Revised systematics and biogeography of ‘Quediina’ of sub-Saharan Africa: new phylogenetic insights into the rove beetle tribe Staphylinini (Coleoptera: Staphylinidae)

Systematic Entomology (2009), 34, 443–466 DOI: 10.1111/j.1365-3113.2008.00468.x

Revised systematics and biogeography of ‘Quediina’ ofsub-Saharan Africa: new phylogenetic insights into therove beetle tribe Staphylinini (Coleoptera: Staphylinidae)

ALEXEY SOLODOVN IKOV and ANDREA SCHOMANNDepartment of Entomology, Natural History Museum of Denmark/University of Copenhagen, Denmark

Abstract. Quediina, a mega-diverse conventional subtribe of the rove beetle tribeStaphylinini, is remarkably species rich in the north and south temperate regions ofthe world. Tropical faunas of this group, and the fauna of the entire Afrotropicalbiogeographical region (¼ Ethiopian region,¼ sub-Saharan Africa), in contrast, areremarkably poor. The taxonomic study of the quediine genera of Staphylinini fromthe Afrotropical region reveals misidentifications for many of them. Their phylo-genetic study demonstrates polyphyly of Quediina and reveals a new evolutionarypattern for the entire tribe Staphylinini. In particular, the formerly quediine generaEuristus Fauvel, 1899, Ioma Blackwelder, 1952, Natalignathus Solodovnikov, 2005,all endemic in the Afrotropical region, belong to the non-related ‘Staphylinina’,‘Philonthina propria’ and ‘Tanygnathinina sensu novo’ lineages of Staphylinini,respectively. Contrary to earlier records, the genus Quedius Stephens, 1929 does notoccur in Africa south of Sahara: Quedius angularis Cameron, 1948 and Quediuscinctipennis Cameron, 1951 are moved to the genus Philonthus Stephens, 1829. Thesame is established for the Asian genus Algon Sharp, 1874, formerly for a long timeassociated with Quediina: African species Algon robustus Wendeler, 1928 is movedto the genusMoeocerus Fauvel, 1899 (here in the ‘Philonthina propria’ lineage); andthe misidentification of Algon africanus Bernhauer, 1915, a species that probablybelongs to a new genus, is discussed. The phylogenetic affiliation of AfroquediusSolodovnikov, 2006, a South African endemic, is still ambiguous. Overall, theformerly seen bipolar distribution pattern for the ‘Quediina’ is demonstrated to bean artefact, not a reality to explain. Historical biogeographical explanations areproposed for some of the Afrotropical endemics, partly as an attempt to applybiogeography as an external criterion for the evaluation of the new phylogeneticpattern revealed for Staphylinini. The monotypic genera Euristus and Ioma, as wellas Heterothops megalops Cameron, 1959, the only representative of this widespreadgenus in the Afrotropical region, are redescribed. Limits and synapomorphies of thegenus Heterothops are discussed. The following new combinations and new namesare proposed: Philonthus cinctipennis (Cameron, 1951) comb.n. (preoccupied byPhilonthus cinctipennis Fauvel, 1875), here replaced by Philonthus pseudoquediusSolodovnikov nom.n.; Philonthus angularis (Cameron, 1948) comb.n.; Moeocerusrobustus (Wendeler, 1928) comb.n. [preoccupied by Moeocerus robustus (Gestro,1881)], here replaced by Moeocerus wendeleri Solodovnikov nom.n. A lectotype isdesignated for Heterothops megalops Cameron, 1959.

Correspondence: Alexey Solodovnikov, Department of Entomo-

logy, Zoological Museum, Universitetsparken 15, 2100 Copenhagen,

Denmark. E-mail: [email protected]

# 2009 The AuthorsJournal compilation # 2009 The Royal Entomological Society 443

SystematicEntomology

Introduction

With 211 genera and more than 5300 described speciesworldwide, the tribe Staphylinini is one of the largest groupsof rove beetles (Herman, 2001; A.F. Newton, unpublished

database). The monophyly of Staphylinini is well supported bymodern studies (Solodovnikov&Newton, 2005; Solodovnikov,2006), but the internal systematics of this tribe needs revision,or rather development. The subtribe Quediina is one of the

largest subdivisions in the current conventional system ofStaphylinini, most of its diversity confined to the north andsouth temperate areas of the world. As recently shown,

however (Solodovnikov, 2006), Quediina is a polyphyleticgroup. Because of the very poor state of the alpha-taxonomicknowledge of Quediina outside the Holarctic and obvious

non-monophyly of some other subtribes of Staphylinini, it isfruitless to move genera among existing subtribes hoping toeliminate the polyphyly of Quediina. The taxonomic revisionof the extra-Holarctic faunas of Staphylinini and a new,

phylogenetic, classification for the tribe is badly needed. Thispaper is a step towards this goal, continuing and summarizingthree earlier contributions (Solodovnikov & Newton, 2005;

Solodovnikov, 2005, 2006). For practical reasons, we still use‘Quediina’ as a subtribe category here, until a new system isreally proposed and formally fixed by nomenclature. To

emphasize the polyphyletic nature of ‘Quediina’, however,especially in the phylogenetic and biogeographical discus-sions throughout the paper, we use it in quotation marks.

Similarly, we use informal names for the newly foundmonophyletic groups of Staphylinini, which are not yet fixedin formal nomenclature (e.g. ‘Quediina propria’, ‘Tanygna-thinina sensu novo’, etc.). In those cases, quotation marks

again emphasize their tentative and informal status.Here we focus on the taxonomy of ‘Quediina’ of the entire

continental Africa south of Sahara, i.e. the area constituting

the main part of the Afrotropical biogeographical region [¼Ethiopian region, ¼ sub-Saharan Africa, for limits anddefinitions see, e.g. Darlington (1957), Crosskey & White

(1977) and Smetana (2004)]. This paper does not include theislands of the Indian Ocean and the Atlantic off the Africancoasts, which formally are included into the Afrotropical

region. Also, it does not include Madagascar, which some-times is considered as a separate biogeographical entity (e.g.Darlington, 1957; Herman, 2001; Newton & Thayer, 2005),but more often is included into the Afrotropical region. To

our knowledge, there are no species of Quediina (or thosethat could be related to the here considered continental taxa)known from any of these islands. The phylogenetically

enigmatic genus Descarpentriesiellus Jarrige, 1978, endemicto Madagascar and currently attributed to none of theconventional subtribes of Staphylinini [incertae sedis in

Herman (2001) and in Newton & Thayer (2005)], seems tobe a highly derived member of the lineage to which mostcurrent Anisolinina belong. It will be dealt with in a separatepublication. Finally, some other Afrotropical genera, which

are currently placed in either Philonthina or Xanthopygina(Herman, 2001; Newton & Thayer, 2005) have some re-semblance to Quedina. These are the genera Taxiplagus

Bernhauer, 1915 (¼ Prionophilonthus Scheerpeltz, 1974; ¼Quediosoma Cameron, 1926) (Philonthina); Prianophtalmus

Bernhauer, 1932, Trigonopalpus Cameron, 1951 and Xan-thopygoides Verdcour, 1952 (all in Xanthopygina). Revisionof Xanthopygoides revealed its synonymy with the genus

Philonthus Stephens, 1829 (Philonthina) (Solodovnikov,2009). Preliminary examination of Taxiplagus, Prianophtal-mus and Trigonopalpus suggested that they are also relatedto the current Philonthina (or ‘Philonthina propria’ lineage

here). But the proper identification and phylogenetic assess-ment of the latter genera, requiring a much broaderexamination of various Philonthina, Xanthopygina and

some Anisolinina will be dealt in a separate contribution.

Materials and methods

Specimens and collections

The material examined in the present study is kept in thefollowing institutions: FMNH, Field Museum of NaturalHistory, Chicago, IL, U.S.A. (A.F. Newton, J. Boone);

IRSNB, Institut royal des Sciences naturelles de Belgique,Bruxelles, Belgium (P. Grootaert); MMUE, ManchesterMuseum, University of Manchester, Manchester, U.K.

(D. Logunov); MRAC, Musee royal de l’Afrique centrale,Tervuren, Belgium (M. De Meyer); NHM, The NaturalHistory Museum, London, U.K. (R. Booth); SANC, South

African National Collection of Insects, Pretoria, SouthAfrica (R. Stals); TMSA, Transvaal Museum, Pretoria,South Africa (J. Harrison, R. Muller); ZMHB, Museum furNaturkunde der Humboldt Universitat zu Berlin, Berlin,

Germany (M. Uhlig, J. Frisch); ZMUC, ZoologicalMuseum,University of Copenhagen, Denmark.

Microscopy and illustrations

Beetle specimens were examined using a dissecting micro-scope (LeicaMZ12.5�). They were mostly examined as

pinned dry specimens, but a few were macerated in 10%KOH, rinsed, disarticulated and examined as wet prepara-tions in glycerine. All line illustrations were made using

a camera lucida.

Measurements and their abbreviations

Measurements were made with an ocular linear micro-

meter and are given in millimetres; they are taken andabbreviated as follows: HL, head length (from apex ofclypeus to neck constriction); HW, head width (maximal,

including eyes); PL, pronotum length (along median line);PW, pronotum width (maximal); EL, elytral length (fromhumerus to most distal apical margin; best taken fromlateral view of the elytron); EW, combined width of both

elytra (maximal, elytra closed along suture). The total lengthof the body given in the descriptions was measured from thetip of the labrum to the tip of the abdomen.

444 A. Solodovnikov and A. Schomann

# 2009 The AuthorsJournal compilation # 2009 The Royal Entomological Society, Systematic Entomology, 34, 443–466

Phylogenetic analysis

A data matrix was prepared using Nexus Data Editor forWindows, version 0.5.0 (Page, 2001). The rationale for thetaxon sampling is discussed below in the ‘Phylogenetic

analysis’ section, and all essential information about thetaxa analysed is summarized in Table 1. The matrix wascomputed in TNT (Goloboff et al., 2000) with all characterstreated as unordered and equally weighted. Space for 600

000 trees was reserved in the memory. A traditional searchwas run with the collapsing rules set to maximum length¼ 0,and the following settings: 1000 replicates with 1000 trees

saved per replication. The root was set as Arrowinusphaenomenalis Bernhauer, 1935 (Arrowinini). Suboptimaltrees were found by TNT, the search continuing until the

number of trees checked reached 100 000. Bremer supportvalues were obtained from suboptimal trees up to 20 stepslonger than the shortest trees. For consensus tree and treeexplorations, only optimal trees were used.

Biogeography

Biogeographical considerations are presented here in a nar-

rative manner, they are not quantitatively tested. This ispartly because biogeography is not the primary focus of thispaper and we wanted only to present the biogeographical

problems clearly for future analysis. Also, our input data areinadequate for such a formal analysis. First, for the reasonslisted in the ‘Conclusions and discussion’ section below, thephylogenetic signal of our resulting optimal trees for Staph-

ylinini is weak. Second, we do not have robust data about theage of the tribe Staphylinini and its main subgroups. Only 30described fossil species are now attributed to the tribe

Staphylinini, the earliest being Cretoquedius oculatus Ryvkin,1988 from the Cenomanian stage of the Cretaceous (Ryvkin,1988). The fossil genera Laostaphylinus Zhang, 1988 and

Mesostaphylinus Zhang, 1988 from the late Jurassic, attrib-uted to the subfamily Staphylininae, lack tribal assignment(Zhang, 1988), but display some affinity to the tribe Staph-ylinini. In general, known paleontological data indicate that

some lineages of the tribe similar to recent ones can be tracedback to middle Cretaceous (ca. 100 Mya), and a genericdiversity close to the extant is recorded by Oligocene (ca. 30

Mya) (Herman, 2001). Therefore, in our biogeographicalestimates we vaguely assume that the tribe Staphylininipossibly goes back to the late Jurassic/early Cretaceous.

Molecular data, which could help to date deeper nodes ofthe Staphylinini phylogeny based on the fossil-inferred agesof some recent genera, are currently absent.

Phylogenetic analysis

The primary purpose of the present formal phylogenetic

analysis was to investigate the sister relationships of theAfrotropical endemic genera Afroquedius, Natalignathusand especially Ioma and Euristus. For this analysis we chose

groups broadly representing the diversity of the mainlineages of Staphylinini, with taxa sister to Staphylinini

taken as the outgroup (Table 1). To keep the analysis assimple as possible we did not include any taxa of secondaryinterest in this paper. For example, a simple morphological

examination was enough for the determination of Hetero-thops megalops or Quedius cinctipennis on a regional basis(the latter, as a result, is transferred to the genus Philonthus).But here we came very close to the problem, that the limits

of the respective genera (Heterothops and Philonthus) areblurred on the world basis. A formal cladistic analysis wouldindeed be helpful to clarify these issues on a larger scale, too,

but that would be beyond the scope of this paper, not tomention a significant amount of alpha-taxonomic work forthese and related genera to be done for this purpose. For

similar reasons, an analysis of the proper generic identity ofAlgon africanus, which needs a taxon sample much denserfor various Philonthina and Anisolinina not relevant for theprimary goals of this paper, has also been postponed.

Another purpose of the present formal cladistic analysiswas to have a reference tree for structuring the phylogeneticand biogeographical discussion relevant for the target taxa.

