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Chatzimanolis, S., Cohen, I. M., Schomann, A. & Solodovnikov, A. (2010). Molecular phylogeny of the mega-diverse rove beetle tribe Staphylinini (Insecta, Coleoptera, Staphylinidae). —Zoologica Scripta, 39, 436–449. Phylogeny of the rove beetle tribe Staphylinini is explored by parsimony and Bayesian analyses of sequences of four genes (COI, wingless, Topoisomerase I, and 28S) for 43 ingroup (various genera of Staphylinini) and eight outgroup (two genera of Paederinae, six genera of other tribes of Staphylininae) taxa. Analyses were conducted for each gene independently and for the concatenated data set. Results of the most robust combined analyses were compared with the morphology-based phylogenies of Staphylinini (‘test phylogeny’), and with the conventional classification of this tribe. Molecular results were congruent with the ‘test phylogeny’ in the following: ancestors of Staphylinini were ‘Quediina-like’ lineages; formal subtribe Quediina mixes at least two relatively basal groups, ‘Quediina propria’ and ‘southern Quediina’; specialized subtribe Amblyopinina is an internal clade within ‘southern Quediina’; a relatively deeply nested ‘Staphylinini propria’ that unites current subtribes Staphylinina, Eucibdelina, Anisolinina, Xanthopygina and Philonthina is well supported as a monophyletic group. In strong contrast with morphology, molecular data place the tribes Othiini and Xantholinini nested within Staphylinini. Molecular results strongly conflict with morphology by uniting morphologically very different genera Holisus and Atanygnathus in one clade that has uncertain position within Staphylinini. Consistently with the most congruent areas of the morphology- and molecular-based phylogenies, taxonomic changes are implemented for the formal subtribes Quediina and Amblyopinina.
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Molecular phylogeny of the mega-diverse rove beetle tribe
Staphylinini (Insecta, Coleoptera, Staphylinidae)
STYLIANOS CHATZIMANOLIS,IAN M. COHEN,ANDREA SCHOMANN &ALEXEY SOLODOVNIKOV
Submitted: 20 January 2010
Accepted: 6 June 2010
doi: 10.1111/j.1463-6409.2010.00438.x
Chatzimanolis, S., Cohen, I. M., Schomann, A. & Solodovnikov, A. (2010). Molecular phy-
logeny of the mega-diverse rove beetle tribe Staphylinini (Insecta, Coleoptera, Staphylini-
dae). — Zoologica Scripta,39, 436–449.
Phylogeny of the rove beetle tribe Staphylinini is explored by parsimony and Bayesian
analyses of sequences of four genes (COI, wingless, Topoisomerase I, and 28S) for 43
ingroup (various genera of Staphylinini) and eight outgroup (two genera of Paederinae, six
genera of other tribes of Staphylininae) taxa. Analyses were conducted for each gene inde-
pendently and for the concatenated data set. Results of the most robust combined analyses
were compared with the morphology-based phylogenies of Staphylinini (‘test phylogeny’),
and with the conventional classification of this tribe. Molecular results were congruent
with the ‘test phylogeny’ in the following: ancestors of Staphylinini were ‘Quediina-like’
lineages; formal subtribe Quediina mixes at least two relatively basal groups, ‘Quediina
propria’ and ‘southern Quediina’; specialized subtribe Amblyopinina is an internal clade
within ‘southern Quediina’; a relatively deeply nested ‘Staphylinini propria’ that unites cur-
rent subtribes Staphylinina, Eucibdelina, Anisolinina, Xanthopygina and Philonthina is well
supported as a monophyletic group. In strong contrast with morphology, molecular data
place the tribes Othiini and Xantholinini nested within Staphylinini. Molecular results
strongly conflict with morphology by uniting morphologically very different genera Holisus
and Atanygnathus in one clade that has uncertain position within Staphylinini. Consistently
with the most congruent areas of the morphology- and molecular-based phylogenies, taxo-
nomic changes are implemented for the formal subtribes Quediina and Amblyopinina.
Corresponding author: Alexey Solodovnikov, Natural History Museum of Denmark, Zoological
Museum, Universitetsparken 15, DK-2100 Copenhagen, Denmark. E-mail: asolodovnikov@
snm.ku.dk
Stylianos Chatzimanolis, Department of Biological and Environmental Sciences, The University of
Tennessee at Chattanooga, 615 McCallie Ave. Dept 2653, Chattanooga, TN 37403, USA.
E-mail: stylianos-chatzimanolis@utc.edu
Ian M. Cohen, Department of Biological and Environmental Sciences, The University of Tennessee
at Chattanooga, 615 McCallie Ave. Dept 2653, Chattanooga, TN 37403, USA. E-mail:
ian-cohen@utc.edu
Andrea Schomann, Natural History Museum of Denmark, Zoological Museum, Universitetsparken
15, DK-2100 Copenhagen, Denmark. E-mail: aschomann@snm.ku.dk
Introduction
With 211 genera and more than 5300 described species, the
tribe Staphylinini is one of the largest and most abundant
groups of rove beetles. Except for high latitudes and alti-
tudes, or otherwise extreme land environments, species of
Staphylinini can be found in any terrestrial habitat of the
globe, often in large numbers of individuals. However, the
systematics of this enormous and dominant group of terres-
trial animals is poorly developed, and even a sound supra-
generic classification is still missing. The current division of
Staphylinini into subtribes (e.g. Herman 2001; Newton &
Thayer 2005) is rather a conventional solution than a result
of a focused phylogenetic study. Historically, this classifica-
tion stemmed from superficial morphological examination
of limited faunas, mostly Palearctic. When such a conven-
tional system is depicted in the form of a phylogeny, it gives
an unresolved polytomy representing existing subtribes (e.g.
fig. 1 in Solodovnikov & Newton 2005). Some of these
branches are species poor, while others are extremely
speciose, but they still lack any further subdivision, leaving
dozens of genera and thousands of species to be organized
no better than in alphabetical order.
436 ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449
Zoologica Scripta
In search of a better supra-generic classification of
Staphylininae, a series of studies recently attempted phylo-
genetic reconstructions applicable to this subfamily and its
largest tribe Staphylinini (Assing 2000; Solodovnikov &
Newton 2005; Solodovnikov 2006; Solodovnikov & Scho-
mann 2009). As expected, those specifically focused on
Staphylinini (Solodovnikov 2006; Solodovnikov & Scho-
mann 2009) revealed a phylogenetic pattern for this tribe
that conflicted in many ways with the current system and
showed more hierarchical steps of branching between tribe
and genus levels (fig. 1 in Solodovnikov & Schomann
2009). But many details of the new pattern were neither
clear nor consistent. Morphology alone may not be able to
resolve all phylogenetic questions related to Staphylinini.
Molecular data, despite their triumphal expansion into
modern animal systematics during the last decade, hitherto
has had no impact either on the systematics of the tribe
Staphylinini or on Staphylinidae as a whole (a family of
beetles larger than all classes of vertebrate animals taken
together). This is largely due to lack of expertise and
resources currently available for the study of rove beetles.
But it seems likely that molecular approaches can provide
good additional data for rigorous exploration of the phy-
logeny of Staphylinidae and Staphylinini in particular.
Given the enormous size of this group and the fact that
gathering molecular data remains a rather expensive and
time- and labour-consuming enterprise (especially if seek-
ing and collecting fresh DNA-quality material is also taken
into account), a pilot study is needed before larger
resources are invested in such research. Such a project is
relevant also because even commonly used molecular
phylogenetic markers have not yet been explored for
Staphylinidae. With the mentioned morphology-based
phylogenies in hand as some evaluation criteria, it is an
appropriate moment for conducting a pilot molecular phy-
logenetic study of Staphylinini.
Reciprocally, molecular data with even a limited number
of taxa and genes analysed would be adequate to test the
newly emerging morphology-based phylogeny of Staphyli-
nini. Recent morphological results, if translated into a new
classification system, would lead to a significant reclassifica-
tion of the tribe. Therefore, testing them as much as possi-
ble prior to implementing any large-scale changes in
classification seems highly desirable. We present here a first
molecular phylogenetic analysis of the tribe Staphylinini,
compare its results to the earlier morphology-based phylo-
genetic pattern, and implement some taxonomic changes
implied by the congruent parts of the morphology-based
and molecular phylogenies. This study is a stepping stone
towards further morphological, molecular and holistic
large-scale phylogenetic studies of Staphylinini, paving the
way for a new classification of this mega-diverse tribe.
There are few published molecular phylogenetic analy-
ses of Staphylinidae. Ballard
et al.
(1998) published a phy-
logeny of Staphylinidae using cytochrome
b
and 12s genes,
Maus
et al.
(2001) reconstructed a phylogeny of the genus
Aleochara
(subfamily Aleocharinae) using COI and COII
genes, and Thomas (2009) presented a preliminary phylog-
eny of Aleocharinae. All of these analyses have no use in
our study as the first included only 23 species of Staphy-
linidae (including only one species of Staphylinini) and
intended to test different methodologies and data sets,
whereas the others involved relationships within a subfam-
ily distantly related to Staphylinini. Some higher level
molecular phylogenetic studies on Staphyliniformia or
Coleoptera (Korte
et al.
2004; Caterino
et al.
2005; Hunt
et al.
2007) have included Staphylinini, but these are far
too fragmentary to be useful for our work.
Materials and methods
Taxon sampling
Table 1 in the online supplement depicts all taxa used in
the analyses along with the GenBank accession numbers.
We assembled a sample of species that, insofar as possible,
represented supra-generic diversity of Staphylinini glob-
ally, and was compatible with the taxon sampling of the
morphology-based analyses underlying the ‘test phylog-
eny.’ Naturally, DNA-quality specimens were not available
for all taxa used in previous morphological analyses.
