Entomologische Abhandlungen 61 (2)(2): 119–172© Museum für Tierkunde Dresden, ISSN 0373-8981, 10.12.2003

Proceedings of the 1st Dresden Meeting on Insect Phylogeny:“Phylogenetic Relationships within the Insect Orders”(Dresden, September 19–21, 2003)
Edited by KLAUS-DIETER KLASS
Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, 01109 Dresden,
Germany [klaus.klass@snsd.smwk.sachsen.de]Foreword
KLAUS-DIETER KLASS1, NIELS PEDER KRISTENSEN2 & MATTHIAS NUSS1
1

Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, 01109 Dresden,
Germany [klaus.klass@snsd.smwk.sachsen.de; matthias.nuss@snsd.smwk.sachsen.de] – 2 Zoological Museum, Universitetsparken 15, 2100 Copehagen Ø, Denmark [npkristensen@zmuc.ku.dk]

Recent years have seen remarkable progress in reconstructing the phylogenetic relationships of the Insecta, based ona flood of new morphological and molecular evidence andrapid development of analytical methods. Whereas the relationships among the different ‘orders’ are still far frombeing robustly resolved, hypotheses about the basal relationships within a number of these ‘orders’ are now approaching maturity. Still, this is true to a very different extent
for the various orders, and it does not mean that there are nolonger disputes about diametrically conflicting hypotheses.
Long-standing questions such as the relationships between
Zygoptera and Anisoptera in Odonata, between Caelifera
and Ensifera in Orthoptera, and between the four subordersof the Coleoptera are cases in point, as are the position of
the Isoptera inside or outside the Blattaria, of the Siphonaptera inside or outside the Mecoptera, and of the Phthiraptera inside or outside the Psocoptera.
These basal phylogenetic relationships within the various
insect orders were the subject of the meeting the proceedings of which are here presented – a subject that would belaborious to comprehensively access through a search of theliterature. The meeting, which took place in the ‘Blockhaus’
in the historical center of Dresden immediately after theannual meeting of the German ‘Society for Biological
Systematics’ (GfBS), was designed and organized byKlaus-Dieter Klass, with much support from Matthias Nussand Niels Peder Kristensen.
In 35 invited talks, 55 speakers (including coauthors) who
have been deeply involved in research on the basal phylogenetic relationships of the individual insect ‘orders’ presented the state-of-the-art in their fields; many of themincluded yet unpublished results from their laboratories.
The program also included the recently (in 2002) discovered
order Mantophasmatodea, which up to now has grown to
include 13 described species, rendering it worthwhile todiscuss its intraordinal phylogeny. Unfortunately, on theother hand, both of the announced talks on Odonata had tobe cancelled briefly before the meeting; still, a written contribution on Odonata phylogeny by Günter Bechly could be
included in these proceedings.

In the talks, evidence from both the morphological and themolecular field was included according to the availabilityof data, and partly results from the two fields were presented in separate talks by speakers specialized on the evaluation of the one or the other kind of data; conflicts between
different data sets were indicated; and it was explained
which phylogenetic hypothesis is presently considered tohave the strongest support. A particularly impressive aspect
was that due to the rapid development of sequencing andanalytical methods for some insect orders molecular data
sets are meanwhile available that include hundreds of species. In contrast, in some other orders, such as Archaeognatha and Zygentoma, the compilation of molecular data
sets is still at a very early stage; and it became also clearthat in a number of insect orders there is still a very limited
knowledge of morphological data.
Speakers came on their own expenses from various institutions in Germany, Austria, Denmark, France, the United
Kingdom, Czechia, Italy, the USA, Australia, Singapore,and Argentina. Including the ca. 50 additional visitors, themeeting was altogether attended by about 100 persons.
A volume presenting the current state of understanding of
the basal splitting events in the various insect orders isplanned for completion within the next two years (editors:
Rudolf Meier, Klaus-Dieter Klass, Niels-Peder Kristensen& Michael Whiting). The team of authors preparing thebook chapters is largely congruent with the group of speakers invited to present a talk at the meeting.
Motivated by the great success of this meeting, the organizers are intending to arrange similar meetings on high-rank
insect phylogeny in Dresden that should take place every
second year. Focal issues of the meeting scheduled for
September 2005 should be: (1) morphological und molecular character systems that are particularly relevant in analysing the relationships among the insect orders; (2) the different and partly competing methods used in the analysis ofmolecular data; and (3) the results of the working groups
that have conducted comprehensive analyses on the relationships among the insect orders.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Participants of the 1st Dresden Meeting on Insect Phylogeny (photo: F. Höhler)

For the following talks presented at the meeting no written
contributions for the ‘Proceedings’ were submitted:“Hemiptera morphological and molecular (18S RNA) phylogenies: real conflict or unfinished work?” by THIERRYBOURGOIN1, DAVID OUVRARD1 & BRUCE C. CAMPBELL1
1
Museum National d’Histoire Naturelle, Entomologie &ESA 8043 du CNRS, 45, Rue Buffon, 75005 Paris,
France [bourgoin@mnhn.fr; ouvrard@mnhn.fr] – 2 Western
Regional Research Center, USDA-ARS, 800 BuchanannSt., Albany, CA 94710-1100, USA [bcc@pw.usda.gov]“The Systematics of the Heteroptera” by WARD C.
WHEELER & RANDALL T. SCHUHDivision of Invertebrate Zoology, American Museum of
Natural Histrory, Central Park West at 79th St., New York, NY
10024-5192, USA [wheeler@amnh.org; schuh@amnh.org]“Molecular phylogenetics in Lepidoptera: resolution andradiation” by BRIAN M. WIEGMANN1, JEROME C. REGIER2 &CHARLES MITTER3
1
Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613, USA [bwiegman@unity.ncsu.edu] – 2 Center for Biosystems Research,
Maryland Biotechnology Institute, University of Maryland,College Park, MD 20742, USA – 3 Department ofEntomology, University of Maryland, College Park, MD
20742, USA“Phylogenetic relationships among the Mecoptera based onmorphological evidence” by RAINER WILLMANN
Institut für Zoologie und Anthropologie, Abteilung für Morphologie, Systematik und Evolutionsbiologie, Berliner Str.
28, 37073 Göttingen, Germany [rwillma1@gwdg.de]Character Evolution in the Archaeognatha: Consensus and Conflict
MARKUS KOCH
Free University of Berlin, Institute of Biology/Zoology, Königin-Luise-Str. 1–3, 14195 Berlin, Germany [mkoch@zoosyst-berlin.de]Introduction
The current view on character evolution in the Archaeognatha (= Microcoryphia) is based on a phylogenetic system
inferred by STURM & BACH DE ROCA (1993) from an evaluation of “a wider spectrum of characteristics”. Cladistic
analyses, however, are still missing for the persisting difficulty to compile a reliable data matrix. This is partly due toinappropriate descriptions of many genera and the need forrevision of other genera (STURM 1991), partly due to thelack of relevant data especially on internal anatomy as well
as molecular sequences, and generally due to the uniformity of archaeognathan external morphology. Differences in
reproductive traits accordingly still form the central arguments for phylogenetic conclusions. Main progress in thepast decade primarily concerns the description of nearly
150 new species, which increased the number of species
known to date to 493 in total (MENDES 2002). STURM &MACHIDA (2001) integrated the new species into the current
system under maintenance of a distinction between so-

called paleoforms, remaining Machilidae, and Meinertellidae (Fig. 1A). This concept leaves three major problems
unresolved: (i) the integration of the fossil record, (ii) theassessment of the phylogenetic position of the ‘paleoforms’and (iii) a high number of parallelisms. Recent insights into
early hexapod evolution permit to revise the early branching
events.

†Dasyleptidae as extinct sister group of Machiloidea
The extinct Dasyleptidae (= †Monura Sharov, 1957) are
generally attributed to the Archaeognatha due to “strong
phenetic resemblance” (KRISTENSEN 1998). Support for thisview no longer only relates to the (1) presence of longmaxillary palps, which seem to be hypertrophic (e.g.,
BITSCH 1956) and functionally arranged like a second pairof antennae. Comparative analyses of the head morphology
in primarily wingless hexapods (KOCH 2001) in addition
favour (2) a sucking mode of food uptake, structurallyEntomologische Abhandlungen 61 (2)121

Fig. 1. Current hypotheses on the phylogeny of the Archaeognatha. A: After STURM & MACHIDA (2001). B: After BITSCH & NEL(1999). C: Present conclusions; numbers refer to potential synapomorphies as explained in the text.

based on the presence of narrow, elongate mouthparts forming a feeding cone, as a further potential synapomorphyof †Dasyleptidae and the extant subgroups (commonlyunited as Machiloidea). The same is true for the mandible
morphology: outgroup comparison (Diplura, Zygentoma,
Ephemeroptera) dictates the interpretation as the mostparsimonious assumption that the inability of biting in
extant archaeognathans is a derived state; the (3) monocondylic state of the mandibles shared by †Dasyleptidae andMachiloidea accordingly represents another potential synapomorphy. Previous support of this view by the attributionof an extinct species with allegedly dicondylic mandibles –inappropriately designated as “Dasyleptus” sp. (cf.
KUKALOVÁ-PECK 1987) – to the Archaeognatha (BITSCH &NEL 1999; Fig. 1B: †Cercopodata), however, seems to be
obsolete: according to the revision of this fossil by
WILLMANN (2003), its phylogenetic position presently
remains as unclear as – among others – the mandible structure of this organism. The same is also true for the extinct,
monotypic †Triassomachilidae, which according to
RASNITSYN (2002) “proved to be mayfly nymph”. All thispresently speaks in favour of a basal splitting ofArchaeognatha into †Dasyleptidae and Machiloidea, since
the monophyly of the latter is beyond any doubt. The mostobvious synapomorphies of the extant species are (4) large
compound eyes being contiguous in the midline; (5) thethorax is strongly arched in correlation with the jumping
ability; and (6) spiracles are lost on the first abdominal segment. A plenty of further potential synapomorphies weresummarized by STURM & MACHIDA (2001), but these sufferfrom being known from only a few species.
Arguments for the alternative view of a sistergroup relationship between †Dasyleptidae and Archaeognatha +Dicondylia (WILLMANN 2003) are less convincing. Potential
synapomorphies of the latter (with the plesiomorphic statepresumed to be maintained in †Dasyleptidae) mainly referto characters showing high variability within basal subgroups of the Ectognatha (subdivision of the tarsus, lengthof the ovipositor valves, enhancement of the thorax as atagma) or imply parallel evolution in Diplura (paired pretarsal claws), respectively. The latter may also concern thegreater reduction of the 11th abdominal segment. This trait
seems to correlate with the peculiarity that its paired appendages, the cerci, are moved – if at all – by longitudinal
muscles of the 10th abdominal segment. Due to the possibility that this is also true in Diplura (SNODGRASS 1931),

their cerci may be homologous appendages of the 11th
abdominal segment, despite its indistinctness both inembryonic and postembryonic stages (for different interpretations see KLASS 2001: 293–294). This tentatively permits to favour the (7) loss of cercal mobility in
Archaeognatha (with subsequent (8) loss of the cerci in†Dasyleptidae) as the more parsimonious assumption over
the alternative view of an independent gain of cerci and/or
cercal mobility in Diplura and Dicondylia. For final decisions, however, the homology of the cercal movers in
Diplura and Dicondylia needs to be clarified.
The cuticular head endoskeleton provides the possibility tofurther test the hypothesis of a sistergroup relationship between †Dasyleptidae and Machiloidea. In the extant species, the metatentorial plate is strongly enlarged in correlation with the jumping ability. The clarification of its shapein †Dasyleptidae accordingly could provide (presently missing) evidence for their jumping ability. With regard to thecurrent view that insects may be terrestrial crustaceans,however, studies on the homology of the muscles involvedin jumping in Caridoidea (Crustacea: Malacostraca) andArchaeognatha are required to clarify whether the jumping
ability was newly acquired or only optimized inArchaeognatha.

Ditrigoniophthalmus oreophilus as sister speciesof the remaining Machiloidea
Among the so-called paleoforms within Machiloidea, only
the basal position of Ditrigoniophthalmus oreophilus presently seems to be justified: all other extant species are characterized by the (9) loss of the styli on the first abdominal
segment. Their presence in D. oreophilus, however, stillneeds to be reconfirmed. The exclusion of the remaining‘paleoforms’ (Charimachilis and Mesomachilis) from the
Machilidae remains questionable as the monophyly of thelatter is still left unsupported by any synapomorphic character. One noteworthy trait restricted to Machilidae within
Machiloidea is the carrier-thread method of sperm transfer,but outgroup comparison (Chilopoda, Progoneata, Collembola, Diplura, Zygentoma) indicates that this method assuch is the plesiomorphic state within Machiloidea (WITTE& DÖRING 1999; conditions in D. oreophilus unknown).
Specifics of this method are variable within Machilidae andapparently characteristic for subgroups only (GOLDBACH

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2000; STURM & MACHIDA 2001). Sperm transfer via stalked
spermatophores in the Meinertellidae, in contrast, correlatewith (10) corresponding transformations of the male genitalia (strongly reduced penis length, no ‘parameres’ = gonapophyses VIII and IX, special glands connected with thevasa deferentia) and maxillary palps (2nd palpomere with
hook-like projection).

Conclusions
Based on the available data, the possibility still cannot be
excluded that the so-called paleoforms (except
Ditrigoniophthalmus) and remaining Machilidae form aparaphyletic assemblage with respect to Meinertellidae(Fig. 1C). The high number of parallelisms resulting fromthe current view on archaeognathan phylogeny may accordingly be artificial. Its explanation by “discontinuously
active genes” (STURM 1994) in any case seems to be premature, since the most parsimonious character distribution
still remains to be inferred.

Acknowledgements
I am grateful to the organizers of this meeting for their invitation. Comments and discussion by Thomas Bartolomaeus(Berlin) and Klaus-Dieter Klass (Dresden) are also gratefully
acknowledged.

References
BITSCH, J. 1956. Considerations anatomiques sur le palpe
maxillaire des Machilidae. – Travaux du Laboratoire deZoologie et de la Station Aquicole Grimaldi de la Facultédes Sciences de Dijon 15: 1–9.
BITSCH, J. & A. NEL 1999. Morphology and classification of
the extinct Archaeognatha and related taxa (Hexapoda). –Annales de la Société Entomologique de France (N.S.) 35:
17–29.
GOLDBACH, B. 2000. The evolutionary changes in the reproductive biology of the Archaeognatha (Insecta). –Zoologischer Anzeiger 239: 215–229.
KLASS, K.-D. 2001. The female abdomen of the viviparous earwig Hemimerus vosseleri (Insecta: Dermaptera:
Hemimeridae), with a discussion of the postgenital abdo-

Proceedings of 1st Dresden Meeting on Insect Phylogeny

men of Insecta. – Zoological Journal of the Linnean
Society 131: 251–307.
KOCH, M. 2001. Mandibular mechanisms and the evolution ofhexapods. – Annales de la Société Entomologique deFrance (N.S.) 37: 129–174.
KRISTENSEN, N.P. 1998. The groundplan and basal diversification of the hexapods. In: R.A. FORTEY & R.H. THOMAS(eds.), Arthropod Relationships. – Systematics
Association Special Volume 55: 281–293. Chapman &Hall, London.
KUKALOVÁ-PECK, J. 1987. New Carboniferous Diplura,
Monura, and Thysanura, the hexapod ground plan, and therole of thoracic side lobes in the origin of wings (Insecta).
– Canadian Journal of Zoology 65: 2327–2345.
MENDES, L. 2002. Taxonomy of Zygentoma andMicrocoryphia: historical overview, present status andgoals for the new millennium. – Pedobiologia 46:
225–233.
RASNITSYN, A.P. 2002. Subclass Lepismatona LATREILLE, 1804.
The wingless insects (= Thysanura LATREILLE 1796, s.l.).
– Pp. 69–74 in: A.P. RASNITSYN & D.L.J. QUICKE (eds.),
History of Insects. – Kluwer Academic Publ., Dordrecht,
Boston, London.
SNODGRASS, R.E. 1931. Morphology of the insect abdomen I:
general structure of the abdomen and its appendages. –Smithsonian Miscellaneous Collections 85: 1–128.
STURM, H. 1991. Three new genera of Machilidae from North
America and Mexico: Leptomachilis, Meximachilis andMesomachilis, with description of two new sense organs
in Mesomachilis males (Insecta, Archaeognatha). –Steenstrupia 17: 53–80.
STURM, H. 1994. Diskontinuierlich aktive Gene und Evolution.
Eine Diskussion am Beispiel der Archaeognatha (Insecta).
– Journal of Zoological Systematics and Evolutionary
Research 32: 241–263.
STURM, H. & C. BACH DE ROCA 1993. On the systematics of theArchaeognatha (Insecta). – Entomologia Generalis 18:
55–90.
STURM, H. & R. MACHIDA 2001. Archaeognatha. In: N.P.
KRISTENSEN & R.G. BEUTEL (eds.), Handbook of Zoology
IV/37: 1–213. – De Gruyter, Berlin, New York.
WILLMANN, R. 2003. Die phylogenetischen Beziehungen derInsecta: offene Fragen und Probleme. – Verhandlungendes Westdeutschen Entomologentags 2001: 1–64.
WITTE, H. & D. DÖRING 1999. Canalized pathways of change
and constraints in the evolution of reproductive modes of
microarthropods. – Experimental and Applied Acarology
23: 181–216.

Towards a Phylogenetic System of the Zygentoma
MARKUS KOCH
Free University of Berlin, Institute of Biology/Zoology, Königin-Luise-Str. 1–3, 14195 Berlin, Germany [mkoch@zoosyst-berlin.de]Introduction
Since ESCHERICH’s (1905) monographic treatment of thesystematics of the Zygentoma, the taxonomic work in thepast 50 years especially by Pedro Wygodzinsky and Luis
Mendes provided a much more differentiated view on thephylogenetic relationships among the 470 silverfish species
known to date. Yet, however, a considerable number of characters were included into phylogenetic analyses of two subgroups only: Lepismatidae, which comprise most of the‘free-living’ representatives, and Nicoletiidae, in which all
cave dwellers and deep edaphic forms are subsumed(MENDES 1991, 1994). Their composition and monophyly is
traditionally a matter of debate. This especially concerns theuncertainty upon whether the generally myrmeco- or termitophilous representatives of the ‘Ateluridae’ form a sub-group of the Nicoletiidae, in as much as nidicolous forms
are also known from the Lepismatidae. The character combination in the Maindroniidae also renders a decision difficult upon whether this taxon is more closely related to
Nicoletiidae or Lepismatidae.
These uncertainties may hardly surprise as only few morphological characters are presently considered to be significant at high-rank level. MENDES (2002) concluded from his
latest analysis that the eyeless representatives (Nicoletiidae
and Ateluridae), in contrast to the traditional view, form aparaphyletic assemblage. Data raised from recent studies on
the head morphology of Tricholepidion gertschi, the only
extant species of the Lepidotrichidae, furthermore strengthened previous doubts upon whether the Lepidotrichidae
and remaining zygentoman subgroups form a monophylum(STANICZEK 2000; BEUTEL & GORB 2001). Recent revision

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Entomologische Abhandlungen 61 (2)Fig. 1. Present conclusions on the phylogenetic interrelationships among the subgroups of the Zygentoma. Numbers refer
to apomorphic characters as explained in the text.
of the structure of the spermatozoa (DALLAI et al. 2001,
2002) in addition rendered the previous view questionable
that these provide support for zygentoman monophyly. New
data especially on the morphology of the Maindroniidae
contribute to a clarification.

New arguments in favour of zygentoman
monophyly
Previous arguments unambiguously in support of zygentoman monophyly concern the (1) loss of the superlinguaeand (2) certain transformations of the walking legs (horizontal orientation of broadened, dorsoventrally flattenedcoxa and femur; WILLMANN 2003) that seem to correlatewith the (3) dorsoventral depression of the body. New evidence for the monophyly of the Zygentoma (incl.
Lepidotrichidae) is derived from the (4) structure of theabdominal cerci: a unique hook-like condyle at their baseproved to be present in representatives of all high-ranking
subgroups except ‘Ateluridae’. Different from what isfound in any other insect subgroup, the condyle is formed
as a lateral protuberance of the cercal base and articulateswith the posterior margin of the 10th abdominal tergum,
which bridges the 11th tergum and overhangs the cercal
base. The shape of this condyle, however, differs within
Zygentoma: it is composed of a single, externally visible
hook in Tricholepidion and Nicoletia, while this hook faces
an additional, adjacent hook hidden by the 10th abdominal
tergum in Maindronia neotropicalis as well as Lepisma andThermobia spp. (and Ctenolepisma longicaudata as illustrated by ESCHERICH 1905: fig. 41). The absence of this condyle in Atelura formicaria (‘Ateluridae’) may correlate withthe great shortening of the cerci. As illustrated by MENDES
et al. (1994: fig. 36), however, the condyle seems to be
maintained on the likewise shortened cerci of Trichotriuroides boneti (Nicoletiidae).
Further new support for zygentoman monophyly is derived
from the cuticular head endoskeleton of Maindronia
neotropicalis. In this zygentoman species, the anterior tentorium is similarly fused with the posterior tentorium as inpterygotes; in Tricholepidion gertschi and the lepismatids
yet studied, in contrast, these two tentorial components haveremained discrete. On the other hand, the paired anteriortentorial apodemes in M. neotropicalis share a peculiar feature with those in Lepismatidae: presence of a wing-likeprotuberance, from which components of the tentorial mandibular ‘adductor’ muscles originate. This protuberance is apotential synapomorphy of Maindroniidae and Lepismatidae (and Nicoletiidae?) as it is not only missing in
T. gertschi, in which the homologous muscles still arise
from the connective transverse mandibular tendon, but also

seems to be absent in the Pterygota. In this group, the origin of the homologous mandibular muscles at the anteriortentorial apodemes accordingly differs in detail from thestate in Maindroniidae and Lepismatidae. All this presently
permits to favour the view that (nearly) correspondingtransformations of the head endoskeleton and coincident
changes of the mandibular musculature happened within
Zygentoma and Pterygota in parallel.

Basal splitting events within Zygentoma
Due to missing data on the state in Nicoletiidae and‘Ateluridae’, the transformation of the head endoskeleton(5: loss of the connective transverse mandibular tendon andcoincident shift of the origin of the individualized bundlesof the mandibular ‘adductor’ muscle to the anterior tentorial apodemes or hypopharnyx, respectively) still remains atentative argument for the view that the Lepidotrichidae
form the adelphotaxon of the remaining Zygentoma. Any
further support for a sistergroup relationship of this kindconcerns a considerable number of reductive traits. Amongthese, the (6) loss of the median eyes (ocelli) is especially
noteworthy. Their apparent absence in the type species of
the Lepidotrichidae, the extinct †Lepidotrix pilifera(CARPENTER 1992: 17), however, questions the previous
view that this species and the extant Tricholepidion gertschi
form a monophylum. The only potential synapomorphy ofthese two species concerns the (7) presence of 5 tarsomeres,
but this feature may rather already belong to the ground pattern of the Dicondylia (Zygentoma + Pterygota; e.g.,
MENDES 2002).
Accordingly, the possibility cannot be excluded that the
Lepidotrichidae form a paraphyletic assemblage, with
Tricholepidion gertschi as sister species to a unit composedof all other zygentomans (Fig. 1). According to this view,the exinct †Lepidotrix pilifera branches from the stem lineof the remaining extant zygentomans characterized by a (8)reduced number of tarsomeres. Further characteristics (withthe plesiomorphic state maintained in T. gertschi and, as far
as known, in †L. pilifera) concern the (9) reduced numberof ommatidia, the (10) reduced number of pectinate appendages (prostheca) on the lacinia, the (11) reduced number ofovarioles and the (12) reduced size of the abdominal pregenital sterna (KRISTENSEN 1998). The states of these characters (9–12), however, are variable in extant zygentomans,
which renders their phylogenetic significance difficult toassess. In this respect, the structure of the male genital
coxosterna (KRISTENSEN 1998: character 68) may provide amore reliable argument, but a comparative analysis of theircomposition is still wanting.

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Nicoletiidae (incl. ‘Ateluridae’) as sistergroup of Protrinemuridae + Maindroniidae +Lepismatidae?
Based on the available data, the exclusion of theProtrinemurinae (now with family rank) from Nicoletiidae(MENDES 2002) seems to contribute a more parsimonious
interpretation of the character evolution within Zygentoma.
The current hypothesis that Protrinemuridae are more closely related to Maindroniidae and Lepismatidae is not only
supported by the (13) loss of the maxillary prostheca andthe (14) total absence of abdominal vesicles (MENDES
2002). Protrinemuridae also correspond with Maindroniidae and Lepismatidae in that (15) all coxosternal plates ofthe pregenital abdomen (I–VII) are no longer subdivided bysutures into a median ‘sternite’ and lateral ‘coxites’. The
significance of this potential synapomorphy, however,remains questionable for being paralleled in ‘Ateluridae’and Subnicoletiinae within Nicoletiidae as well as in the
Pterygota.
The hypothesis of a unit comprising Protrinemuridae,
Maindroniidae and Lepismatidae also permits to favour theview that features common to ‘Ateluridae’ and remaining
Nicoletiidae evolved only once within Zygentoma: (16)secondary sexual modification of the male antennal pedicel, (17) transformation of the penial opening into a thin,longitudinal slit, (18) gain of a spinulated area at the innerdistal wall of the female gonapophyses IX, and probably
also the (19) formation of spermatolophids (MENDES 2002).
The (20) presence of a subgenital plate formed by the hindpart of venter VII and covering the base of the ovipositor in‘Ateluridae’ and Nicoletiidae may also support their closeaffinity. The polarity of this character, however, remains
unclear: a similar plate is formed in the few Pterygota thathave retained both a fully developed ovipositor and the original position of the genital opening at the hind rim of segment VII, at least Dictyoptera and Ensifera (KLASS 1998a),
which indicates that a subgenital plate may already belong
to the ground pattern of the Dicondylia. A small lobe present in Lepismatidae in corresponding position (“languette”
in Thermobia; ROUSSET 1973) may accordingly represent a
remnant of this plate. This presently leaves the hypothesisof a close relationship between ‘Ateluridae’ and Nicoletiidae mainly based on correspondences in reproductive
traits. As a subgroup of the Nicoletiidae, however, themonophyly of the ‘Ateluridae’ seems to be only supportedby the presence of entire coxosterna in the pregenital abdomen (cf. character 15) and hence remains questionable.
Conflicts resulting from the view that Protrinemuridae forma clade with Maindroniidae and Lepismatidae not only concern the implication of a parallel (21) loss of the compoundeyes in Nicoletiidae and Protrinemuridae. This hypothesis
also strongly weakens the support for the monophyly of theProtrinemuridae, with the absence of eyes being left as the
only potential synapomorphy of its two subgroups. In addition, the monophyly of a clade Maindroniidae + Lepismatidae is no longer supported by the available data if thesefamilies form a clade with Protrinemuridae. Under thishypothesis, the only potential synapomorphy seems to be
the (22) reduction of the compound eyes to (constantly?) 12
ommatidia on each side. Due to the complete reduction of
the eyes in the closest relatives (Nicoletiidae andProtrinemuridae), however, the significance of this character is dubious.
The assessment of the phylogenetic relations of theMaindroniidae is further hampered by the persisting difficulty to reason the monophyly of the Lepismatidae. The significance of the (23) presence of a proventriculus in (all?)representatives of this group still remains ambiguous. In
part, this is due to claims that denticles are also present in
the posteriormost part of the foregut in Nicoletia (Nicoleti-

Proceedings of 1st Dresden Meeting on Insect Phylogenyidae) (GRASSI 1886). Detailed structural correspondences ofthe proventriculi in some Pterygota furthermore indicatethat this organ may already belong to the ground pattern of
the Dicondylia (KLASS 1998b). The monophyly of theMaindroniidae, in contrast, is beyond any doubt. The unusually large body size and peculiarities of the head andmouthparts – (24) prognathous head posture, (25) mandibular gnathal lobe composed of incisor teeth only (molar
plate lost) and the (26) transformation of the maxillary lacinia into a strong hook bare of any appendages – indicatethat these may be predators.