And finally, this analysis is an extension (with modification)of the analysis in Solodovnikov (2006), these efforts explor-ing taxa and characters relevant for the phylogeny recon-

struction of the entire tribe Staphylinini. The current datamatrix is a snapshot of our still continuing, larger-scaleexploration of the morphological characters to be used in

phylogeny reconstruction of the tribe Staphylinini. Forexample, some characters in the present study (e.g. 57)combine both neomorphic and transformational compo-nents, which is not advisable (Sereno, 2007). To avoid this

we should have used ‘contingent’ coding of such characters[terminology according to Forey & Kitching (2000)]. How-ever, such coding in this particular data marix largely

increases the amount of inapplicable character states and,according to some experiments, lowers the final resolutionof the analysis.

Characters

1. Antennae, shape of the first segment: 0. antennomere Inot elongate, at most as long as combined length ofantennomeres II and III; 1. first segment more or lesselongate, distinctly longer than antennomeres II and III.

2. Antennae, pubescence: 0. lacking on antennomere I(with only longer sparse setae), starting on antennomereII; 1. lacking on antennomeres I–III (with only sparse

longer setae), starting on antennomere IV; 2. lacking onantennomeres I–IV (with only longer sparse setae),starting on antennomere V; 3. lacking on antennomeres

I–V (with only longer sparse setae), starting on anten-nomere VI; 4. lacking on antennomeres I–VI (with onlylonger sparse setae), starting on antennomere VII; 5.dense on all antennomeres.

3. Head capsule, neck constriction: 0. distinct at sides only;1. fully developed, distinct all around; 2. neck regionindistinct.

‘Quediina’ of sub-Saharan Africa 445


Table

1.Taxonsampleforthephylogeneticanalysis.Currentsubtribes

are

given

accordingto

Herman(2001)andNew

ton&Thayer

(2005).Fortherecentlydescribed

generaNatalignathus

andAfroquedius,subtribes

are

given

from

therespectiveoriginalpublications(Solodovnikov,2005,2006).

Genus(subgenus)

Genusdistribution

Speciesin

analysis

Speciesdistribution

Curentsubtribe

Cladein

Fig.1

ArrowinusBernhauer,1935

South

Africa

Arrowinusphaenomenalis

Bernhauer,1935

South

Africa

Arrowinini(outgroup)

PlatyprosopusMannerheim,1830

World,exceptAustralia

andPacific

PlatyprosopusmexicanusSharp,

1887

Nearctic

Platyprosopini(outgroup)

AcylophorusNordmann,1837

World

Acylophorusglaberrimus(H

erbst,

1784)

Palearctic

Quediina

‘Quediinapropria’

AfroquediusSolodovnikov,2006

South

Africa

Afroquediussexpunctatus

(Bernhauer,1917)

South

Africa

Incertaesedis

‘Tanygnathininasensu

novo’

AlgonSharp,1874

East

Palearctic,

Oriental

AlgongrandicollisSharp,1874

East

Palearctic

Xanthopygina

‘Philonthinalineage’

AnchocerusFauvel,1905

Oriental,northeast

Australia

AnchocerusbirmanusFauvel,1905

NorthOriental

Quediina


AnisolinusSharp,1889

PalearcticandOriental

AnisolinustsurugiensisSawada,

1961

Japan

Anisolinina


AstrapaeusGravenhorst,1802

WestPalearctic

Astrapaeusulm

iRossi,1790

WestPalearctic

Quediina

BasallineageofStaphylinini

AtanygnathusJakobson,1909

World

Atanygnathusterm

inalis(Erichson,

1839)

Palearctic,

world

(introduced,or

misidentified)

Tanygnathinina


novo’

CheilocolpusSolier,1849

Neotropical(tem

perate)

Cheilocolpuspyrostomus(Solier,

1849)

Chile

Quediina


novo’

CreophilusLeach,1819

World

Creophiluserythrocephalus

(Fabricius,1775)

Australian,Pacific,

Neotropical

(introduced)

Staphylinina

‘Staphylinina’

DinothenarusThomson,1858


andPacific

DinothenaruscapitatusJ.

Bland,

1864

Nearctic

Staphylinina

‘Staphylinina’

EdrabiusFauvel,1900

Neotropical(tem

perate)

EdrabiusalticolusSeevers,1955

Peru,Bolivia

Amblyopinina


novo’

Elm

asBlackwelder,1952

Neotropical

Elm

asstrigella

(Bernhauer,1915)

Brazil

Xanthopygina


EucibdelusKraatz,1859

PalearcticandOriental

EucibdellusjaponicusSharp,1874

Japan,Korea

Staphylinina

‘Staphylinina’

EuristusFauvel,1899

WestAfrica

EuristusglobusFauvel,1899

Gabon,Cameroon

Quediina

‘Staphylinina’

EuryporusErichson,1839

WestPalearctic

Euryporuspicipes

(Paykull,1800)

WestPalearctic

Quediina


GlyphesthusKraatz,1858

Africa(A

frotropical)

Glyphesthusneavi

Bernhauer,1927

Kenya,Malawi

Philonthina

‘Philonthinapropria’

HasumiusFairmaire,

1891

Africa(A

frotropical)

HasumiuslamoraliBernhauer,1933

Congo

Xanthopygina


Hem

iquediusCasey,1915

Nearctic

Hem

iquediusferox(LeC

onte,1878)

Nearctic

Quediina


HesperusFauvel,1874

World

HesperusapicialisSay,1830

Nearctic

Philonthina


HeterothopsStephens,1829

World

Heterothopsbinotatus

(Gravenhorst,1802)

Palearctic

Quediina


novo’

HeterothopsdolichocephalusLea,

1925

Australia

Quediina


novo’

HolisusErichson,1839

Neotropical,Nearctic,

Afrotropical

Holisussp.

Mexico

Hyptiomina


IndoquediusBlackwelder,1952

East

Palearctic,

Oriental

Indoquediussikkim

ensis(C

ameron,

1932)

India,Bhutan

Quediina


IomaBlackwelder,1952

Africa(A

frotropical)

Iomasetigera(Fauvel,1899)

Afrotropical

Quediina




Table

1.Continued.

Genus(subgenus)

Genusdistribution

Speciesin

analysis

Speciesdistribution

Curentsubtribe

Cladein

Fig.1

LoncoviliusGermain,1903

Neotropical(tem

perate)

Loncoviliussemiflavus(Fairmaire

andGermain,1861)

Chile

Quediina


novo’

MoeocerusFauvel,1899

Africa(A

frotropical)

MoeocerusoverlaetiBernhauer,

1935

Congo,Ethiopia

Philonthina


NatalignathusSolodovnikov,2005

South

Africa

NatalignathusolgaeSolodovnikov,

2005

South

Africa

Quediina


novo’

NordusBlackwelder,1952

Neotropical

Nordusfungicola

Sharp,1884

CentralAmerica

Xanthopygina


ParisanopusBrethes,1900

Neotropical

ParisanopuscastaneipennisBrethes,

1900

Brazil

Quediina


PhilonthusStephens,1829

World

Philonthuspolitus(Linnaeus,1758)

Palearctic,

world

(introduced)

Philonthina


PlatydracusThomson,1858


andPacific

Platydracusmaculosus

(Gravenhorst,1802)

Nearctic

Staphylinina

‘Staphylinina’

QuediocafusCameron,1945

New

Zealand,

Antipodes

Islands

Quediocafusinsolitus(Sharp,1886)

New

Zealand,

Antipodes

Islands

Quediina


novo’

Quediomim

usCameron,1948

New

Zealand

Quediomim

usbrookesiCameron,

1948

New

Zealand

Quediina


novo’

QuediusStephens,1829

World

Quediuscalogaster

Lea,1929

Australia

Quediina


novo’

(QuediusStephens,1829)

Holarctic

Quediusfuliginosus(G

ravenhorst,

1802)

Palearctic

Quediina

‘Quediinasensu

stricto’

(CyrtoquediusBernh.,1917)

Neotropical

QuediuslabiatusErichson,1840

CentralandSouth

America

Quediina


(QuedionuchusSharp,1884)

Holarctic,

northern

Neotropical

QuediusplagiatusMannerheim,

1843

Holarctic

Quediina


(RaphirusStephens,1829)

Holarctic,

northern

Oriental

QuediusumbrinusErichson,1839

Palearctic

Quediina


ScelotrichusBernhauer,1915

PapuaNew

Guinea

ScelotrichuselegansBernhauer,

1915

PapuaNew

Guinea

Philonthina


SmilaxLaporte,

1835

Neotropical,

Afrotropical,Oriental

Smilaxpilosa

Fabricius,1787

Neotropical

Quediina/

Xanthopygina


SphingoquediusBernhauer,1941

New

Zealand

Sphingoquediusstrandi(Bernhauer,

1941)

New

Zealand

Quediina


novo’

SternotoxusBernhauer,1916

Australia

SternotoxusflavicornisBernhauer,

1916

Australia

Philonthina


Tolm

erinusBernhauer,1923

Palearctic,

Oriental,

Afrotropical

Tolm

erinussp.

Oriental

Incertaesedis/

Anisolinina


TriacrusNordmann,1837

Neotropical

TriacrusdilatusNordmann,1837

Brazil

Xanthopygina


TympanophorusNordmann,1837


andPacific

Tympanophoruspuncticollis

(Erichson,1840)

NorthAmerica

Xanthopygina/

Anisolinina


Valdiviodes

Smetana,1981

Neotropical(south

temperate)

Valdiviodes

ashworthiSmetana,

1981

Chile

Quediina


XanthopygusKraatz,1857

Nearcticand

Neotropical

Xanthopygusxanthopygus

(Nordmann,1837)

CentralandSouth

America

Xanthopygina




4. Frontoclypeal (epistomal) suture: 0. present; 1. absent.5. Infraorbital ridge: 0. absent; 1. present.

Note: Here and below, infraorbital and other head ridgesare considered strictly according to Smetana & Davies(2000).

6. Infraorbital ridge, degree of development: 0. very short,ventrally not extending further than very basal part ofhead; 1. moderately long, ventrally extending to aboutmiddle of distance from neck to base of mandibles; 2.

long, distinct from base of head to base of mandibles.7. Infraorbital and nuchal ridges: 0. separate; 1. confluent.8. Nuchal ridge: 0. absent; 1. present.

9. Nuchal ridge, degree of development: 0. distinct dor-sally and laterally; 1. distinct dorsally, laterally and,more or less, ventrally; 2. distinct only dorsally, dis-

appearing laterally; 3. distinct only laterally and, moreor less, ventrally, entirely obsolete dorsally; 4. distinctonly laterally, entirely obsolete dorsally and ventrally.

10. Postgenal ridge: 0. absent; 1. present.

11. Postmandibular ridge: 0. absent; 1. present.12. Postmandibular ridge, shape: 0. short, extending only

ventrally, sometimes confluent with long ventral ex-

tensions of nuchal or infraorbital ridges; 1. long,extending ventrally and turning more or less dorsallybehind eyes.

13. Infraorbital ridges and postgenal ridges: 0. separate, or,if running together, still always distinct as separateridges along their joint extension; 1. part of infraorbital

ridge confluent with part of postgenal ridge, formingessentially one ridge along their joint extension.

14. Head, dorsal basal ridge: 0. absent; 1. present.15. Labrum, transparent apical membrane: 0. absent; 1.

present (at least laterally).16. Mandibles, dorsolateral groove: 0. absent; 1. present.17. Mandibles, dorsolateral groove, shape: 0. well devel-

oped; 1. rudimentary.18. Mandibles, position of dorsolateral groove: 0. more or

less dorsal on mandible (visible in dorsal view); 1. more

or less lateral on mandible (not visible in dorsal view).19. Maxillary palps, apical segment, shape: 0. more or less

fusiform, not very elongate, at base only slightly

narrower than penultimate segment; 1. more or lessconical, thin, at base distinctly narrower than penulti-mate segment; 2. more or less parallel sided, withtruncate apex, at base only slightly narrower than

penultimate segment; 3. more or less widening apically,with more or less truncate apex; 4. very elongate,slightly fusiform, at base only slightly narrower than

penultimate segment; 5. more or less conical and short,at base narrower, and overall considerably shorter thanpenultimate segment; 6. more or less fusiform, relatively

short and wide; 7. very elongate, apically pointed.20. Maxillary palps, apical segment, setation: 0. glabrous,

with some very sparse setae at most; 1. medium denselysetose; 2. densely setose.

21. Ligula: 0. more or less bilobed, with variously developedrounded lobes; 1. small, entire (or at most slightlynotched medially); 2. strongly reduced, indistinct.

