Revision of the Pseudomorphinae of the Australian region. 2. The genera Pseudomorpha Kirby, Adelotopus Hope, Cainogenion Notman, Paussotropus Waterhouse, and Cryptocephalomorpha Ritsema. Taxonomy, phylogeny, zoogeography. (Insecta, Coleoptera, Carabidae)

508 Pages·1997·224.1 MB·

by  Martin Baehr

Revision of Ihe Pseudomorphinae of the Australian Region
2. The genera PseiidoinorpjJta Kirby, Aäelotopus Hope,
Caiuogenion Notman, Paiissotropus Waterhouse,
and Cn/ptoceplialottiorpha Ritsema.
Taxonomy, phylogeny,zoogeography
(Insecta, Coleoptera, Carabidae)
Martin Baehr
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2. Thegenera Pseudomorpha Kirby, Adelotopus Hope, Cainogenion Notman,
PaussotropusWaterhouseand Cryptocephalomorpha Ritsema
:
taxonomy, phylogeny, zoogeography (Insecta, Coleoptera, Carabidae)
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SPIXIANA
Zeitschrift für Zoologie
Supplement 23
Revision of the Pseudomorphinae
of the Australian Region
2. The genera Pseudomorpha Kirby, Adelotopus Hope,
Cainogenion Notman, Paussotropus Waterhouse,
and Cryptocephalomorpha Ritsema.
Taxonomy, phylogeny, zoogeography
(Insecta, Coleoptera, Carabidae)
Martin Baehr
Gedruckt mit Unterstützung derDeutschen Forschungsgemeinschaft
Verlag Dr. Friedrich Pfeil • München, Ol. April 1997 • ISSN 0177-7424 • ISBN 3-931516-20-2
Ädelotopiis rufozoimfiis, spec. nov. from near EdithCreek, far Northern Territory.

SPIXIANA Supplement23 1-508 München, Ol. 04. 1997 ISSN 0341-8391 ISBN 3-931516-20-2
Revision of the Pseudomorphinae of the Australian Region
2. The genera Pseudotnorpha Kirby, Adelotopus Hope,
Cainogenion Notman, Patissotropus Waterhouse,
and Cryptocephalomorpha Ritsema.
Taxonomy, phylogeny, zoogeography
(Insecta, Coleoptera, Carabidae)*
Martin Baehr
Baehr, M. (1997): Revision of the Pseudomorphinae of the AustraUan Region
2.The genera Pseudomorpha Kirby, Adelotopnis Hope, Cainogenion Notman, Paus-
sotropms Waterhouse, and Cryptocephalomorpha Ritsema. Taxonomy, phylogeny,
zoogeography (Insecta, Coleoptera,Carabidae).-SpixianaSuppl. 23: 1-508
As the second part of a general revision of the Pseudomorphinae of the
Australian, Oriental, and Aethiopian regions the genera Pseudoiuorplm Kirby,
AdelotopiisHope,CainogenionNotman,PaussotropnsWaterhouse,andCri/ptocepha-
lomorplia Ritsema arerevised.
The 117valid speciesand additional 9 subspeciesofAdelotopns aregrouped in
26species-groupseachofwhichconfidentlyrepresentsamonophyleticunit. Keys
areprovided forthespecies-groupsofthegenusAdelotopus, forthespeciesofthe
genera Adelotopus, Cainogenion, and Cryptocephalomorpha, and for the Australian
speciesofthegenus Pseudomorpha. For the latter species a new subgenus Austro-
pseudomorpha, subgen. nov., and for the Argentinian and south Brazilian Pseudo-
morpha laevissima Chaudoirand P.glabraOgueta a newsubgenus Notopseudomor-
pha,subgen.nov.ofthegenusPseudomorphaiserected.ForCainogenionephippiatum
(Newman) a newsubgenus Procainogenion, subgen. nov. iserected.
Cainogenion cylindricum (Chaudoir) is transferred to Paussotropns Waterhouse
and Adelotopus insignis Sloane is transferred to Pseudomorpha Kirby, subgenus
Austropseudomorpha subgen. nov.
,
Following species names are synonymized (valid name second): Adelotopus
bijugusDarlington-A.rnfogidtatus(Blackburn);A.brunneusCastelnau=A.gyrinoides
Hope; A. castaneus Castelnau - A. rubiginosus Newman; A. filiformis Castelnau =
A.nemosomoides Westwood; A. longiphmnis Macleay - A. elongatuhis Macleay;
A.mastersii Macleay = A. dytiscides Newman; A. micans Blackburn = A. paroensis
Castelnau; A. niger Notman = A. paroensis Castelnau; A. occidentalis Castelnau -
A.gyrinoides Hope; Cainogenion bicolor (Castelnau) = C. ef)hippiatum (Newman);
Paussotropnsparalleliis Waterhouse= Paussotropnscylindricns (Chaudoir).