Therefore some taxa were replaced by other closely related
species. We sampled 51 species belonging to 44 genera
representing all subtribes of Staphylinini, as well as most
other tribes in the subfamily Staphylininae: Arrowinini,
Diochini, Othiini, Platyprosopini and Xantholinini. Two
species of Paederinae that is the sister subfamily to Staph-
ylininae, were included as the primary outgroups and used
to root the resulting trees. DNA voucher specimens are
deposited at the Zoological Museum, Natural History
Museum of Denmark (ZMUC), the University of Tennes-
see at Chattanooga (UTCI) and the Santa Barbara
Museum of Natural History (SBMNH) (for deposition
details see Table 1 in the online supplement).
Gene sampling
We amplified the following genes: mitochondrial cyto-
chrome oxidase I (COI); nuclear protein-coding wingless
(Wg), topoisomerase I (TP), as well as the ribosomal 28S
rDNA (28S) (detailed primer information for each respec-
tive gene fragment is shown in Table 2 in the online sup-
plement). These genes were selected for several reasons:
COI is commonly used in molecular phylogenetic studies,
is easy to amplify, and has widely reported success in
detecting shallow phylogenetic patterns (Caterino et al.
2000); Wg and TP are relatively easy to amplify and have
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 437
demonstrated good phylogenetic performance at the taxo-
nomic level between genus and family (e.g. Wild & Madd-
ison 2008; Maddison et al. 2009; Miller et al. 2009); 28S is
relatively easy to amplify, with good performance in recov-
ering deeper phylogenetic divergences; difficulties in align-
ment are somewhat offset by its wide use in beetle
phylogenetic studies and resultant availability of broad
comparative data. With this set of genes, we aimed to get
phylogenetic resolution across a wide gradient of diver-
gences: from subfamily to species level. Additionally, we
used genes employed by the NSF Beetle Tree of Life pro-
ject (Wild & Maddison 2008) to promote sharing and
accumulation of beetle molecular data.
Molecular protocols
All specimens were collected in 95%–100% ethanol and
stored at )80 C. Total genomic DNA was extracted
using DNeasy Tissue extraction kit (Qiagen). The amplifi-
cation profile for COI consisted of an initial denaturation
for 5 min at 94 C, 35 cycles of: 45 s at 94 C, 30 s at
45 C and 1 min at 72 C; followed by a 2 min final
extension at 72 C. For both wingless and topoisomerase
I, we used a nested PCR approach using Platinum Taq
(Invitrogen, Carlsbad, CA, USA) that significantly
improved the performance of the reaction. The amplifica-
tion profile for Wg using the external primers consisted of
an initial denaturation for 1 min at 94 C, 35 cycles of:
30 s at 94 C, 30 s at 59 C and 3 min at 72 C; followed
by a 5 min final extension at 72 C. The PCR profile for
the internal primers was similar with the exception that
the cycles had extension of 1 min at 72 C. The amplifica-
tion profile for TP using the external primers consisted of
an initial denaturation for 1 min at 94 C, 35 cycles of:
45 s at 94 C, 45 s at 59 C and 3 min at 72 C; followed
by a 5 min final extension at 72 C. For the internal pri-
mer reaction, the PCR profile was modified based on the
recommendation of Wild & Maddison (2008) to avoid the
presence of multiple bands: we started with an initial
denaturation of 1 min at 94 C, seven cycles with short
extension time (45 s at 94 C, 45 s at 59 C and 35 s at
72 C) and 27 cycles with longer extension (45 s at 94C,
45 s at 59C and 1.5 min at 72 C). In some cases when a
double band was still present a gel fragment containing
the target band was excised and purified using QIAquick
Gel Extraction kit (Qiagen) and subsequently re-amplified
before sequencing. The amplification profile for 28S con-
sisted of an initial denaturation for 2 min at 94 C, 35
cycles of: 45 s at 94 C, 30 s at 50 C–58 C and 1 min at
72 C; followed by a 2 min final extension at 72 C. PCR
products were purified using ExoSAP-IT (USB, Cleveland,
OH, USA) or QIAquick PCR Purification kit (Qiagen,
Valencia, CA, USA). Sequencing was performed either
commercially by Macrogen Inc. (Seoul, Korea) or at the
Natural History Museum of Denmark Molecular Labora-
tory using an ABI 3130XL sequencer. All fragments were
sequenced in both directions.
Alignment
Sequences were originally edited using Sequencher 4.8
(GeneCodes Corp., Ann Arbor, Michigan, USA), then
imported in Se-Al v2.0a11 (Rambaut 2002) for initial man-
ual alignment and finally exported to Mesquite 2.71
(Maddison & Maddison 2009) for further alignment and
other analyses. Alignment of COI sequences was trivial
because no indels were present. The sequences for wing-
less and topoisomerase were aligned using Opal (Wheeler
& Kececioglu 2007) within Mesquite. No significant het-
erozygosity was observed and in rare occasions where mul-
tiple peaks were present at a single position for both
reads, the bases were coded as ambiguous. The 5¢and 3¢
ends of wingless were easy to align, but a central region of
about 50 nucleotides contains multiple insertions and dele-
tions of amino acids and was more challenging to align.
That region of wingless is consistently hard to align in
beetles (e.g. Wild & Maddison 2008; Maddison et al.
2009). Alignment of the TP sequences was straightforward
as there were few indels. To align the 28S sequences, we
used the SILVA comprehensive ribosomal RNA database
(Pruesse et al. 2007), utilizing the online SINA (Web)
Aligner (http://www.arb-silva.de/aligner/).
Study design and phylogenetic analyses
Parsimony and Bayesian analyses were conducted for the
same taxon sample, first for each gene independently, and
then for the concatenated data set. Only results of the
combined analyses are presented in detail; individual gene
analyses are referred to only when it is instructive to
explore their respective input and individual gene trees are
presented as online supplement Figures.
First, phylogenies obtained by Bayesian and parsimony
analyses of the molecular dataset are compared to the
morphology-based ‘test phylogeny’. Then, Bayesian and
parsimony analyses of the molecular data are compared
with each other. Finally, based on the summary of the
shared and conflicting clades of both molecular analyses,
we assessed them against the morphology-based ‘test
phylogeny’. We discuss which groups of the newly
emerging phylogeny of Staphylinini are likely to be real
and which might be possible artefacts of data shortage or
method deficiency by performing pairwise comparisons of
results from two methods of molecular analyses, and of
the molecular phylogenetic results with those that are
morphology-based. Parsimony analyses were performed
in PAUP 4.0b10 (Swofford 2002), with the following set-
Molecular phylogeny of Staphylinini dS. Chatzimanolis et al.
438 Zoologica Scripta, 39, 5, September 2010, pp 436–449 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters
tings: all bases were weighted equally (although several
weighting schemes were tried for the third codon posi-
tion of COI), gaps treated as 5th base, heuristic search
with TBR branch swapping and 1000 replicates. Non-
parametric bootstrap support was calculated with 500
pseudoreplicates and 10 addition sequences per replicate.
The maximum likelihood models for the three genes
were calculated using jModelTest 0.1.1 (Posada 2008)
and MrModeltest2 (Nylander 2004) using the Akaike
information criterion (AIC). Bayesian analyses were per-
formed with MrBayes 3.1.2 (Huelsenbeck & Ronquist
2001). For all analyses we used two runs with 3 ·10
7
generations, each having four Markov chains, heating
equal to 0.2 and sampling every 1000 generations. To
assess burn-in we used the program AWTY (Wil-
genbusch et al. 2004) as well as the convergent diagnostic
tools in MrBayes.
Morphology-based ‘test phylogeny’; formal and
informal groups of Staphylininae
With minor disagreements among each other, hitherto
existing morphology-based phylogenies of Staphylininae
(Assing 2000; Solodovnikov & Newton 2005), Staphylinini
(Solodovnikov 2006; Solodovnikov & Schomann 2009)
reveal the pattern illustrated in Fig. 3. At least the major
branches of this phylogenetic scheme form a plausible evo-
lutionary scenario when tested against biogeographical
(Solodovnikov & Schomann 2009) or paleontological
(A. Solodovnikov, S. Chatzimanolis, unpublished data) evi-
dence. For our purposes a consensus topology was derived
from those studies and is called here the ‘test phylogeny’
(tree in Fig. 3). This ‘test phylogeny’ does not question
the monophyly of Staphylinini and proposes its sister rela-
tionship with the tribe Arrowinini. Together with Arrowi-
nini, Staphylinini form a clade that in turn is sister to the
lineage consisting of the tribes Platyprosopini, Maorothi-
ini, Othiini, Diochini and Xantholinini (the so-called
‘Xantholinine-lineage’). The phylogeny of the ‘Xantholi-
nine-lineage’ is not entirely clear (for example, consider
incongruence between Assing (2000) and Solodovnikov &
Newton (2005)), but it is less relevant for the focus of this
study.
According to this ‘test phylogeny’, ancestral Staphylinini
were some ‘Quediina-like’ rove beetles which now are rep-
resented by monotypic or species poor genera like
Valdivi-
odes
Smetana, 1981
, Astrapaeus
Gravenhorts, 1802
, Lonia
Strand, 1943 and
Quediomacrus
Sharp, 1884. Early in their
evolution some of those ancestral groups diverged into
two species-rich stocks: ‘southern Quediina’ (or ‘Tanygna-
thinina sensu novo’ in Solodovnikov & Schomann 2009)
now dominating in the south temperate areas of the globe;
and ‘Quediina propria’, abundant in, and mostly restricted
to, the north-hemisphere landmasses. South American and
Australian members of ‘southern Quediina’ independently
gave rise to extremely specialized forms of mammal
symbionts, artificially united in the current system as the
subtribe Amblyopinina.