Conclusions
While the monophyly of the Zygentoma presently remainsa reasonable hypothesis, the interrelationships of its subgroups are still far from being clear. This is due to the deficiency that current conclusions remain based on reductive
traits only and/or arguments implying parallelisms within
Zygentoma and Pterygota. Unique acquisitions are presently known only from Nicoletiidae (excl. Protrinemuridae,
incl. ‘Ateluridae’) and Maindroniidae. These are accordingly the only high-ranking subgroups that at present canbe accepted as monophyla. For any further decisions, available data are insufficient.

Acknowledgements
I am grateful to the organizers of this meeting for the invitation to present the new results. Thanks are also due to Hans
Strümpel (Zoological Museum of Hamburg) for the loan ofMaindronia specimens. Comments and discussion by Thomas
Bartolomaeus (Berlin), Gonzalo Giribet (Cambridge, MA),
Klaus-Dieter Klass (Dresden) and Luis Mendes (Lisboa) arealso gratefully acknowledged.

References
BEUTEL, R.G. & S.N. GORB 2001. Ultrastructure of attachment
specializations of hexapods (Arthropoda): evolutionary
patterns inferred from a revised ordinal phylogeny. –Journal of Zoological Systematics and Evolutionary
Research 39: 177–207.
CARPENTER, F.M. 1992. Superclass Hexapoda. In: Treatise on
Invertebrate Paleontology, Part R, Arthropoda 4, Vol. 3–4,
xxii + 655 pp.
DALLAI, R., P. LUPETTI, A. CARAPELLI, F. FRATI & B.A.
AFZELIUS 2002. Sperm structure and spermiogenesis in
Atelura formicaria Heyden (Zygentoma, Insecta). – Acta
Zoologica (Stockholm) 83: 245–262.
DALLAI, R., P. LUPETTI, F. FRATI, F. NARDI & B.A. AFZELIUS
2001. Binucleate and biflagellate spermatozoa in
Tricholepidion gertschi WYGODZYNSKY (Insecta, Zygentoma). – Tissue & Cell 33(6): 606–613.
ESCHERICH, K. 1905. Das System der Lepismatiden. –Zoologica (Stuttgart) 43: 1–164.
GRASSI, B. 1886. I progenitori dei Miriapodi e degli Insetti. IV.
Cenni anatomici sul genere Nicoletia. – Bollettino della
Società Entomologica Italiana 18: 173–182.
KLASS, K.-D. 1998a. The ovipositor of Dictyoptera (Insecta):
homology and ground-plan of the main elements. –Zoologischer Anzeiger 236: 69–101.
KLASS, K.-D. 1998b. The proventriculus of Dicondylia, withcomments on evolution and phylogeny in Dictyoptera andOdonata. – Zoologischer Anzeiger 237: 15–42.
KRISTENSEN, N.P. 1998. The groundplan and basal diversification of the hexapods. In: R.A. FORTEY & R.H. THOMAS(eds.), Arthropod Relationships. – The Systematics
Association Special Volume Series 55: 281–293.
Chapman & Hall, London.
MENDES, L.F. 1991. On the phylogeny of the genera of
Lepismatidae (Insecta: Zygentoma). Pp. 3–13 in: G.K.
VEERESH, D. RAJAGOPAL & C.A. VIRAKTAMATH (eds.),
Advances in Management and Conservation of Soil
Fauna. – New Delhi, Oxford & IBH Publ.

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Entomologische Abhandlungen 61 (2)MENDES, L.F. 1994. Evolutionary relationships among the Nicoletiidae (Insecta, Zygentoma). – Acta Zoologica Fennica
195: 98–103.
MENDES, L.F. 2002. On the status of the “protrinemurid” and “atelurid” thysanurans (Zygentoma: Insecta). – Boletim da Sociedade Portuguesa de Entomologia 199(VII–17): 201–212.
MENDES, L.F., C. BACH DE ROCA, M. GAJU-RICART & R.
MOLERO-BALANÁS 1994. Trichotriuroides boneti gen. n.,
sp. n. (Zygentoma, Nicoletiidae) and new data on
Zygentoma in the collection of the Museo Nacional deCiencias Naturales in Madrid. – Eos 69: 21–29.
ROUSSET, A. 1973. Squelette et musculature des regions genita-

les et postgenitales de la femelle de Thermobia domestica(PACKARD). Comparaison avec la region genitale deNicoletia sp. (Insecta: Apteryogta: Lepismatida). –International Journal of Insect Morphology &Embryology 2(1): 55–80.
STANICZEK, A.H. 2000. The mandible of silverfish (Insecta:
Zygentoma) and mayflies (Ephemeroptera): its morphology and phylogenetic significance. – Zoologischer
Anzeiger 239: 147–178.
WILLMANN, R. 2003. Die phylogenetischen Beziehungen derInsecta: offene Fragen und Probleme. – Verhandlungendes Westdeutschen Entomologentags 2001: 1–64.

Ephemeroptera Phylogeny and Higher Classification: Present Status
and Conflicting HypothesesTOMÁ√ SOLDÁN
Institute of Entomology, 31 Brani√ovská, 370 05 Çeské Budπjovice, Czech Republic [soldan@entu.cas.cz]

Ephemeroptera relationships to other insects
The order can be characterized by the following principal
characters: (1) bristle-like adult antennal flagellum, (2) specific arrangement of tentorial muscles (STANICZEK 2000);(3) ‘sliding’ articulation of mandibles (KUKALOVÁ-PECK1991); (4) ‘true’ hypopharyngeal superlinguae; (5) galealacinia fusion; (6) unique arrangement of thoracic tracheal
trunks (only a single tracheal trunk coming from the leg trachea, corresponding to the trachea of the paranotal lobes ofZygentoma or Archaeognatha; LANDA 1948); (7) universally lacking occlusor muscles of the abdominal spiracles; (8)well developed, long terminal filament in larvae; (9) retention of winged subimaginal stage; (10) fore wings with prominent basal subcostal brace, and (11) anal brace ending on
CuP at a bulla; (12) aquatic mode of life; (13) telotrophic
ovarioles.
Apparently, some characters are shared with primitive
Neoptera, such as Plecoptera (4, 12), and some with
Odonata (e.g., 1, 7, 12, partly 10, 11), and one (6) is sometimes considered neotenic.
Three hypotheses on ephemeropteran affinities are available: (a) Ephemeroptera + Odonata (= Paleoptera, perhaps
including extinct paleodictyopteran orders) are the sistergroup of Neoptera; (b) Ephemeroptera are the sister groupof Odonata + Neoptera; (c) Odonata are the sister group of
Ephemeroptera + Neoptera. The problem still remains open
to a general discussion. From the neontological-morphological point of view, hypothesis (b) seems to be most parsimonious (BEUTEL & GORB 2001), but requires, e.g., parallelism in the antennal flagellum structure, and the nonhomology of all neopteran hypopharyngeal lobes withsuperlinguae is perhaps disputable. In contrast, a detailed
study of fossil material does lend support to hypothesis (a);according to my opinion this hypothesis is well corroborated by characters of wing venation (e.g., media always withbasal stem) and, in general, by the wing-from-leg-base-exite
theory (KUKALOVÁ-PECK 1983). Hypothesis (c) seems to beunlikely (see, e.g., discussions by KUKALOVÁ-PECK 1991
and KRISTENSEN 1991).

Suborder relationships
The original subordinal classification by MCCAFFERTY &EDMUNDS (1979), based mostly on thoracic morphology
and wing pad position, comprised the holophyletic Pannota
and the paraphyletic suborder Schistonota indicating theretention of certain plesiomorphic traits. Later MCCAFFERTY(1991) suggested a reasonable re-classification into 3 suborders: Rectracheata (including unchanged original Pannota
as an infraorder), Setisura, and Pisciforma. Although heused numerous characters including internal anatomy data
by LANDA & SOLDÁN (1985), the Pisciforma still required
some revision through abandoning the use of plesiomorphic
characters.
Since then some progress has been achieved, but it seems to
be extremely difficult to reach a parsimonious solution ofthis problem because some groups (e.g., the superfamily
Siphlonuroidea) are defined by plesiomorphies, and new
characters are hardly available. Independently, N.Yu. Kluge(see KLUGE 2000 for a complete list of his contributions)defined suborders Costatergalia and Furcatergalia on thebasis of presence or absence of certain gill structures (therein called “tergalia” to emphasize wing origin; however,‘gills’ in mayflies might be of a rather different origin).
Later, the same author suggested two suborders again:
Anteritorna and Posteritorna based on two-fold emergenceof anteromotority. Although partly including clearly holophyletic taxa (e.g., Posteritorna), both subordinal classifications are essentially based on a single character.

Infraorder relationships
MCCAFFERTY’s (1991) suborders Rectracheata and Pisciforma comprise 3 (namely Vetulata, Lanceolata, and Pannota)
and 2 (namely Arenata and Imprimata) infraorders, respectively. This classification seems to be generally accepted,
except for the above mentioned paraphyly of the Pisciforma
and the position of some ‘critical’ genera. For instance, thegenus Oniscigaster (and the respective monotypic family)
requires more attention showing, e.g., dramatic autapomorphic increase in ventral tracheal anastomoses. It is treated in
the monotypic infraorder Vetulata by MCCAFFERTY (1991)
but is left in the Siphlonuridae-like taxa in his Tridentiseta
by KLUGE (2000), probably on the basis of the mouthpart“dentisetae” synapomorphy and bordered gills of thisgenus, which gives to Kluge’s Costatergalia a rather paraphyletic status. Similarly, the genus Pseudiron (and the respective monotypic family) deserves further critical evaluation, being treated in the otherwise well defined Setisuralike lineage (KLUGE 2000, his Brachytergaliae) but in
Pisciforma-Arenata by MCCAFFERTY (1991).
A rather different higher classification using “consistently
non-ranking taxonomy” has been developed by KLUGE(2000, see therein for earlier citations, and on

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http://www.bio.pu.ru/win/entomol/KLUGE/EPH/Contents.
htm for more details). I am definitively not in a position toevaluate the scientific invention of this undoubtedly sophisticated classification, but there are some evident restrictions concerning its general acceptance: in many respects, itinterferes with the ICZN (for instance, rather arbitrary handing with synonymy, e.g., Arthropleidae Balthasar, 1937 isdefinitively not a synonym of Heptageniidae Needham,1901); it is still not applicable to any other insects except
for the Ephemeroptera; and it seems to be extremely complicated to be used in other than strictly taxonomic papers(I do not understand why to use “Turbanoculata
Anteropatellata Baetis/fg7 Acentrella/g1” instead of simply
the genus (subgenus) name Acentrella of the family
Baetidae, e.g., in faunistic lists and ecological papers).
Moreover, as many as 20 suprageneric names (7 of them
newly erected as “taxon nov.”) are introduced in this treatment (KLUGE 2000) and to trace their relationships to widely used sub- and infraordinal, super- and subfamilial, familial or even generic names requires a long time and maximal concentration.

Superfamily and family relationships
Contrary to the situation in the sub- and infra-ordinal ranks,
there is a general consensus in the definitition of taxa of(super-)family rank as well as in the application of taxon
names. Five superfamilies (namely Leptophlebioidea,
Behningioidea, Ephemeroidea, Caenoidea and Neoephemeroidea) have been introduced by MCCAFFERTY (1991) in
addition to the Siphlonuroidea, Baetoidea, and Heptagenioidea, approximately corresponding to his infraorders
Arenata and Imprimata, and the suborder Setisura, respectively. At present, 37 families have been recognized, which
include 376 genera and approximately 3,100 species – contrary to 20 families defined, e.g., by MCCAFFERTY &EDMUNDS (1979). However, taxonomic changes at this level
are restricted mostly to formal shifts in rank (from subfamily to family status). As far as I know, only the genus
Siphluriscus (originally incertae sedis within the Siphlonuroidea) will be treated in a separate family soon.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

References
BEUTEL, R.G. & S.N. GORB 2001. Ultrastructure of attachment
specializations of hexapods (Arthropoda): evolutionary
patterns inferred from a revised ordinal phylogeny. –Journal of Zoological Systematics and Evolutionary
Research 39: 177–207.
KLUGE, N.YU. 2000. Contemporary Systematics of Insects.
Principals of the systematics of living organisms andgeneral system of insects with classification of primary
apterygotes and primitive pterygotes. – Lan Publishing
House, St. Petersburg, 336 pp. [in Russian].
KRISTENSEN, N.P. 1991. Phylogeny of extant hexapods. Pp.
125–140 in: I.D. NAUMANN (ed.), The Insects of Australia.
– CSIRO, Melbourne Univ., Carlton.
KUKALOVÁ-PECK, J. 1983. Origin of the insect wing and wingarticulation from the arthropodan leg. – Canadian Journalof Zoology 61: 1618–1669.
KUKALOVÁ-PECK, J. 1991. Fossil history and evolution of hexapod structures. Pp. 141–179 in: I.D. Naumann (ed.), The
Insects of Australia. – CSIRO, Melbourne Univ., Carlton.
LANDA, V. 1948. Contribution to the anatomy of ephemerid larvae. I. Topography and anatomy of tracheal system. –Vπstník Çeskoslovenské Spoleçnosti Zoologické 12:
25–82.
LANDA, V. & T. SOLDÁN 1985. Phylogeny and higher classification of the order Ephemeroptera: a discussion from thecomparative anatomical point of view. – Studie ÇSAV 4.
Academia, Prague, 121 pp.
MCCAFFERTY, W.P. 1991. Toward a phylogenetic classificationof the Ephemeroptera (Insecta): a commentary on systematics. – Annals of the Entomological Society of America
84: 343–360.
MCCAFFERTY, W.P. & G.F. EDMUNDS JR. 1979. The higher classification of the Ephemeroptera and its evolutionary
basis. – Annals of the Entomological Society of America
72: 5–12.
STANICZEK, A.H. 2000. The mandible of silverfish (Insecta:
Zygentoma) and mayflies (Ephemeroptera): its morphology and phylogenetic significance. – Zoologischer
Anzeiger 239(2): 147–178.

Acknowledgements
My sincere thanks are due to Klaus-Dieter Klass and Niels P.
Kristensen for valuable comments on the manuscript.

Phylogeny of Ephemeroptera: Molecular Evidence
T. HEATH OGDEN & MICHAEL F. WHITING
Department of Integrative Biology, Brigham Young University, Provo, Utah 84602, USA [Heath_Ogden@byu.edu;
Michael_Whiting@byu.edu]

Mayflies are unique among insects by the presence of a
non-reproductive winged stage (subimago) that molts tobecome the reproductive adult. Phylogenetic relationships
among mayfly families are poorly known, and current
hypotheses are based on untested scenarios of character
evolution, which lack any sort of rigorous phylogenetic analysis.
In our ongoing study we use data based on five genes (18S
rDNA, 28S rDNA, 16S rDNA, 12S rDNA, and histone 3) toestimate mayfly phylogenetic relationships. Nearly 100
genera of mayflies, representing the majority of lineages,
are included. Previous hypotheses for higher-level relationships are discussed and tested in light of these data. In thefollowing explanations the use of taxon names refers to theclassification of MCCAFFERTY (1991).

Our analysis supports the family Baetidae as sister to allother mayflies, though this result is sensitive to outgroupand optimization alignment parameter selection. The pisciform mayflies are supported as grossly paraphyletic.
Potamanthidae is nested outside the clade Scapphodonta (=Ephemeroidea + Leptophlebiidae + Behningiidae) andBehningiidae is nested within the Ephemeroidea, rendering
this group of burrowing mayflies non-monophyletic. These
results suggest that mandibular tusks were gained on multiple occasions with a secondary loss in the lineage
Behningiidae. Additionally, the large family Heptageniidae
is not supported as monophyletic because the genera
Pseudiron and Arthroplea are nested within this family. The
families Baetidae, Leptohyphidae, Nesameletidae,
Oligonuridae, Potamanthidae and Ephemerellidae are sup-

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Entomologische Abhandlungen 61 (2)ported as monophyletic groups. While our analysis is in its
preliminary stages, it represents the first formal cladistic
analysis across the major lineages of Ephemeroptera.

References
MCCAFFERTY, W.P. 1991. Toward a phylogenetic classificationof the Ephemeroptera (Insecta): a commentary on systematics. – Annals of the Entomological Society of America
84: 343–360.

The Phylogenetic Relationships of the Three Extant Suborders of OdonataGÜNTER BECHLY
Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany [bechly.smns@naturkundemuseum-bw.de]

The order Odonata includes three extant suborders (Zygoptera, Anisozygoptera: Epiophlebiidae, and Anisoptera) withtotally 5538 described species. The fossil record is relatively
well documented with more than 600 fossil species and reaches from Tertiary representatives of extant families back toprimitive protodonates of the lowermost Upper Carboniferous (320 mya).
The system of Odonata is still largely based on the typological classification by FRASER (1957), but within the lasttwo decades there have been several attempts towards aphylogenetic system of odonates (CARLE 1982; TRUEMAN
1996; LOHMANN 1996; BECHLY 1999; REHN 2003). Differences between the results of these attempts are based on
different selection of characters or even more so on different methodological approaches (e.g., traditional Hennigian
Phylogenetic Systematics versus computer-based numericalanalysis).
There is a broad consensus that Epiophlebiidae and Anisoptera are both monophyletic, while ‘Anisozygoptera’ is aparaphyletic assemblage of Epiophlebiidae and fossil stemgroup representatives of Anisoptera.
The monophyly of Anisoptera is supported by numerous
morphological autapomorphies (sperm vesicle developed
as copulatory organ, wing venation with hypertriangle, triangle, subtriangle and anal loop, larval locomotion by jetpropulsion) and this is also the case for Epiophlebiidae(hamuli posteriores developed as copulatory organ, interocellar lobe, ovoid pedicel, hairy eye tubercle, larval stridulation organs). It is also undisputed that Epiophlebiidae isthe sister group of Anisoptera, because there are several
good synapomorphies (discoidal cell distally distinctly
widened in hindwing, male hindwing with anal angle,
males with a secondary epiproctal projection, synthorax
with the dorsal portion of the interpleural suture suppressed,larvae with anal pyramid).
Concerning Zygoptera, most recent authors consideredthem as monophyletic, while TRUEMAN (1996), in a cladistic
analysis of wing venational characters, suggested that
Zygoptera is a highly paraphyletic group, as already indicatedin FRASER (1957). However, the monophyly of Zygoptera is
supported by several strong putative autapomorphies, suchas the transverse head, the more oblique pterothorax, abdominal sternites with triangular cross-section and longitudinal keel, formation of an ovipositor-pouch by the enlarged
outer valves (valvula 3 = gonoplacs) of the 9th abdominal
sternite, and of course the highly specialized ligula that isdeveloped as copulatory organ. The presence of caudal
gills, even though uniquely present in Zygoptera among
extant odonates, has been demonstrated to be a symplesiomorphy by the finding of a fossil dragonfly larva. This larva
has wing sheaths that clearly show the characteristic veinal
features of the isophlebiid stemgroup representatives of
Anisoptera, but still possesses three caudal gills.
A detailed phylogenetic system of fossil and extant odonates with all synapomorphies, based on my results, is available at http://www.bechly.de/phylosys.htm.

A recent cladistic study of 122 morphological characters by
REHN (2003) basically confirmed this phylogeny; this includes the sistergroup relationship of Tarsophlebiidae and crowngroup Odonata, the monophyly of Zygoptera, a lestinoid +coenagrionoid clade which is sister group to Calopterygoidea, the position of the relict damselfly Hemiphlebia at thevery base of lestinoid zygopteres, the position of Petaluridae at the base of Anisoptera, and the sistergroup relationshipof African Coryphagrionidae to the Neotropical Pseudostigmatidae. The only clear differences concern the positions ofamphipterygid and megapodagrionid damselflies, which
REHN (2003) proposes to represent a paraphyletic basal grade
towards the lestinoid + coenagrionoid clade. However, thewide separation of the zygopteran genera Diphlebia andPhiloganga in this phylogeny appears doubtful, because
these two genera are united by very strong larval synapomorphies and some synapomorphies of the imagines.
Recent molecular studies on the higher phylogeny of odonates (MISOF & RICKERT 1999a, b) did not resolve theZygoptera problem and did partly even conflict with monophyla like Cavilabiata (including Cordulegastridae,
Neopetaliidae, Chlorogomphidae and libelluoids) that arevery well-established by morphological evidence beyond
reasonable doubt. Methodological artefacts like longbranch-attraction and noise seem to be prevalent.
The interpretation of the three different types of secondary
copulatory organs as autapomorphies of the three extant
suborders respectively is based on a groundplan reconstruction of the male secondary genital apparatus. This groundplan includes small hamuli anteriores and posteriores, asmall unsegmented ligula and an undifferentiated vesicula
spermalis. This hypothetical reconstruction has been confirmed by the discovery of a well-preserved male specimenof Tarsophlebiidae, the putative fossil sister group of all
extant odonates, which shows exactly this type of genitalia(Fig. 1). Consequently, none of the substructures was suited
as intromittent organ for sperm transfer in the ground plan,so that there still must have been a mechanism involving
external spermatophores. This is confirmed by the findingthat the primitive protodonate Namurotypus sippeli fromthe Carboniferous of Germany did not yet posses a secondary male genital apparatus, but primary genitalia that are
most similar to those of Zygentoma, which deposit spermatophores. The curious odonate mating wheel probably evolved by attaching the spermatophore on the sternites of thebasal male abdomen.
Concerning the phylogenetic position of the order Odonata
in the tree of insects the evidence is ambiguous. Fossil evidence and some morphological and molecular characters
support the monophyly of Palaeoptera (= Palaeodictyopteroida + Ephemeroptera + Odonata), while rather strong
characters of the extant head morphology (STANICZEK 2000)
and some molecular data support the monophyly of
Metapterygota (= Odonata + Neoptera). Consequently, thisissue still has to be considered as more or less unresolved.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Fig. 1. Tarsophlebia eximia, Upper Jurassic Solnhofen Limestone, male specimen no. SOS 1720 at the Jura Museum in Eichstätt(Germany), camera lucida drawing of ventral side of abdomen with secondary genital apparatus.

References
CARLE, F.L. 1982. The wing vein homologies and phylogeny of
the Odonata: a continuing debate. – S.I.O. Rapid Communications 4: x+66 pp.
BECHLY, G. 1999. Phylogeny and systematics of fossil dragonflies (Insecta: Odonatoptera) with special reference to some
Mesozoic outcrops. – Ph.D. thesis, Eberhard-Karls-University Tübingen; x+755 pp., 4 tabs., 143 textfigs., 70 pls.
FRASER, F.C. 1957. A reclassification of the order Odonata,based on some new interpretations of the venation of the
dragonfly wing. – Handbook Royal Zoological Society of
New South Wales 12: 1–133.
LOHMANN, H. 1996. Das Phylogenetische System der Anisoptera (Odonata). – Entomologische Zeitschrift 106(6):209–252 (first part); 106(7): 253–296 (second part);
106(9): 360–367 (first postscript).
MISOF, B. & A. RICKERT 1999a. A molecular approach to thephylogeny of extant Odonata. – Zoology 102, suppl. II: 9.
MISOF, B. & A. RICKERT 1999b. A molecular phylogeny of
Anisoptera. – Zoology 102, suppl. II: 9.
REHN, A.C. 2003. Phylogenetic analysis of higher-level relationships of Odonata. – Systematic Entomology 28:
181–239.
STANICZEK, A.H. 2000. The mandible of silverfish (Insecta:
Zygentoma) and mayflies (Ephemeroptera): its morphology and phylogenetic significance. – Zoologischer
Anzeiger 239(2): 147–178.
TRUEMAN, J.W.H. 1996. A preliminary cladistic analysis ofodonate wing venation. – Odonatologica 25(1): 59–72.