Adelotopus rufoguttatns (Blackburn) and Cainogenion suhopacum (Macleay) have
beenrevaluated from synonymy to füll speciesStatus.
Lectotypes, and eventuallv paralectotypes, are designated for the following
SupportedbyfourtravelgrantsandagrantforprintingexpensesoftheDeutscheForschungsgemeinschaft
(DFG).
species (including those names synonymized in present work): Pseudoiiiorplia
iiisiguis (Sloane), Adclotopiis analis Macleay, A. apicalis Macleay, A. bimaculatus
Macleay, A.brevipennis Macleay, A. bninneus Castelnau, A. castmieus Castelnau,
A.celeripes Lea, A. dytiscides Newman, A. elongatulus Macleay, A.fasciatus Castel-
nau,A.filiformisCastelnau,A.forfmnniHope,A.gi/rinoidesHope,A.haemorrhoidalis
Erichson,A.laevisMacleay,^4.linearisMacleay,A.loiigipcnnisMadeay,A.maculipennis
Macleay, A. mastersii Macleay, A. inicans Blackburn, A. ncmosoinoides Westwood,
A.occidentalisCastelnau,A.paroensisCastelnau,A.politusCastelnau,A.rubiginosus
Newman, A. rufogiittatus (Blackburn), A. tasimni Blackburn, A. viciinis Castelnau,
A.zonatus Castelnau, Cni)wgcnioii bicolor (Castelnau), C. creberrimum (Blackburn),
C.epbippiatiiiu(Newman),C. ipsoidcs(Westwood),C.obscunim(Castelnau),C.sub-
opacum (Macleay), Pniissotwpus cyliiuiricus (Chaudoir), P. pnrnik'iiis Waterhouse,
Cn/ptoccpliahmwrplmcollaris (Waterhouse),and C. marginata (Waterhouse).
A neotype isdesignated for AdclotopiisaffinisCastelnau.
The following 113 new taxa are described: Pseudomorpha brevis, spec. nov.,
P.insignispilosa,subspec.nov.,P.siibangiilata,spec.nov.;Adelotopusadelaideae,spec.
nov., A. adustus, spec. nov., A. aequiis, spec. nov., A. augustatiis, spec. nov.,
A.aterrimus, spec. nov., A.atrorufiis,spec. nov., A. bncilliis,spec. nov., A. bamagae,
spec.nov.,/4.basirufus,spec.nov.,/4.bimnciilatiisangustior,suhspec.nov.,A.brevior,
spec. nov., A. brittoni, spec. nov., A. browni, spec. nov., A. calims, spec. nov.,
A.cauiae,spec.nov.,A.ciliatiis,spec.nov.,A.ciliatiistenuipunctatiis,subspec.nov.,
A.clcpsydra,spec.nov.,A.conicollis,spec.nov.,A.convexicollis,spec.nov.,A.convexiis,
spec. nov., A. coriaceiis,spec. nov., A. crassus, spec. nov., A. cribricollis, spec. nov.,
A. crucis,spec.nov., A.ciineatus,spec. nov.,A.distiiigiiciidiis,spec. nov.,A.doyeni,
spec.nov.,A.dubius,spec.nov.,A.diibiiisglaber,subspec.nov.,A.dubiushobartensis,
subspec. nov., A. editluie, spec. nov., A.flavescens, spec. nov., A.ßavus, spec. nov.,
A.foliaceus,spec.nov.,A.geminus,spec.nov.,A.gibbosus,spec. nov.,A.gippslandi-
ciis,spec. nov.,A.grossepunctatus,spec.nov.,A.gyrinoidesorientalis,subspec.nov.,
A.houstoni,spec.nov.,y4.howdenorum,spec.nov.,y4.kathcrinei,spec.nov.,A.kurandae,
spec.nov.,/4.languidus,spec.nov.,/4.Inticnudatus,spec.nov.,/4.laticollis,spec.nov.,
A. lafior,spec. nov.,A. latipalpis,spec. nov., A. lawrencci,spec. nov., A. longifoniiis,
spec. nov., A. longus,spec. nov.,A. loiigiis tropicus,subspec. nov., A. lunatus,spec.