22. Labial palps, apical segment: 0. more or less fusiform, atbase slightly narrower than penultimate segment, api-

cally slightly truncate; 1. more or less parallel sided,apically slightly narrowing and truncate; 2. more or lesswidening apically, with truncate apex; 3. very narrow,

more or less conical, much narrower than penultimatesegment; 4. very elongate, more or less parallel sided,slightly narrowing apically, at base only slightly nar-rower than penultimate segment; 5. very elongate,

apically pointed.23. Pronotum, punctation of disc: 0. disc smooth, without

setiferous punctures (if punctured, punctures without

setae); 1. more or less densely punctured, setiferouspunctures not arranged in longitudinal rows; 2. verysparsely punctured, these large setiferous punctures in

some symmetrical arrangement, most often in longitu-dinal rows (sometimes also with distinctly smaller anddenser punctation not arranged in rows); 3. verysparsely punctured, these large setiferous punctures

not arranged in rows.24. Pronotum, superior marginal line of pronotal hypomer-

on: 0. developed through its whole length, not deflexed

under anterior angle of pronotum; 1. developed throughits whole length, deflexed under anterior margin ofpronotum; 2. ending at anterior corners of pronotum,

not deflexed under them.25. Pronotum, superior and inferior marginal lines of

hypomeron: 0. inferior line shorter than superior line

and not meeting it; 1. inferior line subcontiguous orfused to superior line behind anterior angles of prono-tum; 2. inferior line (sometimes partly obsolete) fusedwith superior line in front of anterior angles of prono-

tum; 3. inferior line longer than superior line and notmeeting it.

26. Pronotum, front angles: 0. not strongly produced

beyond (anteriad of) anterior margin of prosternum;1. strongly produced beyond (anteriad of) anteriormargin of prosternum.

27. Pronotal hypomeron: 0. not fully inflexed (visible inlateral view of prothorax); 1. slightly inflexed (hardly,but visible in lateral view of prothorax); 2. strongly

inflexed (not visible in lateral view of prothorax).28. Pronotal hypomeron: 0. with weakly sclerotized post-

coxal process; 1. without weakly sclerotized postcoxalprocess.

29. Pronotosternal suture: 0. well developed, clearly visible,distinctly membranous; 1. clearly visible as weaklysclerotized strip, fine groove or carina, but not mem-

branous; 2. indistinct at least in middle (prosternumfused to pronotum), sometimes distinct at either or bothends.

30. Prosternum, mid-longitudinal carina: 0. absent (onlymedial prominence, not carinate, not longitudinal); 1.developed only along furcasternum; 2. developed alongfurcasternum and at least part of basisternum; 3.

developed only along basisternum.31. Prosternum, shape of longitudinal carina: 0. carina, at

least in its basal part, sharp, forming well-defined ridge;



1. carina more or less rounded, from obtuse ridge tosmooth longitudinal prominence of prosternum.

32. Prosternum, basisternum: 0. without conspicuous mac-rosetae; 1. with one pair of macrosetae (often nearanterior margin of prosternum).

33. Elytral sub-basal ridge: 0. present; 1. absent.34. Elytral sub-basal ridge, shape: 0. immediately adjacent

to elytral articulation, short, extending anteriad (notextending laterad to humerus); 1. long, extending from

level of middle of scutellum to elytral humerus.35. Elytral disc: 0. without punctures or setiferous pores; 1.

with several punctures bearing macrosetae forming

loose rows; 2. more or less densely punctured, puncturesnot arranged in rows.

36. Elytra, spines or spine-like setae (distinctly stronger than

other setae of elytra) on humerus: 0. absent; 1. present.37. Mesosternum: 0. with more or less irregular group of

several macrosetae (in some combined with shortergeneral setation); 1. with one pair of macrosetae; 2.

with two pairs of macrosetae; 3. almost asetose.38. Sternopleural (anapleural) suture: 0. transverse, or

nearly transverse (very slightly oblique); 1. distinctly

oblique (medial end of suture anterior to its lateral end).39. Mesosternum, structure of medial part (with respect to

the position of mesocoxae): 0. disc of mesosternum

situated more or less in one plane with, or only slightlymore ventrally than its median (mesosternal part ofmesocoxal acetabula); 1. disc of mesosternum situated

distinctly more ventrally than median part (mesosternalpart of mesocoxal acetabula), which is usually carinate.

40. Mesosternal intercoxal process: 0. rounded or, ifpointed, forming more or less obtuse angle; 1. narrow,

usually pointed, forming more or less sharp angle; 2.truncated; 3. metasternum fused with mesosternum sothat mesosternal intercoxal process indistinct.

41. Mesocoxae: 0. contiguous; 1. more or less separated byelevated part of metasternum.

42. Mesoscutellum: 0. with one transverse carina, separat-

ing scutellum from prescutum; 1. with two transversecarinae (one posterior, separating scutellum from pre-scutum; another anterior, extending between anterior

notal wing processes).43. Protarsi, shape: 0. tarsomeres I–IV more or less cylin-

drical, not transversely widened and not flatteneddorsoventrally; 1. tarsomeres I–IV more or less dorso-

ventrally flattened and transversely widened.44. Middle and posterior tarsi, shape: 0. elongate (at most

slightly shorter than tibiae), tarsomeres I–IV more or

less longer than wide, not enlarged or distinctly dor-soventrally compressed; 1. shorter, distinctly shorterthan tibiae, tarsomeres I–IV not elongate, more or less

compressed dorsolaterally, II–IV more or less enlarged.45. Protarsal adhesive setae, females: 0. absent; 1. present.46. Mesotarsal adhesive setae, males: 0. absent; 1. present.47. Mesotarsal combs, males: 0. absent; 1. present.

48. Metatarsal adhesive setae: 0. absent; 1. present.49. Tarsal empodia: 0. with one pair of setae; 1. glabrous; 2.

with one seta.

Note: In Acylophorus glaberrimus anterior tarsi withoutempodial setae, but overall the character is coded as ‘0’ for

this species.50. Tarsal empodial setae: 0. on all tarsi about as long as, or

longer than, claws; 1. on all tarsi much shorter (about

half as long or less) than claws; 2. much shorter (abouthalf as long) than claws on anterior tarsi, about as longas, or longer than, claws on middle and posterior tarsi.

51. Tarsal dorsal apical flat seta (located dorsally near apex

of last tarsomere): 0. absent; 1. present.52. Hind coxae, shape: 0. broader concave basal part more

or less gradually transforming into narrower and flatter

apical part; 1. broader concave basal part abruptly andsharply separated from narrower and flatter apical partby carina or strong groove (sometimes distinct only at

lateral parts of coxae).53. Hind wing venation, CuA and MP4: 0. completely

separate; 1. fused in one vein (although often its originfrom two veins still very obvious).

54. Hind wing venation, MP3: 0. vein MP3 present,although sometimes faint; 1. vein MP3 absent.

55. Protergal glands (osmeteria), cuticular manifestation: 0.

absent; 1. present.56. Protergal glands (osmeteria), morphology of cuticular

manifestation: 0. shallow impressions; 1. well-developed

acetabula; 2. more or less invaginated capsules withsmaller openings.

57. Abdominal tergite II, basal longitudinal carina: 0.

present, entire (connecting anterior margin of tergitewith transverse basal carina); 1. present, reduced (notreaching transverse basal carina); 2. present, extended(basally crossing transverse basal carina and extending

posteriad); 3. absent.58. Abdominal tergite III, basal transverse carinae: 0.

anterior and posterior complete; 1. anterior complete,

posterior incomplete; 2. anterior complete, posteriorabsent.

59. Abdominal tergite IV, basal transverse carinae: 0.

anterior and posterior complete; 1. anterior complete,posterior incomplete; 2. anterior complete, posteriorabsent.

60. Abdominal tergite V, basal transverse carinae: 0. ante-rior and posterior complete; 1. anterior complete,posterior incomplete; 2. anterior complete, posteriorabsent; 3. anterior incomplete, posterior absent.

61. Abdominal tergite VI, basal transverse carinae: 0.anterior and posterior present; 1. anterior present,posterior absent; 2. anterior and posterior absent.

62. Abdominal sternite III, basal transverse carina, medialpart: 0. straight, rounded, or slightly pointed; 1. sharplypointed.

63. Male sternum VIII, apex: 0. medially straight to veryslightly concave; 1. with single, variably developedmedian emargination; 2. with two emarginations.

64. Male sternum IX, shape: 0. more or less symmetrical; 1.

basally weakly to strongly asymmetrical.65. Male and female lateral tergal sclerites IX, shape: 0.

produced into more or less inflated, apically sharp



processes; 1. produced into more or less inflated,apically obtuse or rounded processes; 2. produced into

broad, flat, apically pointed processes.66. Ovipositor: 0. consisting of paired proximal and distal

gonocoxites, the latter bearing styli; 1. consisting of only

paired proximal and distal gonocoxites, styli absent.67. Female tergal sclerites IX, setation: 0. with setae only; 1.

apically and externally also with spines.68. Ovipositor, distal gonocoxites: 0. with setae only; 1.

with setae and spines.69. Aedeagus, parameres: 0. paired, well separated; 1. fused

into a single lobe (sometimes this lobe more or less

secondarily bifurcate); 2. absent.70. Aedeagus, sensory peg setae of the paramere(s): 0.

absent; 1. present.

71. Aedeagus, basal part of median lobe: 0. more or lesssymmetrical; 1. more or less asymmetrical.

72. Aedeagus, apical part of median lobe: 0. symmetrical; 1.more or less asymmetrical.

73. Aedeagus, paramere(s), degree of symmetry: 0. sym-metrical; 1. more or less asymmetrical.

74. Aedeagus: 0. paramere(s) is (are) not (or at most

slightly) produced over apex of median lobe, usuallynarrower or at most as wide as median lobe: medianlobe appearing as the larger part of the aedeagus; 1.

paramere strongly produced over apex of median lobe,mostly as large as or larger than median lobe: paramereappearing as a larger part of the aedeagus; 2. paramere

small (short and/or thin), obviously strongly reduced.75. Aedeagus, paramere(s): 0. fused to median lobe only at

base, at place of its (their) attachment to median lobe,otherwise paramere(s) distinctly separated from median

lobe along entire length; 1. fused to median lobe only atbase, at place of its (their) attachment to the former, andvery closely appressed to median lobe along entire

length; 2. fused to median lobe along its (their) entirelength, paramere(s) and median lobe hardly distinguish-able from each other.

76. Aedeagus, position of dorsal (‘parameral’) side inabdomen in repose: 0. facing dorsally; 1. facing ven-trally; 2. facing left laterally.

Results

The cladistic analysis revealed 40 most-parsimonious treesof 437 steps in length, with the consistency and retentionindexes 0.281 and 0.549, respectively. The majority-rule

consensus of these trees (Fig. 1) is largely resolved, withsome polytomies present in the clades ‘Quediina propria’and ‘Philonthina lineage’ (the rationale for the informal

names used for the newly revealed clades is given below inthe ‘Discussion and conclusions’ section). The consensustopology largely disagrees with the current system of thetribe Staphylinini. The better-sampled subtribe Quediina

comes out as largely polyphyletic. Sampled taxa currentlyattributed to the subtribes Xanthopygina and Anisolininado not form respective monophyletic groups, either. Only

Staphylinina in this analysis holds as a monophyletic group(clade ‘Staphylinina’). The subtribe Philonthina also re-

mains largely monophyletic (‘Philonthina propria’ clade),but includes the genus Holisus, currently forming a separatesubtribe Hyptiomina.

Conventional Quediina falls apart into several lineages: (i)various basal Staphylinini [here the genera Valdiviodes,Astrapeus and the South American Quedius (Cyrtoquedius)labiatus and Parisanopus]; (ii) ‘Quediina propria’, which

here includes two subclades: one with the north temperaterepresentatives of the genusQuedius from various subgeneraand the genus Indoquedius, the other with the genera

Acylophorus, Anchocerus, Hemiquedius and Euryporus; and(iii) ‘Tanygnathinina sensu novo’, which includes southtemperate species of ‘Quedius’ and other south temperate

genera of the current ‘Quediina’. The latter also encom-passes the genus Atanygnathus (currently in the monotypicsubtribe Tanygnathinina) and the subtribe Amblyopinina,here represented by the genus Edrabius.

However, there is no strong statistical confidence in therevealed tree topology overall, as manifested by the lowBremer support values (1) for most of the clades. The clades

with the strongest support are: Platyprosopini þ Staph-ylinini (Bremer support 10), Staphylinini (Bremer support5); Acylophorus þ Anchocerus (Bremer support 5); and

Atanygnathus þ Natalignathus (Bremer support 6). Otherrelatively well-supported clades (Bremer support 2–3) arethe large subclade of Staphylinini sister to Valdiviodes, the

clade includingAcylophorus and allies, ‘Staphylinina’ and itssubclades, and the clade Tolmerinus þ Tympanophorus.This analysis agrees only partially with the previous

analysis (Solodovnikov, 2006); the two datasets are not

identical, but are largely comparable in the scope of taxaand characters investigated. It is beyond the goals of thepresent paper to compare both analyses in detail, but we

should stress the main points of such a comparison. Bothanalyses agree on the non-monophyly of ‘Quediina’ and‘Quedius’, and on the non-monophyly of ‘Xanthopygina’.