The core group of Staphylinini, called here ‘Staphyli-
nini propria’, is hypothesized to be monophyletic and is
the most species-rich lineage. It consists of several
clades, which, in the current formal system, are united in
the subtribes Anisolinina, Staphylinina, Eucibdelina,
Philonthina and Xanthopygina. Within ‘Staphylinini
propria’, with single exceptions (e.g. Solodovnikov &
Schomann 2009), morphology-based analyses confirm the
monophyly of the formally recognized subtribes Staphy-
linina, Philonthina and Xanthopygina (the latter only as
far as the Neotropical fauna is concerned, see Chatzi-
manolis 2008; Chatzimanolis & Ashe 2009; Solodovnikov
2009). But it casts doubts on the monophyly of Anisolin-
ina, and it does not provide enough data to conclude
about the status of the subtribe Eucibdelina that presum-
ably is nested within Staphylinina. The ‘test phylogeny’
suggests that
Holisus
Erichson, 1839 with its highly mod-
ified morphology for subcortical habitats (now forming
its own subtribe Hyptiomina), is just a specialized mem-
ber of subtribe Philonthina.
The respective terminology for informal groups that
arose from recent morphology-based phylogenetic studies
of Staphylininae and Staphylinini, and necessary comments
on formal tribes and subtribes of Staphylininae are sum-
marized below (names in alphabetical order).
‘Acylophorus-lineage’.
A group within the formal subtribe
Quediina, that includes the genera
Acylophorus
Nordmann,
1837
, Acylochsellus
Smetana, 1995
, Anchocerus
Fauvel, 1905
,
Anaquedius
Casey, 1915
, Hemiquedius
Casey, 1915
, Austra-
lotarsius
Solod. & Newton, 2009 and probably also
Eurypo-
rus
Erichson, 1839. According to morphology, this group
most likely belongs to the
Quediina propria
lineage of
Quediina.
Amblyopinina
Seevers, 1944
.
A subtribe of Staphylinini
uniting a few highly specialized genera from South Amer-
ica, and one genus (
Myoptyphlus
Fauvel, 1883) from Aus-
tralia, all living in the fur of small mammals as
commensals. Recent morphology-based phylogenetic data
(Solodovnikov & Schomann 2009, and Solodovnikov,
unpublished) indicated that the South American genera
and
Myoptyphlus
do not form a monophyletic group, and
forming at least two lineages within the newly recovered
group ‘southern Quediina’.
Anisolinina
Hayashi, 1993
.
A highly problematic formal
subtribe of Staphylinini that does not appear to be
monophyletic in any of the morphology-based phyloge-
netic analyses of this tribe. The most up-to-date compo-
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 439
sition of Anisolinina can be found in Newton & Thayer
(2005).
Arrowinini
Solodovnikov & Newton 2005
.
A recently
described monotypic tribe of Staphylininae, a plesiomor-
phy-rich sister-group to the tribe Staphylinini (Solodovni-
kov & Newton 2005).
Diochini
Casey, 1906
.
A small tribe of Staphylininae with
uncertain sister relationships, member of the ‘Xantholinini-
lineage’.
Eucibdelina
Sharp, 1889. A formal subtribe of Staphyli-
nini, uniting several genera of the so-called
Eucibdelus
-line-
age, which seem to be members of Staphylinina modified
for arboreal mode of life (Hayashi 2005). Although
formally valid, this subtribe was not widely used and it
was overlooked in the recent catalogues (Herman 2001;
Newton & Thayer 2005).
Hyptiomina
Casey, 1906
.
A monotypic subtribe of Staph-
ylinini that includes the flat subcortical genus
Holisus
Erichson, 1839. Recent morphology-based phylogenetic
analysis (Solodovnikov & Schomann 2009) suggested that
Holisus
is a morphologically derived member of the sub-
tribe Philonthina.
Othiini
Thomson, 1859
.
A tribe of Staphylininae, mem-
ber of the ‘Xantholinini-lineage’, presumably sister-group
to Xantholinini (Solodovnikov & Newton 2005).
Platyprosopini.
A rather peculiar monotypic tribe of
Staphylininae with unclear sister relatonships, presumably
a basal member of the ‘Xantholinini-lineage’ (Solodovni-
kov & Newton 2005).
Philonthina
Kirby, 1837
.
A large subtribe of Staphylinini,
its composition as given in Newton & Thayer (2005), Sol-
odovnikov (2009) and Solodovnikov & Schomann (2009).
According to the morphology-based phylogenetic analysis
of Solodovnikov & Schomann (2009) the subtribe Hyptio-
mina is nested within Philonthina.
‘Southern Quediina’ (= ‘Tanygnathinina sensu novo’).
A
large part of the formal subtribe Quediina that includes
south temperate members of the currently polyphyletic
genus
Quedius
Stephens, 1829 (Solodovnikov 2006)
,
most
of the south temperate genera of Quediina, as well as the
global genera
Heterothops
Stephens, 1829 and
Atanygnathus
Jakobson, 1909. Also, Amblyopinina, according to recent
studies (see above), are placed within this group. Because
the family-group name Tanygnathinina is older than
Amblyopinina, the former (but in a new sense =
sensu novo
)
is also used to refer to this newly recovered group.
Staphylinini
Latreille, 1802
.
The largest tribe of Staphy-
lininae, well supported as a monophyletic group (e.g. Solo-
dovnikov & Newton 2005).
‘Staphylinini-lineage’.
An informal group within the sub-
family Staphylininae that includes the tribes Staphylinini
and Arrowinini. Staphylinini was formerly considered in
the rank of the subfamily Staphylininae (Staphylininae
sensu stricto,
for details see Solodovnikov & Newton 2005).
Staphylinini propria.
A group of relatively derived mem-
bers of Staphylinini, well supported as a monophyletic
lineage. It includes formal subtribes Anisolinina, Staphy-
linina, Eucibdelina, Philonthina and Xanthopygina.
Tanygnathinina
Reitter, 1909
.
This monogeneric sub-
tribe of Staphylinini was established for the genus
Atany-
gnathus
. The discovery of the genus
Natalignathus
Solod.,
2005 (Solodovnikov 2005) and recent phylogenetic studies
(Solodovnikov 2006; Solodovnikov & Schomann 2009) led
to the conclusion that
Atanygnathus
is a rather derived
member of ‘
southern Quediina
’.
‘Tanygnathinina sensu novo’.
See
‘Southern Quediina’.
Quediina
Kraatz, 1857
.
A formal subtribe of Staphylinini
that is undoubtedly polyphyletic. According to recent
studies (Solodovnikov 2006; Solodovnikov & Schomann
2009; and Solodovnikov, unpublished) it consists of two
larger lineages ‘Quediina propria’ and ‘southern Quediina’,
a few rather isolated genera apparently representing basal
lineages of Staphylinini (e.g.
Valdiviodes, Lonia, Quedioma-
crus
) and some genera with still unclear phylogenetic rela-
tionships (e.g.
Parisanopus
Brethes, 1900
, Bolitogyrus
Chevrolat, 1842).
‘Quediina propria’ A large part of the formal subtribe
Quediina that includes its type species. The core of this
group includes the north temperate species of the cur-
rently polyphyletic genus
Quedius
(Solodovnikov 2006) and
several other genera mostly restricted to the northern
hemisphere.
Xantholinini
Erichson, 1839
.
A tribe of Staphylininae,
the largest among ‘Xantholinini-lineage
, presumably sister
to Othiini.
‘Xantholinini-lineage’.
An informal group within the sub-
family Staphylininae that includes tribes formerly placed
in the subfamily Xantholininae (for details see Solodovni-
kov & Newton 2005). Of the tribes used in the analyses in
this paper, Diochini, Platyprosopini, Othiini and Xantholi-
nini belong to this group.
Xanthopygina
Sharp, 1884
.
A formal subtribe of Staphyli-
nini. Although its monophyly has not been rigorously
tested, preliminary morphological data suggest that the
Neotropical core of this subtribe may be a monophyletic
group. Non-Neotropical genera that hitherto have been
placed in Xanthopygina likely belong to other lineages of
Staphylinini (Solodovnikov 2009).
Results
Sequence data
The maximum likelihood model selected was GTR+I+C
for COI, Wg and TP and GTR+Cfor the 28S dataset.
COI sequences were 798-bp long with 363 parsimony
Molecular phylogeny of Staphylinini dS. Chatzimanolis et al.
440 Zoologica Scripta, 39, 5, September 2010, pp 436–449 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters
informative characters, 65 variable parsimony uninforma-
tive characters, and 370 constant characters and average
base composition. Wg sequences were 364–396 bp long
with 173 parsimony informative characters, 39 variable
parsimony uninformative characters and 175 constant
characters. TP sequences were between 718 and 726-bp
long with 337 parsimony informative characters, 60 vari-
able parsimony uninformative characters and 329 constant
characters. 28S sequences were between 678 and 1244-bp
long with 536 parsimony informative characters, 452 vari-
able parsimony uninformative characters and 256 constant
characters. The combined dataset was 3164-bp long. We
sequenced 41 out of 51 taxa for 28S (see Table 1 in the
online supplement for more details) and all taxa for the
other three genes. The corresponding GenBank accession
numbers (GU377314–GU377506) are listed on Table 1 in
the online supplement.
The Bayesian and parsimony trees for the concatenated
datasets are shown in Figs 1 and 2. Bayesian and parsi-
mony trees for the individual gene analyses are available in
the online Supplementary materials. We describe in detail
the results from the combined Bayesian and Parsimony
analyses and refer to the single gene-based topologies only
when necessary to explore the influence of various genes
in more depth. In all cases when we refer to a parsimony
analysis we discuss the 50% majority rule consensus tree.