Morphological Support of the Major Clades of Plecoptera
PETER ZWICK
Limnologische Fluss-Station Schlitz des Max-Planck-Instituts für Limnologie, Damenweg 1, 36110 Schlitz, Germany[pzwick@mpil-schlitz.mpg.de]

The monophyly of the order Plecoptera is beyond doubt, but
the position of Plecoptera among the Neoptera continues tobe debated. Ordinal apomorphies are few: the loop-shaped
gonads, loop-shaped seminal vesicles, absence of an ovipositor, presence of a cercus heart, and oblique intersegmental
musculature supporting the laterally (!) undulating larvalswimming movements.
Since the earliest studies, two clades were distinguishedwithin Plecoptera, at various levels between genus (Perlaversus Nemoura) and suborder (Setipalpia versus Filipalpia,
or Systellognatha versus Holognatha) (KLAPÁLEK 1905;
ENDERLEIN 1909). The competing suborder designations
reflect differences of opinion that were actually formalistic
and marginal; the disagreement is easily overcome by consistent phylogenetic methodology.
Contemporary systems are based on a large number of characters of external and internal morphology that were studied across all families of the order, and by some cytological and behavioural characters (ZWICK 1973, 2000). The

relationships of the four endemic southern hemispherefamilies had previously not been satisfactorily recognized.
Several of the traditionally recognized major taxa proved tobe monophyletic and persist in the present classification.
The Systellognatha are supported, among other, by the
reduction of the adult mandibles, by a complex set of gills
in particular positions on the thorax, complicated male epiprocts involving numerous movable components, and profound modifications of abdominal tergite 10; variations canbe followed across all families. Eggs are hard-shelled, witha suction-disk-like attachment organ, the anchor, at theposterior pole. The first larval instar lives on yolk remains
in the gut, only the 2nd instar is actively feeding. The female receptacle carries a number of accessory glands along its
stalk-like base. The vast majority of all Systellognatha
belong to the Perloidea, which share carnivorous larvae
with a modified, slender type of mandible (some adults inthis group with secondarily functional mandibles possess
the same derived type of mandible), tiny glossae that areEntomologische Abhandlungen 61 (2)129

Fig. 1. Cladogram of extant Plecoptera (from ZWICK 2000). Apomorphic characters supporting monophyly of taxa numbered.
For character descriptions see the original paper.

much smaller than the paraglossae, and long, slender palpi.
The embryo hatches through a defined opercle at the anterior egg pole. The number of free abdominal ganglia is
reduced, etc. Within Systellognatha, the monophyly of the
Perlodidae is still weakly supported. Consequently, the relations of Perlodidae with the other two families in superfamily Perloidea are unclear. Only the Perlidae have also colo-

nized large parts of the southern hemisphere, all other
Systellognatha are holarctic endemics.
The main apomorphies shared by the Euholognatha are an
unpaired corpus allatum fused to the aorta, and the reduction of chorionic sclerotization: euholognathan eggs are
soft, with a delicate sticky shell. The Euholognatha include
a few wingless species of Scopuridae, and the large com-130

plex of the Nemouroidea. The latter are distinguished bynumerous modifications of the thorax, especially through atiny furcasternum on which the furcal bases and spina arelocated and interconnected by sclerite bridges. There are
extra tergopleural flight muscles. Also, a number of attachment points of other muscles have shifted away from ancestral positions. Abdominal sternite 10 is reduced, the paraprocts therefore lie directly behind sternite 9. Males have acharacteristic pedunculate ventral vesicle with soft surface
at the base of sternite 9 that is used in drumming (seebelow). The mode of sperm transfer via accordingly modified paraprocts, epiprocts, or directly the gonopore on theelongated sternite 9 provide means to convincingly distinguish and group the included families as shown in Fig. 1.
Some Nemouridae and Leuctridae advance into the Oriental
region but otherwise the Euholognatha are of holarctic distribution, except the disjunct, exclusively southern hemisphere Notonemouridae. These pose presently unsolved problems. In all probability they are a paraphyletic assemblageof ancient surviving lines of early Nemouroidea.
A syndrome of structural and behavioural characters related
to sexual communication via substrate vibrations strongly
supports the sistergroup relationship between Systellognatha
and Euholognatha, which together from the suborder Arctoperlaria.
The southern hemisphere fauna comprises (in addition to
the Arctoperlaria: Perlidae and Notonemouridae, both ofwhich occur also in Africa) four endemic monophyleticfamilies (none of them extant in Africa). For a long time,family interrelations were doubtful. Two of the families
were for some time regarded as a separate suborder,
Archiperlaria, the alleged archaic sister group of all other
Plecoptera (ILLIES 1960, 1965), while the other two families
were supposedly related to the Filipalpia. However, all fourfamilies share unique details of front leg musculature,
namely absence of a tergal and presence of an additional
sternal depressor of the trochanter. Also, all four families
possess the complicated floriform osmoregulatory cells,
which occur only in them. Apparently, the four families inquestion represent a monophylum, the Antarctoperlaria(ZWICK 1973, 1980). Since their members show none of theconstitutive characters of any of the families or higher-ranked subgroups of the Arctoperlaria, Arctoperlaria andAntarctoperlaria are obviously sister groups. They are pre-Proceedings of 1st Dresden Meeting on Insect Phylogeny

sently recognized as the two largely vicariant suborders of
Plecoptera (Fig. 1).
Superfamily Eusthenioidea includes the largest
Antarctoperlaria. They have gills on basal abdominal segments that stand in ventrolateral positions and appear to bea plesiomorphy. Pteronarcyidae and several other relatively
primitive Arctoperlaria have similar gills. Details of thestructure and musculature of these gills are characteristic ofeach of the two families, Eustheniidae (carnivorous) and
Diamphipnoidae (detritivorous).
Members of superfamily Gripopterygoidea range from verylarge species with reticulate wing venation to tiny ones withmuch reduced venation; they are essentially detritivores andlack paired abdominal gills. Gripopterygidae andAustroperlidae instead have gills at the abdominal tip, eachfamily of a characteristic kind. The close relationship of thetwo families is mainly evidenced by unique modificationsof inner genitalia, namely reduction of a seminal receptaclein females, and development of large accessory maleglands.

References
ENDERLEIN, G. 1909. Klassifikation der Plecopteren sowie
Diagnosen neuer Gattungen und Arten. – Zoologischer
Anzeiger 34: 385–419.
ILLIES, J. 1960. Archiperlaria, eine neue Unterordnung der
Plecopteren (Revision der Familien Eustheniidae und
Diamphipnoidae) (Plecoptera). – Beiträge zur
Entomologie 10: 661–697.
ILLIES, J. 1965. Phylogeny and zoogeography of the
Plecoptera. – Annual Review of Entomology 10:
117–140.
KLAPÁLEK, F. 1905. Conspectus Plecopterorum Bohemiae. –Çasopis Çeské Spoleçnosti Entomologické 2: 27–32.
ZWICK, P. 1973. Insecta: Plecoptera. Phylogenetisches System
und Katalog. – Das Tierreich 94: i-xxxii, 1–465. – De
Gruyter, Berlin, New York.
ZWICK, P. 1980. 7. Ordnung Plecoptera (Steinfliegen). Pp.
1–115 in: J.G. HELMCKE, D. STARCK & H. WERMUTH (eds.),
Handbuch der Zoologie 4 (2) 2/7. – De Gruyter, Berlin,
New York.
ZWICK, P. 2000. Phylogenetic system and zoogeography of the
Plecoptera. – Annual Review of Entomology 45: 709–746.

Phylogeny of Plecoptera: Molecular Evidence and Evolutionary Trends
MATTHEW D. TERRY & MICHAEL F. WHITING
Department of Integrative Biology, Brigham Young University, Provo, Utah 84602, USA [Matthew_Terry@byu.edu;
Michael_Whiting@byu.edu]

Phylogenetic analysis of morphological data (ZWICK 2000)
and six molecular markers (12S, 16S, 18S and 28S rDNA;
and cytochrome oxidase II and histone 3) including 173
Plecoptera species from all extant families and multiple
outgroup exemplars supports a monophyletic Plecoptera
basal to all remaining Neoptera. Subordinal relationships
within Plecoptera are largely congruent with previous morphological hypotheses with a few notable exceptions (M.D.
Terry & M.F. Whiting in prep.: “Phylogeny of Plecoptera:evidence from a combined analysis of six molecular markers and morphology”). Systellognatha and Antarctoperlaria
are monophyletic; however, Euholognatha is paraphyletic

due to its placement as the first several basal clades within
Plecoptera. Under sensitivity analysis this paraphyly is always
supported, however, the relationships within Euholognatha
vary slightly. Nemouridae form the basal most clade, followed by two clades of the remaining euholognathan families.
Notonemouridae (exemplars from Africa, Australia/New
Zealand, and South America) are supported as a monophyletic group and not a “gradotaxon” sensu Zwick. Gripopterygidae appear to be paraphyletic with the Austroperlidae nesting within. Megaleuctra is not supported as amember of the Leuctridae and deserves recognition as its
own family (‘Megaleuctridae’). These results conflict with

131

Entomologische Abhandlungen 61 (2)several morphological features, particularly the paraphylyof Euholognatha in light of the presence of an unpaired corpus allatum; and future studies will need to more fully
examine molecular support for these relationships in lightof morphological characters. Patterns of geographical distribution are consistent with early diversification of basal
lineages prior to the break up of Pangea, the survival of only
two lineages (progenitors of Notonemouridae and Antarcto-

perlaria) on Gondwanaland, and the more recent invasion ofSouth America and sub-Saharan Africa by small groups ofPerlidae.

References
ZWICK, P. 2000. Phylogenetic system and zoogeography of the
Plecoptera. – Annual Review of Entomology 45:
709–746.

Phylogenetics of Embioptera (= Embiidina)
CLAUDIA SZUMIK1, JANICE EDGERLY-ROOKS2 & CHERYL Y. HAYASHI3
1
CONICET, Instituto Miguel Lillo, Tucumán, Argentina [instlillo@infovia.com.ar] – 2 Department of Biology, Santa Clara
University, Santa Clara, USA [jedgerlyrooks@scu.edu] – 3 Department of Biology, University of California, Riverside, USA[chayashi@citrus.ucr.edu]

Introduction
Embioptera is an interesting order comprising relatively
uncommon, relictual, pseudosocial insects. Phylogenetic
studies of the Embioptera are still in their infancy. Despite
the small size of the order – with close to 350 described species (ROSS 2001, 2003a, 2003b) – there are no explicit proposals about the relationships between families and there is
no reason to believe that currently recognized families aremonophyletic units. Rather, a preliminary cladistic study on
the classification of the order (SZUMIK 1996) and subsequent analyses at the family level (SZUMIK 1994, 1998)
show that many of the currently recognized higher groups
are not monophyletic.
Most of the groups in the order have been recognizedalmost solely on the basis of male terminalia. These display
an enormous variety of processes, modified for copulation,and this has kept other potentially informative structures
almost completely ignored in traditional classifications of
Embioptera.
Given this situation, a morphological and molecular analysis including all the potentially useful characters and manytaxa representing all the major groups is strongly needed.

of the 8 families result clearly as monophyletic:
Teratembiidae, Anisembiidae and Australembiidae.
Notoligotomidae (from Java and Australia) results includedin the American-African Archembiidae, with these two
families together forming a monophyletic group which is
ambiguous in the combined data set but supported in themorphological analysis. Embiidae appears non-monophyletic in both analyses; some of its African genera are groupedwith the Australembiidae.
The major difference between the morphological and the
combined analyses is that in the former Teratembiidae +Oligotomidae result as the sister group of the remaining
non-clothodid Embioptera, while in the latter theAnisembiidae take this position.

Conclusion
The results indicate that many non-male-terminalia characters are also very informative; many of them support majorgroups. Given that some groups (like Archembiidae) are notwell supported by the molecular data, the combined data set
needs to be expanded in several ways (e.g., including moretaxa in the DNA data set, and including internal morphology and maternal behavior in the morphological data set).

Data and scope
Our morphological analysis includes 57 genera from 8
families, represented by 157 species. This sample represents about 70 % of the described embiid genera, and about
45 % of the known species. 182 morphological characters
were scored: 29 cephalic, 53 thoracic and abdominal, and100 from terminalia. These are characters that vary between
apparently distant groups as well as characters that only
distinguish genera or groups of genera that seem to be closely related. The molecular data set we analyzed consists ofsequences for 16S, 18S and 28S rDNA and COI for 22 species from 6 families and 15 genera. The trees were rooted
on the ‘primitive’ embiopteran Clothoda; given this, themonophyly and phylogenetic position of Clothodidae isobviously not tested here.

Results
The resulting trees from the morphological and from the
combined data set have well resolved basal clades; in otherwords, they have a good resolution of the higher groups. In
both cases (morphological and combined analyses) only 3

References
ROSS, E.S. 2001. The Embiidae of the Americas (Order
Embiidina). EMBIA, Part 3. – Occasional Papers of the
California Academy of Sciences 150: 1–86.
ROSS, E.S. 2003a. Andesembiidae, a new Andean family of
Embiidina. EMBIA, Part 4. – Occasional Papers of the
California Academy of Sciences 153: 1–13.
ROSS, E.S. 2003b. A review of the family Anisembiidae withdescriptions of new taxa. EMBIA, Part 5. – Occasional
Papers of the California Academy of Sciences 154: 1–123.
SZUMIK, C.A. 1994. Oligembia vetusta, a new fossil teratembiid (Embioptera) from Dominican amber. – Journal of the
New York Entomological Society 102: 67–73.
SZUMIK, C.A. 1996. The higher classification of the order
Embioptera: a cladistic analysis. – Cladistics 12: 41–64.
SZUMIK, C.A. 1998. Two new Neotropical genera of Embiidae(Embioptera, Insecta). – Journal of the New York
Entomological Society 105: 140–153.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Basal Diversification and the Evolution of Wings within Stick Insects(Phasmatodea)
SVEN BRADLER1, MICHAEL F. WHITING2 & REBECCA KLUG1
12Institut für Zoologie und Anthropologie, Berliner Str. 28, 37073 Göttingen, Germany [sbradle@gwdg.de; rklug@gwdg.de] –Department of Integrative Biology, Brigham Young University, Provo, Utah 84602, USA [Michael_Whiting@byu.edu]

The stick and leaf insects or Phasmatodea have recently
been subject to several phylogenetic analyses, based on
both morphological and molecular data. A sistergroup relationship of the wingless Nearctic genus Timema (17 described species) to all remaining Phasmatodea (ca. 3000 species), the Euphasmatodea, has appeared uncontroversial
including molecular studies (KRISTENSEN 1975; TILGNER etal. 1999; TILGNER 2002; BRADLER 2000, 2003; WHITING etal. 2003). Autapomorphies of the latter are hard-shelled,
seed-like eggs, the fusion of metatergum and abdominal tergum 1 and the reduced prothoracic sternal apophysis(furca), while a fusion of the three basal tarsomeres and thepresence of a mesal lobe on the right cercus of the male areapomorphies supporting the monophyly of Timema.
The primary internal splits of the Euphasmatodea remain
debatable (Fig. 1; BRADLER 2000, 2003; TILGNER 2002;
WHITING et al. 2003; ZOMPRO 2003). Molecular data suggest that several clades of presumably non-monophyletic‘Diapheromerinae’ form the basal lineages of extant
euphasmatodeans (Fig. 1B; WHITING et al. 2003). Taking
into consideration the basal position of Timema, this well
supported finding implies that the ancestral phasmid lacked

wings, followed by a long diversification of wingless taxa.
Consequently, wings must have been recovered in those clades that contain fully winged or brachypterous species.
Performing a cladistic analysis of morphological charactersof the adult female and egg capsule, TILGNER (2002) presents reasonable evidence that the Aschiphasmatinae, which
comprise several winged and wingless species and have notbeen included in any molecular analysis so far, form themost basal branch of extant euphasmatodeans (Fig. 1A).
The apomorphic characters supporting the sister group of
Aschiphasmatinae are the presence of a galealobulus, thefusion of coxopleurite and anapleurite in the prothorax, andthe absence of a pro-spina (character dot = cd 1 in Fig. 1;
TILGNER 2002).
Both studies contradict the view of BRADLER (2000, 2003)
that the wingless South American genus Agathemera is themost basal branch within the Euphasmatodea and sistertaxon to the Neophasmatidae (Fig. 1C). Agathemera exhibits a number of plesiomorphic characters such as an orthognathous head and relatively short thoracic and abdominal
segments with longitudinal musculature spanning the entire
segments as in Timema. In all Neophasmatidae the longitu-

Fig. 1. Alternative hypotheses of the basal splitting events within the Phasmatodea. A: After TILGNER (2002). B: After WHITING etal. (2003). C: After BRADLER (2000, 2003). D: After ZOMPRO (2003). The squares represent apomorphic (black) and plesiomorphic (white) characters resp. character dots (cd) referred to in text. Note that in Fig. 1D the name Phasmatodea is replaced by
Phasmatomorpha, Euphasmatodea by Phasmatodea, and Neophasmatidae by Verophasmatodea.

133

Entomologische Abhandlungen 61 (2)dinal muscles do not span the entire length of an abdominal
segment but are restricted to the posterior part of each segment (cd 2 in Fig. 1); at least on the ventral side this is likely due to a loss of the internal longitudinal muscles, only theshort external ventral muscles remaining (KLASS 1999;
BRADLER 2000, 2003). The latter apomorphy is also foundin the pregenital abdomen of Abrosoma festinatum (Fig. 2),
a Bornean member of the Aschiphasmatinae, thus contradicting the basal position of Aschiphasmatinae as proposed
by TILGNER (2002).
The mesonotal lobes of Agathemera have recently been
homologized with wings, incorrectly with hind wings (alae)
instead of tegmina (ZOMPRO 2003). As these lobes are positioned dorsally on the hind margin of the mesonotum, lackvenation and articulation, they are here not consideredhomologous with tegmina, in agreement withCAMOUSSEIGHT (1995). Hence, the possible basal placementof Agathemera would even strengthen the hypothesis ofwing recovery in phasmid evolution since it would form
another basal apterous lineage. However, according tomolecular data Agathemera is a subordinate taxon within
the Euphasmatodea (WHITING et al. 2003).

sister group of all extant Verophasmatodea (= Phylliidae +remaining Phasmatodea). The only character supporting thelatter is the possession of shortened tegmina (cd 3 in Fig. 1).
In the majority of (winged) phasmids the tegmina are reduced to small wing pads that cover the base of the hindwings. However, at least in some members, e.g., the male of
Malaysian Heteropteryx dilatata, the tegmina are almost as
long as the hind wings and reach the tip of the abdomen.
Hence, the minimal difference in length between fore andhind wings in Archipseudophasmatidae and other phasmids
can hardly be considered as a convincing apomorphy.
Within the extant Verophasmatodea ZOMPRO (2003) places
the Phylliinae (= Phylliidae sensu Zompro; walking leaves)as the basal side branch (Fig. 1D). While the Phylliinae
form a well supported clade based on the presence of a stridulatory organ in the antennae of females and juvenile
males, dilated sternites in the abdomen, and laterally fringed micropylar plate of the eggs (cd 4 in Fig.1), its surmised sister group is only founded on plesiomorphic characters: the absence of a stridulatory organ in the antennae,
sternites not dilated, and micropylar plate not fringed laterally (cd 5 in Fig. 1). Hence, the basal placement of the
Phylliinae appears unsupported. On the other hand, according to morphological (TILGNER 2002) and molecular data
(WHITING et al. 2003) the Phyllinae are a rather subordinate taxon among the Euphasmatodea (= Phasmatodea sensu
ZOMPRO).
In summary, the controversial assumptions presented hereon basal splitting events in the Euphasmatodea highlight theneed for future investigations of phasmid phylogeny. These
additional phylogenetic analyses must include a larger selection of taxa, additional molecular markers, and morphological data interpreted via a rigorous phylogenetic analysis.
The next step surely is the inclusion of Aschiphasmatinae inmolecular studies.

Acknowledgements
We like to thank Bernd Baumgart (Göttingen) for technicalsupport.

References➝

Fig. 2. Muscles and nerves of female abdominal segment 5 ofAbrosoma festinatum (Euphasmatodea: Aschiphasmatinae).
Internal view of right half of abdominal segment 5; orientation:➝ anterior, ↕ dorsal. The longitudinal muscles are shaded grey.
G5 = abdominal ganglion 5; A = dorsal nerve; B = lateralnerve; C = ventral nerve (R. Klug, original).

ZOMPRO (2003) agrees with BRADLER (2000, 2003) in theposition of Agathemera (Fig. 1D). In addition, ZOMPRO(2003) presents a phylogeny beyond the level of
Neophasmatidae (= Verophasmatodea therein, Fig. 1D).
The fossil Archipseudophasmatidae are supposed to be the

BRADLER, S. 2000. On the systematic position of AgathemeraStål 1875 within the Phasmatodea. – Zoology 103, Suppl.
III (Abstracts 93.1): 99.
BRADLER, S. 2003. Phasmatodea, Gespenstschrecken. Pp.
251–261 in: H.H. DATHE (ed.), Lehrbuch der Speziellen
Zoologie, I/5 Insecta. – Spektrum, Heidelberg, Berlin.
CAMOUSSEIGHT, A. 1995. Revision taxonomica del genero
Agathemera (Phasmatodea: Pseudophasmatidae) en
Chile. – Revista Chilena de Entomología 22: 35–53.
KLASS, K.-D. 1999. The pregenital abdomen of a mantid and a
cockroach: musculature and nerve topography, with comparative remarks on other Neoptera (Insecta:
Dictyoptera). – Deutsche Entomologische Zeitschrift 46:
3–48.
KRISTENSEN, N.P. 1975. The phylogeny of hexapod “orders”. A
critical review of recent accounts. – Zeitschrift für
Zoologische Systematik und Evolutionsforschung 13:
1–44.
TILGNER, E.H. 2002. Systematics of Phasmida. – Doctoral
Thesis, University of Georgia, USA.
TILGNER, E.H., T.G. KISELYOVA & J.V. MCHUGH 1999. A morphological study of Timema christinae VICKERY with
implications for the phylogenetics of Phasmida. –Deutsche Entomologische Zeitschrift 46: 149–162.
WHITING, M.F., S. BRADLER & T. MAXWELL 2003. Loss andrecovery of wings in stick insects. – Nature 421: 264–267.
ZOMPRO, O. 2003. Eine generische Revision derInsektenordnung Phasmatodea: Areolatae, einschließlich
der Einführung einer neuen Ordnung der Insekten. –Doctoral Thesis, University of Kiel, Germany.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Relationships among the Principal Lineages of Dictyoptera Inferredfrom Morphological Data
KLAUS-DIETER KLASS
Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, 01109 Dresden, Germany [klaus.klass@snsd.smwk.sachsen.de]Introduction
The Dictyoptera include three major subgroups (usuallyconsidered orders), each phenetically distinctive: Blattaria,
Isoptera, and Mantodea. While the monophyly of Dictyoptera is generally accepted, phylogenetic relationships within Dictyoptera have been strongly disputed in recent years.
This concerns mainly the relationships among the three‘orders’ and the basal splitting events in Blattaria.

Monophyly of Dictyoptera
Monophyly is well supported by some striking apomorphies: (1) the perforation in the tentorium through which
the ganglionic connectives pass; (2) the formation of afemale subgenital plate by coxosternum 7 (as in
Dermaptera) that bears a discrete dorsal vestibular sclerite(this is in addition to another, smaller subgenital plate formed by the hind part of venter 7: laterosternal shelf); (3) the
packing of the eggs in an ootheca that is built by secretions
from morphologically and biochemically asymmetricalaccessory glands of abdominal segment 9; and probably (4)the insertion of the abdominal spiracle dilators on the paratergite (rather than on the coxosternum). It should be notedthat in the Isoptera (4) has been demonstrated only for
Mastotermes, and (3) is present only in this genus (see also
NALEPA & LENZ 2000).

Important character systems
For reconstructing dictyopteran phylogeny three charactersystems have been used extensively that are very rich ininformative characters: the male genitalia (phallomeres),female genitalia (including the ovipositor), and proventricle(gizzard; the hindmost, cuticulized part of the foregut).
The phallomeres are in all Blattaria and Mantodea very
complex (due to their composition of many sclerites, formative elements such as processes, apodemes, tendons,pouches, etc., and muscles) and strongly asymmetrical, andacross both groups they show a great structural diversity(especially in Blattaria). This diversity requires – prior to
the definition of characters – an extensive analysis aimed at
the identification of corresponding parts among taxa (corresponding to alignment in molecular analysis; see KLASS
2001b). Reliable ‘alignment’ hypotheses require the inclusion of muscles and formative elements into phallomere
studies, because these provide additional landmarks for thealignment.
Phallomere characters are not applicable to Isoptera, whichhave strongly simplified male genitalia (KLASS 2001a;
KLASS et al. 2000). There is no outgroup comparison possible for phallomere characters, because corresponding phallomere components could not yet be identified between
Dictyoptera and other insects; trees based on phallomere
characters can thus only be rooted by including non-phallomere characters.
Phallomere characters were used in three extensive phylogenetic studies: MCKITTRICK (1964; non-numerical; additi-

ons in MCKITTRICK & MACKERRAS 1965), GRANDCOLAS(1994, 1996; numerical), and KLASS (1995, 1997; numerical re-analysis currently in preparation by K.-D. Klass & R.
Meier). However, in MCKITTRICK’s and especially
GRANDCOLAS’ contributions the study of cuticular morphology is very superficial, muscles are not considered, and thealignment is tentative (see KLASS 1997, 2001b for a detailed scrutiny). With the more detailed comparative analysis
by KLASS, which includes many additional cuticular elements and, partly, the muscles, a very different ‘alignment’
hypothesis results, which leads to a very different set of
informative characters, and eventually to a different phylogenetic hypothesis.
The female genitalia show a complicated pattern of sclerites, articulations, projections (e.g., the gonapophyses andgonoplacs), apodemes, the spermatheca, and the genital
opening. The area lateral to the bases of gonapophyses 8
and 9, where several sclerites contact each other, and thearea comprising the laterosternal shelf and the genital opening are especially rich in characters. The numerous
muscles of the female genitalia show a fairly constant arrangement and thus help in aligning the sclerites; on the other
hand, minor differences in the musculature yield informative characters.
Characters of the female genitalia are fully applicable to
Mastotermes among the termites, while their application toother termites is limited due to reductions. Outgroup comparison is possible for many female genitalic characters of
Dictyoptera but is ambiguous in some of these.
MCKITTRICK (1964) and MCKITTRICK & MACKERRAS(1965) illustrated the female genitalia of many species ofBlattaria, partly including the muscles. However, many
structural details remain unclear, and the descriptions are
altogether too incomplete for composing a reliable character matrix. The more detailed study of the exoskeleton ofeight dictyopteran species in KLASS (1998a) allows implications on the relationships among the three ‘orders’, but
for analysing relationships within these groups additional
taxa have to be studied.
The proventricle shows a pattern of longitudinal, partly
sclerotized folds (plicae) of different rank: wide primary
plicae usually bearing heavy sclerites and teeth; much narrower secondary and tertiary plicae. The proventricular plicae are essentially arranged following a hexaradial symmetry, which, however, is overlain by a bilateral symmetry. The
distinctness of the bilateral component provides the mostobvious point of variation: it varies from very strong (e.g.,
in Blattidae) to completely absent (e.g., in Isoptera), andseveral characters can be drawn from this.
It has long been assumed that a complete radial symmetryof the proventricle is plesiomorphic for Dictyoptera, and thebilateral component has developed within the group.
However, the proventricle shows a very similar bilateral pattern in many Blattaria and Zygentoma (KLASS 1998b).
Using this for outgroup comparison, strongly bilateral proventricles appear plesiomorphic; proventricle characters arethen much more congruent with the phallomere characters,
so this reversed polarity assumption appears supported.

Entomologische Abhandlungen 61 (2)135

Fig. 1. Phylogenetic relationships in Dictyoptera. Relationships in Blattaria and Mantodea based mainly on phallomere studies(mental analysis by KLASS 1995, 1997 and ongoing numerical analyses by K.-D. Klass & R. Meier). Inclusion of Isoptera basedon KLASS (1995) and DEITZ et al. (2003). ‘Blattodea’ is used for comprising Blattaria and Isoptera.

Phylogenetic relationships among Blattaria,
Isoptera, and Mantodea
The point of dispute is here the placement of the eusocial
Isoptera as the sister group (1) of Blattaria + Mantodea, or(2) of Blattaria alone, or (3) of the subsocial blattarian
genus Cryptocercus, which shows many similarities in life
history with the termites. A numerical analysis by THORNE& CARPENTER (1992) suggested the topology Isoptera +(Blattaria + Mantodea). However, a recent revision of thesame data set by DEITZ et al. (2003) demonstrated that in
THORNE & CARPENTER (1992) numerous misscorings oftaxa, disputable polarity assumptions, and cases of misinterpreted literature data were included. DEITZ et al. (2003)
obtained Mantodea + (Blattaria excluding Cryptocercus +(Cryptocercus + Isoptera)), with all nodes strongly supported, thus confirming the hypothesis of KLASS (1995). In
DEITZ et al. (2003) seven characters were added to THORNE& CARPENTER’s (1992) data set, but the latter topology also
resulted using the revised original data set alone.
Due to the vast incompleteness of the available data, in bothanalyses Blattaria excluding Cryptocercus constitute asingle terminal, while actually each major blattarian lineage
should be included as a separate terminal. This has theeffects that the support for a clade Cryptocercus + Isoptera
may be weakened, and the result cannot tell whether thisclade, if existing, is the sister group or a subgroup of theremaining Blattaria.
The monophyly of Blattaria + Isoptera (= Blattodea) is supported by: (1) the loss of the unpaired ocellus; (2) the fat
body harbours symbiotic Eubacteria of the genus
Blattabacterium (only Mastotermes among termites; see LO
et al. 2003); (3) the cardiac valve at the transition between
proventricle and midgut is strongly elongated; (4) the proventricle bears sclerites on the secondary plicae. Two additional characters from the female genitalia depend on thefact that Blattaria and Mastotermes among Isoptera build
the ootheca in the vestibulum (space above subgenital platecoxosternum 7), while Mantodea build it on the substrate:(5) the ovipositor in Blattaria and Isoptera is shorter than in
Mantodea (ending in the vestibulum); (6) as a casting mould
for the ootheca Blattaria and Mastotermes have a pair of
discrete membranous folds on the subgenital plate.
The monophyly of Cryptocercus + Isoptera is supported bycharacters of the proventricle and life history, e.g.: (1) Cryptocercus has the bilateral symmetry component in the proventricle strongly reduced, closely approaching the purely
radial conditions in the Isoptera (partly paralleled in someother blattarian lineages); (2) the primary pulvilli (soft cushions formed by the posterior part of the primary plicae)
form huge bulbs; (3) the teeth are laterally compressed, ±blade-like; (4) the hindgut harbours a unique diversity of136

Proceedings of 1st Dresden Meeting on Insect Phylogeny

oxymonadid and hypermastigid flagellates, with several flagellate taxa shared between Cryptocercus and basal
Isoptera (see KLASS 2001b with regard to GRANDCOLAS &DELEPORTE’s 1996 hypothesis on this issue); (5) the flagellates are transfected to the progeny by anal trophallaxis.
Further similarities in life history add to this.
The monophyly of Mantodea is mainly supported by: (1)specializations of the raptorial forelegs, including the presence of a femoral brush of setae (cleaning device); (2)some peculiarities in the strongly derived proventricle (e.g.,loss of one primary tooth; presence of extensive system of
elevated grooves; KLASS 1998b); (3) presence of a transverse apodemal ridge across the pronotum, which externally isusually visible as a ‘supracoxal groove’; (4) fusion of the
2nd and 3rd abdominal ganglia to the 1st abdominal andmetathoracic ganglia (KLASS 1999).