They agree on revealing the core of ‘Quediina propria’ [cladeC in Solodovnikov (2006)], the large lineage ‘Tanygna-thinina sensu novo’, which also includes Atanygnathus

(Tanygnathinina) and Amblyopinina [clade B in Solodov-nikov (2006)], and on placing some current Quediina asvarious non-related basal groups within Staphylinini [gen-era Valdiviodes, Afroquedius, Quedius (Cyrtoquedius) and

others]. But the relationships among those main lineages aredifferent in the two analyses. In particular, ‘Quediinapropria’ and ‘Tanygnathinina sensu novo’ are sister clades

here but not in Solodovnikov (2006; clades B and C,respectively). Also, the genus Afroquedius was found to beone of the very basal lineages of Staphylinini in Solo-

dovnikov (2006), but here it comes out as a member of the‘Tanygnathinina sensu novo’ clade. Consistent with Solo-dovnikov (2006), Natalignathus is as a sister group toAtanygnathus. The genus Euristus comes out as a basal

member of the ‘Staphylinina lineage’. The genus Ioma,together with other African endemic genera, Moeocerus,Glyphestus and Hasumius (all three placed in Quediini until



very recently, for more details see below), forms a sisterlineage to current members of the subtribe Philonthina (plusHolisus, Hyptiomina).

Review of genera, with notes on taxonomy, phylogenetic

relationships and biogeography

1. Acylophorus Nordmann, 1837. Type species: A. gla-

berrimus (Herbst, 1784) (¼Acylophorus ahrensii Nordmann,1837)This globally distributed genus currently includes 129

described species, of which 24 occur in sub-Saharan Africa

[Herman, 2001; for the latest additions see Solodovnikov(2008)]. Our examination of diverse material from Africaand other parts of the world suggest that this genus, as

defined now (Smetana, 1971, 1988, 1995a; Herman, 2001), ismonophyletic. Phylogenetically, Acylophorus is closelyrelated to the mostly Oriental genus Anchocerus (Smetana,

1988; Solodovnikov, 2008), and is placed in the clade‘Quediina propria’ by the present analysis. The biogeo-graphical history of the widespread genus Acylophorus,

including its African representatives, cannot be addressedbefore the needed species-level revision of this genus iscompleted. We assume that the genus could originate in

Southeast Asia from an Anchocerus-like ancestor anddispersed around the world and to the Afrotropical region.

2. Afroquedius Solodovnikov, 2006. Type species: Que-

dius sexpunctatus Bernhauer, 1917This small genus (four described species) endemic to

South Africa, originally known as Quedius sexpunctatus

Fig. 1. Results of the parsimony-based cladistic analysis of the phylogenetic relationships of some genera of Staphylinini. Majority-rule

consensus of the 40 shortest trees obtained (tree length¼ 437; consistency index¼ 0.281; retention index¼ 0.549). 1–10: Bremer support for the

respective clades under which these numbers are plotted, as computed in TNT (clades with support ‘1’ collapse at the strict consensus).

Afrotropical endemic genera of special interest are marked in bold. Clades of interest are colour coded; distributions of the sampled genera and

their current formal systematic position are summarized. Afrotropical endemic distributions and current formal affiliations to Quediina are

colour coded by red.



Bernhauer, 1917, is rather peculiar morphologically, andis either one of the very basal lineages of Staphylinini

(Solodovnikov, 2006), or a basal member of the species-richlineage ‘Tanygnathinina sensu novo’ (present analysis). Therecent distribution of the ‘Tanygnathinina sensu novo’ is

mostly confined to the disjunct south temperate areas of theworld. Such a pattern suggests association of the origin ofthis group with the former supercontinent Gondwana, aswas earlier suspected by Newton (1985) for the ‘south

temperate Quedius’. The position of Afroquedius as a sistertaxon to the rest of ‘Tanygnathinina sensu novo’ is congru-ent with the classical vicariance-based phylogenetic pattern

where the African taxa are expected to diverge first due toearlier separation of the respective landmass fromGondwana(Sanmartın & Ronquist, 2004). Therefore, the biogeograph-

ical evidence seems to support the present phylogeneticresult rather than that in Solodovnikov (2006). However,the occurrence of Afroquedius only in South Africa, alsoa renowned area of survival for older Laurasian relict taxa

(Werger & Van Bruggen, 1978), can as well be interpretedsimilarly to Arrowinus (Solodovnikov & Newton, 2005), asa very old, pre-Gondwanan relict. Without at least some

evidence about the possible age of Afroquedius, it is hard toapply a biogeographical criterion for the choice between twophylogenetic hypotheses about this genus. The problem can

be further addressed using molecular data.

3. Atanygnathus Jakobson, 1909. Type species: Atanyg-

nathus terminalis (Erichson, 1839)This genus currently includes 49 described species

(Solodovnikov, 2005), of which 18 occur in sub-SaharanAfrica and one is known from Central Sahara (Drugmand,

2004). The phylogenetic position of Atanygnathus withinStaphylinini was a source of controversy in the systematicsof Staphylinidae, until the genus Natalignathus was recently

discovered in South Africa (Solodovnikov, 2005). Thediscussion in Solodovnikov (2005) and the formal analysesin Solodovnikov (2006) and here provide evidence to link

a highly derived Atanygnathus via the plesiomorphy-richNatalignathus with the morphologically diverse ‘Tanygna-thinina sensu novo’ lineage. The mentioned analyses suggest

that Atanygnathus is a derived member of the ‘Tanygnathi-nina sensu novo’ clade and originated in some of the formerGondwanic landmasses with later dispersal around theworld. However, it is premature to address the biogeograph-

ical history of Atanygnathus in detail before its species-levelrevision is completed.

4. Natalignathus Solodovnikov, 2005. Type species: Na-talignathus olgae Solodovnikov, 2005Since the original description, this genus, endemic of

South Africa, was associated with the ‘south temperateQuediina’ (Solodovnikov, 2005) or ‘Tanygnathinina sensunovo’ (Solodovnikov, 2006; present study). Because it isassumed that the diversification of ‘Tanygnathinina sensu

novo’ took place in the period of changing land connectionsamong the landmasses of breaking-up Gondwana, similar tothe case discussed above for Afroquedius, one would expect

a more or less basal position of Natalignathus within the‘Tanygnathinina sensu novo’ clade. But, contrary to such

prediction, in both formal analyses (Solodovnikov, 2006;present study) the monotypic Natalignathus comes out asa relatively terminal lineage of ‘Tanygnathinina sensu novo’.

A rather terminal position of the Natalignathus þ Atanyg-nathus clade within ‘Tanygnathinina sensu novo’, and theearlier discussed (Solodovnikov, 2005) similarity of Nata-lignathus to some of the Australian ‘Quedius’ (Q. nothus Lea,

1925, Q. stenocephalus Lea, 1925, and allies) suggest a dis-persal event that possibly brought the Natalignathus þAtanygnathus ancestor from Australia to Africa some time

after these landmasses had any connection. Although long-distance transoceanic dispersal for the small terrestrialbeetles seems unlikely, it cannot be ruled out. Alternatively,

there is some evidence of stepping-stones between Africaand Australia as late as ca. 45 Mya (Jønsson & Fjeldsa,2006).

5. Heterothops Stephens, 1829. Type species:Heterothopsbinotatus (Gravenhorst, 1802)According to Coiffait (1978), Smetana (1971, 1988) and

others, Heterothops is a globally distributed genus with 149species described from the world (Herman, 2001). Moderndiagnoses of the genus (Smetana, 1971; Coiffait, 1978) are

based on the study of the Holarctic representatives, its limitsbecoming blurred if the world fauna is considered. Inparticular, our studies (in progress) suggest that ‘Hetero-

thops’ from Australia, Lord Howe Island and Papua NewGuinea are not congeneric at least with the Holarcticmembers of this genus (including the type species), andmay be closely related to (and some of them probably

congeneric with) the temperate South American genusCheilocolpus Solier, 1849. Such suspicion about the non-monophyly of the current Heterothops is supported in the

present analysis: the AustralianHeterothops dolichocephalusdoes not form a group with the Palearctic Heterothopsbinotatus (Fig. 1). However, unlike the strongly polyphyletic

‘Quedius’, where current members of the genus are widelyscattered across very different groups of Staphylinini(Solodovnikov, 2006; present analysis), all ‘Heterothops’ at

least seem to be members of the ‘Tanygnathinina sensunovo’ monophylum. With respect to the proper delineationof the generic limits of Heterothops, two misconceptionsconcerning its morphology must be eliminated. Smetana

(1971, 1988) believed that the aedeagus of Heterothops (andAtanygnathus) lacks parameres due to their complete reduc-tion. Coiffait (1978), in contrast, suggested that the

‘absence’ of the parameres in both of these genera is dueto their fusion to the median lobe of the aedeagus. Ourearlier discussion about the structure of the aedeagus of

Atanygnathus (Solodovnikov, 2005) and examination of theaedeagi of the entire ‘Heterothops’ and ‘Tanygnathininasensu novo’ complex leaves no doubt that it is the mor-phological interpretation of Coiffait that is correct. The

species of Heterothops indeed have an aedeagus with a well-developed paramere fused with a highly reduced medianlobe. Another diagnostic character state indicated for



Heterothops (Smetana, 1971, 1988; Coiffait, 1978) is thepresence of the infraorbital ridges on the head capsule.

However, under the more precise definition of the ‘infraor-bital ridges’ (Smetana & Davies, 2000; Solodovnikov, 2006;present analysis), the homology of the ‘infraorbital ridges’

across the entire ‘Heterothops complex’ is not confirmed.Whether all species of Heterothops have infraorbital ridgessensu Smetana & Davies (2000) or something else needsfurther investigation. On the basis of the character states

listed below, Heterothops megalops fits the existing defini-tion of Heterothops well, and is probably congeneric withthe Holarctic lineage of the ‘Heterothops complex’, i.e. with

the genus Heterothops in a possibly restricted sense of thefuture. Although any biogeographical speculations are pre-mature until the entire ‘Heterothops complex’ is sufficiently

studied, the fact that this relatively species-rich lineage isrepresented in the Afrotropical region by only a singlespecies is noteworthy. Seven species of Heterothops knownfrom Northern Africa and the Canary Islands (Smetana,

2004) are apparently of Palearctic origin, based on theirmorphology and distributions. This cannot be said forcertain forHeterothops megalops. This species is redescribed

below.

Heterothops megalops Cameron, 1959. Cameron 1959:117

Type material examined. Republic of Angola: Lectotype(here designated): male, ‘type [circle]/MUSEU DO DUN-DOAnned. Vila Luso [Luena] 2.I.1949 Reg. ANG. 1261.16/

Ang. 1261.16 [in pencil]/M. Cameron. Bequest. B.M. 1955-147./Heterothops megalops Cam. Type [handwritten]/syn-type [circle]/Lectotype Heterothops megalops Cameron,

1959, A. Solodovnikov des. 2008’ (NHM); paralectotype:female, ‘paratypes [circle]/1261.16 [in pencil]/Heterothopsmegalops Cam. Cotype [handwritten]/M. Cameron.

Bequest. B.M. 1955-147./Syntype [circle]/ParalectotypeHeterothops megalops Cameron, 1959, A. Solodovnikovdes. 2008’ (NHM).

Additional material examined. Republic of Rwanda: 42males, 40 females, Bugesera, Biharagu, 27.II.1960, leg. N.Leleup, in dead leaves on termite nest, B. 141; 22 males, 19females, Bugesera, Kibugabuga, 25.II.1960, N. Leleup, in

dead leaves on termite nest, B. 137 (MRAC); Republic of

South Africa: Gauteng: one male, one female, Cullinan,Premier Mine Game Reserve, 18.I.2002, sifting leaf litter,

leg. Endrody-Younga (EY:3483) (FMNH); 30 males, 32females, Cullinan, Windy Brow, 8.I.2002, sifting leaf litter,leg. TMSA staff (EY: 3470) (FMNH, TMSA); Mpumalan-

ga: one male, one female, Goedehoop Farm (The BrookHiking Trail), 15 kmWof Lochiel, 1400 m, rocky outcrop ingrassland, 28.XI.2002, in leaf litter under trees and shrubs,leg. R. Stals (SANC); Transvaal: two males, seven females,

Berlin, gorge edge, 10.XII.1986, in litter between rocks, leg.Endrody-Younga (EY:2373) (TMSA, FMNH); five males,four females, Blyderiver Canyon, 5.V.1981, sifting leaf litter,

leg. Endrody-Younga (EY: 1780) (TMSA, FMNH); onefemale, Hekpoort, Cyara, 9.I.1987, leg. Van Viegen (VV:

245) (TMSA); one male, Hylsvley, hill base, 29.II.1976, inleaf litter, leg. Endrody-Younga (EY: 1050) (TMSA); onemale, four females, Mariepskop, cloud forest, 5.V.1981, in

leaf litter, leg. Endrody-Younga (EY: 1776) (TMSA); onefemale, Nelshoogte, gallery forest, 2.XII.1986, in leaf litter,leg. Endrody-Younga (EY: 2341) (TMSA); three males, onefemale, Nelspruit National Reserve, rivulet valley,

18.XII.1986, in leaf litter, leg. Endrody-Younga (EY:2397) (TMSA); one male, two females, Nylsvley Meteoro-logical Station, 28.V.1976, in leaf litter, leg. Endrody-

Younga (EY: 1162); two females, same data but29.III.1976 (EY: 1112); one male, same data but 27.I.1976(EY: 1015); one female, same data but 30.IV.1976 (EY:

1152) (TMSA); one male, one female, Nylsvley, hill base,29.III.1976, sifting leaf litter, leg. Endrody-Younga (EY:1113); one male, same data but 8.I.1976 (EY: 990); threemales, two females, same data but 28.V.1976 (EY: 1161)

(TMSA, FMNH); one male, Rhenosterpoort NationalReserve, 16.XI.1975, sifting leaf litter, leg. Endrody-Younga(EY: 937); one male, environs of Utsoek Forest Station,

13.XII.1983, in horse dung, leg. Endrody-Younga (EY:2383) (TMSA).

