EP)
erichsoni-group (H)
balkei (FERY & NlLSSON, 1993:103) {Agabus), Siberia (EP)
erichsoni (GEMMINGER & HAROLD, 1868:454) {Agabus), [IN; nigroaeneus MARSHAM sensu ERICHSON
1837],
Germany
(H)
= cincticollis (MÄKLIN,
1881:22)
{Agabus)
= lutosus (CROTCH, 1873:419) {Gaurodytes) [HN]
subtilis (ERICHSON, 1837:157) {Agabus), Germany
(P)
= altaicus (GEBLER, 1848:73) {Agabus)
opacus-group
(H)
austrodiscors (LARSON, 1996:635) {Agabus), California (NA)
confertus
(LECONTE,
1861:340) {Agabus), California (NA)
discors (LECONTE, 1861:341) {Agabus), Washington (NA)
euryomus (LARSON, 1996:639) {Agabus), California (NA)
hypomelas (MANNERHEIM, 1843:221) {Agabus), Alaska (NA)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

32
Koleopt.
Rdsch.
70 (2000)
= irregularis (MANNERHEIM, 1853:159) {Agabus)
jimzim (LARSON, 1996:643) (Agabus), Arizona (NA)
lineellus (LECONTE 1861:340) (Agabus), California (NA)
opacus (AUBÉ, 1837:173) (Agabus), Finland (H)
= gelidus (FALL, 1926:142) (Agabus) [HN]
= mimmi (J.SAHLBERG, 1875:182) (Gaurodytes)
=
pseudoconfertus (WALLIS, 1926:90) (Agabus)
= sachalinensis (KAMIYA, 1938:37) (Agabus)
=
sincarpi
(JAKOBSON, 1908:430) (Agabus) [RN]
= sibericus (SHARP, 1882:519) (Agabus) [HN]
vancouverensis (LEECH, 1937:146) (Agabus), British Columbia (NA)
vandykei (LEECH, 1942:129) (Agabus), California (NA)
verisimilis (BROWN, 1932:4) (Agabus), British Columbia (NA)
walsinghami (CROTCH, 1873:419) (Gaurodytes), Oregon (NA)
wasastjernae (C.R.SAHLBERG, 1824:167) (Dytiscus), Finland (H)
= kenaiensis (FALL, 1926:141) (Agabus)
=
palustris
(WALLIS,
1926:92) (Agabus)
subaeneus-group (H)
aenescens THOMSON, 1870:125, Scandinavia (P)
= kiesenwetteri KRAATZ, 1871:166
angustior (GYLLENHAL, 1808:500) (Dytiscus), Sweden (H)
anjae NlLSSON, 1999:36, Sakhalin (EP)
apicalis SHARP,
1873:51,
Japan (EP)
= intermediate FENG, 1936:10
ater (DE GEER, 1774:401) (Dytiscus), Sweden (P)
= ater (PANZER, 1797:38/15) (Dytiscus) [HN]
= ungularis
(LECONTE,
1862:521
)
(Colymbetes)
biguttulus (GERMAR, 1824:29) (Dytiscus), North America (NA)
= laramaeus
LECONTE,
1859:4
cinctus SHARP, 1878:169, Central Asia (EP)
= angustulus RÉGIMBART 1899:289
= chinensis CSIKI 1901:102
= deplanatus (STEVEN, 1829:26) (Colymbetes) [NO]
chishimanus
KÔNO,
1944:80, Kuril Islands (EP)
= weymarni J.BALFOUR-BROWNE, 1947:446
churchillensis WALLIS, 1939:195, Manitoba (NA)
confusus AUBÉ, 1838:280, North America (NA)
= denikei WALLIS, 1933:271
crassus THOMSON, 1856:224, Sweden (WP)
discedens
SHARP,
1882:557, Hudson Bay (NA)
fenestratus (FABRJCIUS, 1781:294) (Dytiscus), Germany (P)
- aeneus (PANZER, 1797:38/16) (Dytiscus) [HN]
= ciliatus (OLIVIER,
1791:311
)
(Dytiscus)
= evanescens DALLA TORRE, 1877:63
=
prescotti (MANNERHEIM, 1829:21) (Colymbetes)
fraterculus
LECONTE,
1862:521,