+ Blaberidae. These issues are comprehensively discussed
in KLASS (1997, 2001b).

Basal splitting events in Blattaria and Mantodea

References

The phylogenetic results in KLASS (1995, 1997) are basedon a mental evaluation of phallomere characters. A numerical analysis using the same data set (though with some additions) is presently conducted by K.-D. Klass & R. Meier andleads to a nearly identical hypothesis (Fig. 1). In the
Blattaria sample (20 species) all ‘families’ from the classification of MCKITTRICK (1964) are represented (Blattidae,
Cryptocercidae, Polyphagidae, Blattellidae, Blaberidae) as
well as many ‘subfamilies’. In the Mantodea sample (4 species) the three monogeneric families are represented thathave been considered most primitive by BEIER (1968)(Mantoididae, Chaeteessidae, Metallyticidae); Sphodromantis represents the strongly derived Mantidae, while theremaining families of the system by BEIER (1968) were notincluded (Eremiaphilidae, Amorphoscelidae, Empusidae,
Hymenopodidae). Of the 107 phallomere characters included in the matrix, roughly equal partitions come from thesclerites, formative elements, and muscles. The numericalanalyses yielded four most parsimonious trees, of which thestrict consensus is shown in Fig. 1 (with resolution lacking
only in the clade comprising the Blattellinae, Nyctiborinae,
and Blaberidae).
In the Mantodea, Mantoida constitutes the most basal clade,followed by Chaeteessa (both genera neotropical); both
nodes are strongly supported. Mantoida shows several‘blattarian’ features in its phallomeres that are uniqueamong Mantodea. Two of the autapomorphies of the clade
Metallyticus + Sphodromantis were also consistently foundin the phallomeres of various Eremiaphilidae, Amorphoscelidae, Empusidae, Hymenopodidae, and Mantidae
described by LAGRECA (1954) (other autapomorphies
mostly not visible therein), supporting the view that
Mantoida and Chaeteessa are actually the most basal
Mantodea.
In the Blattaria, the Blattidae s.str. (excluding Tryonicinaeand Lamproblattinae) is obtained as the sister group of theremaining cockroaches. The next branch is the blattid subfamily Tryonicinae. The remaining Blattaria fall into two
clades, Cryptocercidae + (Lamproblattinae + Polyphagidae)
and Blattellidae + Blaberidae. In the latter clade,Anaplectinae are basal, followed by paraphyletic
Plectopterinae (= Pseudophyllodromiinae) and a clade
comprising Blattellinae, Nyctiborinae, and the ovoviviparous Blaberidae (and probably Ectobiinae). The placementof the Isoptera in Fig. 1 (as the sister group of Cryptocercus) is based on the results of DEITZ et al. (2003).
The phylogenetic hypothesis of GRANDCOLAS (1994, 1996,1997) differs from that in KLASS (1995, 1997) mainly by a
deeply subordinate position (1) of Tryonicinae andLamproblattinae in Blattidae and (2) of Cryptocercus in
Polyphagidae, and by (3) monophyletic Plectopterinae (=Pseudophyllodromiidae) and (4) monophyletic Plectopterinae

BEIER, M. 1968. Mantodea (Fangheuschrecken). Pp. 1–47 in:
J.G. HELMCKE, D. STARCK & H. WERMUTH (eds.),
Handbuch der Zoologie 4 (2) 2/12. – De Gruyter, Berlin.
DEITZ, L.L., C.A. NALEPA & K.-D. KLASS 2003. Phylogeny ofDictyoptera re-examined. – Entomologische Abhandlungen 61: 69–91.
GRANDCOLAS, P. 1994. Phylogenetic systematics of the subfamily Polyphaginae, with the assignment of Cryptocercus
Scudder, 1862 to this taxon (Blattaria, Blaberoidea,
Polyphagidae). – Systematic Entomology 19: 145–158.
GRANDCOLAS, P. 1996. The phylogeny of cockroach families: acladistic appraisal of morpho-anatomical data. –Canadian Journal of Zoology 74: 508–527.
GRANDCOLAS, P. 1997. Systématique phylogénétique de lasous-famille des Tryonicinae (Dictyoptera, Blattaria,
Blattidae). In: J. NAIT & L. MATILE (eds.), Zoologia
Neocaledonica, vol. 4. – Mémoires du Muséum Nationald’Histoire Naturelle Paris 171: 91–124.
GRANDCOLAS, P. & P. DELEPORTE 1996. The origin of protistan
symbionts in termites and cockroaches; a phylogenetic
perspective. – Cladistics 12: 93–98.
KLASS, K.-D. 1995. Die Phylogenie der Dictyoptera. – Ph.D.
dissertation, University Munich; Cuvillier, Göttingen.
KLASS, K.-D. 1997. The external male genitalia and the phylogeny of Blattaria and Mantodea. – Bonner Zoologische
Monographien 42: 1–341.
KLASS, K.-D. 1998a. The ovipositor of Dictyoptera (Insecta):
homology and ground-plan of the main elements. –Zoologischer Anzeiger 236: 69–101.
KLASS, K.-D. 1998b. The proventriculus of Dicondylia, withcomments on evolution and phylogeny in Dictyoptera andOdonata. – Zoologischer Anzeiger 237: 15–42.
KLASS, K.-D. 1999. The pregenital abdomen of a mantid and a
cockroach: musculature and nerve topography, with comparative remarks on other Neoptera (Insecta:
Dictyoptera). – Deutsche Entomologische Zeitschrift 46:
3–42.
KLASS, K.-D. 2001a [dated 2000]. The male abdomen of therelic termite Mastotermes darwiniensis (Insecta: Isoptera:
Mastotermitidae). – Zoologischer Anzeiger 239:
231–262.
KLASS, K.-D. 2001b. Morphological evidence on blattarian
phylogeny: “phylogenetic histories and stories” (Insecta:
Dictyoptera). – Deutsche Entomologische Zeitschrift 48:
223–265.
KLASS, K.-D., B.L. THORNE & M. LENZ 2000. The male postabdomen of Stolotermes inopinus: a termite with unusually well-developed external genitalia (Dictyoptera:
Isoptera: Stolotermitinae). – Acta Zoologica (Stockholm)
81: 121–130.
LAGRECA, M. 1954. Sulla struttura morfologica dell’apparato
copulatore dei Mantodei. – Annali dell’Istituto Superiore
di Scienze e Lettere di Santa Chiara, Napoli, 1953/54:
1–28.
LO, N., C. BANDI, H. WATANABE, C.A. NALEPA & T. BENINATI
2003. Evidence for cocladogenesis between diverse dictyopteran lineages and their intracellular endosymbionts. –Molecular Biology and Evolution 20: 907–913.

Future tasks
The most important tasks in the morphology-based reconstruction of dictyopteran phylogeny are the extension ofphallomere studies to a larger taxon sample, and the studyof the female genitalia in all species sampled for the phallomere studies. Here it is required that morphology is analysed in great detail and that the musculature is also included. This secures a reliable identification of correspondingparts among taxa and also yields a much greater amount of
informative characters. The search for new characters in allparts of the body is important with regard to the inclusionof Isoptera, which have strongly reduced genitalia.

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Entomologische Abhandlungen 61 (2)MCKITTRICK, F.A. 1964. Evolutionary studies of cockroaches.
– Cornell University, Agricultural Experiment Station
Memoir 389: 1–197.
MCKITTRICK, F.A. & J.M. MACKERRAS 1965. Phyletic relationships within the Blattidae. – Annals of the Entomological Society of America 58: 224–230.

NALEPA, C.A. & M. LENZ 2000. The ootheca of Mastotermes
darwiniensis Froggatt (Isoptera: Mastotermitidae): homology with cockroach oothecae. – Proceedings of the Royal
Society of London, Series B, 267: 1809–1813.
THORNE, B.L. & J.M. CARPENTER 1992. Phylogeny of the
Dictyoptera. – Systematic Entomology 17: 253–268.

Molecular Phylogenetics of Dictyoptera: Insights into the Evolution ofTermite Eusociality and Bacterial Endosymbiosis in Cockroaches
NATHAN LO
DIPAV, Sezione di Patologia Generale e Parassitologia, Universitá degli Studi di Milano, Milan 20133, Italy [nathan.lo@unimi.it]

The Dictyoptera is a well-accepted monophyletic group,
consisting of three easily recognized insect types: cockroaches, termites and praying mantids. Cockroaches are wellknown for their relatively detailed fossil record, though theearly roach-like fossils dating back to the Upper
Carboniferous (~300 million years ago), which dominatedthe terrestrial fauna of this ancient era (CARPENTER 1980),
were almost certainly stemgroup representatives of the
Dictyoptera. Termites have the relatively rare traits among
animals of wood-feeding and highly eusocial behaviour, thelatter of which is present in all species. Finally, praying
mantids are famous for their striking morphological appearance, and their strict predatory diet.
As is the case for many insect groups, relationships within
the Dictyoptera have been the subject of controversy forseveral decades (NALEPA & BANDI 2000; DEITZ et al. 2003),
and this controversy has extended into recent molecularstudies. The questions of whether praying mantids and termites are each monophyletic have never been in doubt;
however, the monophyly of the cockroaches – which mightseem reasonable based on the overall uniformity in the bodyplan of all cockroaches – has been disputed. The two main
rival hypotheses of basal dictyopteran relationships havebeen: (1) a close relationship between praying mantids andall cockroaches, with termites as the outlier (e.g., THORNE& CARPENTER 1992); (2) a close relationship between termites and wood-feeding cockroaches of the genus
Cryptocercus, with this clade being either separate to, or
nested within, other cockroaches. In the latter scenario,
Mantodea are considered the outlier to cockroaches and termites (NALEPA & BANDI 2000; DEITZ et al. 2003).
Four genes have now been sequenced from various representatives of all three insect ‘types’: the mitochondrial
genes 12S rDNA, 16S rDNA and cytochrome oxidase II,
and the nuclear 18S rDNA gene. Initial studies of 12S
rDNA and 16S rDNA supported hypothesis (1) above(KAMBHAMPATI 1995). A study of cytochrome oxidase II
that did not include Cryptocercus indirectly supported
hypothesis (2) (MAEKAWA et al. 1999). Following this, a
study of 18S rDNA, cytochrome oxidase II as well as endogenous cellulase genes of termites and cockroaches supported hypothesis (2) (LO et al. 2000). In a recent study, all fourgenes available for mantids, termites and cockroaches werecombined and shown to support hypothesis (2) (LO et al.
2003). Included in this study was an examination of thephylogeny of bacterial symbionts present in one primitive
termite (Mastotermes darwiniensis) as well as all cockroaches. These symbionts are apparently transmitted strictly
vertically and are expected to be obligate mutualists, conditions which should lead to phylogenetic congruence between host and symbiont. Evidence for such congruence
was indeed found, and the phylogeny of the endosymbionts
was shown to support a termite-Cryptocercus relationship.
Estimates of divergence times between the main lineages of

cockroaches indicated that these occurred at the beginningof the Cretaceous period (~130 mya), not the Carboniferous.
Thus there is now growing molecular evidence that termites
are most closely related to the cockroach genus
Cryptocercus, which would render the cockroaches paraphyletic. There is also some evidence for Mantodea being
the most basal lineage of the Dictyoptera, although this is
not strong. With regard to the evidence for a termiteCryptocercus grouping, two things should be considered.
The first is that for two of the genes that show strong support for this grouping (endogenous cellulases and endosymbiont 16S rDNA), no mantid sequences are available,
because their diet does not contain cellulose, and because
they do not harbour the relevant endosymbionts. Thus it isconceivable that the absence of mantids in these analyses
might lead to incorrect relationships being estimated fortermites and cockroaches. The second consideration istaxon sampling for the cockroaches. Although representatives of each of the five traditional families of cockroaches
have been examined, it is possible that these morphologybased family classifications are misleading, and that several other important high-level (basal) cockroach lineages
exist. Recent morphological studies by KLASS (1995, 1997)
indeed suggest that this is the case. Inclusion of extra
cockroach taxa in molecular studies may show that one ofthese is more closely related to termites than Cryptocercus,
although several shared characteristics between these twogroups make this unlikely. One distinct possibility thatshould be explored in future studies is that mantids are thesister group of another lineage of cockroaches, which
would render modern cockroaches polyphyletic with respect to both termites and mantids.
Future molecular studies of basal dictyopteran relationships
should focus on deciphering the number of deep branching
cockroach lineages that exist, perhaps using genes such as
cytochrome oxidase or 18S rDNA. Following this, many
additional nuclear genes will likely be required to understand relationships between these main lineages, and between these and termites and praying mantids.

References
CARPENTER, F.M. 1980. Studies of North American
Carboniferous insects. 6. Upper Carboniferous insectsfrom Pennsylvania. – Psyche 87: 109–119.
DEITZ, L.L., C.A. NALEPA & K.-D. KLASS 2003. Phylogeny ofDictyoptera re-examined. – Entomologische Abhandlungen 61: 69–91.
KAMBHAMPATI, S. 1995. A phylogeny of cockroaches and related insects based on DNA sequence of mitochondrial
ribosomal RNA genes. – Proceedings of the National
Academy of Sciences, USA, 92: 2017–2020.

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KLASS, K.-D. 1995. Die Phylogenie der Dictyoptera. – Ph.D.
dissertation, University Munich; Cuvillier, Göttingen.
KLASS, K.-D. 1997. The external male genitalia and the phylogeny of Blattaria and Mantodea. – Bonner Zoologische
Monographien 42: 1–341.
LO, N., C. BANDI, H. WATANABE, C.A. NALEPA & T. BENINATI
2003. Evidence for cocladogenesis between diverse dictyopteran lineages and their intracellular endosymbionts. –Molecular Biology and Evolution 20: 907–913.
LO, N., G. TOKUDA, H. WATANABE, H. ROSE, M. SLAYTOR, K.
MAEKAWA, C. BANDI & H. NODA 2000. Evidence frommultiple gene sequences indicates that termites evolved
from wood-feeding cockroaches. – Current Biology 10:
801–804.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

MAEKAWA, K., O. KITADE & T. MATSUMOTO 1999. Molecularphylogeny of orthopteroid insects based on the mitochondrial cytochrome oxidase II gene. – Zoological Science
16: 175–184.
NALEPA, C.A. & C. BANDI 2000. Characterizing the ancestors:
paedomorphosis and termite evolution. Pp. 53–75 in: T.
ABE, D.E. BIGNELL & M. HIGASHI (eds.), Termites:
Evolution, Eusociality, Symbioses, Ecology. – Kluwer
Academic Publishers, Dordecht, The Netherlands.
THORNE, B.L. & J.M. CARPENTER 1992. Phylogeny of the
Dictyoptera. – Systematic Entomology 17: 253–268.

Phylogeny of Mantodea Based on Molecular Data:
Evolution of a Charismatic Predator
GAVIN J. SVENSON & MICHAEL F. WHITING
Department of Integrative Biology, Brigham Young University, Provo, Utah 84602, USA [empusid@email.byu.edu;
Michael_Whiting@byu.edu]

Phylogenetic relationships among mantids are poorly
known, and a formal quantitative character analysis has yet
to be performed. We inferred phylogenetic relationships for
Mantodea from DNA sequence data. Five genes (16S
rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase II, andhistone 3) were sequenced for 63 taxa representing major
mantodean lineages and outgroups. The monophyly of
mantodean families and subfamilies was tested undervarying parameter settings using parsimony and bayesiananalyses.
In regards to the classification of EHRMANN (2002), analyses supported the non-monophyly of Hymenopodidae,
Iridopterygidae, Mantidae, and Thespidae and the monophyly of the Amorphoscelidae subfamily Paraoxypilinae.
All represented subfamilies of Iridopterygidae andMantidae appear non-monophyletic. Mantoididae is sistergroup to the rest of the sampled mantid taxa, and is so placed basally on the topology. The other presumably basal

families (Chaeteessidae, Metallyticidae, and Eremiaphilidae)
were not represented in our analysis. Lineages congruent
with current subfamilial taxonomy include Paraoxypilinae,
Hoplocoryphinae, Hymenopodinae, Acromantinae, andOligonicinae.
We defined mantid hunting strategy as either generalist,
cursorial, or ambush predators. Mapping hunting strategy
onto our phylogenetic hypothesis reconstructs the ancestral
predatory condition as generalist hunting, with three independent shifts to cursorial hunting and one shift to ambush
hunting. The single origin of ambush hunting strategy is
associated with the largest radiation of mantodean species.

References
EHRMANN, R. 2002. Mantodea: Gottesanbeterinnen der Welt. –Verlag: Natur und Tier.

The Basal Phylogenetic Relationships in the Dermaptera
FABIAN HAAS1 & KLAUS-DIETER KLASS2
1

Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany [haas.smns@naturkundemuseum-bw.de] –Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, 01109 Dresden,
Germany [klaus.klass@snsd.smwk.sachsen.de]2

Introduction
The Dermaptera, which include ca. 2000 species, have long
been divided into three ‘suborders’ (e.g., HAAS 2003).
While the Forficulina comprise the ‘typical’ earwigs (Fig.
2C), the Hemimerina (Hemimerus, Araeomerus; Fig.
2D,E,F) and Arixeniina (Arixenia, Xeniaria; Fig. 2A,B)
include few, highly specialized species living in close association with certain rats (Muridae) and bats (Molossidae),respectively (NAKATA & MAA 1974). The most primitive
Forficulina are comprised in the Karschiellidae (Karschiellaand Bormansia), Diplatyidae (e.g., Haplodiplatys andDiplatys), and Pygidicranidae, the two latter possibly being
paraphyletic, and all three often comprised in a single family,

Pygidicranidae (s.l.) (e.g., KLASS 2003). The Pygidicranidae
include a number of ‘subfamilies’: Anataelinae, Challinae,
Esphalmeninae (Esphalmenus), Blandicinae, Pyragrinae(e.g., Pyragra), Pygidicraninae (e.g., Tagalina and Dacnodes), Cylindrogastrinae, and Echinosomatinae (e.g., Echinosoma).
Following the classical studies of Burr, Verhoeff andZacher, dermapteran phylogeny was re-assessed in some
contributions by POPHAM (e.g., 1985). He used charactersand character state transformation, however, no outgroup
and no numerical procedure was used for reconstructing thetree. Both was introduced to the study of Dermaptera by
HAAS (1995) and HAAS & KUKALOVÁ-PECK (2001). Whilethe latter authors focus on wing structure, the morphology

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Entomologische Abhandlungen 61 (2)Fig. 1. Cladistic hypothesis of dermapteran relationships, modified after HAAS & KUKALOVÁ-PECK (2001). For the purpose of the
discussions in this contribution the Arixeniina, Hemimerina, and Forficulina are regarded as separate higher taxa of Dermaptera,although this view is critisized here. The lack of hind wings is a major issue in the presented discussion, and hence it is mapped
onto the tree; filled square: only wingless species in taxon; empty square: only winged species in taxon; filled and empty square:
winged and wingless species in taxon.

of the female genitalia (ovipositor and surrounding structures) and other parts of the abdomen (e.g., basis of cerci) was
studied by KLASS (2001, 2003). The two character systems
most elaborately studied so far (hindwings, ovipositor) have
limitations in terms of their evidence on dermapteran phylogeny, because Hemimerina, Arixeniina, and several subgroups of basal and derived Forficulina lack hindwings(Fig. 1), and Hemimerina, Arixeniina, and all ‘higher
Forficulina’ have strongly reduced ovipositors.

Monophyly of Dermaptera
Autapomorphies so far suggested for Dermaptera sufferfrom many sources of uncertainty. Some apomorphic conditions present throughout Dermaptera, such as the prognathous head, the presence of apodemes on the anterior rim of
the abdominal coxosterna, and the formation of a female
subgenital plate from coxosternum 7, could as well be (largely conflicting) synapomorphies of Dermaptera andvarious other Pterygota. The same is true for reductive apomorphies in the female genitalia: gonapophyses 9 (if present) much shorter than gonapophyses 8, without a medianfusion, and shifted posteriorly; olistheter absent; fusion of
extreme lateral parts of abdominal terga 9 and 10. The
reduction to three-segmented tarsi, the lack of ocelli, theone-segmented cerci, and the lack of styli on male abdominal coxosternum 9 (subgenital plate) are likely dermapteranautapomorphies, since five- or four-segmented tarsi, twoocelli, many-segmented adult cerci, and male styli are present in ‘Protelytroptera’ (close extinct relatives of Dermaptera) and some Jurassic Dermaptera (VISHNYAKOVA 1980;
HAAS & KUKALOVÁ-PECK 2001; WILLMANN 2003). However,
due to the frequent homoplasy of these reductive traits in Pterygota, their support for dermapteran monophyly is limited.
The highly derived condition of both forewings (as shorttegmina) and hindwings (with many unique structural adap-

tations serving for folding; HAAS 1995; HAAS &KUKALOVÁ-PECK 2001) in Dermaptera appears as a particularly rich source of ordinal autapomorphies. However, conclusiveness is limited because Hemimerina, Arixeniina,
Karschiellidae, and several subgroups of Pygidicranidae(i.e., most of the taxa crucial in the issue of dermapteranmonophyly) lack hindwings or both pairs of wings. The
clasper-shape of the cerci applies to all Forficulina; its
absence in Hemimerina (thread-like cerci) and indistinctness in Arixeniina is plausibly explained as a secondary loss
resp. reduction (KLASS 2001). Holocentric chromosomes
may also be autapomorphic for Dermaptera (including
Hemimerina), but data are said to be doubtful for
Arixeniina, and only a Dacnodes has been studied among
the primitive Forficulina (WHITE 1976). The X-shaped invagination on the stipes, peculiar sensory papillae on the
maxillary and labial palps, the accessory lateral cervicalsclerite, the location of the mesosternal pits on an infolding,a metapleural accessory plate, and the virgae of the male
genitalia may be further dermapteran autapomorphies(POPHAM 1985) but have likewise been studied only in alimited sample of basal Forficulina.
The presence of a manubrium on the male coxosternum 9(subgenital plate) is not a dermapteran autapomorphy(KLASS 2001). The overlapping of the abdominal terga andcoxosterna probably is a dermapteran (+ ‘protelytropteran’)autapomorphy (HAAS & KUKALOVÁ-PECK 2001) though asimilar abdominal structure occurs in the distantly related
Zygentoma (KLASS 2001).

Relationships within Forficulina
The presently most elaborate phylogenetic hypothesis (Fig.
1) is based on HAAS (1995) and HAAS & KUKALOVÁ-PECK(2001), who focus on wing characters: Karschiellidae +a(Haplodiplatys + b(Diplatys + c(Pyragra + d(Echinosoma +

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A

Proceedings of 1st Dresden Meeting on Insect Phylogeny

C

B

E

F

D
Fig. 2. The three major taxa of the Dermaptera – Arixeniina, Hemimerina and Forficulina – are shown in comparison to demonstrate their different appearance. A, B: Arixenia esau Jordan, 1909, dorsal view (A) and lateral view (B). C: Two specimens ofAnataelia canariensis Bolivar, 1899, dorsal view. D, E: Hemimerus bouvieri Chopard, 1934, dorsal view (D) and lateral view (E).
F: Cerci of Hemimerus deceptus Rehn & Rehn, 1936, showing no annulation. Not to scale. Copyright for all images except C at
NHM; C taken at ZMUC.

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Entomologische Abhandlungen 61 (2)e(Tagalina + f‘higher Forficulina’))))). Hemimerina andArixeniina are not included because of their winglessness,and basal Forficulina are represented by Karschiellidae(Karschiella), two genera of Diplatyidae (Haplodiplatys,
Diplatys), and three genera of Pygidicranidae belonging to
Pyragrinae, Echinosomatinae, and Pygidicraninae, respectively. The ‘higher Forficulina’ (clade f), comprising all
Forficulina not belonging to one of the three latter ‘families’, is very well supported through 12 autapomorphies,
while nodes basal to this clade are less strongly supported
and partly ambiguous.
Clade a is supported by the presence of a device locking thetegmina to the thorax (longitudinal groove on metanotum;
ridge bearing macrotrichae on tegmen). Clade b, rendering
Diplatyidae paraphyletic, is supported by the presence of a
spiny ridge on each side of the metanotal groove (anotherpart of the locking device) and the asymmetry of the tegmina. However, evidence from the tegmina locking device isambiguous, because reductions may have accompanied theloss of the hindwings in Karschiellidae, as indicated by thereport of tegminal macrotrichae as well as an indistinct
metanotal groove and spiny ridge from some species of
Karschiella (HAAS & KUKALOVÁ-PECK 2001; VERHOEFF
1902). On the other hand, the species of Diplatys andHaplodiplatys studied by KLASS (2003) display four apomorphic features in the female genitalia that are quite unique in insects and suggest them to form a monophyletic
Diplatyidae: the (paired) spermathecae are strongly branched and are internally widened into bulbs; gonapophyses 9(2nd valves of ovipositor) are shaped as broad, flat lobes
and hug into a membranous mould behind their bases.
Moreover, a clade Karschiellidae + Diplatyidae is supported by paired spermathecal openings, which are unambiguously apomorphic by outgroup comparison (for a detailed description of the structures see KLASS 2003).
Autapomorphies (all in the hindwing) of clade c are the specialized fustis area (the antero-basal tenth of the dermapteran hindwing), a short-and-broad or long-and-broadcondition of the costal area (instead of long and slender),the position of the end of cubital vein CuA3+4 anterior to
the 8th branch of anal vein AP1+2, the flat condition of thetail of the ano-jugal arm FAJ, and the broad separation of
the proximal branches of anal vein AP1+2. However, allthese hindwing characters are neither applicable to thewingless Karschiellidae nor to the outgroup (Dictyoptera),and therefore with monophyletic Diplatyidae being a reasonable alternative based on new evidence from the female
genitalia, the characters here under consideration may proveas yielding additional autapomorphies for Diplatyidae (withthe states listed above being the plesiomorphic ones).
Clade d is supported by the presence of two virgae in themale genitalia, and clade e by the small size of the jugal
prong of the 3rd axillare of the hindwing articulation.
Nonetheless, in these characters there are problems in terms
either of character polarity or coding of states, leaving thesenodes ambiguous (KLASS 2003).
Incidently the monophyly of the remaining ‘families’ of theDermaptera is not well supported. Notable exceptions are
the Apachyidae, which are highly autapomorphic in many
characters, while Forficulidae, Chelisochidae and Spongiphoridae are supported by few characters. The Anisolabididae, at the moment, appear as an assemblage of superficiallysimilar Dermaptera that can hardly be supported at all, andthe Labiduridae proved almost as difficult to support (HAAS& KUKALOVÁ-PECK 2001).