Redescription (Fig. 2A–C). Measurements (15 specimens):HL: 0.43–0.49 (0.46); HW: 0.49–0.55 (0.52); PL: 0.55–0.65

(0.60); PW: 0.59–0.72 (0.66); EL: 0.61–0.72 (0.68); EW:

0.73–0.88 (0.80); total length of body: 3.7–4.8 (4.2) mm.Slender species with round, large-eyed head, relatively

broad pronotum and short elytra. Head black; pronotumbrown; elytra pale brown, often with darker discal area;

abdomen usually dark brown to black with pale, reddishapical margins of tergites; legs, antennae and mouthpartspale brown. Body glossy, with long yellowish hairs on elytra

and abdomen (Fig. 2A).Head slightly wider than long, with large broadly rounded

eyes (longitudinal diameter of eye more than six times as

long as tempora). Dorsal chaetotaxy of head at each side asfollows: one seta near base of antenna; four setae alonginner margin of eye: one on frons anteriorly [‘anterior

frontal’ seta in Smetana (1971)], one in middle, two inposterior part of inner margin of eye [of which posterior setais called ‘posterior frontal’ seta in Smetana (1971)]; and oneseta posteromedially, near neck constriction [‘vertical’ seta

in Smetana (1971)]. Surface of head with transverse wavymicrosculpture. Antennae: antennomere I slightly shorterthan antennomeres II and III combined; antennomeres II–X

gradually becoming shorter towards antennal apex: anten-nomeres II–V longer than wide, VI–X about as long as wide,last antennomere (XI) slightly shorter than antennomeres

IX and X combined.Pronotum slightly wider than long, distinctly wider than

head, widest at area of its broadly rounded posteriorcorners, at sides converging to its rounded anterior corners.

Disc of pronotum with two setae in each dorsal row, withoutsetae in sublateral area (between dorsal row and large lateralseta). Microsculpture of pronotum as that on head.



Elytra relatively short, but slightly longer than pronotum,with shallow sparse seta-bearing punctation. Elytral surfacevery glossy. Surface and punctation of scutellum as onelytra. Wings highly reduced.

Abdominal tergites with shallow seta-bearing punctationsimilar to that of elytra and gradually becoming sparserfrom base towards apex of abdomen. Tergal surface with

distinct transverse microsculpture. Tergite VII with thinwhitish margin apically.Male. Anterior tarsi slightly enlarged, as wide as apex of

anterior tibiae, wider than in female. Abdominal sterniteVIII with shallow median emargination apically. Aedeagus(Fig. 2B, C) with long apically pointed paramere firmlyattached to short and more weakly sclerotized median lobe;

apical portion of paramere slightly curved ventrad (param-eral side ¼ dorsal side).Female. Anterior tarsi slightly narrower than apex of

anterior tibiae, distinctly narrower than tarsi in males.Abdominal sternite VIII with simple, concave apicalmargin.

Comparison. Without a broad-scale revision of the genus,it is difficult to judge about the sister relationships of

Heterothops megalops. However, compared with all othersix species recorded for the African continent [the wide-spread Palearctic species H. binotatus (Gravenhorst, 1802),H. dissimilis (Gravenhorst, 1802), H. minutus Wollaston,

1860 and H. praevus Erichson, 1839; as well as NorthernAfrican H. marocanus Coiffait, 1977 and H. ripariusPeyerimhoff, 1929], Heterothops megalops has remarkably

large eyes and thus extremely short tempora (longitudinaldiameter of the eye nearly six times as long as tempora).Thus, it can be easily distinguished from all those species bythe shape of the head. Also, all the listed species have a more

or less straight paramere, but the paramere of Heterothopsmegalops is slightly curved ventrad.

Remarks on the generic position. Although, as explainedabove, the limits of the genusHeterothops on the world scaleneed correction, Heterothops megalops fits the core concept

of this genus, as it shares the diagnostic combination ofcharacters with its Holarctic representatives: maxillary palpswith the last segment very small, thin and sharp; pronotumwith two punctures in the dorsal rows; punctuate scutellum;

pronotal hypomera without a translucent postcoxal process;aedeagus with the long paramere firmly attached to thereduced median lobe.

Distribution and bionomics. Currently, the species isknown from three areas in Central and Southern Africa.

One is the type locality in eastern Angola, another twolocalities in Rwanda, and the third area, with most of therecent records, broadly covers the northeastern part of the

Republic of South Africa. Heterothops megalops has pre-sumably a broader distribution still unknown due to a lackof material. The fragmentary biotope data available fromthe labels suggest that this species lives in a variety of

habitats. Most of the available specimens were collected inforest leaf litter at low elevations. But, for example, twospecimens from Mpumalanga (Goedehoop Farm in South

Fig. 2. Morphology of some Afrotropical Staphylinini. A–C, Heterothops megalops, habitus (A), aedeagus (parameral side) (B), aedeagus

(lateral) (C); (D) Philonthus pseudoquedius, habitus.



Africa) were collected in litter under trees and shrubs ingrassland at an elevation of 1400 m. One specimen (from

near the Utsoek Forest Station) was found in horse dung.

Lectotype designation. In the original description,

Cameron (1959) indicated three syntypes. Of the twosyntypes from the NHM, a male specimen with the fullgeographical labels and Cameron’s label ‘type’ is designatedas a lectotype. The two other syntypes, one of which we

could not locate and the other (female) examined here,become paralectotypes. A circle label ‘paratype’ earlierattached to this syntype is incorrect. Lectotype designation

is provided to fix the identity of that species.

6. Euristus Fauvel, 1899. Type species: Euristus globus

Fauvel, 1899, by monotypy

Type material examined. Gabon:Holotype: male, ‘LoangoGabon/Euristus Fauvel/globus Fvl./R.I.Sc.N.B. 17.479 Coll.

et det. A. Fauvel/Type’ (IRSNB).

Additional material examined. Republic of Cameroon:

One male, ‘Cameroun/Coll. et det. A. Fauvel Euristus globusFauv. R.I.Sc.N.B. 17.479’ (IRSNB); one female, ‘KamerunRohd’/’Quediini’ Euristus globus Fvl. Det. A.F. Newton

1989’ (ZMHB); one male, Neu-Kamerun Sardi b. Dengdeng2.IV.1914 Mildbraed S.G. (ZMUC).

Redescription (Fig. 3). Measurements (four specimens)HL: 1.53–1.66; HW: 2.55; PL: 3.45–3.60; PW: 4.65–

4.95; EL: 4.20–4.73; EW: 4.58–4.80; total length of body12–15 mm.

Head capsule slightly oblong; neck constriction indicatedonly laterally; eyes well developed, slightly protruding overcontour of head, their longitudinal diameter equalling

length of tempora (Fig. 3A). Head capsule ridges in generalrather weakly indicated: postmandibular ridge weak, dis-tinct from base of mandible through under eyes, then

turning dorsad behind eyes and completely disappearing;nuchal ridge absent, but nuchal constriction present, indi-cated laterally only, entirely obliterated dorsally; infraorbi-

tal ridge absent, very slightly indicated laterally, confluentwith nuchal constriction ventrally, not extending anteriorly;postgenal ridge absent; ventral basal ridge slightly indicated;dorsal basal ridge absent; gula broad, parallel sided along

most of its length, without transverse basal impression.Submentum with smooth surface as gula, without setae orsetiferous punctures. Head capsule rugosely punctured and

setose except smooth apical portion of frons, gula and neckregion. Antennal insertions situated at anterolateral marginsof frons, anterior to eyes; distance between them only

slightly larger than distance from antennal insertion toeye. Antennae: first three antennomeres elongate and gla-brous, with long setae only; antennomere I as long asantennomeres II and III combined; antennomeres IV–VI

each longer than wide; VII–IX each slightly longer thanwide; antennomeres IV–VII distinctly moniliform. Mandi-bles (Fig. 3E, H) moderately long and curved, apically

narrow and sharp, with highly developed basolateral ridge,which is located distinctly on dorsal side of mandible; each

mandible internally with three strong teeth, rather symmet-rical; prostheca rather short, broadly attached to mandible.Labrum (Fig. 3H) large, transverse but relatively long,

deeply bilobed. Maxilla (Fig. 3B) with thick brush ofthinner setae on lacinia and sparser brush of stouter setaeon galea, the latter also having a dense cluster of darker andsmaller setae. Maxillary palp moderately long, with stout,

fusiform, setose apical segment. Labium (Fig. 3C, D)relatively short; mentum very short, with two pairs ofmacrosetae near anterior margin at sides; ligula with slight

incision apically forming two distinct glossae, its laterallobes (paraglossae) with dense stout setae dorsally; labialpalp with two short basal segments and relatively long and

broad, securiform, setose apical segment; subapical segmentwith numerous macrosetae at inner margin.Pronotum (Fig. 3A) roundish, slightly transverse, disc

evenly punctured leaving only middle longitudinal band

impunctate; anterior angles of pronotum significantly pro-jecting anteriad over apical margin of prosternum; pronotalhypomera (Fig. 3G) very wide, strongly inflexed inwards

(not visible in lateral view), without translucent postcoxalprocess but with short distinct postcoxal process sclerotizedsimilarly to rest of hypomeron (not delimited from rest of

hypomeron); superior marginal line well developed throughits whole length separating hypomeron from disc, notinflexed under anterior angles of pronotum; inferior mar-

ginal line also well developed through its whole lengthconverging with superior line over anterior corners ofpronotum. Prosternum (Fig. 3G) strongly transverse, firmlyjoined to pronotum (notosternal sutures distinct, but not

membranous); basisternum strongly protruding and cari-nate posteriad, without macrosetae, with scattered micro-setae; furcasternum very short and wide, without carina;

sternacostal ridge well developed. Mesosternum (Fig. 3F)transverse, firmly joined to metasternum (mesosternal–metasternal suture very weak through its whole length),

without medial carina, with very weak, very obtuse andbroad mesosternal process having few macrosetae; sterno-pleural sutures transverse. Elytra (Fig. 3A) moderately long

with rounded humerus covered by thick setae but not spines;sub-basal ridge immediately adjacent to elytral articulation,short, extending anteriad; deflexed epipleural part of elytranarrow, no epipleural ridge; lateral length of elytra exceeds

their length at suture, sutural angle obtuse; elytral surfacefinely and densely punctured, setose, setae along lateral sidesand apical margin of elytra longer. Mesothoracic scutellum

(Fig. 3J) with two transverse carinae in basal part. Meta-thorax (Fig. 3F) wider than long, but longer than mesotho-rax; mesocoxal acetabula (and mesocoxae themselves)

rather widely separated from each other. Wings well devel-oped. Legs moderately long, posterior legs longest; anteriortarsi in both sexes widened, with white adhesive setaeventrally; empodium with two setae on all tarsi. Abdomen

(Fig. 3A, I) elongate, conical (i.e. strongly narrowed api-cally and not flattened dorsoventrally), its entire surfaceevenly densely punctured and setose. Abdominal tergite I



(Fig. 3K) laterally with small roundish shallow indentations

and several setae nearby [apparently manifestations of eitherfunctioning or reduced protergal glands (osmeteria)]. Ter-gite II with basal part medially crossed by longitudinal

carina. Tergites III and IV with two pairs of paratergites.Tergites V and VI with only one pair of paratergites, eachparatergite of tergite V being basally divided into two closely

attached lobes. Tergites VII and VIII without paratergites(but lateral portions of sternite VII somewhat bent dorsad,appearing as narrow paratergites). Tergites III–VIII each

with pair of narrow spiracles basally at sides, with one basalcarina.Male. Sternite VIII apically with shallow median emar-

gination. Sternite IX oblong, with emarginate apex and

short, slightly asymmetrical basal portion. Lateral sclerites

of tergite IX relatively short, apically obtusely pointed,

dorsally widely separated by tergite X. Tergite X graduallyconverging apicad, with pointed apex (Fig. 3L, dashed line).Apical portions of abdominal segment VIII, sternite IX,

laterosclerites IX and tergite X with long macrosetae.Aedeagus (Fig. 3M, N) in lateral left position (parameralside faced left) in abdomen in repose; median lobe elongate

with broadly rounded (in dorsal or ventral view), somewhatshovel-like (in lateral view) apex; basal portion moderatelylarge, rather symmetrical; dorsal side of median lobe

(contrary to stronger sclerotized ventral side) relatively soft;paramere relatively small (by far not reaching apex ofmedian lobe, much narrower than median lobe), dorsoven-trally flattened, with obtuse, very slightly incised apex

covered by several large macrosetae, without sensory peg

Fig. 3. Morphology of Euristus globus. (A) Habitus; (B) maxilla (left, ventral view); (C) labium (ventral view); (D) labium (lateral view); (E)

mandible (right, dorsal view); (F) meso- and metathorax (ventral view, right leg removed); (G) prothorax (ventral view, legs removed); (H)

labrum and mandible (dorsal view, only left side fully illustrated); (I) abdomen, segments II–VII (lateral view); (J) scutellum (thoracic

mesoscutellum, dorsal); (K) abdominal tergite I (left side, with spiracle and cuticular manifestation of the protergal gland shown); (L) male

genital segment (ventral view); (M) aedeagus (parameral view); (N) aedeagus (lateral view).



setae; internal sac (as seen when inverted) with fields ofmicrostructures of stronger sclerotization.