North Red River (NA)
foiliginosus fuliginosus (FABRICIUS, 1792:191) (Dytiscus), Germany (P)
=foetidus (O.F.MÜLLER, 1776:71) (Dytiscus) [NO]
- lacustris (PANZER, 1797:38/14) (Dytiscus) [HN]
= /w>imcw.sGUÉORGUIEV, 1957:25
foiliginosus turcestanicus GSCHWENDTNER, 1934:74, Kyrgyzstan (EP)
guttiger (GYLLENHAL, 1808:499) (Dytiscus), Sweden (P)
= immunis (STEPHENS, 1828:81) (Colymbetes)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON:
Generic classification
of
Agabus-group
of
genera
of
Agabini
(DYTISCIDAE)
33
=
kiesenwetteri
WEHNCKE,
1872:136
[HN]
= quadrinotatus (STEPHENS, 1828:83) {Colymbetes)
ignarus (LECONTE, 1862:521) {Colymbetes), Lake Superior (NA)
incarinatus ZIMMERMANN, 1928:181, Illinois (NA)
lateralis (GEBLER, 1832:40) {Colymbetes), Dauria (EP)
= limbatus SHARP, 1882:557
meridionalis AUBÉ, 1837:126, France (WP)
= hispanicus SHARP, 1872:260
nakanei
NlLSSON,
1994:58, Sakhalin (EP)
oblitus SHARP, 1882:560, North America (NA)
obtusus
SHARP,
1882:558, Siberia (EP)
ovalis GSCHWENDTNER, 1934:74, Siberia (EP)
picipes (KiRBY, 1837:71) {Colymbetes), North America (H)
pleuriticus
LECONTE,
1850:213, Lake Superior (NA)
= inversus SHARP, 1882:552
poppimi ZAITZEV, 1907:208, Siberia (EP)
quadriguttatus (LACORDAIRE, 1835:316) {Colymbetes), France
(P)
= obscurus (MARSHAM, 1802:414) {Dytiscus) [HN]
= sexdentatus SCHIÖDTE, 1841:487
quadrimaculatus AUBÉ, 1838:274, North America (NA)
similis THOMSON, 1856:225, Sweden
(P)
= ovatus HOCHHUTH, 1871:237
subaeneus ERICHSON, 1837:156, Germany (H)
= badenii WEHNCKE, 1871:164
= chalybeatus THOMSON, 1860:48
= lapponicus SCHOLZ, 1917:251
= suffusus
CROTCH,
1873:411
= viridiaeneus
CROTCH,
1873:411
vittiger (GYLLENHAL, 1827:379) {Dytiscus), Sweden (H)
Genus Plat ambus
THOMSON,
1859 (H, O)
Autapomorphy: Prosternai process with lateral bead broadly inflated posterior of procoxae and/or
mesocoxae widely separated.
[Syn.: Agabinus
CROTCH,
1873;
Anagabus
JAKOVLEV,
1897;
Colymbinectes
FALKENSTRÖM,
1936;
Stictogabus GuiGNOT,
1948;
,4//ogaOKs GuiGNOT,
\95\\
Agraphis GuiGNOT,
1954;
Neoplatynectes
VAZIRANI,
1970;
Paraplatynectes
VAZIRANI,
1970]
americanus-group
(NA)
americanus (AUBÉ, 1838:334) {Agabus), Mexico (NA)
= obscurior (ZIMMERMANN, 1919:212) {Gaurodytes) [HN]
confusus-group
(NA)
confusus (BLATCHLEY, 1910:229) {Rhantus), Indiana (NA)
= amplus (FALL, 1922:12) {Agabus)
glabrellus-group
(NA)
glabrellus (MOTSCHULSKY, 1859:171) {Colymbetes), California (NA)
= morulus
(LECONTE,
1861:340) {Agabus)
sculpturellus (ZIMMERMANN, 1919:205) {Agabinus), California (NA)
maculâtus-group
(O, P)
angulicollis (RÉGIMBART, 1899:273) {Agabus), Tibet (EP)
balfourbrownei VAZIRANI 1965:28, Assam (EP)
biswasi
VAZIRANI,
1965:32, Nepal (EP)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

34
Koleopt.