Placement of Hemimerina and Arixeniina
Historically, the Hemimerina and Arixeniina were placed at
the same hierarchical level as the Forficulina – usually ‘suborders’, sometimes ‘orders’ – because in their structure andnatural history they differ fundamentally from all
Forficulina (e.g., HAAS 2003). Both taxa have a strongly
modified physical appearance due to their epizoic habits
and are not easily recognized as Dermaptera (which apartfrom these taxa are rather uniform in appearance), and bothare viviparous. The related features are unique among
Dermaptera and are not found in the Forficulina (with thenotable exception of viviparity in Marava arachidis), andso, by stressing differences instead of common features in a
non-phylogenetic methodology, the Hemimerina andArixeniina were not perceived as potentially nested in the
Forficulina. This view on difference and common features
has changed and to date the most likely hypothesis suggests,
yet supported only by weak evidence, for Hemimerina andArixeniina a position deeply within Forficulina. For
Hemimerina support comes from some apomorphies sharedwith higher Forficulina (KLASS 2001): advanced immobilization (not fusion) of the shortened female abdominal
terga 8–10 and specific reductions in the female genitalia; aclose relationship with Apachyidae is furthermore indicatedby the shared absence of a dorsal telson sclerite (or perhaps
its fusion with tergum 11) and fusion of terga 10 and 11 in
the female postabdomen. For Arixeniina a placement within
Forficulina is supported by apomorphies shared with theSpongiphoridae (= Labiidae in POPHAM 1985), mainly thepresence of auxiliary sclerites associated with the virga inthe male genitalia (POPHAM 1985); the characters concerned, however, need re-examination.

Conclusions
The relationships at the basis of Forficulina are presently
only tentatively resolved, and the same is true for the placement of Hemimerina and Arixeniina. Most important forforthcoming morphology-based analyses is the inclusion ofrepresentatives of the remaining pygidicranid ‘subfamilies’(see above and Fig. 1), including wingless ones. This should
be based on detailed studies of male and female genitalia
and wing structure. Another requirement is the explorationof new character systems such as the structure of the tarsi
and their adhesive devices, investigations on which havebeen begun by HAAS & GORB (in press). An extensive molecular data set is also presently being established, using
sequences of 18S rDNA, 28S rDNA, and histone 3 (manuscript in preparation by K.J. Jarvis, F. Haas & M.F.
Whiting). Thus, considerable progress in the analysis of
intra-ordinal relationships in Dermaptera is expected for thenear future.

Acknowledgements
Even shorter scientific contributions may be the result ofextensive collaborations. We therefore thank for the permission of The Natural History Museum London (NHM) and the
Zoological Museum of the University of Copenhagen (ZMUC)
to photograph their specimens. The visits at these institutions
would have not been possible without the EU-IHProgramme(Improving Human Ressources Programme), called SYSRESOURCE in the NHM and COBICE in the ZMUC, and weare grateful for that. We would also like to thank our scientifichosts Mrs J. Marshall (NHM) and Dr N.M. Andersen (ZMUC)
for their support during the visits.

References
HAAS, F. 1995. The phylogeny of the Forficulina, a suborder ofthe Dermaptera. – Systematic Entomology 20: 85–98.
HAAS, F. 2003. 12. Ordnung Dermaptera, Ohrwürmer. Pp.
173–180 in: H.H. DATHE (ed.), Lehrbuch der Speziellen
Zoologie, Band I: Wirbellose Tiere, 5. Teil: Insecta, 2ndedn. (founded by A. Kaestner). – Spektrum Akademischer
Verlag, Heidelberg, Berlin.

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HAAS, F & S. GORB in press. Evolution of tarsal attachment
structures in the Dermaptera (Insecta). – Arthropod
Structure and Development.
HAAS, F. & J. KUKALOVÁ-PECK 2001. Dermaptera hindwing
structure and folding: new evidence for familial,ordinal and superordinal relationships within Neoptera (Insecta). – European Journal of Entomology 98:
445–509.
KLASS, K.-D. 2001. The female abdomen of the viviparous earwig Hemimerus vosseleri (Insecta: Dermaptera:
Hemimeridae), with a discussion of the postgenital abdomen of Insecta. – Zoological Journal of the Linnean
Society 131: 251–307.
KLASS, K.-D. 2003. The female genitalic region in lower-gradeearwigs (Insecta: Dermaptera: Pygidicranidae s.l.). –Entomologische Abhandlungen 61: 173–225
NAKATA, S & T.C. MAA 1974. A review of the parasitic earwigs. – Pacific Insects 16: 307–374.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

POPHAM, E.J. 1985. The mutual affinities of the major earwig
taxa (Insecta, Dermaptera). – Zeitschrift für Zoologische
Systematik und Evolutionsforschung 23: 199–214.
VERHOEFF, K.W. 1902. Über Dermapteren; 1. Aufsatz: Versuch
eines neuen, natürlicheren Systems auf vergleichend-morphologischer Grundlage und über den Mikrothorax derInsekten. – Zoologischer Anzeiger 25: 181–208.
VISHNYAKOVA, V.N. 1980. Earwigs (Insecta, Forficulida) fromthe Upper Jurassic of the Karatau Range. – Palaeontological Journal 1980: 78–94.
WHITE, M.[J.D.] 1976. Blattodea, Mantodea, Isoptera,
Grylloblattodea, Phasmatodea, Dermaptera andEmbioptera. – Animal Cytogenetics 3(2): i–v, 1–75. –Gebrüder Borntraeger, Berlin.
WILLMANN, R. 2003. Die phylogenetischen Beziehungen derInsecta: Offene Fragen und Probleme. – Pp. 1–64 in:Verhandlungen Westdeutscher Entomologentag 2001,
Löbbecke Museum, Düsseldorf.

Phylogeny and Evolution of Acoustic Communication in Orthoptera
MANDA CLAIR JOST1 & PIOTR NASKRECKI2
1Harvard University, USA (currently at Department of Integrative Biology, Patterson Laboratories, University of Texas at Austin,
Austin, TX 78751, USA) [mandaclair@mail.utexas.edu] – 2 University of Connecticut, USA (currently with Conservation
International at Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA)[p.naskrecki@conservation.org]

Based on morphology and the molecular analyses in FLOOK& ROWELL (1997) and ROWELL & FLOOK (1998), a sistergrouprelationship between the suborders Caelifera and Ensifera ispresently the best supported hypothesis on the basal splitting event in Orthoptera. Relationships among subgroups of
Caelifera are considered in FLOOK & ROWELL (1997) and
ROWELL & FLOOK (1998).
The Ensifera (crickets, katydids, and related insects) is
usually classified into the families Gryllidae; Gryllotalpidae;
Rhaphidophoridae; Haglidae and/or Prophalangopsidae;
Schizodactylidae; Stenopelmatidae and/or Anostostomatidae
and/or Gryllacrididae; and Tettigoniidae. Representatives of
Ensifera are well known for acoustic signals produced in
the contexts of courtship and mate recognition. However,there is still controversy regarding the monophyly of major
lineages, the relationships between them, and whether the
acoustic structures and behaviors of Ensifera are homologous (plesiomorphic) or multiply derived. To address thesequestions, M. Jost works on a phylogenetic estimate of theEnsifera for a sample of 51 systematically diverse exemplars,
using sequences from 18S rRNA, 28S rRNA and 16S rRNA,
and three exemplary species of Caelifera as outgroup taxa.
The results support a basal split between monophyletic
Grylloidea (comprising Gryllidae, Gryllotalpidae, andSchizodactylus) and monophyletic Tettigonioidea (comprising Tettigoniidae, Rhaphidophoridae, Cyphoderris, and the‘gryllacridoid’ families) (Fig. 1). Support was also foundfor a monophyletic Tettigoniidae and for a monophyletic‘Gryllacridoidea’ including the Gryllacrididae, Stenopelmatidae, Anostostomatidae, and the genus Lezina, and theresults provide a robust estimate of phylogeny within amonophyletic Gryllidae.
Tree comparison tests with the molecular data using parsimony found Jost’s estimate to be significantly better thanthe phylogenetic hypotheses of SHAROV (1968) and
GOROCHOV (1995) although the data could not significantly
reject the result of the only numerical analysis of Ensifera
to date, GWYNNE (1995), which used morphological data.
Conversely, the most parsimonous tree from GWYNNE(1995) was not significantly better than Jost’s fully resolved
estimate when tested using Gwynne’s data matrix and cha-racter assumptions, although it was significantly better thanthe estimates from SHAROV (1968) and GOROCHOV (1995).
Molecular data were then used to estimate phylogenetic
relationships within the Gryllidae, and the resulting phylogeny was used to test the hypothesis that losses of acoustic
structures are usually coupled with flightlessness, microptery, or aptery. Jost presents a phylogenetic estimate for
the Gryllidae using 18S rRNA and 16S rRNA sequences fora diverse sample of 49 species, representing 46 genera from
13 subfamilies. Parsimony and Bayesian results strongly
support a monophyletic Gryllidae, as well as the monophyly of most recognized subfamilies including the Gryllinae,
Podoscirtinae, Oecanthinae, Phalangopsinae, Nemobiinae,
and Trigonidiinae. Tree comparison tests using parsimony
found Jost’s most parsimonious estimate to be significantly
better at explaining the molecular data than the phylogenetic hypotheses from two prior studies: GOROCHOV (1984)and DESUTTER (1987). Two tests of character state distributions in the Gryllidae (which accounted for phylogeny) provided significant support for the hypothesis that losses of
acoustic traits are usually coupled with hindwing microptery and/or aptery.
Close examination of wings, stridulatory tegmina, and tibial tympana in gryllids reveals a continuum of adult phenotypes that closely resemble the observed stages of mid- tolate-instar nymphs. Acoustic structures and wings are strictly adult characters in crickets, and might be coupledthrough similar post-embryonic developmental constraints.
To test the strength and significance of these correlations,character states for hindwings, forewings, and tibial auditory tympana were coded as an ontogenetic series andscored for adults of a diverse sample of gryllid taxa.
Felsenstein’s method of independent contrasts was used totest the hypothesis that adult-like character states are positively correlated with other adult-like character states, andjuvenile-like character states are positively correlated withother juvenile-like character states. Two phylogenetic hypotheses were used to compute node contrasts: one estimatedby a Bayesian analysis of 16S and 18S sequence data, andanother from a previously published phenetic study(DESUTTER 1987). Regardless of the hypothesis used,

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Entomologische Abhandlungen 61 (2)Fig. 1. Phylogenetic relationships among major lineages of Ensifera. From ongoing work by M. Jost based on analyses of
18S rRNA, 28S rRNA and 16S rRNA in 51 ensiferan exemplars, simplified. Caelifera used as outgroup. * Gryllacridoidea resulting as subordinate within Tettigonioidea; ** monophyly of taxon not strongly supported.

Spearman’s rank correlation (with ties) on the node contrast
values showed that forewing length and hindwing length aresignificantly correlated with one another, and that forewing
length is significantly correlated with tibial tympana development. Hindwing length was significantly correlated withtibial tympana development using Jost’s Bayesian estimate,
but not using the hypothesis from DESUTTER (1987); however, the sum of wing length scores was significantly correlated with tibial tympana development, regardless of thephylogenetic hypothesis used. It is suggested that this pattern has resulted from a common mechanism such asheterochrony, and that many independent losses of wings
and acoustic structures are due to paedomorphosis.
P. Naskrecki’s ongoing work relates to the phylogeny andclassification of Tettigoniidae, which contains more than
6400 species distributed worldwide. Naskrecki has reconstructed phylogeny in the Tettigoniidae using 195 taxa and331 morphological characters, many newly identified frominformative structures on the head, sternal endoskeleton,
ovipositor, and eggs. The strict consensus of most parsimonious trees found most currently recognized tettigoniid subfamilies to be paraphyletic (including the large subfamilies
Conocephalinae and Pseudophyllinae) or polyphyletic(including Bradyporinae and Listroscelidinae). Naskrecki ispreparing to introduce a new classification based on themonophyletic lineages found in this analysis. He then uses
hypotheses on phylogeny to examine the evolution of female stridulation in the Tettigoniidae.
Naskrecki’s phylogenetic hypothesis supports at least sevenindependent origins of female stridulation, which he classifies into several types. In the ‘bradyporoid’ female stridulatory mechanism (considered to be the most primitive), themale and female sound producing structures are fully
homologous. In the ‘ephippigeroid’ type, male and female
sound producing structures are functionally similar, but nothomologous. In the ‘phaneropteroid’, ‘polyancistroid’, and‘pterophylloid’ types, male and female sound producing
structures are functionally different and not homologous. In
the phaneropteroid type, females have multiple stridulatory
files on lateral branches of 2A on the dorsal side of the righttegmen. In the ‘polyancistroid’ type, the female stridulatory
file is on a secondary branch of Cu1 on the dorsal side of
the right tegmen. In the ‘pterophylloid’ type, females have a

stridulatory ‘net’ on the lateral branches of 2A on the ventral side of the left tegmen, and a multi-tooth scraper on the
inner edge of the right tegmen. In all of these except for thephaneropteroid and pteryphylloid types, females are capable of producing calls similar to those of males, and often
pure tone calls. The phylogenetic distribution of female stridulation among the Tettigoniidae may suggest that bidirectional phonotaxis between the sexes may evolutionarily precede unidirectional, female-to-male phonotaxis.
The authors also wish to call attention to the Orthoptera
Species File online, a complete taxonomic and synonymy
database of all Orthoptera described prior to 1998.
The URL for this public database is http://140.247.119.145/
Orthoptera/.

References
DESUTTER, L. 1987. Structure et evolution du complexe phallique des Gryllidea (Orthopteres) et classifiation des genres
Neotropicaux de Grylloidea. Premiere partie. – Annales
de la Société Entomologique de France (N.S.) 23(3):213–239.
FLOOK, P.K. & C.H.F. ROWELL 1997. The phylogeny of the
Caelifera (Insecta, Orthoptera) as deduced from mtrRNA
gene sequences. – Molecular Phylogenetics and Evolution
8(1): 89–103.
GOROCHOV, A.V. 1984. A contribution to the taxonomy ofmodern Grylloidea (Orthoptera) with a description of new
taxa. – Zoologicheskii Zhurnal 63(11): 1641–1651.
GOROCHOV, A.V. 1995. System and evolution of the suborder
Ensifera. – Proceedings of the Zoological Institute of theRussian Academy of Sciences, Vol. 260 [in Russian].
GWYNNE, D.T. 1995. Phylogeny of the Ensifera (Orthoptera): a
hypothesis supporting multiple origins of acoustical signalling, complex spermatophores and maternal care incrickets, katydids, and weta. – Journal of Orthoptera
Research 4: 203–218.
ROWELL, C.H.F. & P.K. FLOOK 1998. Phylogeny of the
Caelifera and the Orthoptera as derived from ribosomal
gene sequences. – Journal of Orthoptera Research 7:
147–156.
SHAROV, A.G. 1968. Phylogeny of the Orthopteroidea (B.B.
ROHDENDORF ed.). – Akademie Nauk SSSR (Trudy)
Paleontological Institute, Vol. 188. Israel Program for
Scientific Translations, Jerusalem.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Species Diversity and Intraordinal Phylogenetic Relationships ofMantophasmatodea
KLAUS-DIETER KLASS1, JAKOB DAMGAARD2 & MIKE D. PICKER3
1

Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, 01109 Dresden,
Germany [klaus.klass@snsd.smwk.sachsen.de] – 2 Zoological Institute and Zoological Museum, University of Copenhagen,
Universitetsparken 15, 2100 Copenhagen Ø, Denmark [jdamgaard@zi.ku.dk] – 3 Zoology Department, University of Cape Town,
Rondebosch 7700, Cape Town, South Africa [mpicker@botzoo.uct.ac.za]

The description of the new insect order Mantophasmatodea
by KLASS et al. (2002) was based on two lines of morphological evidence: (1) these wingless insects lack the sets ofapomorphies that characterize the other insect orders; (2)evidence on the sistergroup relationship of Mantophasmatodea to some other insect order was weak and conflicting. The ordinal status of Mantophasmatodea was doubted
by TILGNER (2002), but his argumentation was refutedby KLASS (2002). While the assignment of Mantophasmatodea to the Pterygota-Neoptera is supported by several
apomorphies (KLASS et al. 2002), the identification of its
sister group on a morphological basis has remained doubtful due to character conflicts (KLASS et al. 2003). Molecularstudies suggest a sistergroup relationship to the Notoptera(= Grylloblattodea; M.D. Terry & M.F. Whiting, pers.
comm.).
Presently 13 extant (all African) and 1 fossil species(European) of Mantophamatodea are known. The probably
earliest report of mantophasmatodeans in the literature isthat in ARILLO et al. (1997), which relates to immature
Baltic amber specimens that were not formally described
therein. The first species to be described (prior to thedescription of the order!) was Raptophasma kerneggeri
Zompro, 2001 from Baltic amber, classified as “Orthoptera
incertae sedis” (ZOMPRO 2001). Mantophasma zephyra
Zompro et al., 2002 from Namibia and Tanzaniophasma
subsolana (Zompro et al., 2002; as Mantophasma) from
Tanzania were described along with the ordinal description
in KLASS et al. (2002); both are known from a single specimen and were probably not recovered since (the species
identity of the specimens considered “Mantophasma zephyra” in DALLAI et al.’s 2003 study of sperm structure is unresolved, see KLASS et al. 2003: 48). Another Namibian species, Praedatophasma maraisi Zompro & Adis, 2002, wasdescribed in ZOMPRO et al. (2002).
A short paper by PICKER et al. (2002) was the first to indicate that probably several Mantophasmatodea species occur
in western South Africa. Subsequent work by our team led
to the description of five new genera including eight new
species from this area (KLASS et al. 2003): Austrophasma(A. rawsonvillensis, A. caledonensis, A. gansbaaiensis),
Lobophasma (L. redelinghuysensis), Hemilobophasma (H.
montaguensis), Karoophasma (K. biedouwensis, K. botterkloofensis), and Namaquaphasma (N. ookiepensis). In addition, another new genus and species from Namibia has been
described by KLASS et al. (2003), Sclerophasma (S. paresisensis), and our collecting efforts in 2003 led to finding
some additional South African species still waiting for taxonomic treatment. Eventually, a further, spiny Namibian species was mentioned as “Tyrannophasma gladiator Zompro,
2003” in DALLAI et al. (2003) and will probably soon be formally described.
Thus, extant Mantophasmatodea are only known fromAfrica south of the equator, and in South Africa and possibly in Namibia the group shows strong endemism, with atleast many species restricted to some limited distribution
area. The approximately 45 million year old Raptophasma
fossils from Baltic amber, however, show that in the early
Tertiary mantophasmatodeans also occurred in Europe.
KLASS et al. (2003) discuss characters that support themonophyly of Mantophasmatodea, and they gave the firstdetailed description of taxonomically significant and phylogenetically informative character systems in this order,
most importantly the exoskeleton of the male and female
postabdomen (including the genitalia) and antennae. A formal morphology-based cladistic analysis of phylogenetic
relationships within the order, however, has not yet beenpossible due to problems in outgroup comparison (absenceor doubtful identification of many character-bearing structures of Mantophasmatodea in other Neoptera, see KLASS et
al. 2003: 57f; influential but doubtful choice of outgrouptaxa due to lacking resolution of basal neopteran phylogeny). KLASS et al. (2003) present a phylogenetic tree of
Mantophasmatodea based on two mitochondrial genes
encoding COI and 16S rRNA; considering the later inclusion in our data set of a specimen that likely belongs to“Tyrannophasma gladiator”, our molecular analysis presently includes all abovementioned species except
Mantophasma zephyra, Tanzaniophasma subsolana,
Praedatophasma maraisi, Namaquaphasma ookiepensis,
and the fossil Raptophasma kerneggeri (Fig. 1). Some of
the included species were represented by specimens fromdifferent localities.
The molecular and the partly tentative morphologicalresults on Mantophasmatodea phylogeny were our basis fora classification of Mantophasmatodea (KLASS et al. 2003).
Apart from the definition of genera this includes a subdivision into three families, Tanzaniophasmatidae, Mantophasmatidae, and Austrophasmatidae, which so far are congruent
with Tanzanian, Namibian, and South African Mantophasmatodea, respectively (Praedatophasma and Raptophasma,with neither genitalia nor molecules then studied, left
“incertae sedis”; Tyrannophasma then unpublished). With
the forthcoming refinement of our molecular tree, we will
use the results on basal dichotomies within the order forpolarizing morphological characters in order to make theseavailable for cladistic analyses.
The monophyly of Mantophasmatodea is morphologicallywell-founded by (1) a very specialized structure of theantennae, which are composed of a varied number of basiflagellomeres that have no constrictions between them andhave only coarse setation, and the distal ones of which show
a secondary subdivision; and of distiflagellomeres thatalways count seven, the first being very long and the second
very short, are separated by discrete constrictions, and havevery fine setation in addition; (2) by the strongly angled
course of the subgenal sulcus on the head, which depends
on a dorsal shift of the origins of the anterior tentorial arms;(3) by a medioventral projection on the male subgenital
plate (used for drumming; also present in many Plecoptera);(4) by a conspicuously blunt tip of gonapophyses 8 in theovipositor; and (5) by a small dorsal projection in the membrane beyond the 3rd tarsomere. Monophyletic Mantophasmatodea are also strongly supported in our molecularanalysis, where, however, the apparently basal Tanzaniophasma and Raptophasma as well as notopterans (as animportant outgroup) are lacking.
The monophyly of Austrophasmatidae is strongly supportedby the molecular analysis; the inclusion of Namaquaphasma
in this family is based on its male genitalia being very simi-

Entomologische Abhandlungen 61 (2)145

Fig. 1. Molecular-based phylogenetic hypothesis for Mantophasmatodea using mitochondrial COI and 16S rRNA genes;
Namaquaphasma, Mantophasma, Praedatophasma, and Tanzaniophasma (with 1 species each) not included. All sequenced specimens included. Locality given in brackets behind species name (see KLASS et al. 2003: 11 for abbreviations; Pa, Br in northern
Namibia; Bi, Bo, Ca, Cl, Dr, Ga, Ko, Mo, Ra, Re, Tw, Wo in Western Cape Province, and Cv in Northern Cape Province of South
Africa), ‘2x’ in front of locality abbreviation means that 2 specimens were studied that had identical sequences.

lar to those in other austrophasmatid genera. The relationships among the five genera of Austrophasmatidae haveremained unresolved due to low support values. In our analysis we obtain the Namibian Sclerophasma as the sistergroup of Austrophasmatidae. This taxon has genitalia quite
different from all Austrophasmatidae, with, e.g., presenceof two additional sclerites but absence of two small sclerotized hooks in the male, and presence of a paired vaginal
sclerite and a paired row of strong setae ventrally between
the two gonoplacs of the ovipositor in the female. Since inthe mentioned female characters Mantophasma (males unknown) conforms with Sclerophasma, the two genera weretentatively combined in the same family (‘Mantophasmatidae’ due to nomenclatural priority). Nonetheless, among
the characters characterizing Mantophasmatidae, only thesetae between the gonoplacs are quite surely apomorphic.
Tyrannophasma results as the sister group of Austrophas-matidae + Mantophasmatidae in our molecular analysis andwould thus deserve the status of a separate family; genitalia
in this taxon have remained unstudied. Tanzaniophasma
differs fundamentally from the other Mantophasmatodea
with male genitalia known by the simple structure and bilateral symmetry of these (very complicated and asymmetrical in others); we thus classified it into a separate family
Tanzaniophasmatidae, though it is not entirely certain
whether the configuration of its genitalia is actually plesiomorphic for Mantophasmatodea. Raptophasma, according
to ZOMPRO et al. (2002), lacks the strong tibial spines present in all other species; this is likely to be a plesiomorphic
condition, and a position as the sister group of the remaining Mantophasmatodea (making it deserve the status of aseparate family) is the best guess at the moment – which
would nicely match with the isolated occurrence of thisgenus in the northern hemisphere.

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Acknowledgements
We are grateful to those who helped in collecting Mantophasmatodea (Jonathan Colville, Anthony Roberts, Geoff Hoy, Elke
Klass, and Simon van Noort) or supplied specimens to ourteam (Eugéne Marais, Margie Cochrane, Michael Ohl, Leonie
Pretorius, and Tharina Bird). This project was supported by travel grants from the Deutsche Forschungsgemeinschaft to K.-D.
Klass (DFG: GZ 445 SUA-111/3/02 and 111/3/03), grants fromthe Carlsberg Foundation and Villum Kann-Rasmussen
Foundation (both Denmark) to J. Damgaard, and URC andWWF research grants allowed to M. Picker.

References
ARILLO, A., V.M. ORTUÑO & A. NEL 1997. Description of an
enigmatic insect from Baltic amber. – Bulletin de laSociété Entomologique de France 102: 11–14.
DALLAI, R., F. FRATI, P. LUPETTI & J. ADIS 2003. Sperm ultrastructure of Mantophasma zephyra (Insecta,
Mantophasmatodea). – Zoomorphology 122: 67–76.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

KLASS, K.-D. 2002. Mantophasmatodea: a new insect order?
[Response to technical comment by Erich Tilgner] –Science 297: 731.
KLASS, K.-D., O. ZOMPRO, N.P. KRISTENSEN & J. ADIS 2002.
Mantophasmatodea: a new insect order with extant members in the Afrotropics. – Science 296: 1456–1459.
KLASS, K.-D., M.D. PICKER, J. DAMGAARD, S. VAN NOORT & K.
TOJO 2003. The taxonomy, genitalic morphology, andphylogenetic relationships of southern African Mantophasmatodea. – Entomologische Abhandlungen 61: 3–67.
PICKER, M.D., J.F. COLVILLE & S. VAN NOORT 2002.
Mantophasmatodea now in South Africa. – Science 297:
1475.
TILGNER, E. 2002. Mantophasmatodea: a new insect order?
[Technical comment] – Science 297: 731.
ZOMPRO, O. 2001. The Phasmatodea and Raptophasma n. gen.,
Orthoptera incertae sedis, in Baltic amber (Insecta:
Orthoptera). – Mitteilungen des Geologisch-Paläontologischen Instituts der Universität Hamburg 85: 229–261.
ZOMPRO, O., J. ADIS & W. WEITSCHAT 2002. A review of theorder Mantophasmatodea (Insecta). – Zoologischer
Anzeiger 241: 269–279.