Female. Sternite VIII apically simple, without emargina-tion. Ovipositor consisting of paired proximal and distalgonocoxites, the latter bearing styli.

Distribution and bionomics. Euristus globus is known onlyfrom the above-listed localities in western tropical Africa.No data have been recorded about the habitat requirements

of this species, or about the methods used to collect thespecimens.

Comparison. The relatively large size and limuloid shapeof the body, including the conical abdomen with only onepair of paratergites on the segments V and VI and without

paratergites on segments VII and VIII, allow this genusto be easily distinguished from any other genera ofStaphylinini.

Notes on type material. In the original description, Fauvel(1899) explicitly mentioned one type specimen (male fromLoango). Of the two specimens identified by him as Euristus

globus, and kept at IRSNB, the one from Loango and withFauvel’s mark ‘type’ (for details see ‘Material examined’section) is here marked as holotype.

Remarks on the systematic position. In the originaldescription of Euristus, Fauvel (1899) emphasized the

peculiar habitus of this genus and, without discussion ofsupporting characters, placed it in ‘Quedii’ (now Quediina)near the genus Euryporus. The general habitus and thesecuriform apical segment of maxillary palps in Euristus

probably led Fauvel to such a conclusion. The characterstates Euristus shares with Quedius [in the narrow sense,sensu Solodovnikov (2006)] or with ‘Quediina propria’ here

are the strongly deflexed hypomera and anteriorly pro-longed anterior corners of the pronotum. But Euristus doesnot have infraorbital ridges. Strongly autapomorphic fea-

tures of Euristus (reduced head capsule ridges and neck;conical abdomen with reduction of paratergites) make itreally difficult to infer sister relationships of this genus

among Staphylinini. The structure of the pronotum inEuristus, with the inferior marginal lines extending alongthe entire length of the pronotum over its anterior angles,affiliates this genus with the subtribe Xanthopygina. How-

ever, Xanthopygina is probably not a monophyletic group(Chatzimanolis, 2004; present study). The present cladisticanalysis places Euristus in the ‘Staphylinina’ lineage, corre-

sponding to the current subtribe Staphylinina.

7. Philonthus Stephens, 1829. Type species: Philonthus

splendens (Fabricius, 1793)With more than 1000 described species, Philonthus is one

of the largest genera of Staphylinini and Staphylinidae. Areview of the taxonomic history of this genus and its latest

diagnosis can be found in Smetana (1995b). On a worldbasis, the limits of Philonthus are unclear. With a narrowergeneric concept applied to Philonthus in the future, the size

of this genus may shrink (Newton et al., 2000). However, asfar as our current knowledge of the world material of

Staphylinini and Philonthina allows such a conclusion tobe made, the current concept of Philonthus, although notperfect, is far better than the currently accepted concept of

Quedius. At least Philonthus now includes species that aremuch more closely related to each other than the speciescurrently listed in Quedius (e.g. Herman, 2001). A closerexamination of two species of Quedius from sub-Saharan

Africa (see below) reveals that they match the current de-finition of Philonthus and must be transferred accordingly.

Philonthus pseudoquedius Solodovnikov nom.n. (Fig. 2D)Quedius cinctipennis Cameron, 1951: 406Philonthus cinctipennis (Cameron, 1951) comb.n. (junior

secondary homonym of Philonthus cinctipennis Fauvel, 1875)

Type material examined. Holotype: female, ‘‘type [redcircle label]/Mac Arthur & Museum Staff Kanzeko. 1-42/

Quedius cinctipennis Cameron Type/Pres by Com Inst Ent BM 1952-575/syntype [blue circle label]/Holotype Quediuscinctipennis Cameron, 1951. A. Solodovnikov rev. 2008 [red

label]/Philonthus pseudoquedius Solod. nom.n. A. Solodov-nikov det. 2008’’ (NHM).

Additional material examined. One female (Fig. 2D),‘‘Mac Arthur & Museum Staff Kanzeko. 1-42/Quediuscinctipennis Cameron Cotype/M. Cameron Bequest B.M.

1955-147./syntype [blue circle label]/paratype [yellowcircle label]/not a type A. Solodovnikov rev. 2008/Philonthus pseudoquedius Solod. A. Solodovnikov det.2008’’ (NHM).

Comments. In the original description of Quedius cincti-pennis (Cameron, 1951) there is a note ‘Type in B.M. [British

Museum]’, which, according to Article 73 of the ICZNmeans that there is only one type specimen (holotype) forthis species. There are two specimens at the NHM London

marked as various kinds of types by Cameron and sub-sequent curators (for details see above material examined).Both specimens are conspecific females with identical geo-

graphical labels matching the type locality data indicated inthe original description (‘Kenya Colony: Kanzeko; January,1942’). However, one female is marked by Cameron as‘type’, and the other as ‘cotype’. Naturally, we consider the

‘type’ specimen as the holotype, whereas the ‘cotype’ laterincorrectly marked as ‘syntype’ and ‘paratype’ does nothave a formal type status.

Quedius cinctipennis has no empodial setae [synapomor-phy of the subtribe Philonthina according to Schillhammer(1998) and Smetana & Davies (2000)]. Also, contrary to the

members of the genus Quedius, especially in its narrowersense [clade C in Solodovnikov (2006)] and ‘Quediinapropria’ here, but in agreement with the characteristicsof Philonthus and other Philonthina (Smetana, 1995b;

Smetana & Davies, 2000), it lacks infraorbital ridges, hasthe prosternum fused with the pronotum (no notosternalsutures), has the entire ligula, and it has no translucent



postcoxal process. Quedius cinctipennis is in fact a speciesof the genus Philonthus.

Apparently, Cameron’s misidentification was caused bythe superficial similarity of this species with the genusQuedius. Namely, Quedius cinctipennis has a smooth pro-

notal disc (without dorsal rows of punctures characteristicof most Philonthus species) and a Quedius-like structure ofthe pronotum with deflexed hypomera.The new combination Philonthus cinctipennis (Cameron,

1951) is preoccupied by Philonthus cinctipennis Fauvel,1875. Therefore, it is replaced by the new name Philonthuspseudoquedius Solodovnikov. The new name refers to the

misleading Quedius-like appearance of this species, and isformed as noun in apposition.

Philonthus angularis (Cameron, 1948) comb.n.

Quedius (Sauridus) angularis Cameron, 1948: 41

Comments. The original description of Quedius angularisdoes not contain exact data about the type material

(Cameron, 1948). The only available information is the typelocality: ‘West of Fort Archambault, Sara-Dai (Type). Rivesdu Moyen Chari, Mont des Niellims’. We were unable to

locate the type material at the NHM London or elsewhere.In the course of several years, Solodovnikov has examinedextensively large collections of Staphylininae from Africa in

several museums of the world, all of which containednumerous Philonthus specimens (representing more thanone species), which more or less match the habitus outlinedin the original description of Quedius angularis. At the same

time, no Quedius material from the Afrotropical region hasever been found. Lack of Quedius there seems to be a bio-geographical fact (see discussion below). Although not

being able to examine the type material, based on theoriginal description and indirect data, we are confident thatQuedius angularis, similar to Quedius cinctipennis, is another

misidentified species of Philonthus, and thus we proposea respective new combination.It is important to stress the fact that the genus Quedius is

not just poorly represented in the Afrotropical region as itappeared before, but it does not occur there at all. Given therevealed polyphyly of the current ‘Quedius’ (Solodovnikov,2006; present study), and the emerging new concept instead

[primarily north temperate ‘true’ (type bearing) Quedius vsnon-related south temperate ‘Quedius’, now members of the‘Tanygnathinina sensu novo’], the lack of ‘Quedius’ in the

Afrotropical region is a noteworthy biogeographical factthat needs close attention.

8. Glyphesthus Kraatz, 1858: 364. Type species: Glyph-esthus picipennis Roth, 1851 (¼G. rufipennis Kraatz, 1858)Since the original description (Kraatz, 1858), this African

genus (12 species) was affiliated with Quediini, untilSmetana & Davies (2000: 15) briefly stated that Glyphesthusbelongs to Philonthina. This taxonomic solution is followedin the catalogue of Herman (2001). The only character states

that Glyphesthus shares with Quedius and Quediina arerelated to the shape of the prothorax with deflected hypo-mera and anterior angles of pronotum protruding over the

anterior margin of the prosternum. At the same time,Glyphesthus lacks infraorbital ridges, it has well-developed

postmandibular ridges, its pronotum is fused with theprosternum (but weak notosternal sutures are still visible),and it has no empodial setae. Obviously it is not related to

Quedius and its placement in the subtribe Philonthina issupported by the present analysis.

9. Ioma Blackwelder, 1952 (replacement name for

Tachinopsis Fauvel, 1899). Type species: Ioma setigera(Fauvel, 1899), by monotypy.Tachinopsis setigera: Scheerpeltz, 1974: 25.

Type material examined. Syntypes: One female, ‘Bogos1870 Sciotel, O. Beccari/Khayes, Mt. Senegal/Coll. Et. det.

A. Fauvel Tachinopsis setigera Fauv. R.I.Sc.N.B. 17.479/Syntype’; one female, ‘Bogos 1870 Sciotel, O. Beccari/Tachinopsis Fauvel./setigera Fvl./R.I.Sc.N.B. 17.479 Coll.Et det. A. Fauvel/Syntype’ (IRSNB).

Other material examined. Federal Republic of Nigeria:

Two females, Kano Distr. N. Nigeria June 1954, W.E.S.

Merrett; one male, same data but May 1953; one female,same data but July 1953 (MMUE); one male, Zaria, light,3.IV.1933 (NHM); Republic of Cameroon: one male,

Kamerun int. Bg Sidderi 30.VII.09 Riggenbach S.G.(ZMHB); Republic of Chad: one female, N.E. de FortArchambault, Mara-Goulfez Mission Chari-Tchad, D.J.

Decorse 1904, septembre (NHM); Republic of Niger: onefemale, ‘. . .der’ [illegible] Niger, 15.VI.1981 Paul Bouchard(ZMUC); two females, ‘Agadez, Air sud. 525 m, 1.VIII/IFAN-1947 L. Chopard A. Villers (NHM); one female, Mt.

Baguezans 1500–1600 m, 31.VIII-4.IX/IFAN-1947 L. Cho-pard A. Villers (NHM); Republic of Senegal: two males,Senegal (FMNH, ZMHB); Republic of Sudan: two females,

Sudan Govt. Kurgul, R.S.M. Darling, 3.II.31, on light(FMNH); two females, Sudan, Prov. N. Darfur, El Geneina,18.VI.1979 leg. I.W. Abuzinid (ZMHB).

Redescription (Fig. 4). Measurements (12 specimens)HL: 0.70–0.76 (0.74); HW: 1.14–1.22 (1.18); PL: 1.11–1.20

(1.15); PW: 1.62–1.90 (1.80); EL: 1.27–1.35 (1.32): EW:1.79–1.99 (1.91); total length of body: 5.9–6.3 (6.0) mm.Head capsule transverse, with large eyes; tempora

strongly converging basad; relatively narrow neck very

distinct laterally but without constriction dorsally; eyesnot significantly protruding over head contour, their ante-rior margins adjacent to base of mandibles (Fig. 4A). Head

capsule ridges as following: postmandibular ridge welldeveloped, extending from base of mandible along ventralborder of eye to lateral side of head capsule; infraorbital

ridge absent; postgenal and ventral basal ridge well devel-oped, not confluent with each other; nuchal and dorsal basalridge absent. Gular sutures widely separated from eachother, slightly converging posteriad; gula with slight trans-

verse basal impression. Submentum with pair of macro-setae. Antennal insertions at anterolateral margins of frons,distance between them much larger than distance from



insertion to eye. Antennae relatively short, with antenno-meres IV–XI strongly flattened dorsoventrally, antenno-meres IV–X with somewhat attenuate apical corners;

antennomeres I–III glabrous, with sparse macrosetae; an-tennomere III slightly shorter than IV. Mandibles (Fig. 4C)moderately large, with sharp apex, slightly curved, with one

tooth internally; with well-developed dorsolateral ridgesituated dorsally on mandible; both mandibles rather sym-metrical; prostheca broadly attached to mandible, with

feather-like setae. Labrum (Fig. 4E) large, transverse,deeply bilobed. Maxilla with thick brushes of setae onlacinia and galea, the latter having a setal brush in apical

portion; maxillary palpi moderately long, with apical seg-ment fusiform, glabrous. Labium (Fig. 4B) with broadmentum having a pair of macrosetae at sides; ligula small,entire; labial palpi moderately long, with glabrous, fusiform

apical segment.