Rdsch.
70 (2000)
excoffieri
RÉGIMBART,
1899:281, China (EP)
fimbriatus
SHARP,
1884:445, Japan (EP)
= kansouis
FENG,
1936:9
fletcheri ZIMMERMANN, 1928:176, India (O)
guttulus (RÉGIMBART, 1899:283) (Platynectes), China (EP)
incrassatus GSCHWENDTNER, 1935:62, Burma (O)
lindbergi GuÉORGUIEV, 1963:218, Afghanistan (EP)
= guignoti VAZIRANI, 1965:27
lunulatus (STEVEN, 1829:26) (Agabus), Caucasus (P)
= sinuatus (AUBÉ, 1837:148) (Agabus)
maculatus (LINNAEUS, 1758:412) (Dytiscus), Europe (P)
= biocellatus (O.F.MÜLLER, 1776:72) (Dytiscus)
= cantalicus PlC, 1912:57
= caucasicus ZAITZEV, 1927:18
= escalerai
RÉGIMBART,
1900:121
= glacialis (GRAËLLS, 1858:42) (Agabus)
[UN]
= graellsi (GEMMINGER & HAROLD, 1868:455) (Agabus) [RN]
= hebraicus (GEOFFROY in FOURCROY, 1785:70) (Dytiscus)
= inornatus SCHILSKY, 1888:183
= ornatus (HERBST, 1784:125) (Dytiscus)
=
praetextus (DALLA
TORRE,
1877:63) (Agabus)
=
pulchellus (HEER, 1839:149) (Colymbetes) [HN]
nepalensis (GuÉORGUIEV, 1968:42) (Stictogabus), Nepal (EP)
= bhutanensis WEWALKA, 1975:157
pictipennis (SHARP, 1873:49) (Agabus), Japan (EP)
satoi BRANCUCCI, 1982:226, Nepal (EP)
schaefleini BRANCUCCI, 1988:188, China (EP, O)
strbai HENDRICH & BALKE, 1998:107, Laos (O)
wittmeri WEWALKA, 1975:158, Bhutan (EP)
optatus-group (H, O)
ater (FALKENSTRÖM, 1936:97) (Colymbinectes), China (EP)
coriaceus (RÉGIMBART, 1899:282) (Platynectes), Assam (O)
ikedai (NlLSSON, 1997:635) (Agabus), Japan (EP)
insolitus (SHARP, 1884:444) (Agabus), Japan (EP)
jilanzhui WEWALKA & BRANCUCCI, 1995:98, China (EP)
koreanus (NlLSSON, 1997:636) (Agabus), Primorye (EP)
nakanei (NlLSSON, 1997:631) (Agabus), Japan (EP)
obtusatus (SAY, 1823:99) (Colymbetes), North America (NA)
= nitidus
(SAY,
1823:98) (Colymbetes)
optatus (SHARP, 1884:445) (Agabus), Japan (EP)
= miyamotoi (NAKANE, 1959:95) (Gaurodytes)
planatus
(SHARP,
1882:503) (Agabus), North America (NA)
princeps (RÉGIMBART, 1888:615) (Platynectes), Burma (EP, O)
schillhammeri WEWALKA & BRANCUCCI, 1995:99, China (EP)
stygius (RÉGIMBART, 1899:279) (Agabus), China (EP)
ussuriensis (NlLSSON, 1997:632) (Agabus), Primorye (EP)
sawadai-group (EP, O)
kempi (VAZIRANI, 1970:330) (Agraphis), India (O)
khukri BRANCUCCI, 1990:240, Nepal (EP)
punctatipennis BRANCUCCI, 1984:153, China (EP)
regulae BRANCUCCI, 1991:345, Vietnam (O)
sawadai (KAMIYA, 1932:6) (Agabus), Japan (EP)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

NlLSSON: Generic classification of
Agabus-gwup
of genera of Agabini (DYTISCIDAE) 35
= confossus (GUIGNOT 1954:200) (Agraphis)
semenowi-group (EP)
lineatus GSCHWENDTNER, 1935:62, India (EP)