New Insights in Grylloblattodean Phylogeny
KARL J. JARVIS & MICHAEL F. WHITING
401 WIDB, Brigham Young University, Provo, UT 84602, USA [karljarvis@byu.edu; michael_whiting@byu.edu]

Introduction
Grylloblattodea (= Notoptera) is one of the most enigmaticand diverse lineages of extremophile insects. Also known as
ice crawlers, this order of 25 species inhabits ice caves, glacial margins, talus slopes, forest floors, and similar habitats.
North American grylloblattids generally require temperatures near 0˚C for survival, thus the common name ‘ice crawlers’. Populations of Grylloblatta are distributed throughout the northern Rockies and Sierra Nevadas in numerous
disjunct populations. KAMP (1979) postulated that this distribution is due to post-Pleistocene migration from south of
the Pleistocene glaciers.
Order Grylloblattodea is comprised of five genera in asingle family (Grylloblattidae). Grylloblatta is the mostwidespread genus (11 spp.), found exclusively in the
Pacific Northwestern USA and western Canada. The remaining four genera are found in Asia, with Galloisiana (11spp.), Grylloblattina (1 sp., 2 subsp.) and Namkungia (1sp.) in eastern Asia in and around Japan, and Grylloblattella(1 sp.) in central Siberia. STOROZHENKO (1996) presented
the only hypothesis thus far on the phylogenetic relationships among these genera, based on an informal examination of characters. He proposed that the North American
Grylloblatta is sister to the Asian grylloblattids and that
Galloisiana and Grylloblattella are sister taxa.

Materials and methods
This phylogenetic analysis includes four Grylloblatta species, two Galloisiana species and the two subspecies of the
only species of Grylloblattina. Five additional representatives of undescribed Grylloblatta and Galloisiana species
were also included, and individuals from multiple populations of the same species were included for a total of 15 grylloblattid taxa. Recent analyses of extensive data (in a manu-

script submitted by M.D. Terry & M.F. Whiting) indicatethat mantophasmatodeans are sister to grylloblattodeans.
Therefore, two mantophasmatodean taxa and one species
each from Blattaria, Mantodea and Isoptera were included.
Trees were rooted to Mantodea.
Sequence data were generated from cytochrome oxidase II
mitochondrial DNA (650 bp), 18S nuclear ribosomal DNA(2000 bp), 28S nuclear ribosomal DNA (2400 bp), andhistone-3 nuclear protein-coding DNA (370 bp) for all
exemplar taxa. STOROZHENKO (1996) coded a morphological and ecological matrix of 12 characters within
Grylloblattodea, which was adapted to the taxon samplingof this analysis. The data were analyzed simultaneously in
POY (GLADSTEIN & WHEELER 2001) using direct optimization
under a 1:1:1 gap:transversion:transition ratio. TBR branchswapping was performed on 200 random sequence additions. Bootstrap values were calculated in PAUP* (SWOFFORD
2002) using the implied alignment output from POY.

Results
Phylogenetic reconstruction based on these data indicates a
Mantophasmatodea + Grylloblattodea relationship with abootstrap value of 100, and analyses of the genes separately also recover this relationship. These data also support themonophyly of the genera Grylloblatta, Grylloblattina andGalloisiana. Contrary to the only previous hypothesis ofgrylloblattid phylogeny (STOROZHENKO 1996), our data
indicate a basal Galloisiana, rather than a basal
Grylloblatta. This result is supported by a bootstrap valueof 100 for the clade Grylloblatta + Grylloblattina. The phylogeny within the North American grylloblattodeans issomewhat discordant with the distribution of the populations. This indicates that the colonization of modern populations was not a simple dispersion, but rather there may havebeen multiple migration events.

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Entomologische Abhandlungen 61 (2)Acknowledgements

References

We thank the BYU DNA Sequencing Center, the BYUIntegrative Biology Department, the Whiting Insect Molecular
Systematics Lab, Hiroshi Ando, Sergei Storozhenko, Rod
Crawford, Ed Lisowski, Toru Kishimoto, Riley Nelson,
Matthew Terry and the National Science Foundation formaking this research possible.

GLADSTEIN, D. & W.C. WHEELER 2001. POY: Phylogeny
Reconstruction via Optimization of DNA Data, version
2. – American Museum of Natural History.
KAMP, J.W. 1979. Taxonomy, distribution, and zoographic evolution of Grylloblatta in Canada (Insecta: Notoptera). –Canadian Entomologist 111: 27–38.
STOROZHENKO, S.Y. 1996. On origin and historical development of the family Grylloblattidae (Insecta:
Grylloblattida). – A.I. Kurentsov’s Annual Memorial
Meetings VI: 13–19.
SWOFFORD, D.L. 2002. PAUP*: Phylogenetic Analysis Using
Parsimony (and Other Methods), version 4.0b10. –Sinauer Associates, Inc. Publishers.

Phylogeny of the Zoraptera
MICHAEL S. ENGEL
Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, Snow Hall, 1460
Jayhawk Boulevard, University of Kansas, Lawrence, Kansas 66045-7523, USA [msengel@ku.edu]

Introduction

Discussion of basal Zoraptera

Among the ‘smaller’ insect orders, a few stand out as truly
depauperate in number of species within a class famed for
its overwhelming diversity. The Zoraptera is one of a few
insect orders whose diversity does not exceed 100 described
species; 38 living and fossil species are presently known.
Zorapterans are minute, enigmatic insects superficiallyresembling booklice (Psocoptera) and termites (Isoptera).
Individuals live gregariously under the bark of decaying
logs or within termite nests, where they principally feed on
fungal hyphae as well as nematodes or minute arthropods(e.g., mites, collembolans) (ENGEL 2003, in press). Adults
occur in two morphs within each species – eyed, winged
forms (i.e., alates), which eventually shed their wings, andblind, apterous forms. The order occurs pantropically with
only four species found distributed slightly outside of thetropical zone. Six fossil species are known, the oldest being
four (only three included in the present study) from Middle
Cretaceous (lowermost Cenomanian) amber from Myanmar(ENGEL & GRIMALDI 2002).
Zoraptera monophyly is based on a peculiar wing venation(see figures in ENGEL & GRIMALDI 2002; ENGEL 2003); 2segmented tarsi (the more elongate 2nd segment probably
results from fusion of two segments); peculiar mating via‘mating hook’ (also evident in Cretaceous fossils); unsegmented cerci (2-segmented in one derived fossil species);
stout metafemoral spines; and moniliform, 9-segmented
antennae (additional characters discussed by ENGEL &GRIMALDI 2000; ENGEL in press). Placement of the orderamong other hemimetabolous insects has been troubling
and the Zoraptera have been placed in a plethora of phylogenetic positions (reviewed in ENGEL & GRIMALDI 2002).
The placement of the order will not be discussed further
herein but to note that outgroup comparisons for polarity
matter little whether Embiodea or Psocoptera are used, i.e.,
whether the Polyneoptera (WHEELER et al. 2001; clade comprising Anartioptera = Plecoptera, Embiodea, Orthoptera,
Phasmatodea, Grylloblattodea, Mantophasmatodea,
Dermaptera,
theirextinct
allies†Titanoptera,†Caloneurodea, and various families of the unnatural‘Protorthoptera’, and possibly the Zoraptera) or the
Eumetabola (HENNIG 1969; KRISTENSEN 1995; clade comprising the hemipteroid orders, Endopterygota, and possibly the Zoraptera) hypothesis for placement is preferred.

Overall species of the order are remarkably homogenous.
Only one taxon departs from overall plan of zorapteranmorphology, albeit, only slightly so. The Middle Cretaceous†Xenozorotypus burmiticus plesiomorphically retains anadditional vein in the hind wing (M3+4), thereby suggestingthat it is sister to all other Zoraptera (the loss of this vein
being a putative event that occurred in the common ancestorof all other Zoraptera; Fig. 1). All remaining species of theorder are placed in the genus Zorotypus.
Basal divisions within Zorotypus remain problematic (Fig.
1). The Middle Cretaceous species †Zorotypus acanthothorax and †Z. nascimbenei (both placed here in Octozoros
subgen.n., see below) are perhaps basal within the genus, as
indicated by the plesiomorphic retention of the strong andslightly expanded empodium of the meta-pretarsus (reduced to a single, slender seta or absent in Zorotypus s.str.;
Fig. 1). Together these two species form a monophyletic
group supported by the apomorphic reduction of the antenna to eight segments (nine in all other Zorotypus and in†Xenozorotypus) and the presence of jugate setae along the
middle third of the posterior border of the forewing (absent
in all other Zoraptera).
ENGEL & GRIMALDI (2000, 2002) highlighted a few of thepotential character systems that may be of utility in the
Zoraptera (e.g., the terminal spine present on the apex of thecerci of some species may eventually unite a clade).
However, phylogenetic studies within the order are compromised by the heterogeneity of material and information
available for individual species. For instance, of the 38
described living and fossil species merely 13 are known on
the basis of both sexes and only 7 are known from bothapterous and winged morphs (not including the 6 known
only from apterous and deälated individuals, where wing
morphology is therefore unknown). Thus, characters fromwing structure are available for less than 20 % of the species in the order, and male genitalic characters are available
for just less than 45 % of the species. This dramaticallylimits the amount of comparable data between species. The
greatest strides toward resolving phylogenetic relationships
within the order will come from intensive fieldwork. Field
studies are needed to obtain useful material for molecularstudies, additional morphs and/or sexes, and to discover yet
unknown species (e.g., the order likely has a much wider

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Fig. 1. Preliminary phylogenetic hypothesis for zorapteran relationships (including only a few representatives of Zorotypus s.str.),with emphasis on basal divergences. Relationships in Zorotypus are difficult to discern owing to paucity of comparable data acrossspecies. A putative node “A” is indicated for representative species with uniform series of metafemoral spines; a putative node“B” is indicated for species with elongate cerci. For apomorphies of †Xenozorotypus and the entire order refer to ENGEL &GRIMALDI (2002), and ENGEL & GRIMALDI (2000) and ENGEL (2003, in press), respectively.

distribution with additional species in Southeast Asia andtropical Africa). In addition, more data on the biology andethology of zorapterans are required to thoroughly evaluate
the utility of these characters within the order.

†Octozoros subgen.n.
Diagnosis. This subgenus is overall similar to Zorotypuss.str. but differs by reduction of the antenna to eight segments, presence of jugate setae along middle third of forewing posterior border, and empodium of meta-pretarsus
strong and slightly expanded (reduced to a single, slenderseta or absent in Zorotypus s.str.).
Type species. Zorotypus nascimbenei Engel & Grimaldi,
2002.
Other included species. Zorotypus acanthothorax Engel &Grimaldi, 2002.
Derivatio nominis. A combination of the Greek ‘okto’(meaning ‘eight’) and ‘zoros’ (meaning ‘pure’); a reference
to the 8-segmented antenna. The name is masculine.

Acknowledgements
I am grateful to K.-D. Klass for inviting me to participate in the
symposium “The Phylogenetic Relationships within the Insect

Orders” and to M.G. Rightmyer for comments on this shortmanuscript.

References
ENGEL, M.S. 2003. Zoraptera. Pp. 1201–1203 in: V.H. RESH &R.T. CARDÉ (eds.), Encyclopedia of Insects. – Academic
Press, New York.
ENGEL, M.S. in press. Zoraptera. In: J. LLORENTE-BOUSQUETS& J.J. MORRONE (eds.), Biodiversidad, Taxonomía yBiogeografía de Artrópodos de México, vol. 4. – UNAM,
Mexico D.F.
ENGEL, M.S. & D.A. GRIMALDI 2000. A winged Zorotypus in
Miocene amber from the Dominican Republic (Zoraptera:
Zorotypidae), with discussion on relationships of and within the order. – Acta Geologica Hispanica 35: 149–164.
ENGEL, M.S. & D.A. GRIMALDI 2002. The first Mesozoic
Zoraptera (Insecta). – American Museum Novitates 3362:
1–20.
HENNIG, W. 1969. Die Stammesgeschichte der Insekten. –Senckenbergbuch (Frankfurt a. M.) 49: 1–436.
KRISTENSEN, N.P. 1995. Forty years’ insect phylogenetic systematics: Hennig’s “Kritische Bemerkungen...” and subsequent developments. – Zoologische Beiträge, N.F., 36:
83–124.
WHEELER, W.C., M. WHITING, Q.D. WHEELER & J.M.
CARPENTER 2001. The phylogeny of the extant hexapod
orders. – Cladistics 17: 113–169.

149

Entomologische Abhandlungen 61 (2)Phylogenetics of Psocoptera and the Origins of Lice
KEVIN P. JOHNSON
Illinois Natural History Survey, 607 East Peabody Drive, Champaign, IL 61820, USA [kjohnson@inhs.uiuc.edu]

The close relationship between Psocoptera (bark lice andbook lice) and Phthiraptera (parasitic lice) has long beenrecognized. Together these two orders comprise the insectgroup Psocodea. The monophyly of Psocodea is supported
on the basis of 18S rDNA sequence data (WHITING et al.
1997) and by several morphological autapomorphies, suchas reduction of the ovipositor and development of an atmospheric water-vapor uptake system, among others.
Recent studies of both morphological and molecular data
have begun to investigate higher level phylogenetic relationships within Psocodea. LYAL (1985) used morphologicalcharacters to show that the chewing lice (‘Mallophaga’) are
not a monophyletic group, but rather form a grade within
Phthiraptera; this has been confirmed by recent molecularstudies (JOHNSON & WHITING 2002; BARKER et al. 2003; seealso the following article by Vincent Smith on Phthiraptera
phylogeny). Most importantly, LYAL (1985) identified the
psocopteran family Liposcelididae as the sister taxon ofPhthiraptera rendering the order Psocoptera paraphyletic.
This result was based on the following apomorphic characters shared between Liposcelididae and Phthiraptera: dorsoventral compression of the head, reduction of the compound
eye to two ommatidia, loss of abdominal spiracles I and II,
and shortening of legs. Data from the 12S and 16S mitochondrial genes confirm a monophyletic clade Liposcelididae + Phthiraptera (YOSHIZAWA & JOHNSON 2003). With
these data, the monophyly of Phthiraptera is uncertain,
because in many analyses Liposcelididae tends to fall within
the Phthiraptera, albeit with weak support. Liposcelididae
also appears to share a dramatically elevated mitochondrial
substitution rate with lice, so long branch attraction must beseriously considered as a possible confounding factor.
Further molecular data from the nuclear 18S gene (K. Johnson, unpublished) also support a clade uniting Liposcelididae and Phthiraptera, lending independent support to thishypothesis. Analyses of this gene tend to place Liposcelididae as the sister taxon of Amblycera, making both
Psocoptera and Phthiraptera paraphyletic. If this relationship is correct, either parasitism evolved twice in lice or was
lost in Liposcelididae.
Until recently, the higher level phylogenetic relationships
within Psocoptera have received less attention than those inlice. Little attempt has been made to reclassify Psocoptera
taking into account its putative paraphyly. Traditionally theorder Psocoptera is divided into three suborders: Trogiomorpha, Troctomorpha, and Psocomorpha. An early phylogenetic analysis based on morphology for Psocoptera wasconducted by SMITHERS (1972). He suggested that the
Psocomorpha is the sister taxon of Troctomorpha based onsynapomorphies such as a closed phallosome, loss of theparaproct spine, smooth eggs, and shortened antennae.
However, much of the remainder of the classification
implied by Smithers’ phylogenetic hypothesis has generally
been abandoned in favor of more traditional classifications
within each suborder.
The phylogeny of the suborder Psocomorpha has received
recent attention. Traditionally Psocomorpha is divided into
four infraorders: Epipsocetae, Caeciliusetae, Homilopsocidea, and Psocetae. A recent cladistic study of morpho-

logical characters by YOSHIZAWA (2002) suggested that thefamily Archipsocidae should be removed from the
Homilopsocidea, being sister to all other Psocomorpha. In
addition, his analysis suggested that Hemipsocidae beremoved from Psocetae. In other respects the phylogenetic
results of YOSHIZAWA (2002) resembled traditional classification. JOHNSON & MOCKFORD (2003) used data from thenuclear 18S and mitochondrial 16S, 12S, and COI genes totest various classification schemes for Psocomorpha. Using
outgroups within Psocoptera, support was found for monophyly of Psocomorpha. They also found support for a basal
position of Archipsocidae within Psocomorpha, as suggested by YOSHIZAWA (2002). The family Hemipsocidae,
however, was placed within Psocetae with molecular data,
contradicting YOSHIZAWA’s (2002) study, but consistent withtraditional classification. Monophyly of the infraorders
Caeciliusetae and Psocetae was supported (JOHNSON &MOCKFORD 2003), but monophyly of Homilopsocidea could
not be recovered, even with Archipsocidae excluded. Morephylogenetic work is needed to provide a comprehensiveclassification of Psocomorpha. The phylogenetic relationships within Trogiomorpha and Troctomorpha remain to beinvestigated in detail.

Acknowledgements
I thank Edward Mockford and Kazunori Yoshizawa for theirassistance in providing and identifying samples of Psocoptera,
as well as their collaborative efforts.

References
BARKER, S.C., M.F. WHITING, K.P. JOHNSON & A. MURRELL
2003. Phylogeny of the lice (Insecta, Phthiraptera) inferred from small subunit rRNA. – Zoologica Scripta 32:
407–414.
JOHNSON, K.P. & E.L. MOCKFORD 2003. Molecular systematicsof Psocomorpha (Psocoptera). – Systematic Entomology
28: 409–416.
JOHNSON, K.P. & M.F. WHITING 2002. Multiple genes and themonophyly of Ischnocera (Insecta: Phthiraptera). –Molecular Phylogenetics and Evolution 22: 101–110.
LYAL, C.H.C. 1985. Phylogeny and classification of the
Psocodea, with particular reference to lice (Psocodea:
Phthiraptera). – Systematic Entomology 10: 145–165.
SMITHERS, C.N. 1972. The classification and phylogeny of the
Psocoptera. – Memoirs of the Australian Museum 14:
1–349.
WHITING, M.F., J.C. CARPENTER, Q.D. WHEELER & W.C.
WHEELER 1997. The Strepsiptera problem: Phylogeny of
the holometabolous insect orders inferred from 18S and
28S ribosomal DNA sequences and morphology. –Systematic Biology 46: 1–68.
YOSHIZAWA, K. 2002. Phylogeny and higher classification ofsuborder Psocomorpha (Insecta: Psocodea: “Psocoptera”). – Zoological Journal of the Linnean Society 136:
371–400.
YOSHIZAWA, K. & K.P. JOHNSON 2003. Phylogenetic position ofPhthiraptera (Insecta: Paraneoptera) and elevated rate ofevolution in mitochondrial 12S and 16S rDNA. –Molecular Phylogenetics and Evolution 29: 102–114.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Lousy Phylogenies: Phthiraptera Systematics and the Antiquity of Lice
VINCENT S. SMITH
Wellcome Trust Junior Research Fellow, Graham Kerr Building, Institute of Biomedical and Life Sciences, University of Glasgow,
Glasgow, G12 8QQ, United Kingdom [v.smith@bio.gla.ac.uk]

Phthiraptera (parasitic lice) comprise about 5,000 described
species present in four suborders (Fig. 1): Anoplura (colloquially known as sucking lice), Rhynchophthirina (a monogeneric group found on elephants and warthogs),
Ischnocera and Amblycera. Phthirapteran monophyly is
supported by 19 apomorphies, the most significant of
which include haploid reduction in primary spermatogonia,
the presence of a basal hydropile in the egg, dorsoventral
compression of the head and loss of the dorsal tentorial
arms.
Every conceivable relationship between the four phthirapteran suborders has been proposed at some time.
Historically the chewing lice (Amblycera, Ischnocera andarguably Rhynchophthirina) had been subsumed under thename ‘Mallophaga’, which was awarded ordinal statusalong with the Anoplura. Morphological data supporting
the monophyly of ‘Mallophaga’ was proposed by KIM &LUDWIG (1978, 1982), although these results were controversial (HAUB 1980).
LYAL (1985) conducted a detailed review of the morphological data that define basal louse relationships. His study
confirmed the monophyly of all four suborders, althoughischnoceran monophyly was the least well supported. The
subordinal phylogeny established by LYAL (1985) confirmed that ‘Mallophaga’ are paraphyletic and form a grade
within Phthiraptera. Significant synapomorphies given by
LYAL (1985) defining these relationships are (1) for
Rhynchophthirina + Anoplura: head ‘fixed’ in relation tothorax, loss of anterior tentorial pits and partial transfer ofantennal muscle attachment site to dorsum of head; (2) for
Rhynchophthirina + Anoplura + Ischnocera: developmentof saucer-shaped antennal sensilla, partial occlusion of
occipital foramen by a connective tissue septum and development of spiracular glands.
Recent molecular evidence supports LYAL’s (1985) phylogeny. Using nuclear genes CRUICKSHANK et al. (2001),JOHNSON & WHITING (2002) and most recently BARKER etal. (2003) have investigated phthirapteran basal relationships. All but CRUICKSHANK et al.’s (2001) results are completely congruent with those of LYAL (1985), and the discrepancies in CRUICKSHANK et al.’s study are likely to be theresult of an over-reliance on a relatively short fragment of
the faster evolving EF1-␣ gene (SMITH et al. in press).
Familial relationships within each of the suborders areslightly less problematic, with the notable exception of theIschnocera.
Anopluran lice exclusively parasitise mammals and have asignificant medical and veterinary importance. In part, this
explains why they are the best-studied suborder ofPhthiraptera. Between 1920 and 1935 Gordon Ferris provided the foundation for modern taxonomic work on the
Anoplura, and when republished as a monograph FERRIS(1951) recognized 6 families. In the light of new species
descriptions this was expanded to 15 (KIM & LUDWIG
1978). Apomorphies for Anoplura include the developmentof piercing stylets from the hypopharynx and labium, fusion of the pronotum to the mesonotum, and reduction of themeso- and metathoracic terga. A generic level morphological phylogeny for Anoplura was proposed by KIM (1988).
Molecular studies on anopluran relationships are ongoing,but for the moment BARKER et al. (2003) provides an initialaccount of anopluran relationships using 18S rRNA.
Rhynchophthirina comprise just three species in a single
genus. The subordinal status of this taxon was awarded by

FERRIS (1931) on account of the peculiar morphology of its
members. Both monophyly and high rank are supported bymorphological (LYAL 1985) and molecular (BARKER et al.
2003) data. Notable apomorphies for Rhynchophthirina
include an anterior prolongation of the head into a rostrum
with mandibles terminal and rotated 180°, and extension of
the pretarsal apodeme into the femur without a tibial muscle
bundle.
Amblyceran classification has been the subject of several
detailed studies, most notably by CLAY (1970), who didmuch to stabilise their familial groupings. CLAY (1969) also
considered possible relationships of genera in the largest
amblyceran family, the Menoponidae. This study has been
considerably expanded by MARSHALL (2003), who provides
the most comprehensive study of amblyceran relationships
to date covering generic level relationships for almost all
taxa that parasitise birds and Australasian marsupials. Her
study is broadly consistent with the available molecular studies, although the latter are few in number and largely confined to terminal clades of the amblyceran tree. Significant
apomorphies for Amblycera include the development of anantennal fossa concealing the antennae and the presence ofa pedunculate first flagellar segment.
Ischnocera form the largest suborder of Phthiraptera andtheir basal systematics is the subject of intense debate.
HOPKINS & CLAY (1952) recognize just 3 families while
EICHLER (1963) accepts 21. Based on both molecular andmorphological data the most recent studies on Ischnocera
recognize at least three monophyletic groups (LYAL 1985;
MEY 1994; SMITH 2000, 2001; SMITH et al. in press). These
are the Trichodectidae (sensu HOPKINS & CLAY 1952)
restricted to mammalian hosts, the Heptapsogasteridae(sensu SMITH 2000) present on tinamiform birds and theGoniodidae (sensu SMITH 2000) of Galliformes andColumbiformes. A fourth group, the Philopteridae (sensu
EICHLER 1963) comprise some 70 % of ischnoceran species
and are present on almost all families of birds. It is generally accepted that this is a miscellaneous collection ofgenera and is almost certainly para- or polyphyletic.
Despite being the subject of several extensive phylogenetic
studies there is little consensus about basal ischnoceran
relationships (SMITH et al. in press). Indeed, even the monophyly of the Ischnocera is weakly supported with no unequivocal morphological apomorphies defining this clade.
Character convergence, multiple substitutions at high divergences, and ancient radiation over a short period of time
have contributed to the problem of resolving ischnoceran
phylogeny with the data currently available. A monotypictaxon (the Trichophilopteridae) represented by a single species present on Madagascan primates (Lemuridae andIndridae) may be related to the avian ‘Philopteridae’. This
species bears a number of significant morphological characters that are apparently intermediate between the ‘Philopteridae’ of birds and the Trichodectidae of mammals.
Consequently the affinities of this genus are unclear, and it
has been variably placed amongst both these groups or in an
independent family within Ischnocera (SMITH 2001).

Acknowledgements
I thank Dale Clayton, Kevin Johnson, Rod Page, Ricardo
Palma, and Roger Price for their collaborative efforts throughout my research on parasitic lice.

151

Entomologische Abhandlungen 61 (2)Fig. 1. The four suborders of Phthiraptera and their species
content, host range, and phylogenetic relationships.

References
BARKER, S.C., M. WHITING, K.P. JOHNSON & A. MURRELL
2003. Phylogeny of the lice (Insecta: Phthiraptera) inferred from small subunit rRNA. – Zoologica Scripta 32:
407–414.
CLAY, T. 1969. A key to the genera of the Menoponidae(Amblycera: Mallophaga: Insecta). – Bulletin of the
British Museum (Natural History) Entomology 24: 3–26.
CLAY, T. 1970. The Amblycera (Phthiraptera: Insecta). –Bulletin of the British Museum (Natural History)
Entomology 25: 75–98.
CRUICKSHANK, R.H., K.P. JOHNSON, V.S. SMITH, R.J. ADAMS,
D.H. CLAYTON & R.D.M. PAGE 2001. Phylogenetic analysis of partial sequences of Elongation Factor 1 identifies
major groups of lice (Insecta: Phthiraptera). – MolecularPhylogenetics and Evolution 19: 202–215.
DURDEN, L.A. & G.G. MUSSER 1994. The sucking lice (Insecta,
Anoplura) of the world: a taxonomic checklist withrecords of mammalian hosts and geographic distributions.
– Bulletin of the American Museum of Natural History
218: 1–90.