Pronotum (Fig. 4A) transverse, with rounded anteriorand posterior angles, widest behind its middle, with sidesstrongly converging anteriad, with several very large macro-

setae along lateral margins; disc smooth, with three smallermacrosetae in each of two dorsal rows, and a few setaelateral to dorsal rows; pronotal hypomera (Fig. 4F) very

wide, inflected inwards (not visible in lateral view), withoutpostcoxal translucent process; anterior angles of pronotumprojecting anteriad over apical margin of prosternum.

Pronotum strongly fused with prosternum, notosternalsutures indistinct. Prosternum (Fig. 4F) transverse; basi-sternum smooth, without carina, with two small macro-

setae; sternacostal ridge well developed, slightly producedbasad in its middle. Mesosternum (Fig. 4H) transverse,without medial carina, with few irregularly located macro-setae, strongly fused with metasternum, mesosternal–

metasternal suture indistinct. Elytra (Fig. 4A) moderately

Fig. 4. Morphology of Ioma setigera. (A) Habitus; (B) labium (ventral view); (C) mandible (right, dorsal view); (D) maxilla (left, ventral view);

(E) labrum (dorsal view); (F) prothorax (ventral view, legs removed); (G) male genital segment (ventral view); (H) meso- and metathorax

(ventral view, left leg removed); (I) abdominal tergite I (left side, with spiracle and cuticular manifestation of the protergal gland shown); (J)

scutellum (thoracic mesoscutellum, dorsal); (K) aedeagus (parameral side); (L) aedeagus (lateral view).



long, their lateral length only slightly exceeding length alongsuture, with rounded humerus, with sub-basal ridge imme-

diately adjacent to elytral articulation and extended ante-riad, with narrow deflexed epipleural area, withoutepipleural ridge; elytral surface with sparse fine punctation

and short setae, apical margin and especially lateral portionsof elytra with large macrosetae, without spines on humeri.Mesothoracic scutellum (Fig. 4J) with one transverse carinain basal part. Metathorax (Fig. 4H) slightly transverse,

longer than mesothorax; mesocoxal acetabula (and meso-coxae themselves) separated from each other. Wings welldeveloped. Legs moderately long, with rather wide coxae

and femora, anterior legs shortest, posterior legs longest.Anterior tarsi with tarsomeres I–IV widened in both sexes,apical tarsomere about as long as tarsomeres I–IV, which

have long pale adhesive setae on ventral side. Middle andposterior tarsi not enlarged, without pale adhesive setaeventrally, their apical tarsomere about as long as firsttarsomere. All tarsi without empodial setae, with moder-

ately long claws. Anterior tibia with tibial comb near apex;anterior and middle tibiae with few spines externally,posterior tibia with spines only at apex.

Abdomen (Fig. 4A) normally elongate and dorsoven-trally flattened, widest at base, gradually converging api-cally; tergites and sternites sparsely finely punctate and

setose, with large macrosetae at apical and lateral marginsof sclerites. Tergite I with pair of well-developed protergalglands (osmeteria) whose cuticular manifestation is a round-

ish chamber with narrow opening posteriorly (Fig. 4I); thesestructures covered externally by several macrosetae. TergiteII without longitudinal internal carina in middle of basalpart (usually seen through the wall of tergite); tergites III–VI

with two pairs of paratergites, of about equal length; tergiteVII also with two pairs of paratergites, but medial para-tergites much shorter than lateral ones; tergites III–VIII

with one basal carina and a pair of round spiracles situatedlaterally rather distant from base of tergite.Male. Sternite VIII with deep and wide medial emargi-

nation at apical margin. Sternite IX elongate, symmetrical,with emarginated apex and very short basal lobe. Lateraltergal sclerites IX relatively narrow, dorsally broadly sepa-

rated by tergite X. Tergite X gradually converging apically,with broadly rounded apex (Fig. 4G). Apical part of sterniteVIII, as well as apex and external sides of lateral tergalsclerites IX with large macrosetae. Aedeagus (Fig. 4K, L)

elongate; median lobe symmetrical, only its basal portionslightly asymmetrical, its apex narrowly protruding as bluntlobe (in ventral or dorsal view); parameres fused in one

piece, the resulting one paramere asymmetrical, with elon-gate right lobe and very short, barely indicated left lobe,each lobe with two to three small sensory peg setae at its

apex; internal sac (as seen when inverted, through the wallsof median lobe) without significantly stronger sclerotizedstructures. Aedeagus in dorsal position (parameral sidefaced dorsally) in abdomen in repose.

Female. Ovipositor consisting of paired proximal anddistal gonocoxites, the latter bearing styli. Unlike in males,sternite VIII with simple (non-emarginate) apical margin.

Anterior tarsi, as in males, with slightly expanded tarso-meres I–IV.

Distribution and bionomics. Ioma setigera is broadlydistributed in Eastern and Central Africa, north of the

equator. Nothing is recorded about the habitat require-ments of this species. At least two of the examined specimenswere collected when attracted to light. Fauvel (1899) pro-posed that Ioma could be myrmecophilous, which is prob-

ably true given the somewhat limuloid shape of the body inthis species. An additional two specimens of Ioma setigerafrom northeastern Africa, not examined here, were listed

by Scheerpeltz (1974: two females from ‘Sudan, UpperNile, Malakal, 5–20.I.1963, Linnavuiori’ and ‘Sudan,Darfur, Safaha-AbuMatriq, 30.IV.–2.V.1963, Linnavuiori’,

respectively).

Comments on the type material. In the original descrip-tion of Tachinopsis setigera, Fauvel (1899) did not specify

anything about the type material he had under his study.However, from the information about the collecting local-ities he provided [‘Abyssinie: Bogos, Sciotel (O. Beccari);

Haut-Senegal: Kayes (Dr. Nodier)], it is clear that he had atleast two or more syntypes from different localities collectedby different people. Among the material examined (see

above), we were able to find only two syntypes from ‘Bogos,Sciotel’ collected by Beccari and kept in IRSNB. Both ofthese syntypes are females. The lectotype designation is here

refrained from, in the hope that a male specimen may existamong other potential syntypes from ‘Kayes’ collected byNodier, which may be located somewhere in the future.

Comparison. A very characteristic limuloid habitus, themesothoracic scutellum having only one carina and the tarsilacking empodial setae, make Ioma setigera very easily

distinguishable from any other genera of Staphylinini,especially on the African continent.

Remarks on the taxonomic position. In the originaldescription of the genus (as Tachinopsis), Fauvel (1899)mentioned that it was close to Quedius. Without a doubt,

such a decision was made based on habitus similarity only.On the basis of such character states as: no empodial setaeon all tarsi, entire ligula, absent infraorbital ridges, hypo-mera of pronotum meeting prosternum at an obtuse angle,

both sclerites being fused and notosternal suture thereforeabsent, Ioma can be affiliated with Philonthina. Unlike mostof the Philonthina though, Ioma, in addition to its Quedius-

like structure of the pronotum, has a very long postman-dibular ridge (as in many Xanthopygina), an obliteratenuchal ridge, and it does not have a dorsal basal ridge (as

in ‘Quediina propria’ and some other lineages of Staph-ylinini). Interestingly, Ioma has a dorsal position of theaedeagus (parameral side faced dorsally) in abdomen inrepose, a character state highly unusual for Staphylinini.

And similarly to the ‘Tanygnathinina sensu novo’ lineage, ithas only one carina on the mesothoracic scutellum. Overallcladistic assessment of character states throughout the entire



external morphology does not support the affinity of Iomasetigera to ‘Quediina propria’, but places it in the ‘Philon-

thina propria’ lineage, as a sister group to the Moeocerus þGlyphesthus þ Hasumius clade.

10. Moeocerus Fauvel, 1899. Type species: Moeocerusmimus (Fauvel, 1899)Fauvel (1899) characterized this genus (described as

Homoeocerus) as closely related to Glyphesthus (i.e. affiliated

to Quediini at that time). Such placement continued later on(e.g. Bernhauer & Schubert, 1916; Gridelli, 1939), untilSmetana & Davies (2000: 15) briefly mentioned that this

genus in fact belongs to Philonthina. This point of view wasfixed in the most recent catalogue (Herman, 2001). As in theabove-mentioned cases of Glyphesthus and Ioma, the only

character states that Moeocerus shares with ‘Quediina prop-ria’ are those related to the shape of the pronotum withdeflexed hypomera. However, unlike ‘Quediina propria’, thepronotum in Moeocerus is fused with the prosternum (no-

tosternal sutures hardly distinct). Also, Moeocerus has noinfraorbital ridges and no empodial setae. The presentanalysis supports close phylogenetic relationships of Moeo-

cerus with Glyphesthus and Hasumius, and justifies its recentmove (Smetana & Davies, 2000) from Quediina to Philon-thina. Morphologically, strictly African Moeocerus (14 spe-

cies, including new combination) is somewhat similar to theOriental genus Agacerus Fauvel, 1895 and the Neotropicalgenus Flohria Sharp, 1884 (both now in Philonthina); these

presumably sister relationships must be further expored.

Moeocerus wendeleri Solodovnikov nom.n.

Moeocerus robustus (Wendeler, 1928) comb.n. [juniorsecondary homonym ofMoeocerus robustus (Gestro, 1881)].

Algon robustus: Wendeler, 1928: 35.

Comments on the type material and new

combination. Apart from the type locality [Sud-Afrika,Kapland] and the fact that the material was retained byAlexander Heyne, the original description does not specify

any information about the types of Algon robustus. We wereunable to locate this type material; such a situation seems tobe not unusual as Alexander Heyne was an insect dealer

whose material is now very difficult or impossible to track(M. Uhlig, personal communication). Although the descrip-tion of Algon robustus is not detailed enough by modernstandards, we positively identify this species as a member of

the genus Moeocerus based on the following evidence. Theoriginal description of Algon robustus is of a glossy blackspecies of 14–15 mm long with red elytra, tarsi, and apical

segment of the maxillary palps, and partially reddishantennae. Such general habitus, along with some othercharacters indicated in the descripton, largely conflict with

the description of Algon (Schillhammer, 2006), but matchthose of Moeocerus. Also, as pointed out below, the genusAlgon seems to be strictly Asian. Other possible misidenti-fications of this species can also be ruled out based on the

relatively good state of knowledge of the South Africanfauna of Staphylininae, including Solodovnikov’s survey ofseveral large world collections particularly strong for this

region. The name Moeocerus robustus (Wendeler, 1928) ispreoccupied by M. robustus (Gestro, 1881), so the new

patronymic name, M. wendeleri, is here proposed for thisspecies.

11. Algon Sharp, 1874. Type species: Algon grandicollis

Sharp, 1874As summarized in Schillhammer (2006), the systematic

position of Algon at the subtribal level is swinging between

Quediina and Xanthopygina and still awaiting a satisfactorysolution. Placement of Algon in either of these subtribes isnot supported in Solodovnikov (2006) and in the present

analysis. Moreover, these analyses support monophyly forneither Quediina nor Xanthopygina. Sister relationships ofAlgonare still tobe found.Currently,only two species ofAlgonwere recorded from the Afrotropical region: A. africanus

Bernhauer, 1915 and A. robustus Wendeler, 1928. Schillham-mer (2006), on the basis of the examination ofAlgon africanus,stated that this species was not congeneric with the Asian

species ofAlgon. Our examination ofAlgon africanus (severalcongeneric but non-conspecific specimens identified as ‘Algonafricanus’ by Bernhauer and his contemporaries) confirms this

generic misidentification and suggests that this group ofspecies belongs to a new genus of Staphylinini, phylogeneti-cally remote from Algon. Its description is postponed untila proper comparative study is completed. The misidentifica-

tion of Algon robustus and its proper position in the genusMoeocerus is discussed above. So, the genus Algon, confinedto the eastern Palearctic and Oriental regions, does not occur

in Africa.

Discussion and conclusions

This paper is an example of how poor the basic taxonomicknowledge is for the tribe Staphylinini in the areas of theglobe outside the better-studied Holarctic region. Also, it

shows that current systematics of Staphylinini can be verymisleading if used as a reference for any broader-scaleevolutionary or biogeographical inference. Before recent

studies (Solodovnikov, 2005, 2006; Solodovnikov & Newton,2005; present article), Afrotropical ‘Quediina’ consisted ofdrastically different lineages. Of them, genera endemic tosub-Saharan Africa bear important biogeographical infor-

mation, which, however, could not be properly read beforeclarifying their taxonomy and systematic position withinStaphylinini. Although we can formally fix all taxonomic

changes at the genus and species levels here, this cannot bedone at the supra-generic level. Monophyly of the newlydiscovered lineages of Staphylinini must be thoroughly

tested further on, before we are able to suggest new group-ings and confidently move genera among them. As a tem-porary solution, we use informal names for the newlyoutlined groups, all of them presented in Fig. 1 and Table 1.

These names are derived from the oldest valid family-groupnames applicable for the respective clades, and are given inquotation marks to stress their informal status.



The present analysis reveals a phylogenetic pattern for thetribe Staphylinini where most of the former quediine genera

are divided between two large sister clades. One of them isthe north temperate ‘Quediina propria’ (so-called because itcontains the type species the type genus of the family-group

name Quediina Kraatz, 1857). The other is south temperate‘Tanygnathinina sensu novo’ (so called because it includesthe genus Atanygnathus, the type of the family-group nameTanygnathinina Reitter, 1909, now applied in a new sense).