= striatus (ZENG & Pu, 1992:482) (Hydronebrius)
semenowi (JAKOVLEV, 1897:38) (Anagabus), Turkestan (EP)
sogdianus (JAKOVLEV, 1897:39) {Anagabus), Turkestan (EP)
= holzschuhi WEWALKA, 1975:155
=jucundus (GUIGNOT, 1954:222) (Agabus)
= ledouxi
(LEGROS,
1977:14) (Anagabus)
= limbibasis (REITTER, 1900:226) (Anagabus)
= vatelloides (RÉGIMBART, 1899:270) (Agabus)
wewalkai BRANCUCCI, 1982:120, India (EP)
semivittatus-group (NA)
astrictovittatus (LARSON & WOLFE, 1998:40) (Agabus), Florida (NA)
flavovittatus
(LARSON
& WOLFE, 1998:44) (Agabus), Tennessee (NA)
johannis (FALL, 1922:10) (Agabus), Florida (NA)
semivittatus
(LECONTE,
1852:204) (Agabus), Florida (NA)
= spilotus (LECONTE, 1859:5) (Agabus)
= texanus
(SHARP,
1882:505) (Agabus)
stagninus (SAY, 1823:100) (Colymbetes), North America (NA)
= striola (AUBÉ, 1838:308) (Agabus)
texovittatus (LARSON & WOLFE, 1998:46) (Agabus), Texas (NA)
spinipes-group (NA)
apache (YOUNG, 1981:349) (Agabus), Arizona (NA)
spinipes
(SHARP,
1882:32) (Agabus), Mexico (NA)
Acknowledgements
I am grateful to R. Brett, San Francisco, and P.J. Spangler, Washington D.C., who supported this
study by loan of material. Thanks are also due to D.J. Larson, St. John's, for helpful discussions.
References
ALARIE,
Y. & LARSON, D.J. 1998: Larvae of Agabinus Crotch: Generic characteristics, description of
A.
glabrellus (Motschulsky), and comparison with other genera of the subfamily Colymbetinae
(Coleoptera: Adephaga: Dytiscidae). - The Coleopterists Bulletin 52: 339-350.
ALARIE,
Y.,
SPANGLER,
P.J. &
PERKINS,
P.D. 1998: Study of the larvae of Hydrotrupes palpalis Sharp
(Coleoptera: Adephaga, Dytiscidae) with implications for the phylogeny of the Colymbetinae. -
The Coleopterists Bulletin 52: 313-332.
BALFOUR-BROWNE, J. 1939: A contribution to the study of the Dytiscidae. I. (Coleoptera, Adephaga). -
Annals and Magazine of Natural History
(11)3:
97-114.
BALKE, M.,
DETTNER,
K. & HENDRICH, L. 1997: Agabus
("Metronectes")
aubei Perris: Habitat,
morphological adaptations, systematics, evolution, and notes on the phanerofluicolous fauna
(Coleoptera: Dytiscidae). - Aquatic Insects 19: 75-90.
BEUTEL, R.G. 1994: On the systematic position of Hydrotrupes palpalis Sharp (Coleoptera: Dytiscidae).
- Aquatic Insects 16: 157-164.
BRANCUCCI, M. 1988: A revision of the genus Platambus Thomson (Coleoptera, Dytiscidae). -
Entomologica Basiliensia 12: 165-239.
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at

36
Koleopt.
Rdsch. 70
(2000)
BRINCK,
P. 1948: Coleoptera of Tristan Da Cunha. - Results of the Norwegian Scientific Expedition to
Tristan Da Cunha 1937-1938 17: 1-121 +
1
pi.