EICHLER, W.D. 1963. Arthropoda. Insecta. Phthiraptera I.
Mallophaga. In: H. WEBER (ed.), H.G. Bronn’s Klassen
und Ordnungen des Tierreichs. – Akademische Verlagsgesellschaft Geest & Portig K.G., Leipzig, 291 pp.
FERRIS, G.F. 1931. The louse of elephants, Haematomyzus elephantis Piaget (Mallophaga: Haematomyzidae). –Parasitology 23: 112–127.
FERRIS, G.F. 1951. The sucking lice. – Memoirs of the Pacific
Coast Entomological Society 1: 1–320.
HAUB, F. 1980. Letter to the editors: concerning „phylogenetic
relationships of parasitic Psocodea and taxonomic position of the Anoplura“ by K.C. Kim and H.W. Ludwig. –Annals of the Entomological Society of America 73: 3–6.
HOPKINS, G.H.E. & T. CLAY 1952. A Checklist of the Genera
and Species of Mallophaga. – British Museum of Natural
History, London.
JOHNSON, K.P. & M.F. WHITING 2002. Multiple genes and themonophyly of Ischnocera (Insecta: Phthiraptera). –Molecular Phylogenetics and Evolution 22: 101–110.
KIM, K.C. 1988. Evolutionary parallelism in Anoplura andeutherian mammals. Pp. 91–114 in: M.W. SERVICE (ed.),Biosystematics of Haematophagous Insects. – Clarendon
Press, Oxford.
KIM, K.C. & H.W. LUDWIG 1978. The family classification of
Anoplura. – Systematic Entomology 3: 249–284.
KIM, K.C. & H.W. LUDWIG. 1982. Parallel evolution, cladistics,
and classification of parasitic Psocodea. – Annals of the
Entomological Society of America 75: 537–548.
LYAL, C.H.C. 1985. Phylogeny and classification of the
Psocodea, with particular reference to lice (Psocodea:
Phthiraptera). – Systematic Entomology 10: 145–165.
MARSHALL, I.K. 2003. A morphological phylogeny for fourfamilies of amblyceran lice (Phthiraptera: Amblycera:
Menoponidae, Boopiidae, Laemobothriidae, Ricinidae). –Zoological Journal of the Linnean Society 138: 39–82.
MEY, E. 1994. Beziehungen zwischen Larvenmorphologie undSystematik der Adulti bei den Vogel-Ischnozeren (Insecta,
Phthiraptera, Ischnocera). – Mitteilungen aus dem
Zoologischen Museum Berlin 70: 3–84.
SMITH, V.S. 2000. Basal ischnoceran louse phylogeny(Phthiraptera: Ischnocera: Goniodidae and Heptapsogasteridae). – Systematic Entomology 25: 73–94.
SMITH, V.S. 2001. Avian louse phylogeny (Phthiraptera:
Ischnocera): a cladistic study based on morphology. –Zoological Journal of the Linnean Society 132: 81–144.
SMITH, V.S., R.D.M. PAGE & K.P. JOHNSON in press. Data incongruence and the problem of avian louse phylogeny. –Zoologica Scripta.

The Morphological Background to Thysanoptera Phylogeny
LAURENCE A. MOUND1 & DAVID C. MORRIS2
1
Honorary Research Fellow, CSIRO Entomology, GPO Box 1700, Canberra, Australia; and, Scientific Associate, The Natural
History Museum, Cromwell Road, London SW7 5BD, United Kingdom [Laurence.Mound@csiro.au] – 2 School of Botany andZoology, Australian National University, Canberra, ACT 0200 Australia [David.Morris@anu.edu.au]

The order Thysanoptera either forms an unresolved trichotomy with the Hemiptera and Psocodea (Psocoptera +Phthiraptera), or is sister group to the Hemiptera (based on
the structure of the wing sclerites: YOSHIZAWA & SAIGUSA
2001) with Psocodea sister to these two. BHATTI (1988),
whilst accepting the monophyly of Thysanoptera, considered that the level at which this group is recognised shouldbe raised. He thus stressed the many structural differences
between the two traditional suborders, Terebrantia andTubulifera, and suggested that the Thysanoptera should beconsidered a superorder with two orders. However, thischange in taxonomic levels does not affect our understanding of relationships, based on the synapomorphic structureof the maxillary laciniae and pretarsal arolium in all thrips,
and it is not accepted by other workers on the group. In contrast, ZHERIKHIN (2002) reduced the Thysanoptera to subor-

der Thripina and placed this with the fossil suborder
Lophioneurina in an order Thripida; this also is not accepted currently, as the fossils have tarsal claws, extra wingveins, no wing fringes, and apparently two mandibles. In
proposing such relationships, palaeontologists place greatemphasis on the relatively superficial similarities provided
by narrow wings with long fringing setae. They thus ignore
the great diversity of wing form that occurs in extant
Thysanoptera, and take no cognisance of the functional
relationships between wing form and the ways in whichwings are used in flight and particularly the way in whichwings are parked on the abdomen when not in use.
The monophyly of the suborder Tubulifera is uncontested,on the basis of the structure of the wings with no longitudinal veins and the marginal cilia not arising from sockets,
also the tubular condition of abdominal segment 10 and the152eversible chute-like ovipositor of females. However, relationships between Tubulifera and Terebrantia remain unresolved: Tubulifera is either the sister group of Terebrantia,
or it is sister to part of a family within the Terebrantia(MOUND et al. 1980: Panchaetothripinae). An extensive
summary by BHATTI (1988) of the morphological evidence
supports the first hypothesis, all members of the Terebrantia
having three well-developed longitudinal wing veins bearing setae, the marginal cilia in figure-of-eight shaped sockets, and abdominal segment 10 fully divided ventrally withthe ovipositor comprising four saw-toothed valves.
In the Terebrantia, eight families are recognised, with thefour genera comprising the Melanthripidae currently
removed from the Aeolothripidae (cf. fig. 1 of following
article by D.C. Morris & L.A. Mound), although European
workers retain these four genera within the Aeolothripidae(as in the said figure). More recent consideration of the data
has resulted in the Aeolothripidae being defined on two
synapomorphies (loss of abdominal sternite 8 in females;presence of longitudinal sensilla on antennae), neither ofwhich are present in any of the Melanthripidae. Moreover,melanthripids are all flower-feeding, whereas aeolothripids
are either facultative or obligate predators. One problem
involving the Melanthripidae is that it cannot be distinguished satisfactorily from the Merothripidae, both groups
being defined on plesiomorphies, despite the great difference in their body sizes and feeding associations. Similarly,
an analysis of a data matrix for the genera in the
Fauriellidae and Adiheterothripidae provided no support foreither family (Fig. 1), and genera in both of these groups
show a remarkably disjunct distribution between western
USA and the Mediterranean (MOUND & MARULLO 1999).
The 60 species in the Heterothripidae occur only in the New
World, whereas the major Terebrantia family, the Thripidae,
with 2000 species and 260 genera, occurs worldwide. The
monophyly of this large family has never been questioned,
based largely on the presence of slender emergent sensillaon the 3rd and 4th antennal segments. But equally, it has
never been adequately demonstrated, and is accepted largely for reasons of tradition. Unfortunately, within this large
family there are further systematic problems: 50 % of thegenera are monotypic, and the available suprageneric classification has few supporting character states and cannot beapplied satisfactorily to the world fauna.
In the Tubulifera, only one family is usually recognised, thePhlaeothripidae, with two subfamilies, Phlaeothripinae andIdolothripinae. In contrast, BHATTI (1994, 1998) has recognised twelve additional families (Fig. 2), whilst retaining
98 % of the 3200 described tubuliferan species within the
Phlaeothripidae. The Aleurodothripidae was erected for asingle species in which the labro-maxillary complex isamalgamated with the cranium, but this same character
state was used to define two further families, theAdurothripidae and the Urothripidae. The Xaniothripidae
was erected for a single genus in which the wings are sometimes unusually broad and the abdomen bears many spines.
However, this genus is considered, on molecular and behavioural evidence, to be sister to the genus Koptothrips in the
Phlaeothripidae; the species in these two genera are biologically similar in being kleptoparasitic in the domiciles ofother thrips species on Acacia in Australia, but are structurally highly divergent (MORRIS et al. 2002). Of the twelve
families distinguished from the Phlaeothripidae by BHATTI(1998), nine include only a single genus (four with one
species, two with two species), and a tenth includes twogenera (each with one species). Finally, the Hyidiothripidae
and the Urothripidae each include about 30 species,
although both appear to have their closest relatives within
the Phlaeothripidae, and both are best considered as informal genus-groups within the Phlaeothripidae.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

This recognition of a series of small families does little topromote our understanding of relationships amongst the2500 described species of Phlaeothripinae. With 50 % of
the available genera monotypic, and limited understandingof the biological and genetic significance of the structuralpolymorphism amongst so many species, development of asuprageneric classification has not proved possible.
Currently three lineages that are weakly defined morphologically are recognised in this group, the Haplothrips-lineage of flower-feeding species, the Liothrips-lineage of leaffeeding species, and the Phlaeothrips-lineage of fungusfeeding species (MOUND & MARULLO 1996).
Within the second of the Tubulifera subfamilies, the Idolothripinae, systematic relationships appear to be more clearly
defined (MOUND & PALMER 1983). However, monophyly ofthis subfamily itself requires further support. Despite themain character state (maxillary stylets more than 4 μm in
diameter) appearing to be robust, it is related to the fact that
all the species feed by ingesting whole fungus spores. Since
feeding habits among thrips are notoriously opportunistic,
the possibility of homoplasy must be further investigated.
None of the Phlaeothripidae exhibit any character state thatcan be considered to be in a plesiomorphic state, whereas
within the Terebrantia families there is a progressive series
from plesiomorphic to derived character states. Moreover,some fossil Terebrantia exhibit plesiomorphic character
states, but no fossil ‘proto-Tubulifera’ have been described.
There can be little doubt that the Thysanoptera is monophyletic, considering the unique structure of the maxillarystylets enclosing a single channel. But at what point the
tubuliferan lineage separated from the terebrantian lineage
is a problem. If these two suborders are sister groups, theneither the proto-Tubulifera did not live in suitable situations
to be fossilised, or such fossils have yet to be found. The
alternative evolutionary scenario is that the Tubulifera
evolved relatively late, arising through neotenic development from larval Panchaetothripinae within the most highly derived of the Terebrantia families, the Thripidae, andsubsequently radiated rapidly. This suggestion is based on
the presence of a tubular abdominal segment 10 in Panchaetothripinae larvae, and the presence in many adult Panchaetothripinae of sigmoid wing-retaining setae on the abdominal
tergites.
The plesiotypic life-style of thrips is considered to be fungusfeeding in detritus, with the small-bodied, fungus-feeding
species in the Terebrantia family Merothripidae retaining
the largest number of plesiomorphies (antennae 9-segmented; tentorium complete; large pair of trichobothria on
abdomen). However, a sistergroup relationship to theremaining Terebarantia families is not clear, and as indicated above the morphology-based family classification owes
more to tradition than to phylogenetic character state analysis. Clearly, the future of phylogenetic studies on
Thysanoptera must involve alternative sources of data
including molecular data.

References
BHATTI, J.S. 1988. The orders Terebrantia and Tubulifera of thesuperorder Thysanoptera (Insecta). A critical appraisal. –Zoology (Journal of Pure and Applied Zoology) 1:
167–240.
BHATTI, J.S. 1994. Phylogenetic relationships among
Thysanoptera (Insecta) with particular reference to the families of the order Tubulifera. – Zoology (Journal of Pureand Applied Zoology) 4: 93–130.
BHATTI, J.S. 1998. New structural features in the order
Tubulifera (Insecta). 1. Amalgamation of labro-maxillarycomplex with cranium and other cephalic structures. –Zoology (Journal of Pure and Applied Zoology) 5:
147–176.

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Entomologische Abhandlungen 61 (2)

MORRIS, D.C., M.P. SCHWARZ, S.J.B. COOPER & L.A. MOUND
2002. Phylogenetics of Australian Acacia thrips: the evolution of behaviour and ecology. – Molecular Phylogenetics
and Evolution 25: 278–292.
MOUND, L.A., B.S. HEMING & J.M. PALMER 1980. Phylogenetic
relationships between the families of recent Thysanoptera.
– Zoological Journal of the Linnean Society of London 69:
111–141.
MOUND, L.A. & R. MARULLO 1996. The Thrips of Central andSouth America: an Introduction. – Memoirs on Entomology,
International 6: 1–488.
MOUND, L.A. & R. MARULLO 1999. Two new basal-clade
Thysanoptera from California with Old World affinities. –Journal of the New York Entomological Society 106: 81–94.

MOUND, L.A. & J.M. PALMER 1983. The generic and tribal
classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolothripinae). – Bulletin of the British Museum(Natural History). Entomology 46: 1–174.
YOSHIZAWA, K. & T. SAIGUSA 2001. Phylogenetic analysis ofparaneopteran orders (Insecta: Neoptera) based on forewingbase structure, with comments on monophyly of
Auchenorrhyncha (Hemiptera). – Systematic Entomology
26: 1–13.
ZHERIKHIN, V.V. 2002. Order Thripida Fallen, 1914. Pp.
133–143 in: A.P. RASNITSYN & D.L.J. QUICKE (eds.),
History of Insects. – Kluwer Academic Publishers,
Dordrecht, Boston, London.

Fig. 2. Tubuliferan families recognised by BHATTI (1994); it isunclear whether tree dichotomies are intended to propose phylogenetic relationships; see BHATTI (1998) for Adurothripidae,
Aleurodothripidae, and Habrothripidae.

Fig. 1. Relationships among Adiheterothripidae and Fauriellidae according to MOUND & MARULLO (1999), using the ‘ie’
option in Hennig86.

Thysanoptera Phylogeny – the Molecular Future
DAVID C. MORRIS1 & LAURENCE A. MOUND2
1

School of Botany and Zoology, Australian National University, Canberra, ACT 0200 Australia [David.Morris@anu.edu.au] –Honorary Research Fellow, CSIRO Entomology, GPO Box 1700, Canberra, Australia; and, Scientific Associate, The Natural
History Museum, Cromwell Road, London SW7 5BD, United Kingdom [Laurence.Mound@csiro.au]

2

The monophyly of the Thysanoptera is strongly supported
by a number of morphological characters including the
protrusible arolium on the pretarsus and the asymmetricmouthparts (right mandible is absent). Molecular analyses
also strongly support the monophyly of the Thysanoptera(CRESPI et al. 1996).
The current classification of Thysanoptera consists ofapproximately 5500 species in two suborders, Tubulifera
and Terebrantia, and nine families, with Phlaeothripidae
alone constituting the Tubulifera. The only recent attempt toassess the phylogeny of the order as a whole (MOUND et al.
1980) raises several questions about the relationships of thefamilies within the order that are unresolved with morphological methods. One such question was that of the evolutionary origins of the Phlaeothripidae, which possesses somany autapomorphic characters that it could either be sistertaxon to the remaining eight families (i.e., to Terebrantia) or
it could be derived from the subfamily Panchaetothripinae,
making the Thripidae paraphyletic (see Fig. 1). The former
hypothesis is supported by many apomorphic charactersthat suggest that Phlaeothripidae may be independently
evolved from the Protothysanoptera, and several synapomorphic characters that are shared by all of the Terebrantianfamilies (MOUND et al. 1980). The latter hypothesis is basedon a number of apparent synapomorphies uniting Phlaeothripidae and Panchaetothripinae to the exclusion of theremaining families (as well as the remaining Thripidae).
Some of the questions relating to the relationships of thethysanopteran families were subsequently addressed using
molecular methods by CRESPI et al. (1996), who used frag-154

Proceedings of 1st Dresden Meeting on Insect Phylogeny

Fig. 1. Two alternative hypotheses for thysanopteran familial relationships based on morphological characters (redrawn from
MOUND et al. 1980). A: Phlaeothripidae sister to all other families, with the suborders Tubulifera and Terebrantia as sister taxa. B:
Phlaeothripidae as sister taxon to Panchaetothripinae (a subfamily within Thripidae), making the Terebrantia paraphyletic.

Fig. 2. A phylogenetic hypothesis for the order Thysanoptera based on a maximum likelihood analysis (model TrNef+I+G) of 18S
rDNA. Numbers shown above branches are parsimony bootstrap values (10,000 reps using the ‘fast’ stepwise addition algorithm
in PAUP* 4.0). Current family-level classification for the genera is shown with slender vertical bars, with broken lines indicating
the families whose representatives are not monophyletic in this tree, e.g., Thripidae (which also includes Panchaetothripinae) and
Aeolothripidae. The broad, bold vertical bars are indicative of subordinal (in the case of the ingroup Thysanoptera) or ordinal affiliations.

155

Entomologische Abhandlungen 61 (2)ments of the mitochondrial COI gene and the nuclear ribosomal 18S gene to examine the subordinal and familial relationships within Thysanoptera. Unfortunately, the small
number of taxa (approximately 0.1 % of known species)
included in that study limited the utility of the results.
In our study we attempt to expand the data set of CRESPI etal. (1996) to incorporate a broader range of thrips species.
We obtained sequence data for approximately 600 bp of the
18S gene from 52 species of Thysanoptera representing
most of the thysanopteran families. To these data we added
a number of 18S sequences (taken from Genbank) for arange of other paraneopteran taxa to serve as outgroups.
Maximum likelihood analysis of this data set yielded thetree shown in Fig. 2.
This result is not robust due to a lack of informative characters; however, it does suggest some relationships that are
not reflected in the current hypotheses of thysanopteranphylogeny. This analysis recovers only two significant clades with appreciable levels of support among the ingrouptaxa. One clade is that containing all Tubulifera, supportinga hypothesis of the monophyly of this suborder (i.e., of
Phlaeothripidae), but with no resolution between its two
subfamilies (Phlaeothripinae and Idolothripinae). The otherwell-supported clade contains all sampled Panchaetothripinae. This is an unanticipated result, given that all otherterebrantian taxa are unresolved, and no family groupings
are recovered. MOUND et al. (1980) noted a number of morphological characters in common between the Phlaeothripidae and Panchaetothripinae. Should our results be supported by further data, this would suggest that these characters are plesiomorphic for Thysanoptera, especially because
Panchaetothripinae and then Phlaeothripidae are presently
indicated to be the two most basal clades of Thysanoptera(Fig. 2). Our current results suggest that the present classification may not reflect the phylogeny of the order, but further molecular data is required to confirm this.

References
CRESPI, B., D. CARMEAN, L. VAWTER & C. VON DOHLEN 1996.
Molecular phylogenetics of Thysanoptera. – Systematic
Entomology 21: 79–87.
MOUND, L.A., B.S. HEMMING & J.M. PALMER 1980.
Phylogenetic relationships between the families of recent
Thysanoptera (Insecta). – Zoological Journal of theLinnean Society 69: 111–141.

Phylogeny of the Strepsiptera – Morphological Evidence
HANS POHL1 & ROLF G. BEUTEL2
1Allgemeine und Spezielle Zoologie, Universität Rostock, Universitätsplatz 2, 18055 Rostock, Germany [hans.pohl@biologie.unirostock.de] – 2 Institut für Spezielle Zoologie und Evolutionsbiologie, FSU Jena, Erbertstrasse 1, 07743 Jena, Germany [b5bero@rz.uni-jena.de]Introduction
The Strepsiptera are entomophagous insects. There areabout 600 described extant species world-wide and, in a traditional classification, nine or ten families. The minute first
instar larvae are free-living and represent the infectious
stage. The second instar larvae of all species and the neotenic females of the Stylopidia are endoparasites of representatives of various higher taxa of insects including the
Zygentoma, Blattaria, Mantodea, Orthoptera, Hemiptera,Hymenoptera, and Diptera. Due to their parasitism, they
have evolved extremely modified morphology and life
histories. Conspicuous characters of the males include thereduced forewings, which resemble dipteran halteres, andthe fan-shaped hindwings as well as the ‘raspberry’ compound eyes. Females are always wingless and only those of
the basal Mengenillidae are free living, and partly leave
their larval exuviae. In Stylopidia, only the anterior part of
the female body is externally exposed; the females remain
enclosed in the exuviae of earlier stages, and form a functional unit with them within the host.

Basal relationships of the Strepsiptera
The phylogeny of the Strepsiptera proposed by KINZELBACH(1990) was based on a non-numerical analysis (Hennigian
approach) of morphological characters, mostly of adults,
and is generally accepted today (Fig. 1A). KINZELBACH(1990) suggested the division into two major groups, thebasal Mengenillidia with free living females (not confirmed
for the Baltic amber fossil Mengea) and the Stylopidia withendoparasitic females.
POHL (2002) carried out the first cladistic analysis based onmorphological characters of the first instar larvae (Fig. 1B).
This analysis supported the monophyly of Stylopidia, whereas the monophyly of Mengenillidia could not be verified.

The first instar larvae of the Baltic amber fossil Mengea are
not known. Discrepancies with the topology based on adults
pertain to subtaxa of the Stylopidia, especially the placement of Elenchidae, which are almost certainly not thesister group of Myrmecolacidae as proposed by
KINZELBACH (1990), and the paraphyly of Stylopidae. Xenos
and Pseudoxenos turned out as not closely related withother Stylopidae sensu KINZELBACH (1990), and consequently POHL (2002) re-established Xenidae.
Here we summarize our recent results on the basal relationships of the order based on a cladistic analysis of 38 characters of males of 12 ingroup (including a newly discovered Baltic amber fossil) and 3 outgroup taxa (manuscript
submitted by H. Pohl, R.G. Beutel & R. Kinzelbach:“Protoxenidae fam.n. from Baltic amber – a ‘missing link’in strepsipteran phylogeny”) (Fig. 1C).
The monophyly of Strepsiptera s.l., Strepsiptera excl. thenewly discovered fossil (= sp. a in Fig. 1C), and Strepsiptera
excl. this fossil and Mengeidae (including only Mengea) isstrongly supported. Important apomorphies of the males ofStrepsiptera s.l. are the reduction of the tentorium, distinctly separated ommatidia, eight antennomeres, flabellateantennomeres 3–7, absence of the lacinia, 1-segmented
maxillary palps, absence of the labial palps, mesothoracic
halteres, and a strongly sclerotized abdominal segment 9.
Apomorphies of Strepsiptera excl. the newly discovered
fossil are size reduction (less than 7 mm), a dorsomedianfrontal impression of the head, a strongly shortened or
membranised coronal suture, seven antennomeres or less,absence of the galeae, wings wider than long, an elongatedand shield-like metapostnotum, and abdominal sternites
more heavily sclerotised than the tergites. Strepsiptera s.str.
(i.e., excl. stem lineage) is not supported by many autapomorphies, but one important derived feature is the strongly
reduced labrum of males. Strepsiptera s.str. excl. Eoxenos iswell supported and this implies paraphyly of Mengenillidae,

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A

A

Proceedings of 1st Dresden Meeting on Insect Phylogeny

Mengeidae
Mengenillidae
Corioxenidae

Mengenillidia

Halictophagidae
Callipharixenidae
Bohartillidae
Elenchidae

Stylopidia

Myrmecolacidae
Stylopidae

B

B

Eoxenos
Mengenilla
Elenchidae
Corioxenidae
Halictophagidae
Xenidae

Stylopidia

Myrmecolacidae
Stylopidae

C

C

sp. a
Mengeidae
Eoxenos
Mengenilla
Corioxenidae
Elenchidae
Halictophagidae
Xenidae
Stylopidae
Myrmecolacidae

Stylopidia

Fig. 1. Hypotheses on strepsipteran phylogeny. A: Tree modified from KINZELBACH (1990). Dotted lines indicate uncertain
relationships. B: Tree based on characters of first instar larvae;
strict consensus tree of four minimum length trees (modified
from POHL 2002). C: Tree based on characters of males; one oftwo minimum length trees (modified from a manuscript submitted by Pohl et al.); sp. a = newly discovered species fromBaltic amber.

which are mainly characterised by one plesiomorphic feature, the free living females.
The major step in the evolution of Strepsiptera is the rise of
Stylopidia (97 % of all known species), which are well supported by several character transformations of males (modifications of tarsomeres with specialised adhesive hairs to
attach to the host of the parasitic females, loss of abdominal spiracles). The most striking feature however, is theendoparasitism of the adult females, correlated with highly
unusual morphological character transformations, such as
the formation of a unique type of brood organs and of asecondary birth opening in the cephalothorax.

The ‘Strepsiptera problem’
The phylogenetic placement of Strepsiptera is one of themost enigmatic issues of ordinal-level insect systematics(“the Strepsiptera problem”; KRISTENSEN 1981). Strepsiptera
was first described by ROSSIUS (1793), who placed it in the
Hymenoptera near the Ichneumonidae. At the end of the
19th century, the group had been associated with theEphemeroptera, Odonata, Hymenoptera, Lepidoptera, Diptera, and Siphonaptera (PIERCE 1909).
Recently, the phylogenetic placement centered around four
major hypotheses: (1) as a subordinate group of polyphagan
beetles (CROWSON 1960), (2) as the sister group of Coleoptera (KINZELBACH 1990), (3) as the sister group of Diptera(“Halteria”; WHITING et al. 1997), and (4) as having a position outside of Endopterygota (“Neoptera incertae sedis”;
KRISTENSEN 1991).
The first mentioned hypothesis (1) was based on superficial
similarities and can be ruled out with reasonable certainty.
A non-reductive potential synapomorphy with the Coleoptera (2), the presence of densely setose tarsal attachment
devices (BEUTEL & GORB 2001), has evolved within the
Strepsiptera (Stylopidia) and does not belong to the groundplan of the order (see above). Another potential synapomorphy, the stronger degree of sclerotisation of the abdominal sternites (compared to the tergites) is also invalid:
this condition was absent in the most basal strepsipteranfrom Baltic amber. Posteromotorism is also a derived feature shared by Coleoptera and Strepsiptera. However, thiscondition has evolved within several lineages of insects
independently (e.g., Orthoptera, Dermaptera).
The ‘Halteria’ concept (3) is mainly supported by 18S
rDNA sequences, but the clade comprising Strepsiptera and
Diptera may be due to long-branch attraction (e.g.,
HUELSENBECK 1997). 5.8S rDNA, 28S rDNA, and theengrailed homeobox gene do not support a sistergroup relationship between these two groups (HWANG et al. 1998;
ROKAS & HOLLAND 2000).
The position of Strepsiptera outside of Endopterygota (4)was discussed mainly based on the presence of external
wing buds in late instar larvae of the Strepsiptera. Other
potential plesiomorphies of Strepsiptera are the formationof compound eyes in the second instar larvae (not in the
pupal stage, as in Endopterygota) and the presence of a welldeveloped abdominal segment 11 in the first instar larvae.
A sistergroup relationship between Strepsiptera and theremaining Endopterygota should probably be considered as
the most viable working hypothesis.

References
BEUTEL, R.G. & S. GORB 2001. Ultrastructure of attachment
specializations of hexapods (Arthropoda): evolutionary
patterns inferred from a revised ordinal phylogeny. –Journal of Zoological Systematics and Evolutionary
Research 39(4): 177–207.
CROWSON, R.A. 1960. The phylogeny of Coleoptera. – Annual
Review of Entomology 5: 111–134.
HUELSENBECK, J.P. 1997. Is the Felsenstein zone a fly trap? –Systematic Biology 46: 69–74.
HWANG, U.W., W. KIM, D. TAUTZ & M. FRIEDRICH 1998.
Molecular phylogenetics at the Felsenstein zone:
Approaching the Strepsiptera problem using 5.8S and 28S
rDNA sequences. – Molecular Phylogenetics andEvolution 9: 470–480.
KINZELBACH, R. 1990. The systematic position of Strepsiptera(Insecta). – American Entomologist 35: 292–303.
KRISTENSEN, N.P. 1981. Phylogeny of insect orders. – Annual
Review of Entomology 26: 135–157.
KRISTENSEN, N.P. 1991. Phylogeny of extant hexapods. Pp.
125–140 in: I.D. NAUMANN, P.B. CARNE, J.F. LAWRENCE,
E.S. NIELSEN, J.P. SPRADBERRY, R.W. TAYLOR, M.J.
WHITTEN & M.J. LITTLEJOHN (eds.), The Insects ofAustralia: A textbook for students and research workers,
2nd edn. Edited by CSIRO. – Melbourne University Press.
PIERCE, W.D. 1909. A monographic revision of the twisted winged insects comprising the order Strepsiptera Kirby. –Bulletin of the United States National Museum 66: 1–232.
POHL, H. 2002. Phylogeny of the Strepsiptera based on morphological data of the first instar larvae. – Zoologica
Scripta 31: 123–134.

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Entomologische Abhandlungen 61 (2)ROKAS, A. & P.W.H. HOLLAND 2000. Rare genomic changes asa tool for phylogenetics. – Trends in Ecology and
Evolution 15: 433–478.
ROSSIUS, P. 1793. Observation de M. Rossi sur un nouveau
genre d’lnsecte, voisin des Ichneumons. – Bulletin
Société Philomathique 1: 49.