nov., A. lutcus, spec. nov., A. macilcntus, spec. nov., A. mainae, spec. nov.,
A.marginicollis,spec.nov.,A.minor,spec.nov.,A.montisatri,spec.nov.,/4.montorum,
spec. nov., A.multipiinctatus, spec. nov., A. niurrayanus, spec. nov., A. nigricaiida,
spec. nov., A.nitens, spec. nov., A. nitidior, spec. nov., A. obsoletus, spec. nov.,
A.ooldeae,spec.nov.,A.ovatus,spec.nov.,A,palumae,spec.nov.,A.parumpunctatus,
spec. nov., A.pienelopeae, spec. nov., A. piceiis, spec. nov., A. punctatissimus, spec.
nov., A.pinicticollis angustemaculatiis, subspec. nov., A. punctiilifer, spec. nov.,
A.queenslandicus, spec. nov., A. rufescens, spec. nov., A. riifocnudatus, spec. nov.,
A. rufomarginatus, spec. nov., A. rufozonatiis, spec. nov., A. scdlnceki, spec. nov.,
A.semilunatus, spec. nov., A. seminitidus, spec. nov., A. sericcus, spec. nov.,
A.seriepunctatusstriatus, subspec. nov., A. similis,spec. nov., A. sinuaticollis,spec.
nov.,A.sinuaticolliscalliope,subspec.nov.,A.sparsepunctatus,spec.nov.,A.substriatus,
spec. nov., A. iilriclii, spec. nov., A. unicolor, spec. nov., A. victoriensis, spec. nov.,
A.villosus,spec. nov.,A. virgatus,spec. nov.,A.yorkeusis,spec. nov.,A.zborowskii,
spec. nov.; Cainogenion clypeale, spec. nov., C. creberrimumgnaltae, subspec. nov.,
C.depressum,spec. nov., C.glabratiim,spec. nov., C. intcriore,spec. nov., C. ipsoides
occidentale, subspec. nov., C. parumpilosiim, spec. nov., C. rotundicolle, spec. nov.,
C.tropicum, spec. nov.; Cryptocephalomorpha australica, spec. nov., C. gcnieri, spec.
nov., and C. maior, spec. nov.
Followingsixspeciesremaindoubtful,becausethetypesarelostandthespecies
belongtospecies-groupsin whichspeciesdistinction is impossiblewithoutcom-
parison of the types: Adelotopus apliodioides Westwood, A. connitus Castelnau,
A.hydrobioides Westwood, A. inqiiinatus Newman, A. papuaniis Gestro, and
A.scolytidesNewman.
First instar larvae are described for the first time from the oviducts of many
species of the genera Adelotopus, Cainogenion, and Paussotropiis. Their special
charactersare included in the phylogeneticanalysis.
ThepossiblerelationshipsofthegeneraofPseudomorphinae,thesubgeneraof
thegenusPseiidoinorphn,thespecies-groupsofthegenusAddotopus,andthespecies
ofthegenera Cninogeuionand Cn/ptoccphalomorphaarediscussedanddescribedin
cladogramsbasedonareconstructedphylogenyemployingthemethodsproposed
by Hennig. ThephylogeneticrelationsofthespeciesofAdclotopiis and ofPseiido-
inorpJia are discussed. Phylogenetic evidence shows that Sphallouiorplm is the
adelphotaxonofallothergenera,Pscudomorphntheadelphotaxonoftheremaining
genera,thehighlyspecializedgenusCn/ptoccplialoiiiorpliapresumablytheadelpho-
taxonoftheAdt'/ofo^)((s-lineage, inwhichAdelotopusistheadelphotaxonofCaino-
gcnion+Paussotropus, whereas Paussotivpiis is the most evolved genus of this
lineage.
Thereisstrongevidenceforanincreasinggradeofadaptation tomyrmecophi-
lous habits in Pseudomorphinae in general, but also within the larger genera.
WhereasthegeneralstructureinSpilmlloinorphaandevenstillinPsciidoinorpJiamay
beratheradaptativetothesubcorticolouslife, this isreduced infavourofcertain
adaptations to myrmecophily in the other genera with Paussotropus and again
Cn/ptoccplmloiuorplmbeingmostevolvedinthisrespect.Comparableadaptationsto
myrmecophilous habits are also noted in the larvae of Pscudoinorplm, Addotopus,
Cniiiogeuiou, and Paussotropus, while larvae of Crifptocephaloiuorpha are still un-
known.
Thedistributionofthespeciesisdepictedinmaps.About25distinctdistribution
patternsaredistinguishedandmostofthemmayberegardedasrelatedtofaunistic
subregions. Phylogenetic evidence shows that those species and species-groups
thatareplesiomorphicinmanyrespectsconcentrateinthesoutheasternsubregions
ofAustralia,wherealsothemostplesiotypicspeciesofthegenera Sphallomorplm,
Addotopus, Caiuogcuioii, and Pseudomorplia subgenus Austropseudoniorpha occur.