Bayesian analysis: tree topology (Fig. 1)
The tree was well resolved with mostly high posterior
probabilities. Subfamily Staphylininae was recovered as
monophyletic. Tribes Arrowinini and Diochini together
were sister to the rest of Staphylininae. Tribe Staphylinini
was not recovered as monophyletic: tribes Platyprosopini,
Xantholinini and Othiini were placed inside it. Platyproso-
pini was linked as a sister taxon to Hyptiomina + Tany-
gnathinina, the three together forming a rather basal clade
of Staphylininae. The rest were divided further into two
clades: the smaller, where ‘southern Quediina’ (including
Amblyopinina) is sister to the clade Othiini + Xantholinini;
and the larger, in composition corresponding to ‘Quediina
propria’ and ‘Staphylinini propria’ of the ‘test phylogeny’.
‘Quediina propria’, however, was not recovered as mono-
phyletic, its members branched off as two separate clades.
The next clade consisted of the monophyletic ‘Staphylinini
propria’ plus Bolitogyrus as sister lineage. All currently rec-
ognized subtribes of ‘Staphylinini propria’ (Anisolinina,
Staphylinina, Philonthina and Xanthopygina) were recov-
ered as monophyletic, except that the genus Algon Sharp,
1874 formed a separate lineage sister to Anisolinina plus
Staphylinina. Xanthopygina (excluding Algon) was recov-
ered as sister to Philonthina and Anisolinina as sister to
Staphylinina.
Parsimony analysis: tree topology (Fig. 2)
Subfamily Staphylininae was recovered as monophyletic.
Tribes Platyprosopini and Arrowinini plus Diochini are
successive sister-groups to the rest of Staphylininae. Staph-
ylinini was not recovered as monophyletic: tribes Xanthol-
inini and Othiini were placed inside it. The first split of
Staphylinini plus Othiini plus Xantholinini divided the
group into two clades: a smaller one, with Othiini plus
Xantholinini as sister-group to the clade consisting of
‘southern Quediina’ (including Amblyopinina) plus part
of the ‘Quediina propria’; and a larger one including the
rest of the Staphylinini. The larger of these two basal
clades in turn had two clades branching off sequentially:
‘Quediina propria’ (largely monophyletic except that Anch-
ocerus + Acylophorus were placed elsewhere, as discussed
above), and a clade combining Bolitogyrus with Hyptiomina
and Tanygnathinina as sister-groups. The remaining
large sister clade corresponding to the monophyletic
‘Staphylinini propria’ was divided into two major lineages:
one with largely monophyletic Xanthopygina and mono-
phyletic Philonthina sister to each other; and the other
consisting of Algon as sister to a clade with Staphylinina
paraphyletic with respect to Anisolinina.
Bayesian analysis versus ‘test phylogeny’
Consistent with morphology, subfamily Staphylininae was
recovered as monophyletic. The most striking difference
of that tree from the ‘test phylogeny’ was the different
basal topology of Staphylininae, including non-monophyly
of Staphylinini. The position of Arrowinini and Diochini
within Staphylininae was consistent with the morphology-
based results, but their linkage conflicted with that, where
Diochini was a member of the ‘Xantholinini-lineage’. A
rather basal position of Platyprosopini within Staphylini-
nae was consistent with the morphology-based results,
but its sister-group relationships were not. The position
of Atanygnathus (Tanygnathinina) in this clade together
with Holisus (Hyptiomina) was strongly conflicting with
the morphology-based data, where Atanygnathus was a
derived member of ‘southern Quediina’ and Holisus a
derived, specialized member of Philonthina. Monophyly
(with the exception of Atanygnathus) of the ‘southern
Quediina’ clade, inclusion of Amblyopinina and a rela-
tively basal position of that clade was consistent with the
scenario of the ‘test phylogeny’. However, placement of
the Othiini and Xantholinini as a sister-group to ‘southern
Quediina’ and within the Staphylinini of the ‘test phylog-
eny’ contrasting strikingly with the latter and with the
established classification of Staphylininae. The next large
clade included only (though not all) Staphylinini, and its
topology was rather consistent with that recovered by
morphology. Unlike morphology, molecular data did not
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 441
recover ‘Quediina propria’ as a monophyletic group, but,
consistently with the ‘test phylogeny’ they were placed
basally in the tree. Similar to the morphology-based anal-
ysis, subtribes Staphylinina, Philonthina and Xanthopygi-
na (as far as only the Neotropical genera were concerned)
were recovered as monophyletic and rather terminal
groups, with the same hierarchy of sister-group relation-
ships among them: Philonthina sister to Xanthopygina
and both sister to Staphylinina. However, unlike the mor-
phological analysis that linked Anisolinina closer to Philo-
nthina + Xanthopygina than to Staphylinina, molecular
data placed them closer to the latter. The sister relation-
ship of Tolmerinus and Tympanophorus (Anisolinina) in the
molecular analysis was consistent with the results of the
‘test phylogeny’. Unfortunately, our sample of Anisolinina
for the molecular analysis was very limited, so we could
not rigorously test either the monophyly of the subtribe
or its relationships to other lineages. Notably, Anisolinina
Arrowinus relictus (ARROWININI)
Diochus sp. (DIOCHINI)
Atanygnathus acuminatus
Atanygnathus sp
Holisus sp. (Hyptiomina)
Platyprosopus sp. (PLATYPROSOPINI)
Amblyopinus emarginatus (Amblyopinina)
Quedius antipodus (Quediina)
Quediocafus insolitus (Quediina)
Atrecus punctiventris
Othius punctulatus
Nudobius pugetanus (XANTHOLININI)
Algon sp. (Xanthopygina)
Tolmerinus sp.
Tympanophorus sp.
Dinothenarus saphyrinus
Tasgius morsitans
Ocypus olens
Tasgius pedator
Naddia sp.
Ontholestes murinus
Platydracus cinnamopterus
Platydracus sp.
Hadropinus fossor
Hadrotes crassus
Thinopinus pictus Belonuchus mimeticus
Neobisnius occidentoides
Philonthus splendens
Cafius seminitens
Hesperus sp.
Hesperopalpus sp.
Elecatopselaphus sp.
Xanthopygus sp.
Xenopygus analis
Gastrisus sp.
Nausicotus spectabilis
Nordus fungicola
Oligotergus sp.
Bolitogyrus sp. (Quediina)
Quedius cruentus
Velleius dilatatus
Quedius picipes
Quedius suturalis
Quedius molochinus
Indoquedius sp.
Anaquedius vernix
Acylophorus sp.
Anchocerus sp.
Astenus sp.
Paederus sp.
1
0.95
1
1
0.93
1
1
1
1
0.95
1
0.96
0.95
1
1
1
0.99
1
0.82
1
1
0.79
1
0.93
1
1
0.88
1
0.76
1
0.74
0.99
0.96
1
1
1
1
0.96 1
1
1
1
1
0.88
0.87
(Tanygnathinina)
(OTHIINI)
(Anisolinina)
(Staphylinina)
(Philonthina)
(Xanthopygina)
(Quediina)
(Quediina)
[PAEDERINAE]
Informal monophyletic groups revealed in morphology-based phylogeny
(Solodovnikov & Schomann 2009):
“Philonthina lineage”
“Philonthina propria”
“Quediina propria”
“Staphylinina”
“Tan
yg
nathinina sensu novo” (= “Southern Quediina” here)
Fig. 1 Bayesian phylogenetic tree for the partitioned combined analysis of four genes. Only posterior probabilities above 0.75 are shown.
Branches coloured consistently with the informal monophyletic groups recovered in the morphology-based phylogeny (Solodovnikov &
Schomann 2009). Suprageneric taxa of Staphylininae accepted in the current conventional system (Newton & Thayer 2005) are given in
round brackets: tribes in capitols; subtribes of Staphylinini in lower case letters.
Molecular phylogeny of Staphylinini dS. Chatzimanolis et al.
442 Zoologica Scripta, 39, 5, September 2010, pp 436–449 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters
was not supported as monophyletic by the morphological
data. Consistent with the morphological data, molecular
analysis placed Bolitogyrus and Algon in the base of ‘Staph-
ylinini propria’.
Parsimony analysis versus ‘test phylogeny’
The overall topology of the parsimony combined tree was
similar to that of the Bayesian analysis. Subfamily Staphy-
lininae was recovered as monophyletic, although its basal
Acylophorus sp
Anchocerus sp
Quediocafus insolitus
Quedius antipodus
Amblyopinus emarginatus (Amblyopinina)
Atrecus punctiventris
Othius punctulatus
Nudobius pugetanus (XANTHOLININI)
Algon sp (Xanthopygina)
Dinothenarus saphyrinus
Tasgius morsitans
Tasgius pedator
Ocypus olens
Naddia sp
Ontholestes murinus
Platydracus cinnamopterus
Platydracus sp
Tolmerinus sp
Tympanophorus sp
Hadropinus fossor
Hadrotes crassus
Thinopinus pictus
Belonuchus mimeticus
Cafius seminitens
Neobisnius occidentoides
Philonthus splendens
Hesperus sp
Hesperopalpus sp
Elecatopselaphus sp
Xenopygus analis
Gastrisus sp
Nausicotus spectabilis
Nordus fungicola
Oligotergus sp
Xanthopygus sp
Atanygnathus acuminatus
Atanygnathus sp
Holisus sp (Hyptiomina)
Bolitogyrus sp (Quediina)
Anaquedius vernix
Indoquedius sp
Quedius cruentus
Velleius dilatatus
Quedius picipes
Quedius suturalis
Quedius molochinus
Arrowinus relictus (ARROWININI)
Diochus sp (DIOCHINI)
Platyprosopus sp (PLATYPROSOPINI)
Astenus sp
Paederus sp
97
88
85
99
91
97
100
100
93
80
82
93
100
100
100
80
94 97
95
98
97
100
84
Informal monophyletic groups revealed in morphology-based phylogeny
(Solodovnikov & Schomann 2009):
“Philonthina lineage”
“Philonthina propria”
“Quediina propria”
“Staphylinina”
“Tany
g
nathinina sensu novo” (= “Southern Quediina” here)
(Quediina)
(OTHIINI)
(Staphylinina)
(Anisolinina)
(Staphylinina)
(Philonthina)
(Xanthopygina)
(Tanygnathinina)
(Quediina)
[PAEDERINAE]
Fig. 2 50% majority rule consensus tree of four equally parsimonious trees for the combined parsimony analysis. Tree length = 13 386
steps. Only bootstrap values above 75 are shown. Branches coloured consistently with the informal monophyletic groups recovered in the
morphology-based phylogeny (Solodovnikov & Schomann 2009). Suprageneric taxa of Staphylininae accepted in the current conventional
system (Newton & Thayer 2005) are given in round brackets: tribes in capitols; subtribes of Staphylinini in lower case letters.