DE MARZO, L. &
NILSSON,
A.N. 1988: Structure of the "preoral comb" in larvae of European Agabini
(Coleoptera, Dytiscidae). - Entomologica, Bari 23: 191-205.
FERY, H. & NiLSSON, A.N. 1993: A revision of the Agabus chalconatus- and erichsoni-groups
(Coleoptera: Dytiscidae), with a proposed phylogeny. - Entomologica Scandinavica 24: 79-108.
FRANCISCOLO,
M.F. 1979: Coleoptera. Haliplidae, Hygrobiidae, Gyrinidae, Dytiscidae. - Fauna d'Italia
14:
1-804.
GUÉORGUIEV, V.B. 1965: Zoologische Ergebnisse der Österreichischen Iran-Expedition 1949/50
Karakorum-Expedition 1958 und Irak-Expedition 1963 (Coleoptera: Dytiscidae et Gyrinidae). -
Reichenbachia5: 114-118.
GUÉORGUIEV, V.B. 1971: Notes sur les Agabini (Coleoptera, Dytiscidae). I. Les genres Agametrus
Sharp, Leuronectes Sharp et Andonectes gen. n. - Izvestija na Zoologitjeskija Institut s Musei
Sofia
33:
165-176.
LARSON, DJ. 1975: The predaceous water beetles (Coleoptera: Dytiscidae) of Alberta: systematics,
natural history and distribution. - Quaestiones Entomologicae
11:
245-498.
LARSON, D.J. 1987: Revision of North American species of Ilybius Erichson (Coleoptera: Dytiscidae),
with systematic notes on Palaearctic species. - Journal of the New York Entomological Society
95:341-413.
LARSON, D.J. 1989: Revision of North American Agabus Leach (Coleoptera: Dytiscidae): introduction,
key to species groups, and classification of the ambiguus-, tristis-, and arcticus-groups. - The
Canadian Entomologist 121: 861-919.
LARSON, D.J. 1991: Revision of North American Agabus Leach (Coleoptera: Dytiscidae): elongatus-,
zetterstedti-, and confinis-groups. - The Canadian Entomologist 123: 1239-1317.
LARSON, D.J. 1996: Revision of North American Agabus Leach (Coleoptera: Dytiscidae): the opacus-
group. - The Canadian Entomologist 128: 613-665.
LARSON, D.J. 1997: Revision of North American Agabus Leach (Coleoptera: Dytiscidae): the seriatus-
group. - The Canadian Entomologist 129: 105-149.
LARSON, D.J. & WOLFE, G.W. 1998: Revision of North American Agabus (Coleoptera: Dytiscidae): the
semivittatus-group. - The Canadian Entomologist 130: 27-54.
LEECH, H.B. 1942: Key to the Nearctic genera of water beetles of the tribe Agabini, with some generic
synonymy (Coleoptera: Dytiscidae). - Annals of the Entomological Society of America 35: 355-
362.
NiLSSON, A.N. 1992: A revision of Afrotropical Agabus Leach (Coleoptera, Dytiscidae), and the
evolution of tropicoalpine super specialists. - Systematic Entomology 17: 155-179.
NiLSSON, A.N. 1997 (1996): A redefinition and revision of the Agabus optatus-group (Coleoptera,
Dytiscidae); an example of Pacific intercontinental disjunction. - Entomologica Basiliensia 19:
621-651.
NiLSSON, A.N. & LARSON, D.J. 1990: A review of the Agabus affinis group (Coleoptera: Dytiscidae),
with the description of a new species from Siberia and a proposed phylogeny. - Systematic
Entomology 15:227-239.
SEMENOV,
A. 1899: Notes on morphological pecularities of Agabus kessleri Hochh. (Coleoptera,
Dyticidae). - Horae Societatis Entomologicae Rossicae 32: 510-514. (In Russian.)
Anders
N.
NiLSSON
BMG,
Umeâ
University,
S- 901 87
Umeâ,
Sweden
(Anders.Nilsson@bmg.umu.se)
©Wiener Coleopterologenverein (WCV), download unter www.biologiezentrum.at