Especiallyin thenortheastern, northern, western, and central subregionsa larger
number of species shows a higher degree of apomorphic character states. This
pattern of distribution is exemplified by several dichotomous and trichotomous
vicariantrelationships,inwhichtheevolvedvicariant(s)almostinvariablyoccur(s)
in the northern or western part ofthecommon ränge.
Distribution of the apparent basal species of all genera either in southeastern
Australia (Sphalloiiiorpha, Addotopus, Caiuogoiio}!, PseudovwrpJm subgenus Austro-
pseudoiuorpha),orinsouthernSouthAmerica (PscudomorphasubgenusNotopscudo-
iiiorplia), orinSouth Africa (Cryptocephaloinorpha) isevidenceofa Gondwanaland
originofthesubfamilywithoutknowingtheexactplaceoforigin. Evidentlymost
genera apart from Cryptoccphaloiuorplia and perhaps Pscudomorpha originated in
southeasternAustraliaortheadjacentpartofGondwanaland.Cryptoccpimloinorpilia,
however,iscertainlyarecentarrivalintheAustralianregionthatimmigratedfrom
theOrientalregion,whereitprobablyarriveddriftingonterranesoftheprevious
Sundaland. According to chorological, phylogenetic, and paleogeographic evi-
dence the subfamily and most of its genera must have evolved at least during
UpperJurassic oreven earlier.
The further history of the genera in Australia was comparable to that of
Spliallomorpha,buttheirtaxonomicradiationmayhavebeenevenmorerecentthan
in the latter genus and it is certainly not yet finished. It occurred mainly in
Pleistocene, synchronous with the late taxonomic ciiversification of eucalypts
duringthesameperiodandgenerallyincloserrelationtoants.Severallineagesof
Adelotopusand Cainogeuiou thenspread overmostofAustralia,mainlyinananti-
clockwise, northern and western direction. The repeated change of wet pluvials
and dry interpluvials during Pleistocene, with their spreading of suitable plant
communities (namely Eucalyptus forests orsavannahs) and thesubsequent Isola-
tionofthesecommunitiestogetherwiththeirPseudomorphinehostsaccountsfor
the high numberofendemicand highlyevolved speciesespecially in the refugia
ofcentraleasternand northernQueensland,farNorthernTerritory, northwestern
Australia, and central and southwesternWestern Australia.
Dr. M. Baehr, Zoologische Staatssammlung Mi^inchen, Münchhausenstr. 21,
D-81247München,Germany
Contents
1. Introduction 12
2. Material 12
3. Abbreviations of collections mentioned in text 12
4. Methods 14
4.1. Taxonomicprinciples 14
4.2. Phylogeneseprinciples 14
4.3. Descriptions 15
4.4. Types 15
4.5. Origin ofmaterial 16
4.6. Distinguishingcharacters 16
4.7. Chetotaxy 17
4.8. Illustrations 17
4.9. Data ofexamined material 17
4.10. Distributionmaps 18
4.11. Measurements 18
4.12. Magnifications 18
4.13. Abbreviations 18
5. Way of life and collecting methods 19
5.1. Way ofUfe 19
5.2. Myrmecophiloushabits 19
5.3. Collecting methods and preservation 21
6. Larvae 21
6.1. The larvae of the different pseudomorphine genera 22
7. Classification 40
7.1. SubfamilyPseudomorphinae 40
7.1.1. Key to the genera ofPseudomorphinae 40
7.2. Genera 41
7.2.1. GenusPscudoinorpha Kirby 41
7.2.1.1. SuhgenusNotopisciidoiiiorplia, subgen. nov 42
7.2.1.2. SuhgenusAustropseudomorpha, subgen. nov 42
7.2.1.3. DescriptionoiAiistropseiidoniorplm 43
7.2.1.4. Key to theAustralian species ofgenus Pseudoiuorplia Notman, subgenus Aiistwpseu-
domorpha, subgen. nov 43
7.2.1.5. ThespeciesofAustropseudonwvpha, subgen. nov 43
7.3. Genus Adelotopus Hope 51
7.3.1. Description ofAdelotopiis 51
7.3.2. Key to the species-groups of the genus Adelotopus Hope 58
7.3.3. Key to the species of the genus Adelotopus Hope 60
7.3.4. Thespecies ofthe genus Adelotopius Hope 76
7.4. Genus Cabiogenkvi Notman 328
7.4.1. Description of Caiuogeiiiou 329
7.4.2. Key to the species of the genus Caiiiogeuio}i Notman 334
7.4.3. The species of the genus Caiiiogoikvi Notman 336
7.4AA. SubgenusProcainogenion,subgen. nov 336
7.4.4.2. Subgenus Cainogeiüou s. str 340
10