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 443
topology and non- monophyletic Staphylinini did not cor-
respond with the ‘test phylogeny.’ Consistent with mor-
phology, Platyprosopini was a basal lineage followed by
the clade composed of Arrowinini plus Diochini. The
clade branching off next includes Othiini plus Xantholi-
nini, which together were sister to ‘southern Quediina’
(including Amblyopinina), plus part of ‘Quediina propria’.
Notably, ‘southern Quediina’ was paraphyletic with
respect to the clade Acylophorus +Anchocerus. The core of
‘Quediina propria’ was recovered as monophyletic and as a
rather basal clade of Staphylinini, consistently with the
‘test phylogeny’. Contrary to the morphology-based data,
Hyptiomina and Tanygnathinina were grouped in a clade
with Bolitogyrus outside ‘Staphylinini propria.’ The large
clade ‘Staphylinini propria’ uniting subtribes Anisolinina,
Staphylinina, Philonthina and Xanthopygina was con-
sistent with the morphology-based phylogeny, as were
Philonthina and Xanthopygina as monophyletic sister-
groups. Also, most of Staphylinina was supported as
monophyletic except a group of genera (Hadropinus,
Hadrotes and Thinopinus) that forms a sister clade to Aniso-
linina. Thus, contrary to the ‘test phylogeny’ Staphylinina
was paraphyletic with respect to Anisolinina. And, also
contrary to that, Algon has a position deeper inside ‘Staph-
ylinini propria’, as sister to Staphylinina + Anisolinina.
Discussion
Bayesian versus parsimony analysis (Figs 1 and 2)
Although based on exactly the same dataset, the results from
these two methods of phylogenetic inference were not iden-
tical. The main areas of congruence between them con-
cerned major branching events (overall tree topology), as
well as unambiguous monophyly and internal resolution of
the clade ‘Staphylinini propria.’ According to both analyses,
the tribes Platyprosopini, Arrowinini and Diochini were
basal lineages within Staphylininae, and Staphylinini was
Fig. 3 Summary of major congruence and incongruence between molecular and morphology-based phylogeny of Staphylinini and allied
groups. Habitus photos of Staphylininae as examples of various groups: Actinus macleayi Olliff, 1887 as Staphylinini propria (photo by A.
Solodovnikov); Atrecus affinis (Paykull, 1789) as Othiini, Xantholinus tricolor (F., 1787) as Xantholinini (photos by Kiril Makarov);
Atanygnathus terminalis (Er., 1839) as Tanygnathinina (photo by Harald Schillhammer) and Holisus humilis Er., 1839 as Hyptiomina
(photo by Margaret Thayer). Branches coloured consistently with the informal monophyletic groups recovered in the morphology-based
phylogeny (Solodovnikov & Schomann 2009).
Molecular phylogeny of Staphylinini dS. Chatzimanolis et al.
444 Zoologica Scripta, 39, 5, September 2010, pp 436–449 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters
paraphyletic with respect to the tribes Othiini and Xantholi-
nini (which were sister-groups in both analyses). According
to both analyses, the conventional subtribe Quediina was
not monophyletic and was divided at least into ‘Quediina
propria’ (not entirely monophyletic) and ‘southern Quediin-
a’ (without Atanygnathus). In both analyses, taxa constituting
these Quediina lineages (in terms of the current system of
Staphylinini) were rather basal as far as the entire tribe
Staphylinini is concerned. Both analyses recovered the clade
Tanygnathinina + Hyptiomina that was an unexpected
grouping from the standpoint of the ‘test phylogeny’. In
both analyses the genus Algon had an isolated position
within ‘Staphylinini propria’ not matching any of the con-
ventionally recognized subtribes. The main incongruence
between the two types of inference, concerned the resolu-
tion of the ‘Quediina propria’ (especially position of the
Anchocerus +Acylophorus clade); conflicting sister relation-
ships of the Tanygnathinina + Hyptiomina clade; and differ-
ences in the positions of Bolitogyrus and Algon.
Overall, the Bayesian results were more congruent with
the ‘test phylogeny’ than were the parsimony results.
Molecular versus morphological results: congruence (Fig. 3)
Observing some degree of congruence among the morphol-
ogy-based ‘test phylogeny’ and the molecular results were
satisfying. Tribes Platyprosopini and Arrowinini were con-
sistently shown as rather basal lineages of Staphylininae.
Conventional Quediina was shown to be a polyphyletic
assemblage consisting of at least two major species-rich
lineages: ‘southern Quediina’ (‘Tanygnathinina sensu novo’
in Solodovnikov & Schomann 2009) and ‘Quediina
propria’. These two branches were relatively basal within
Staphylinini. However, morphological and molecular data
did not agree entirely on their composition or sister rela-
tionships. The extremely specialized Amblyopinina, accord-
ing to both morphological and molecular data, was a
derived member of ‘southern Quediina.’ ‘Staphylinini pro-
pria’ was a well-supported large monophyletic group within
Staphylinini sensu lato and consists of monophyletic stocks
largely equivalent to the currently recognized subtribes
Staphylinina, Anisolinina, Xanthopygina and Philonthina.
Among those, Xanthopygina and Philonthina were both
monophyletic and sister-groups. The subtribe Staphylinina
was also largely monophyletic, but its composition and rela-
tionships with respect to the current subtribe Anisolinina
were still unclear.
Molecular versus morphological results: incongruence
(Fig. 3)
The most striking disagreement between morphological
and molecular data concerns the position of the ‘Xantholi-
nini-lineage’ taxa: tribes Diochini, Othiini and Xantholi-
nini. In contrary to the ‘test phylogeny’, the Bayesian and
parsimony analyses placed Othiini and Xantholinini inside
Staphylinini, as a subclade sister to ‘southern Quediina’
(the latter being polyphyletic in the parsimony analysis).
This was an unexpected result from the morphological
analysis standpoint, as Othiini and Xantholinini, based on
both adult and larval morphology (Assing 2000; Solo-
dovnikov & Newton 2005), are well-supported members
of the ‘Xantholinini-lineage’ and thus distinct from Staph-
ylinini. It is noteworthy that these tribes were placed
inside the Staphylinini in every single-gene analyses except
that of 28S. However, beyond those tribes always being
placed inside Staphylinini, there was no consistency among
the single-gene analyses regarding their exact placement.
Both Bayesian and parsimony analyses of 28S placed those
tribes within the subfamily Staphylininae, consistently with
the morphology (see online supplement figures).
The next noticeable disagreement between molecular
and morphology-based inferences was the sister-group
relationship of Atanygnathus (Tanygnathinina) and Holisus
(Hyptiomina). Morphology placed Atanygnathus as a
derived member of ‘southern Quediina’ (actually, as the
type of a family-group name, it is Atanygnathus that gives
this group the name ‘Tanygnathinina sensu novo’); and
suggested that Holisus was nothing more than a specialized
flat subcortical member of Philonthina (implying also a
formal synonymy of the subtribes). Molecular data, on the
contrary, consistently placed Atanygnathus outside ‘south-
ern Quediina’ and Holisus outside Philonthina, and united
them in one clade. Morphologically, Atanygnathus and Hol-
isus are strikingly different and from that line of evidence
their sister relationship seems problematic. However, we
must also mention here the results from the morphology-
based study in Solodovnikov & Newton (2005), where
Atanygnathus was grouped with Holisus. That was consid-
ered an artefact of the small taxon sampling (Solodovnikov
2006), but must be explored further. It is thus interesting
that a sister-group relationship between Atanygnathus and
Holisus was consistently supported by most single-gene
analyses (see online supplement figures). Only the parsi-
mony analysis of COI and Bayesian analysis of TP did not
support the clade Holisus +Atanygnathus. Sister relation-
ships of a group Holisus +Atanygnathus shown by various
single-gene analyses vary dramatically. It should be noted
here that our sample of Holisus was limited by larval mate-
rial only (no DNA-quality material of adults are so far
available for this genus). Although misidentification seems
unlikely (morphologically Holisus larvae are quite charac-
teristic, and our samples were identified by association
with respective adults collected together under bark), sam-
ples of the adult specimens of Holisus must be sequenced
before any further conclusions are made.
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 445
A final significant incongruence between morphology-
based and molecular results concerns ‘Quediina propria’.
Neither of the combined molecular analyses recovered
them in the same clade as the ‘test phylogeny’ did. Dis-
agreements here mostly focused on the so-called ‘Acylopho-
rus-lineage’. Morphology-based analyses defines that
lineage consists of at least of the genera Acylophorus, Anch-
ocerus, Hemiquedius and Anaquedius, but does not include
Indoquedius. Molecular data never recovered the ‘Acylopho-
rus-lineage’ with such a composition, and never placed it
as a member of Quediina propria. Additional smaller scale
disagreements concerned sister relationships of genera like
Algon and Bolitogyrus, that are also problematic from the
standpoint of morphology.