Several other former ‘Quediina’ appear as basal lineages ofStaphylinini, which are not provided with any tentativesupra-generic names. The biologically very specialized sub-

tribe Amblyopinina appears nested within the ‘Tanygnathi-nina sensu novo’ clade. (The family-group nameAmblyopinina Seevers, 1944 is younger than Tanygnathini-

na Reitter, 1909, so the informal name of the clade is derivedfrom the latter name). Of the other existing subtribes ofStaphylinini, only Staphylinina and Philonthina stand asmonophyletic, the former including the subtribe Hyptiomi-

na. Overall, the more inclusive clade, here called ‘Philonthi-na lineage’, is very problematic. In particular, it includesgenera of Anisolinina and Xanthopygina, none of these

groups coming as monophyletic. (The informal, respec-tively, more and less inclusive names ‘Philonthina lineage’and ‘Philonthina propria’ are derived from the oldest

family-group name Philonthina Kirby, 1837, the type ofwhich is nested in both of these hierarchical clades).The taxon sample analysed in the present study is indeed

too small for any immediate changes of the system, whichalso was not the purpose of this limited paper. However,some of them are consistent with the views already expresseda number of times in the systematic literature. For example,

Coiffait (1978) recognized the affinity of the genera Hetero-thops and Atanygnathus, proposing a new supra-generictaxon ‘Heterothopsi’ (incorrectly applied family-group

name) for these two genera. Newton (1985) expresseddoubts that the south temperate Quediina are closely relatedto the northern ones. The position of the generaMoeocerus,

Glyphesthus and Hasumius in the sister group to the cladeformed by genera now in the subtribe Philonthina isconsistent with the recent suggestion (Smetana & Davies,

2000) to place the former African genera in Philonthina,which is partially fixed in Herman (2001) and Newton &Thayer (2005). Problems with the systematic position ofAlgon and the polyphyly of Xanthopygina were emphasized

by Schillhammer (2006) and Chatzimanolis (2004), respec-tively. The position of the amblyopinines nested within the‘Tanygnathinina sensu novo’ clade disagrees with the earlier

views of Seevers (1955) on the evolution of this group, butbiogeographically seems to be more simple and plausiblethan his explanations involving assumptions of dispersal

events and extinctions. Newly proposed placement of thespecialized subcortical genus Holisus (subtribe Hyptiomina)within the core of Philonthina does not look so surprisingwhen considering other representatives of Philonthina with

more or less flat bodies and possibly dead-wood related oreven subcortical habitats (some species of Belonuchus,genera Scelotrichus and Sternotoxus). It is noteworthy that

until fairly recently Holisus was affiliated with the wrongsubfamily, merely due to its flat shape (Newton, 1988). The

position of this genus as a sister group to Atanygnathus inSolodovnikov & Newton (2005) is probably an artefact ofthe long branch attraction phenomenon (Bergsten, 2005)

due to a high number of autapomorphies in both genera andlow taxon sampling in that analysis, designed for differentpurposes.The phylogenetic analysis performed demonstrates that

a somewhat similar habitus of the Afrotropical endemicsIoma and Euristus is due to convergence, possibly because oftheir presumed inquiline lifestyle. The placement of the

genus Natalignathus here is consistent with earlier expressedcomparative considerations (Solodovnikov, 2005) and anal-ysis (Solodovnikov, 2006). The phylogenetic position of the

genus Afroquedius remains ambiguous.An overall problem of the present analysis is the low

support for most of the clades. It is difficult to address thisissue in a brief format, but apparently the low taxon sample

and, even more so, unclear (note numerous ‘?’ in the datamatrix, Table 2) or inadequately defined homologies formany characters are responsible for such a result. For

example, many characters used here (e.g. shape of themaxillary and labial palps) stemming from the earliersystematics of the group need more precise delimitation,

which is difficult to achieve so far. As our tree explorationsshow, they are very homoplastic and probably should beabandoned or at least redefined.

One of the tests for a newly emerging phylogenetic patternof Staphylinini is biogeography. For some of the hereconsidered Afrotropical endemic genera we tried to explorewhether their position in the phylogeny in Fig. 1 can be

reasonably explained in terms of historical biogeography.Africa rifted off the Gondwana before 135 Mya (Keast,1996). According to Smith et al. (1994), rifting between

Africa þ South America and the remainder of Gondwanawas already almost completed in the late Jurassic (Tithonian,148 Mya), leaving Madagascar as a part of India and

isolated from Africa. The final separation between SouthAmerica and Africa began in the early Cretaceous and wascompleted in the Albian (Briggs, 1995, after numerous

authors). To the north, Africa was separated from theLaurasian landmasses for a long time by the Eocene, andin the early Miocene it ultimately contacted with Asia viathe Arabian Peninsula (Briggs, 1995). Assuming that the age

of the basal nodes of Staphylinini phylogeny may corre-spond to the early Cretaceous or even late Jurassic (fordetails see above ‘Material and methods: biogeography’

section), the diversification of the clade ‘Tanygnathininasensu novo’ can be associated with the break-up of Gond-wana. If so, the position of Afroquedius as a member of this

clade [contrary to its very basal position in Solodovnikov(2006)] is easier to justify in a vicariance-based biogeograph-ical framework. In the same framework, however, theterminal position of the monotypic genus Natalignathus

within the same clade needs additional assumptions (fordetails, see the section about this genus above). A biogeo-graphically interesting observation coming from the new



Table

2.Data

matrix

forthephylogenetic

analysis.Forthefullnames

ofalltaxa,andexem

plary

speciessampled,seeTable

1.

11111111112

2222222223

3333333334

4444444445

5555555556

6666666667

777777

1234567890

1234567890

1234567890

1234567890

1234567890

1234567890

1234567890

123456

Arrowinus(outgroup)

0200???0?0

0??010??00

0000012002

0101011011

0010100000

00??110000

1000000000

000000

Platyprosopus(outgroup)?1110??120

0??000??20

0220012101

1001210000

00?0100000

00000?1223

2000200000

000000

Acylophorus

10110??111

???010??02

1020012110

?000212111

0100000002

0010110222

1101111011

000001

Afroquedius

02110??111

10?0010001

1020012022

0101201111

0110100001

0100112200

0111100010

000202

Algon

0111???101

10?1110030

1200001022

0100200011

0110100001

0110110222

1011100011

000002

Anchocerus

1111???101

???010??02

2020012112

0000210111

0100000000

0010110222

1101011011

00000?

Anisolinus

01110??101

10?1110000

1011000121

0100200111

0100100001

0110120222

1011100011

111002

Astrapeus

01110??111

0??010??32

1200012010

?100110011

0110100001

0000110222

1011100010

000002

Atanygnathus

0511120101

??0010??70

2520012110

?01?200110

0000000000

1011111002

1100111110

000121

Cheilocolpus

01110??111

10?0110010

1020011120

?00?200110

0010100021

0010111001

1111100010

000112

Creophilus

04110??131

0??1110000

1002300110

?000200100

1110100001

00000?0111

1010100010

000002

Dinothenarus

02110??101

0?01110000

0011100012

1100200100

1110100001

00000?0222

1011100011

111002

Edrabius

01110??111

0??0010010

0300012120

?11?210111

0010100001

00??113222

1011100011

000112

Elm

as

02110??121

10?1110030

1210200120

?100200100

1110100001

0110110222

1010100011

000002

Eucibdelus

03110??121

0??1010000

1011110112

0000200100

1110100001

0000100222

1011100010

000022

Euristus

01010??0?1

10?0110060

0210212113

0000200110

1110100001

0000100222

1010100010

000002

Euryporus

01110??111

???000??32

1220012013

0100210111

0100000001

0000102222

1001000011

000002

Glyphesthus

01010??0?1

11?1110050

0230211123

0100200000

111110001?

0110120002

1010000011

000002

Hasumius

01110??111

0??1110050

0220011120

?000200100

111110001?

0110120000

1010100011

000002

Hem

iquedius

01110??111

???000??02

1020011010

?100110111

0110100001

0000112222

1001100011

000002

Hesperus

0111???121

1011110040

1431100120

?100200100

111010001?

0110120012

1011100010

000002

Heterothopsbin.

0111???111

10?010??10

1320012110

?01?200110

0010100021

0010111002

1011100010

000111

Heterothopsdol.

0111???111

10?0010010

1320012013

1001200110

001000001?

0010103222

1111100010

000002

Holisus

01110??0?1

0??1011010

2331100120

?11?210101

101000001?

0011110022

100011002?

00???0

Indoquedius

?111???111

???0011000

1020011011

100?210111

0110100001

0000102222

1111100011

000002

Ioma

01110??0?1

11?0110000

1020012120

?100200000

101010001?

0110123222

1010100011

001001

Loncovilius

0110???111

10?010??00

1020011010

?01?200110

0010110001

0010103222

1110100011

000112

Moeocerus

02010??0?1

11?1110050

1220012120

?001200011

011110001?

0110120002

1011100010

000002

Natalignathus

0111???111

???0111070

2520012110

?00?200110

0010000001

1010113002

1011111110

100121

Nordus

03110??101

11?1110000

1030200120

?000200100

1110100001

0110110002

1011100010

000002

Parisanopus

01110??111

10?0110000

1020012010

?000100111

0111110101

0000110222

1111100010

000002

Philonthus

01110??121

10?1110040

1421001120

?100200100

111010001?

0110120000

1011100011

100002

Platydracus

02110??101

10?1110000

0011100011

1100200000

1110100000

00000?0222

1011100010

000002

Quediocafus

0111???1?1

10?0110000

1020012013

110?200101

001011001?

0010111222

1011100010

000112

Quediomim

us

01110??111

0??0110000

1020012120

?10?210110

0010101021

0011111000

1011100011

000111

Quediuscal.

01100??111

???010??00

1020012010

?00?200110

0010100021

0010113022

1111100010

000111

Quedius(s.str.)ful.

01111?11?1

1000110000

1020012012

1100210111

0110100001

0000102222

1011100011

000002

Quedius(Cyrt.)

lab

02110??111

???0110000

1000012010

?100110111

0110100001

0000110222

1011100010

000012

Quedius(Quedion.)

pl.

0111100101

1000010000

1020011012

1100110111

0110100001

0000110222

1011100011

000002

Quedius(Raph.)

umb.

01111?11?1

??00011100

1020012013

0100210111

0110100001

0000112222

1011100011

000002

Scelotrichus

01110??0?1

0??1110040

2421100120

?000200102

111011011?

0110120022

1110100010

000002



data in this paper is the complete absence of the northtemperate Quedius in the Afrotropical region, which do not

occur south of the Sahara [according to Smetana (2004), 29Palearctic species of Quedius are known from Africa northof the Sahara]. Explanations for that should perhaps be

sought in the assumption that there were never suitablebiotic conditions providing ‘landscape corridors’ for a dis-persal of the north temperateQuedius south into the Africancontinent. Finally, the striking poverty of the ‘Tanygnathi-

nina sensu novo’ in sub-Saharan Africa is also noteworthy.Atanygnathus, Natalignathus and Afroquedius (the lattergenus being attributed to this group with much less confi-

dence) are the only members of this overall species-richlineage here. Presumably, major diversification events of the‘Tanygnathinina sensu novo’ took place in the period when

Africa was already separated from the rest of Gondwana.

Acknowledgements

We are very obliged to the colleagues from several institu-tions around the world, listed in the ‘Materials and methods’section, who provided material for our study. All of them

were very hospitable, friendly and helpful during the visits ofA. Solodovnikov to study the collections under their care onsite. Some part of the research, which this paper is based on,

was carried out by A. Solodovnikov as a US NSF PEET-based (grant DEB-0118749) postdoctoral fellow at the FieldMuseum of Natural History at Chicago. Field Museum-

based colleagues A. Newton and M. Thayer inspired andlargely facilitated the research programme this paper is partof. Finally, the paper was continued and finished at theEntomology Department of the Zoological Museum at

Copenhagen, where we would like to acknowledge ourcolleagues for the very stimulating and accommodatingwork environment. We thank Stylianos Chatzimanolis

(University of Tennessee at Chattanooga, U.S.A.) for thediscussion about the identity and status of some African‘Xanthopygina’ and helpful review of the manuscript.

Harald Schillhammer (Natural History Museum of Vienna,Austria) also provided a number of suggestions on variousaspects of the study of Staphylinini, and on this project in

particular during A. Solodovnikov’s visit to Vienna museumjust before this paper was finalized. Critical review of thispaper by M. Thayer helped to improve its quality. Financialsupport for the research in this paper, including the intern-

ship for A. Schomann, came from the US NSF grantDEB-0715705.

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11111111112

2222222223

3333333334

4444444445

5555555556

6666666667

777777

1234567890

1234567890

1234567890

1234567890

1234567890

1234567890

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Accepted 8 December 2008

First published online 23 March 2009



Documents

Revised systematics and biogeography of ‘Quediina’ of sub-Saharan Africa: new phylogenetic insights into the rove beetle tribe Staphylinini (Coleoptera: Staphylinidae)