Conclusions
Molecular phylogenetics, consistent with the morphology-
based phylogenetic inference, proposes a new evolutionary
pattern for Staphylinini that is more complex and largely
different from the ‘pattern’ implied by the traditional, cur-
rently used conventional systematics of the tribe. Molecu-
lar data are largely congruent with the morphology-based
results in suggesting that the tribes Arrowinini and Platy-
prosopini are rather plesiomorphy-rich basal lineages of
Staphylininae stemming from a hypothetical ancestor that
likely also gave rise to the tribe Staphylinini. The morphol-
ogy-based conclusion that the current subtribe Quediina is
the most problematic, highly polyphyletic assemblage of
species is fully supported by molecular data: current Quedi-
ina mixes at least two species-rich, relatively old basal lin-
eages: ‘Quediina propria’ and ‘southern Quediina’
(‘Tanygnathinina sensu novo’). Molecular data also support
the morphology-based hypothesis that Amblyopinina, con-
trary to the current formal system, is not an isolated lineage
within Staphylinini, but rather an internal clade within
‘southern Quediina’. Like morphology, molecular data do
not provide a definite answer about the composition of the
‘Quediina propria’, although the core clade of this lineage is
consistently recovered by both datasets as a relatively basal
group of Staphylinini. Both molecular and morphological
data consistently and strongly support monophyly of a rela-
tively nested clade ‘Staphylinini propria’, which unites the
species-rich subtribes Staphylinina, Anisolinina, Xanthopy-
gina and Philonthina. Based on the more terminal position
of this clade, it is likely to be a younger group than either
‘southern Quediina’ or ‘Quediina propria’. Molecular data
are congruent with morphology in revealing roughly the
same internal phylogenetic topology for the ‘Staphylinini
propria’, in particular by supporting monophyly of the tra-
ditionally recognized subtribes Staphylinina, Xanthopygina
and Philonthina. Molecular data unite the latter two sub-
tribes as sister-groups. With respect to the subtribe Aniso-
linina, another internal member of Staphylinini propria,
both morphological and molecular results are insufficient
for making a sound conclusion about monophyly and sister-
group relationships of this group. Molecular data suggest
that the tribes Othiini and Xantholinini are not sister to
Staphylinini, but rather basal members of the latter, thus
making Staphylinini in the current sense a paraphyletic
group. This is the most striking and thought-provoking
conflict with the morphological hypothesis and current sys-
tematics. Considerable disagreements between the single-
gene datasets about sister relationships of Othiini and Xan-
tholinini, as well as relatively short branch lengths of several
basal nodes point to a possible deficiency of our dataset for
deeper level divergences. Our results, however, call for dee-
per studies of the phylogenetic relationships among groups
within the entire subfamily Staphylininae before proposing
any changes in the classification at the tribal level. Molecu-
lar results strongly conflict with morphology with respect to
the phylogenetic placement of the peculiar and morphologi-
cally strikingly different genera
Holisus
(member of mono-
generic subtribe Hyptiomina according to the formal
system, and derived member of the subtribe Philonthina
according to morphology-based phylogenetic analysis;
available only as larvae for this molecular analysis) and
Ata-
nygnathus
(member of monogeneric subtribe Tanygnathin-
ina according the formal system, and derived member of the
‘southern Quediina’ according to morphology-based phylo-
genetic analysis). In the molecular analyses, these two gen-
era are sister-groups within a relatively basal clade of
Staphylinini. The conflict between morphological and
molecular data is so severe that any systematic decisions
concerning these lineages are postponed until the case is
investigated with more data, especially until adult speci-
mens of
Holisus
will be found for molecular analysis. The
overall incongruence among various phylogenetic results
for Staphylinini (morphology
versus
molecules, combined
molecular dataset
versus
single-gene analyses, Bayesian
inference
versus
parsimony) suggests that the quantity and
quality of data used for phylogeny reconstruction of this
mega-diverse group is still insufficient for getting consis-
tent results that would be a sound basis for large-scale
classification changes. Given that additional morphologi-
cal data are generally becoming more difficult to obtain,
additional genes, especially slower evolving ones to target
deeper divergences within Staphylinini and among Staph-
ylinini and allied taxa, are needed in future analyses.
Taxonomic implications: new composition of the
subtribes Quediina and Amblyopinina
In accordance with the most congruent results of this
molecular study and previously published morphology-
based phylogenies of Staphylinini (see above), some
Molecular phylogeny of Staphylinini dS. Chatzimanolis et al.
446 Zoologica Scripta, 39, 5, September 2010, pp 436–449 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters
changes in the classification of Staphylinini are here intro-
duced. The formal subtribe Quediina is reduced to the
monophyletic clade ‘Quediina propria’, whereas simulta-
neously the formal subtribe Amblyopinina is expanded to
accommodate the entire monophyletic clade ‘southern
Quediina’. Most of the formerly Quediine genera not
belonging to Quediina in the new restricted sense are to be
moved to ‘southern Quediina’. But for some former quedi-
ine genera not belonging either to ‘southern Quediina’, or
to ‘Staphylinini propria’, proper formal systematic catego-
ries within Staphylinini are lacking. It is likely that new
supra-generic taxa of subtribal level must be erected in the
future at least for those ‘Quediine’ genera with basal posi-
tion in the newly emerging phylogeny of Staphylinini.
Regarding the rank and formal name for the newly defined
clade ‘southern Quediina’, a subtribe Amblyopinina seems
as the best solution at the moment. Our molecular analysis
was congruent with earlier morphology-based studies in
placing highly derived members of Amblyopinina within
the ‘southern Quediina’. The older family-group name Ta-
nygnathinina would be available for the clade ‘southern
Quediina’ only according to the morphological studies that
also placed the genus
Atanygnathus
inside that clade. How-
ever, and contrary to morphology, molecular data neither
supports the position of
Atanygnathus
inside the ‘southern
Quediina’, nor provides a sound alternative for its position
so far. Therefore, the concept of the subtribe Tanygnathin-
ina is left intact pending future phylogenetic studies. The
proposed taxonomic changes are summarized in Table 1.
Naturally, they involve all genera thoroughly studied mor-
phologically, not only those included in the limited dataset
for the molecular analysis here.
Acknowledgements
We thank K.-J. Ahn, M. S. Caterino, A. F. Newton and
M. K. Thayer for providing specimens for molecular work.
Also, a significant portion of the material from Southeast
Asia was collected by A. Solodovnikov and J. Pedersen
during a recent trip to Laos together with S. Tarasov,
A. F. Newton and M. Thayer; great companionship and
synergism in the fieldwork by these colleagues is appreci-
ated. Natalia Solodovnikova is acknowledged for some
initial help to A. Schomann in the laboratory. Thanks also
to Ole Seberg, Gitte Pedersen and Charlotte Hansen for
providing molecular laboratory facility to A. Solodovnikov
and A. Schomann. A. F. Newton, M. K. Thayer, M.S.
Caterino and one anonymous reviewer are acknowledged
for useful comments which led to the improvement of this
manuscript. We are thankful to Harald Schillhammer,
Kiril Makarov and Gracen Brilmyer for providing some
habitus photos of Staphylininae. Some aspects of this work
were carried out using the resources of the Computational
Table 1 Proposed taxonomic changes in the systematics of the tribe Staphylinini. Below is the new composition of the subtribes Quediina
and Amblyopinina, and several genera formerly in Quediina are now placed as incertae sedis in Staphylinini, pending further analyses. The
composition of the other subtribes remains unchanged
Amblyopinina Seevers, 1944 Quediina Kraatz, 1857
Incertae sedis
Amblyopinodes
Seevers, 1955
Acylohsellus
Smetana, 1995
Afroquedius
Solodovnikov 2006
Amblyopinus
Solsky, 1875
Acylophorus
Nordmann, 1837
Astrapaeus
Gravenhorst, 1802
Cafioquedus
Sharp, 1886
Anaquedius
Casey, 1915
Beeria
Hatch, 1957
Cheilocolpus
Solier, 1849
Anchocerus
Fauvel, 1905
Bolitogyrus
Chevrolat, 1842
Chilamblyopinus
Ashe & Timm, 1988
Australotarsius
Solodovnikov & Newton, 2009
Heinzia
Korge, 1971
Ctenandropus
Cameron, 1926
Euryporus
Erichson, 1839
Lonia
Strand, 1943
Edrabius
Fauvel, 1900
Hemiquedius
Casey, 1915
Parisanopus
Brethes, 1900
Heterothops
Stephens, 1829
Indoquedius
Blackwelder, 1952
Quediomacrus
Sharp, 1884
Loncovilius
Germain, 1903
*Paratolmerus
Cameron, 1932
Quelaestrygon
Smetana, 1999
Megamblyopinus
Seevers, 1955
*Pseudorientis
Watanabe, 1970
Sedolinus
Solodovnikov 2006
Mimosticus
Sharp, 1884 **
Quedius
Stephens, 1829
Strouhalium
Scheerpeltz, 1962
Myotyphlus
Fauvel, 1883
Quetarsius
Smetana, 1996
Termitoquedius
Bernhauer, 1912
Natalignathus
Solodovnikov 2005
Valdiviodes
Smetana, 1981
Philonthellus
Bernhauer, 1939
Quediocafus
Cameron, 1945
Quediomimus
Cameron, 1948
Quediopsis
Fauvel, 1878
Rolla
Blackwelder, 1952
Sphingoquedius
Bernhauer, 1941
*Genera
Paratolmerus
and
Pseudorientis
are pending further in-depth morphological and molecular study to confirm their tentative position within Quediina.
**Although not formally reclassified yet, native species of
Quedius
from the south temperate areas of the globe belong to several genera (including undescribed ones) of the
subtribe Amblyopinina in the new sense.
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 447
Biology Service Unit from Cornell University that is par-
tially funded by Microsoft Corporation. Direct financial
support for the project was provided by NSF DEB-
0741475 to S. Chatzimanolis and M. S. Engel, by NSF
DEB-0715705 to A. Newton and A. Solodovnikov, and by
the Entomology Department Frame Grant from the
National Danish Research Council.
References
Assing, V. (2000). A taxonomic and phylogenetic revision of
Maorothiini trib. n. from the New Zealand subregion
(Coleoptera: Staphylinidae, Staphylininae). Beitra
¨ge zur
Entomologie,50, 3–64.
Ballard, J. W. O., Thayer, M. K., Newton, A. F. & Grismer, E.
R. (1998). Data sets, partitions, and characters: philosophies
and procedures for analyzing multiple data sets. Systematic
Biology,47, 367–396.
Caterino, M. S., Cho, S. & Sperling, F. A. H. (2000). The
current state of insect molecular systematics: a thriving Tower
of Babel. Annual Review of Entomology,45, 1–54.
Caterino, M. S., Hunt, T. & Vogler, A. P. (2005). On the
constitution and phylogeny of Staphyliniformia (Insecta:
Coleoptera). Molecular Phylogenetics and Evolution,34, 655–672.
Chatzimanolis, S. (2008). A revision of the neotropical beetle
genus Isanopus (Coleoptera: Staphylinidae: Staphylinini). Journal
of Natural History,42, 1765–1793.
Chatzimanolis, S. & Ashe, J. S. (2009). A revision of the
neotropical genus Ocyolinus (Coleoptera: Staphylinidae:
Staphylinini). Zootaxa,2162, 1–23.
Hayashi, Y. (2005). Studies on the Asian Staphylininae
(Coleoptera, Staphylinidae) VI. A revision of the genus
Philetaerius, with description of a new species and some notes
on the Eucibdelina. Elytra,33, 547–560.
Herman, L. H. (2001). Catalog of the Staphylinidae (Insecta:
Coleoptera). 1758 to the end of the second millennium. Parts
I-VII. Bulletin of the American Museum of Natural History,265,
1–4218. (in 7 vols.).
Huelsenbeck, J. P. & Ronquist, F. (2001). MRBAYES: Bayesian
inference of phylogenetic trees. Bioinformatics,17, 754–755.
Hunt, T., Bergsten, J., Levkanicova, Z., Papadopoulou, A., John,
O. S., Wild, R., Hammond, P. M., Ahrens, D., Balke, M.,
Caterino, M. S., Go
´mez-Zurita, J., Ribera, I., Barraclough, T.
G., Bocakova, M., Bocak, L. & Vogler, A. P. (2007). A
comprehensive phylogeny of beetles reveals the evolutionary
origins of a superradiation. Science,318, 1913–1916.
Korte, A., Ribera, I., Beutel, R. G. & Bernhard, D. (2004).
Interrelationships of Staphyliniform groups inferred from 18S
and 28S rDNA sequences, with special emphasis on
Hydrophiloidea (Coleoptera, Staphyliniformia). Journal of
Zoological Systematics And Evolutionary Research,42, 281–288.
Maddison, W. P. & Maddison, D. R. (2009). Mesquite: A Modular
System for Evolutionary Analysis. Version 2.71 Available at:
http://mesquiteproject.org [Accessed on 24 October, 2009].
Maddison, D. R., Moore, W., Baker, M. D., Ellis, T. M., Ober,
K. A., Cannone, J. J. & Gutell, R. R. (2009). Monophyly of
terrestrial adephagan beetles as indicated by three nuclear genes
(Coleoptera: Carabidae and Trachypachidae). Zoological Scripta,
38, 43–62.
Maus, C., Peschke, K. & Dobler, S. (2001). Phylogeny of the
genus Aleochara inferred from mitochondrial cytochrome
oxidase sequences (Coleoptera: Staphylinidae). Molecular
Phylogenetics and Evolution,18, 202–216.
Miller, K. B., Bergsten, J. & Whiting, M. F. (2009). Phylogeny
and classification of the tribe Hydaticini (Coleoptera:
Dytiscidae): partition choice for Bayesian analysis with multiple
nuclear and mitochondrial protein-coding genes. Zoologica
Scripta,38, 591–615.
Newton, A. F. & Thayer, M. K. (2005, onwards). Catalog of
Higher Taxa of Staphyliniformia and Genera and Subgenera of
Staphylinoidea. Chicago: Field Museum of Natural History.
Available at: http://www.fieldmuseum.org/peet_staph/db_1a.
html [Accessed on 16 January, 2010].
Nylander, J. A. A. (2004). MrModeltest v2. Program distributed by
the author. Uppsala, Sweden: Evolutionary Biology Centre,
Uppsala University.
Posada, D. (2008). jModelTest: phylogenetic Model Averaging.
Molecular Biology and Evolution,25, 1253–1256.
Pruesse, E., Quast, C., Knittel, K., Fuchs, B., Ludwig, W.,
Peplies, J. & Glo
¨ckner, F. O. (2007). SILVA: a comprehensive
online resource for quality checked and aligned ribosomal RNA
sequence data compatible with ARB. Nucleic Acids Research,35,
7188–7196.
Rambaut, A. (2002). Se-Al: Sequence Alignment Editor. Version
2.0 Available at: http://http://tree.bio.ed.ac.uk/software/seal/
[Accessed on 11 July, 2009].
Solodovnikov, A. Y. (2005). Natalignathus, gen. nov. and larvae
of Atanygnathus: a missing phylogenetic link between subtribes
Quediina and Tanygnathinina (Coleoptera: Staphylinidae:
Staphylininae: Staphylinini). Invertebrate Systematics,19,
75–98.
Solodovnikov, A. Y. (2006). Revision and phylogenetic assessment
of Afroquedius gen. nov. from South Africa: toward new concepts
of the genus Quedius, subtribe Quediina and reclassification of
the tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae).
Annals of the Entomological Society of America,99, 1064–1084.
Solodovnikov, A. Y. (2009). Xanthopygoides niger Cameron, 1951
(Xanthopygina) belongs to the genus Philonthus Stephens, 1829
(Philonthina): systematic and nomenclatural changes for the
African Staphylinini (Coleoptera: Staphylinidae: Staphylininae:
Staphylinini). Zookeys,5, 7–12.
Solodovnikov, A. Y. & Newton, A. F. (2005). Phylogenetic
placement of Arrowinini trib. n. within the subfamily
Staphylininae (Coleoptera: Staphylinidae), with revision of the
relict South African genus Arrowinus and description of its
larva. Systematic Entomology,30, 398–441.
Solodovnikov, A. & Schomann, A. (2009). Revised systematics
and biogeography of ‘Quediina’ of Subsaharan Africa: new
phylogenetic insights into the rove beetle tribe Staphylinini
(Coleoptera: Staphylinidae). Systematic Entomology,34, 443–446.
Swofford, D. L. (2002). PAUP*. Phylogenetic Analysis Using
Parsimony (* and Other Methods), Version. 4.0b10. Sunderland:
Sinauer Associates.
Thomas, J. C. (2009). A preliminary molecular investigation of
aleocharine phylogeny (Coleoptera: Staphylinidae). Annals of the
Entomological Society of America,102, 189–195.
Wheeler, T. J. & Kececioglu, J. D. (2007). Multiple alignments
by aligning alignments. Bioinformatics,23, i559–i568.
Molecular phylogeny of Staphylinini dS. Chatzimanolis et al.
448 Zoologica Scripta, 39, 5, September 2010, pp 436–449 dª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters
Wild, A. L. & Maddison, D. R. (2008). Evaluating nuclear
protein-coding genes for phylogenetic utility in beetles.
Molecular Phylogenetics and Evolution,48, 877–891.
Wilgenbusch, J. C., Warren, D. L. & Swofford, D. L. (2004).
AWTY: A System for Graphical Exploration of MCMC Convergence
in Bayesian Phylogenetic Inference. Available at: http://ceb.csit.fsu.
edu/awty [Accessed on 11 July, 2009].
Supporting Information
Additional Supporting Information may be found in the
online version of this article:
Table S1. Taxon sampling and GenBank numbers for
the species used in this study. All GenBank accessions
(except the COI accession for
Hadropinus fossor
) are novel.
Vouchers deposited include both DNA and pinned speci-
mens.
Table S2. Primers and genes used in this study.
Fig. S1 Bayesian phylogenetic tree for the analysis of
the COI gene. Only posterior probabilities above 0.75 are
shown.
Fig. S2 Fifty percent majority rule consensus tree of
three equally parsimonious trees for the analysis of the
COI gene. Tree length = 3759 steps. Only bootstrap
values above 75 are shown.
Fig. S3 Bayesian phylogenetic tree for the analysis of
the wingless gene. Only posterior probabilities above 0.75
are shown.
Fig. S4 Fifty percent majority rule consensus tree of
682 equally parsimonious trees for the analysis of the
wingless gene. Tree length = 1260 steps. Only bootstrap
values above 75 are shown.
Fig. S5 Bayesian phylogenetic tree for the analysis of
the topoisomerase I gene. Only posterior probabilities
above 0.75 are shown.
Fig. S6 Fifty percent majority rule consensus tree of 14
equally parsimonious trees for the analysis of the topoi-
somerase I gene. Tree length = 3814 steps. Only bootstrap
values above 75 are shown.
Fig. S7 Bayesian phylogenetic tree for the analysis of
the 28S rDNA gene. Only posterior probabilities above
0.75 are shown.
Fig. S8 Fifty percent majority rule consensus tree of 63
equally parsimonious trees for the analysis of 28S rDNA
gene. Tree length = 3288 steps. Only bootstrap values
above 75 are shown.
Please note: Wiley-Blackwell is not responsible for the
content or functionality of any supporting materials sup-
plied by the authors. Any queries (other than missing
material) should be directed to the corresponding author
for the article.
S. Chatzimanolis et al. dMolecular phylogeny of Staphylinini
ª2010 The Authors. Journal compilation ª2010 The Norwegian Academy of Science and Letters dZoologica Scripta, 39, 5, September 2010, pp 436–449 449
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... Regarding phylogenetic affinities, morphologically based studies (Kypke, Solodovnikov, Brunke, Yamamoto, & Żyła, 2019;Solodovnikov & Newton, 2005;Solodovnikov, Yue, Tarasov, & Ren, 2013) have placed Diochini within a lineage formed by Xantholinini, Maorothiini, and Othiini, with Diochini as a sister lineage to remaining taxa. This hypothesis is in strong disagreement with findings from molecular studies that included representatives of the tribe (Brunke, Chatzimanolis, Schillhammer, & Solodovnikov, 2016;Chatzimanolis, Cohen, Schomann, & Solodovnikov, 2010), where Diochini was recovered as an early derived lineage within the subfamily Staphylininae. On the contrary, more recent molecular phylogenetic studies, based on larger multilocus datasets, are fully consistent with the relationships derived from morphological data (Tihelka, Thayer, Newton, & Cai, 2020;Żyła & Solodovnikov, 2020) and led to the recent proposal of the subfamily rank for the lineage including Xantholinini, Maorothiini, Othiini, and Diochini (subfamily Xantholininae sensu Żyła & Solodovnikov, 2020; see also Tihelka et al., 2020 for arguments against the new classification proposed by Żyła & Solodovnikov, 2020). ...
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... Initially, this distinctive genus was assigned to the subtribe Quediina mostly due to its overall similarity to the other members of the subtribe. Later, a series of phyloge-netic studies of Staphylinini were successively conducted by various authors (e.g., Solodovnikov 2006;Solodovnikov and Schomann 2009;Chatzimanolis et al. 2010;Brunke and Solodovnikov 2013;Brunke et al. 2016;Żyła and Solodovnikov 2019;Tihelka et al. 2020). As a result, the subtribe Quediina was confirmed to be highly polyphyletic and redefined as a more restricted group, while many genera were moved out from Quediina sensu Brunke et al. (2016). ...
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A male of the very rare and phylogenetically puzzling species, Quelaestrygon puetzi Smetana, 1999, is described for the first time based on a single specimen from Sichuan Province, China. High quality color images and line drawings of the male external and genitalic traits are provided.
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Despite their familiarity to most naturalists, the large rove beetles of the subtribe Staphylinina are poorly understood systematically and have never been the subject of a focused phylogenetic analysis. Within a total evidence analytical framework, an unusual Staphylinina from China was demonstrated to be the sister group of Ontholestes and was described as Lesonthotes vulneratus Smetana and Brunke, gen. et sp. nov. Based on these results, Ontholestes was morphologically redefined and the Cingulatus group of species was established. The Neotropical genus Leistotrophus was resolved as a member of Ontholestes, within the latter clade but synonymy was avoided until molecular data are available for the former. Ontholestes lowi Rougemont, 2016 syn. nov. is proposed as a junior synonym of Ontholestes borneensis Cameron, 1942. We also confirmed the existence of and morphologically defined three major lineages within Staphylinina, of which two were previously based on molecular data only. This study builds on previous work by Smetana and Davies (2000 Smetana, A., & Davies, A. (2000). Reclassification of the North Temperate Taxa Associated with Staphylinus Sensu Lato, including comments on relevant subtribes of Staphylinini (Coleoptera: Staphylinidae). American Museum Novitates, 3287, 1–88. doi:10.1206/0003-0082(2000)287<0001:ROTNTT>2.0.CO;2[Crossref] , [Google Scholar]) and lays the framework for more extensive analysis of the subtribe. Our results emphasize the continued value of morphological characters in a total evidence framework for resolving the systematics of charismatic but poorly explored groups. www.zoobank.org/urn:lsid:zoobank.org:pub:776F033D-4556-483C-8725-707E1B02C040
Thesis
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The research contained in this thesis explores the phylogenetic systematics of a hitherto very poorly known group of predatory rove beetles, the subtribe Amblyopinina (Coleoptera: Staphylinidae). This subtribe comprises one of the predominant rove beetle groups in the south temperate continents and islands of the world where they are abundant in forest leaf litter and other terrestrial habitats. Of them, only two genera also occur throughout the northern hemisphere and on numerous, remote oceanic islands. Phylogenetically nested within the subtribe are a group of genera that have a unique, mutualistic relationship with mammals in the Neotropics and Australia, where they prey on ectoparasites in the fur and nests of their hosts. The highly disjunct distribution, occurrence on numerous large and small, more and less isolated landmasses, mutualistic relationship with mammals in some and prevalence as a poorly known component of austral biodiversity made Amblyopinina an important target for phylogenetic, systematic and biogeographic investigations, some of which are presented here. To build a framework for future research on amblyopinines in all directions, from the badly needed taxonomic exploration to biogeographic research, the first molecular phylogeny of this group is provided. It gives an overall insight into the internal relationships of Amblyopinina, reveals interesting biogeographic patterns and suggests multiple origins of mammal association within the subtribe. In connection with the revision of the Australian mammal-associated genus Myotyphlus a review of the amblyopininemammal symbiosis is made. With molecular data for rove beetles still in its infancy but growing all the time, morphology-based phylogenetic analyses were used to demonstrate that a peculiar new genus and species of Staphylinini rove beetle, Devilleferus brunkei from the high Andes is a sister group to all other Amblyopinina. Apart from this new taxon and two new species of Myotyphlus, the alpha-taxonomic component of the thesis is focused on one of the remote oceanic islands inhabited by amblyopinines. A systematic review of the entire Staphylinini rove beetle fauna of Lord Howe Island is made which included the description of two new species for Amblyopinina and implementation of several other taxonomic changes. The review highlighted the strong affinity of the Lord Howe Island fauna with the Australian mainland and the biogeographic potential of amblyopines.
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The Staphylinini genus Ocyolinus Sharp, 1884 is revised. Three species are described as new, O. astenos Chatzimanolis & Ashe sp. n., O. dimoui Chatzimanolis & Ashe sp. n. and O. nebulosus Chatzimanolis & Ashe sp. n., all from Panama. A lectotype is designated for O. rugatus Sharp, 1884 and O. vulneratus Bernhauer, 1906. Ocyolinus vulneratus Bernhauer is placed in synonymy with O. rugatus Sharp. Distribution maps, an identification key and illustrations of structural features are provided.
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Discovery of adults and a larva of Natalignathus, a new South African endemic genus, and examination of previously undescribed larvae of Atanygnathus Jakobson shed light on the phylogenetic relationships of the puzzling rove beetle subtribe Tanygnathinina (Staphylinidae:Staphylininae:Staphylinini). Based on plesiomorphic states for many characters of Natalignathus, gen. nov., inferred on the basis of comparison of the new genus with Atanygnathus, various Quediina and other Staphylinini, Natalignathus can be considered either a relatively plesiomorphy-rich member of Tanygnathinina, or a derived genus of Quediina, essentially linking these two subtribes together. Based on morphological and distributional data, it is preliminarily assumed that Atanygnathus evolved from some lineage of southern Quediina, which currently are very poorly known. Comparative study of the adults of Natalignathus and Atanygnathus allows correction of earlier misinterpretations of the structure of the aedeagus and head capsule ridges of Atanygnathus. Detailed morphological descriptions are provided for adults and a larva of Natalignathus and for the larvae of two species of Atanygnathus, A. bicolor (Casey) and A. sp. 1. Special appendages of unknown function are present on the mesothoracic spiracles of the examined larvae of Atanygnathus. No similar structures are known in any other group of Staphylinidae or Coleoptera. Based on the new findings, a review of the state of knowledge of the subtribes Quediina and Tanygnathinina and an updated diagnosis of the latter are provided.
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We compared four approaches for analyzing three data sets derived from staphylinoid beetles, a superfamily whose known species diversity is roughly comparable to that of vertebrates. One data set is derived from adult morphology and the two molecular data sets are from 12Sribosomal RNA and cytochrome b mitochondrial DNA. We found that taxonomic congruence following conditional data combination, herein called compatible evidence (CE), resolved more nodes compatible with an initial conservative hypothesis than did total evidence (TE), conditional data combination (CDC), or taxonomic congruence (TC). CE sets a base of nodes obtained by CDC analysis and then investigates what further agreement may arise in a universe where these nodes are accepted as given. We suggest that CE75–75 may be appropriate for future studies that aim to both generate a well-corroborated tree and investigate conflicts between data sets, partitions, and characters. CE75–75 is a 75% bootstrap consensus CDC tree followed by combinable-component consensus of a 75% bootstrap consensus of each homogeneous set of partitions having hierarchical structure.
Book
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
Article
The Staphylinini genus Ocyolinus Sharp, 1884 is revised. Three species are described as new, O. astenos Chatzimanolis & Ashe sp. n., O. dimoui Chatzimanolis & Ashe sp. n. and O. nebulosus Chatzimanolis & Ashe sp. n., all from Panama. A lectotype is designated for O. rugatus Sharp, 1884 and O. vulneratus Bernhauer, 1906. Ocyolinus vulneratus Bernhauer is placed in synonymy with O. rugatus Sharp. Distribution maps, an identification key and illustrations of structural features are provided.