WHITING, M.F., J.C. CARPENTER, Q.D. WHEELER & W.C.
WHEELER 1997. The Strepsiptera problem: phylogeny of
the holometabolous insect orders inferred from 18S and
28S ribosomal DNA sequenes and morphology. –Systematic Biology 46: 1–68.

Phylogeny of the Neuropterida – Morphological Evidence
and the Molecular Advocatus Diaboli
ULRIKE ASPÖCK1, HORST ASPÖCK2 & ELISABETH HARING1
1Naturhistorisches Museum Wien, Burgring 7, 1014 Wien, Austria [ulrike.aspoeck@nhm-wien.ac.at; elisabeth.haring@nhmwien.ac.at] – 2 Department of Medical Parasitology, Clinical Institute of Hygiene and Medical Microbiology, University of Vienna,
Kinderspitalgasse 15, 1095 Wien, Austria [horst.aspoeck@univie.ac.at]

Neuropterida
Understanding the phylogeny of this basal holometabolan
superorder comprising the orders Raphidioptera,
Megaloptera and Neuroptera, is thought-provokingly correlated with hypotheses on general phenomena: evolution of
aquatic larvae, evolution of complex larval sucking tubes
with extraintestinal digestion, evolution of cryptonephry,production of silk etc. Monophyly of the Neuropterida isbased on rather inconspicuous characters concerning thoracic and abdominal sutures, special musculature of the ovipositor, characters of larval stemmata, and specializations at
the wing-bases (literature summarized in ASPÖCK 2002;
ASPÖCK et al. 2001; KRISTENSEN 1999). A sistergroup relationship with Coleoptera is the preferred hypothesis at present (KRISTENSEN 1999).
Prehennigian and most recent classifications favour
Raphidioptera + Megaloptera against Megaloptera +Neuroptera (the concept of Raphidioptera + Neuroptera isout of the main discussion). Argumentations pro
Raphidioptera + Megaloptera (summarized in WILLMANN
2003) base on symplesiomorpies (e.g., the gula), or appear
to be highly sophisticated (postepimeron-complex, polyintersegmental muscle between abdominal tergites 2–5,
reduction of tergite 2, various ovariole-concepts). Overallsimilarity might have been an inductive factor for thisapproach. Arguments pro Megaloptera + Neuroptera resulted from a morphological approach mainly comprising larval characters (head capsules, mouthparts, primary aquatic
lifestyle; ASPÖCK 1995) and were corroborated by a cladistic computer analysis based on a broad pallet of morphological characters (ASPÖCK et al. 2001). The cladogenesis of
this group as deduced from this analysis is summarized inFig. 1. Additional support comes from unpublished molecular data (Haring & Aspöck in prep.: “Molecular phylogeny of the Neuropterida”). Synapomorphies ofMegaloptera + Neuroptera are rosette-like organized trichobothria on the ectoproct, male gonocoxites of segment 9
modified to appendices of the tergite, aquatic larvae, integration of the cardines (basal parts of the maxillae) into thehead capsule, and elongation of larval stipites (ASPÖCK et al.
2001; ASPÖCK 2002).
The monophyly of Raphidioptera (including families
Raphidiidae and Inocelliidae), which has never been questioned, is reflected by several adult characters: subcosta
running into costa, third tarsale enlarged, tergite and sternite of abdominal segment 9 in the male amalgamated to aring, and ovipositor of the female elongated to form a hoselike structure. Also paraphyly of Megaloptera (including
families Corydalidae and Sialidae) is meanwhile no longer
discussed. Monophyly of this order is supported by specia-

lizations in the organization of somatic ovarian tissues andthe tracheal gills of the larvae. The monophyly of
Neuroptera has never been doubted and is mainly based on
the complex sucking tubes of the larvae and the silken
cocoon of the pupae.

Controversies on splitting events in Neuroptera
The great challenge in Neuropterida are sistergroup relationships within the extremely heterogeneous Neuroptera(17 families). Morphological analyses (commented in
ASPÖCK 2002) revealed three monophyletic groups, either(Nevrorthiformia + Myrmeleontiformia) + Hemerobiiformia, or Nevrorthiformia + (Myrmeleontiformia + Hemerobiiformia) (Fig. 1). The controversial hypotheses derived
from the above cited cladistic analyses concern Myrmeleontiformia: (Psychopsidae + Nemopteridae) + (Nymphidae + (Myrmeleontidae + Ascalaphidae)) versus Psychopsidae + (Nemopteridae + (Nymphidae + ( Myrmeleontidae+ Ascalaphidae))) is still unsettled but of minor relevancecompared with controversies in Hemerobiiformia. This
suborder comprises an assembly of 11 extremely heterogeneous families. Arguments in support for monophyly of theeidonomically primitive basal group Ithonidae (including
Rapismatidae) + Polystoechotidae rest on the morphologyof their scarabaeiform blind larvae (the larvae ofRapismatidae are, however, unknown) and on the hypothesis of phytosuccivory of these larvae as root suckers. Allother families constitute the ‘higher Hemerobiiformia’,with constricted larval cardines as synapomorphy. A sistergroup relationship of Osmylidae + Chrysopidae revealed
from the analyses is only weakly supported, and also
Coniopterygidae + Sisyridae need corroboration. Only thedilarid clade comprising Dilaridae + (Mantispidae +(Rhachiberothidae + Berothidae)), with strong synapomorphies of the larvae (flattened heads, broad insertions of the
cardines, elongation of the postmentum), has never been
doubted.
If we accept aquatic larvae as a synapomorphy of Megaloptera and Neuroptera, terrestrial larvae (and their cryptonephries) must have evolved twice in a concept (Nevrorthiformia + Myrmeleontiformia) + Hemerobiiformia, but only
once in a concept Nevrorthiformia + (Myrmeleontiformia +Hemerobiiformia). However, in the latter case the terrestrial clade of the Hemerobiiformia contains aquatic Sisyridae.
This hypothesis involves secondarily evolved aquatic larvae(with one cryptonephric malpighian tubule as reminiscenceof a terrestrial ‘intermezzo’) in this family. The compact
specialized larval head capsule (with several autapomorphies concerning the tentorium, and the enormous sucking

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Fig. 1. Simplified preferred cladogram of the Neuropterida basedon morphological and biological characters (for details see
ASPÖCK et al. 2001 and ASPÖCK 2002).

Fig. 2. Simplified preferred cladogram of the Neuropterida
based on molecular characters (Haring & Aspöck in prep.).

tubes) of the Myrmeleontiformia apparently evolved from a
simple compact head, similar to recent Nevrorthidae, andnot as a reversal of an open maxillary head, as in recent
Hemerobiiformia, which is not compact at all, but predominantly constructed by basal pieces of the maxillae – therefore the term ‘maxillary head’. In the concept of monophyletic Hemerobiiformia the maxillary head is certainly themost important synapomorphy, symbolizing the splitting of
the two suborders.
Our molecular analysis corroborates the sistergroup relationship of Megaloptera and Neuroptera, the Nevrorthiformia as the sister group of the remaining Neuroptera, andthe monophyly of Myrmeleontiformia. A result in conflictwith our previous (morphology-based) concept is that the
Hemerobiiformia do not turn out as a monophyletic group(Fig. 2). As a consequence we are faced with a parallel
evolution of the maxillary head – which is at least not parsimonious at all. Another conflict concerns straight sucking
tubes in the larvae, a specialization in correlation with certain food strategies, which were considered as a synapomorphy of certain families. Straight sucking tubes could
also have evolved several times independently, which would
be the necessary assumption in the context of the molecularanalysis and avoiding the interpretation of curved sucking
tubes being reversals.
It should be mentioned that in the DNA trees the Ithonidae

are not monophyletic. This primitive but nonetheless specialized group as well as the highly specialized
Coniopterygidae – usually apostrophized as the ‘early offshoots’ in previous classical as well as in recent nonhennigian concepts – do not emerge basally in the two comprehensive analyses, neither in the morphological nor in themolecular approach, but nevertheless remain enigmatic.

References
ASPÖCK, U. 1995. Neue Hypothesen zum System der
Neuropterida. – Mitteilungen der Deutschen Gesellschaft
für Allgemeine und Angewandte Entomologie 10:633–636.
ASPÖCK, U. 2002. Phylogeny of the Neuropterida (Insecta:
Holometabola). – Zoologica Scripta 31: 51–55.
ASPÖCK, U., J.D. PLANT & H.L. NEMESCHKAL 2001. Cladistic
analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida:
Neuroptera). – Systematic Entomology 26: 73–86.
KRISTENSEN, N.P. 1999. Phylogeny of endopterygote insects,
the most successful lineage of living organisms. –European Journal of Entomology 96: 237–253.
WILLMANN, R. 2003. Die phylogenetischen Beziehungen derInsecta: Offene Fragen und Probleme. – Pp. 1–64 in:Verhandlungen Westdeutscher Entomologentag 2001,
Löbbecke Museum, Düsseldorf.

Molecular Phylogeny of Neuropterida with Emphasis on the Lacewings(Neuroptera)
SHAUN L. WINTERTON
Department of Entomology, North Carolina State University, Raleigh, North Carolina, 27695, USA [wintertonshaun@netscape.net]

The Neuropterida, comprising the orders Megaloptera(dobsonflies, alderflies), Raphidioptera (snakeflies) and
Neuroptera (= Planipennia, lacewings), are considered oneof the most basal, or ‘plesiomorphic’, groups ofHolometabola. While numerically they are a small to medium sized group, there is considerable morphological diversity across the ca. 21 families within the clade. This diversity has made estimates of homology difficult to reconcile
across families, and while Neuropterida has undergone
numerous taxonomic and morphological studies, little progress has been made towards developing an overarching
phylogeny. Indeed, the phylogeny of Neuroptera published

by WITHYCOMBE (1925) has served as the basis for our
understanding of the evolution of order until very recently.
Recent important morphological studies using cladistic
methodology (e.g., ASPÖCK et al. 2001) have proposed new
hypotheses of neuropterid relationships. Unfortunately,
these hypotheses are based on relatively few characters andhave left many unanswered questions (or poorly supported
conclusions) regarding the position of several key families
and family groups. Moreover, the relationships of the threeorders, Megaloptera, Raphidioptera and Neuroptera, aresubject to considerable debate.
I am presently examining the phylogeny of Neuropterida,

Entomologische Abhandlungen 61 (2)159

Fig. 1. Phylogeny of Neuropterida: strict consensus tree of 39 most parsimonious trees (length = 1220 steps; CI = 0.62; RI = 0.65)
based on combined morphological characters and 18S rDNA gene sequence.

with emphasis on Neuroptera, using data from 18S ribosomal DNA and adult and larval morphology. A phylogeny
based on preliminary data of 2.2 kb of 18S rDNA and 56
adult and larval morphological characters from 38 genera of
Neuropterida and three coleopteran outgroups is presented
in Fig. 1. There is extensive node resolution throughout the
Neuroptera, with both the major suborders, Myrmeleontiformia and Hemerobiiformia, being resolved (the third suborder, Nevrorthiformia, here not included). Notable exceptions include the positions of Osmylidae, Dilaridae, andSisyridae + Coniopterygidae. There is general agreementwith the morphology-based phylogenetic hypothesis of
ASPÖCK et al. (2001) in regard to the placement of various
families within each suborder, although with some notable

exceptions. Unfortunately, there are fewer than 30 % of thenodes in the tree generated from the equally weighted analysis that have bootstrap values above 50 %. Moreover, theMegaloptera are not recovered, while the Raphidioptera and
Neuroptera are both well supported. Several important taxa
are yet to be included in the analysis, such as Nevrorthidae,
Coniopterygidae, Rhachiberothidae and Polystoechotidae,
as well as more genera of Megaloptera. All of these taxa are
crucial to the final analysis and will likely influence thefinal analysis greatly. Therefore, I consider these results as
only preliminary and indicative of trends in the data, awaiting the addition of further characters and completed taxon
sampling.

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Acknowledgements

References

Thank you to the Neuroptera community for their support forthis project, especially those numerous people who provided
specimens for DNA sequencing. Thank you also to Brian M.
Wiegmann for his support of the project and Brian Cassel forhelp with sequencing. This research is supported by a National
Science Foundation research award (DEB-0236861). Views
and statements presented here do not necessarily reflect thoseof the National Science Foundation.

ASPÖCK, U., J.D. PLANT & H.L. NEMESCHKAL 2001. Cladistic
analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida:
Neuroptera). – Systematic Entomology 26: 73–86.
WITHYCOMBE, C.L. 1925. Some aspects of the biology andmorphology of the Neuroptera. With special reference tothe immature stages and their possible phylogenetic significance. – Transactions of the Entomological Society of
London 15: 303–411.

Basal Splitting Events in Coleoptera
ROLF G. BEUTEL1 & FABIAN HAAS2
1
2

Institut für Spezielle Zoologie und Evolutionsbiologie, FSU Jena, Erbertstrasse 1, 07743 Jena, Germany [b5bero@rz.uni-jena.de] –Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany [haas.smns@naturkundemuseum-bw.de]

Introduction
With more than 360,000 described species Coleoptera areby far the largest group of Metazoa. Considering this unusual diversity, a major issue for coleopterists should be thereasons for this evolutionary success. However, for thedevelopment of possible scenarios, a well founded phylogenetic hypothesis for the major groups is needed.
Surprisingly the crucial question of the interrelationships ofthe four suborders is still discussed controversially.

Current hypotheses
KUKALOVÁ-PECK & LAWRENCE (1993) have postulated asistergroup relationship between Polyphaga and the remaining Coleoptera, and between Archostemata andMyxophaga + Adephaga. This hypothesis was based on
characters of the wing venation and articulation and a nonnumerical character evaluation. It is confirmed in a hitherto unpublished study of these authors with a cladistic analysis of a similar data set. A phylogenetic scheme which was
already proposed by CROWSON (1960) – Archostemata +(Adephaga + (Myxophaga + Polyphaga)) – was supported
by a cladistic analysis of 107 characters of adults and immature stages (BEUTEL & HAAS 2001). A basal position ofArchostemeta was confirmed by an analysis of 18S rDNA
sequences presented by CATERINO et al. (2002). However,Archostemata

Adephaga

Adephaga was placed as the sister group of Polyphaga inthis contribution.

Scenario of splitting events
Coleoptera is characterised by many autapomorphies, notably the rigidly connected sclerites, the absence of exposed
membranes, elytra with epipleura, folded alae, a prognathous, wedge-shaped head, invaginated terminal abdominal
segments, and the loss of several thoracic muscles. The
retention of a considerable number of plesiomorphies inadults of Archostemata (e.g., transverse ridge of mesoventrite present, metatrochantin exposed, presence of M. notosternalis 2) is most easily explained with a basal position of
the group within Coleoptera. The monophyly of the suborder is well supported, however almost exclusively by larvalfeatures, which are correlated with wood-boring habits:
strongly sclerotised wedge-shaped head, shortened or reduced legs, unsclerotised cylindrical body, and presence of
tergal ampullae. Potential synapomorphies of Adephaga,
Myxophaga and Polyphaga are the loss of nine homologous
thoracic muscles, the absence of an exposed metatrochantinus, the loss of the transverse ridge of the mesoventrite, andthe primary absence of the mesothoracic katepisternal joint(similar structure present in some scirtids). The fused tibia
and tarsus of larvae, the single larval claw, the prothoracic
trochantinopleura, and several muscle losses are potential
Myxophaga

Polyphaga

8.1–11.1

5.1–7.1

1.1–4.1

Fig. 1. Hypothesis of basal splitting events in Coleoptera. Apomorphies mapped on cladogram: 1.1 absence of exposed membranes,
2.1 elytra with epipleura, 3.1 wing folding, 4.1 muscle losses; 5.1 loss of nine thoracic mucles, 6.1 external metatrochantinus
absent, 7.1 transverse ridge of mesoventrite absent, 8.1 larval tibiotarsus, 9.1 single claw of larvae, 10.1 muscle losses, 11.1 meso- andmetaventrites firmly connected.

161

Entomologische Abhandlungen 61 (2)synapomorphies of Myxophaga and Polyphaga. The interpretation of firmly fused meso- and metaventrites as a synapomorphy of these suborders implies reversal in somegroups of Polyphaga (most scirtoids, derodontids, few
agyrtids and leiodids, some cantharoids).
A scenario with Archostemata as basal group is better inagreement with the fossil record than the hypothesis of
KUKALOVÁ-PECK & LAWRENCE (1993). Archostemata andAdephaga are well represented before the earliest records of
Polyphaga.

Acknowledgements
We are grateful to Jarmila Kukalová-Peck and John F.

Lawrence for sending us an unpublished manuscript and forvaluable information and stimulating discussions.

References
BEUTEL, R.G. & F. HAAS 2000. Phylogenetic relationships ofthe suborders of Coleoptera (Insecta). – Cladistics 16:
103–141.
CATERINO, M.S., V.L. SHULL, P.M. HAMMOND & A.P. VOGLER
2002. Basal relationships of Coleoptera inferred from 18S
rDNA sequences. – Zoologica Scripta 31(1): 41–49.
CROWSON, R.A. 1960. The phylogeny of Coleoptera. – Annual
Review of Entomology 5: 111–134.
KUKALOVÁ-PECK, J. & J.F. LAWRENCE 1993. Evolution of thehind wing in Coleoptera. – Canadian Entomologist 125:
181–258.

The Basal Relationships of Coleoptera Based on 18S rRNA Sequences
ALFRIED P. VOGLER1 & MICHAEL S. CATERINO2
1

The Natural History Museum, Cromwell Road, London, SW7 5BD, and Imperial College London, Silwood Park Campus, Ascot,
Berkshire SL5 7PY, United Kingdom [apv@nhm.ac.uk] – 2 Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road,
Santa Barbara, CA 93105, USA [mcaterino@SBNATURE2.ORG]

Together with numerous collaborators, our laboratory has
accumulated 18S rRNA sequences which are used here tobuild a comprehensive tree of the Coleoptera. Sequenceswere compiled from Genbank and unpublished data. Afterremoval of incomplete and low quality sequences, a total of
795 sequences representing ca. 130 families (of 160–170
total in Coleoptera) were used in the phylogenetic analysis.
Sequence alignment was achieved in a stepwise procedureusing the ClustalW software, where taxa from undoubtedly
monophyletic groups at the level of superfamilies were aligned separately (including some 30 sequences ofHisteroidea, 150 sequences of Chrysomeloidea, etc.), prior
to aligning them to hierarchically higher groups using the‘profile alignment’ option in ClustalW. The aligned matrix
was subjected to parsimony searches, and yielded shortest
trees of 28,950 steps (CI = 0.133, RI = 0.705).
The basal relationships of suborders were consistently
found to be Archostemata + (Myxophaga + (Adephaga +Polyphaga)), in accordance with previous findings from amuch smaller subset of these data (CATERINO et al. 2002).
Rates of change and number of inferred indels were muchgreater in Adephaga than Polyphaga, and hence both groups
were analysed separately. The analysis of adephagan relationships included 220 taxa and a representative sample of all
major groups. Parsimony searches yielded a tree of 14,166
steps (CI = 0.221, RI = 0.677). The tree showed the wellestablished (on the basis of 18S rRNA) separation of Geadephaga and Hydradephaga taxa, and the unexpected placement of some presumably basal groups (Cicindelidae,
Rhysodidae, Paussidae) within derived carabid groups(SHULL et al. 2001), even with the more comprehensive
sampling in this study. Relationships within Hydradephaga
included the close relationships of Dytiscidae, Aspidytidae,
Hygrobiidae, and Amphizoidae, as described previously(RIBERA et al. 2002).
The analysis of Polyphaga included 575 taxa in over 110
families and produced shortest trees of 14,747 steps (CI =0.173, RI = 0.651). Among the five Series of families in thesuborder, the Elateriformia represented the most basalgroup, as sister to all other Polyphaga. Within
Elateriformia, a clade comprising Eucinetoidea (including
families Scirtidae, Eucinetidae, Clambidae, Decliniidae)

and the bostrichiform Derodontidae was sister to all otherfamilies. The remaining elateriform groups include clearlymonophyletic Dascilloidea and Buprestoidea, and larger
clades roughly corresponding to Elateroidea andByrrhoidea of previous authors, with all aquatic families ofthe latter in close proximity.
The following nodes in the tree correspond to a clade ofmonophyletic Staphyliniformia plus Scarabaeiformia, andBostrichiformia paraphyletic with respect to the latter twogroups. In the Staphyliniformia we find (weak) evidence fortwo major lineages, with the Staphylinidae (including
several smaller families) as sister to a clade of Hydraenidae,
Ptiliidae, Agyrtidae, Leiodidae, Silphidae, plus three families of Histeroidea. The aquatic Hydrophilidae surprisingly
showed affinities to the basal Scarabaeiformia.
The Cucujiformia, the largest Series of Polyphaga, wasrepresented in our analysis by 330 taxa of 51 families andclearly recovered as monophyletic. Among the six superfamilies within Cucujiformia, the ecologically diverse
Cucujoidea was paraphyletic with respect to Cleroidea. The
latter grouped as sister to Kateretidae, with Trogossitidae
occupying the most basal node within the Cleroidea. Fiveclades of 3–5 families each were well supported in
Cucujoidea, but one of these (including families Cryptophagidae, Latridiidae, Phalacridae, and Corylophidae)
grouped within Tenebrionoidea. This is likely due to long
branch attraction, also affecting the placement of Lymexyloidea in this part of the tree. Further we find Curculionoidea plus Chrysomeloidea (the ‘Phytophaga’) to bemonophyletic, with basal relationships mainly confirming
morphological analyses.
In conclusion, we find that morphological classifications ofprevious authors, in particular CROWSON (1960) and
LAWRENCE & NEWTON (1982), are generally confirmed by
the 18S rRNA gene, but our tree needs to be subjected tofurther testing, using more complete searches, and adding
other markers. Methods are needed to assemble large trees
from subsets of closely related sequences, and to assess
support in very large trees. However, this tree is the mostcomplete phylogenetic hypothesis of basal Coleoptera relationships to date, and can be used to test morphological andecological diversification of the group.

162

References
CATERINO, M.S., V.L. SHULL, P.M. HAMMOND & A.P. VOGLER
2002. The basal phylogeny of the Coleoptera inferredfrom 18S rDNA sequences. – Zoologica Scripta 31:
41–49.
CROWSON, R.A. 1960. The phylogeny of Coleoptera. – Annual
Review of Entomology 5: 111–134.
LAWRENCE, J.F. & A.F. NEWTON 1982. Evolution and classification of beetles. – Annual Review of Ecology and
Systematics 13: 261–290.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

RIBERA, I., J.E. HOGAN & A.P. VOGLER 2002. Phylogeny of
hydradephagan water beetles inferred from 18S rRNA
sequences. – Molecular Phylogenetics and Evolution 23:
43–62.
SHULL, V.L., A.P. VOGLER, M.D. BAKER, D.R. MADDISON &P.M. HAMMOND 2001. Sequence alignment of 18S ribosomal RNA and the basal relationships of adephagan beetles: evidence for monophyly of aquatic families and theplacement of Trachypachidae. – Systematic Biology 50:
945–969.

Towards a Consensus: Latest Results from Simultaneous Analysis of theBasal Hymenopteran Lineages
LARS VILHELMSEN
Zoological Museum, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen Ø, Denmark [lbvilhelmsen@
zmuc.ku.dk]

The phylogeny of the basal hymenopteran lineages has been
the subject of intense scrutiny since the groundbreaking
work of A.P. Rasnitsyn (summarized in RASNITSYN 1988).
Although his phylogenetic hypothesis was derived intuitively, many of the relationships proposed in his paper are
upheld by later cladistic analyses, including the reanalysis
of his dataset by RONQUIST et al. (1999). VILHELMSEN (1997,
2001) compiled comprehensive morphological datasetsfrom a number of sources and performed computerized analyses. SCHULMEISTER et al. (2002) and SCHULMEISTER(2003) presented the first simultaneous analyses of basal
hymenopteran phylogeny, expanding the morphologicaldataset of VILHELMSEN (2001) as well as providing molecular data from five genes (12S, 16S, 18S, and 28S rDNA and
COI), including a total of 343 morphological characters andalmost 3,000 basepairs, 1,136 of which were phylogenetically informative. More than half of the informative sites
were provided by COI.
There is substantial agreement between the results of thelatest computerized analyses (Fig. 1). The monophyly of theHymenoptera is strongly supported by a host of morphological characters (VILHELMSEN 2001). Prominent autapomorphies are: laterocervicalia fused with propleuron, articulation between propleuron and profurcal arm present, anterior
apical protibial spur modified into calcar, cenchrus present,
metafurca arising anteriorly on discremen, forewing analveins not reaching wing margin, distal hamuli present,
abdominal spiracles surrounded by sclerotized cuticle, cordate apodeme on female abdominal tergite 9 present, basal
ring and volsella in male genitalia present, and common
cornea in larval eye present. Molecular analyses also provide some support (SCHULMEISTER 2003). The monophyly ofthe order has never really been disputed.
The monophyly of most of the basal superfamilies within
the Hymenoptera as currently defined is strongly corroborated. The relationships between them as suggested by themorphology alone and the simultaneous analyses are practically identical, many nodes being well supported.
However, in some cases there is conflict between the morphological and the molecular data. Under some circumstances, analysis of the morphological data alone does notplace Xyelidae as the sister group to the remaining
Hymenoptera. The presence of a separate mesothoracic
anepisternum and of an unbranched Rs vein in the forewing
provides some support for the monophyly of non-xyelid
Hymenoptera, as does the molecular data; however, othermorphological characters contradict this. The Xyelidae
always comes out as well supported, autapomorphies being
found in the antennal structure (enlarged third segment) andespecially in the mouthparts which have the labrum, mandibles and hypopharynx adapted to grind pollen. However,these features are also found in many fossils and in some
outgroup taxa, respectively. The difficulties with resolving
the basal splitting events within the Hymenoptera might bedue to difficulties with polarizing the characters, caused by
the highly autapomorphic status of the order.
The Blasticotomidae consistently comes out as the sistergroup to all other Tenthredinoidea (the Tenthredinoidea
s.str.). Otherwise, the phylogeny of this superfamily is poorly resolved and weakly supported, with considerable differences between morphological and molecular hypotheses; itis necessary to sample tenthredinoid diversity much more
densely as well as expand the character sets to derive arobust phylogeny. The next superfamily to branch off is thePamphilioidea, then the Cephoidea. The placement of thefamily Anaxyelidae is contentious; molecular data supports
a sistergroup relationship with Siricidae, whereas the morphological data places it as sister to Siricidae + Xiphydriidae + Orussidae + Apocrita. The latter hypothesis is weakly supported, as substantial character conflict occurs. The
sistergroup relationship between Orussidae and Apocrita isvery strongly supported by morphology. However, analysis of
the molecular data alone fails to retrieve this relationship.
In general, the accumulation of data leads to an increasingly
robust phylogeny for the basal hymenopteran lineages. The
consensus phylogeny supports the scenario for the evolution of larval lifestyle in basal Hymenoptera depicted in Fig.
2. The larva of the common ancestor of all Hymenoptera
was herbivorous. The transition from the herbivorous to thecarnivorous/parasitoid lifestyle took place in the common
ancestor of Orussidae + Apocrita. Prior to this transition,
Hymenoptera had invaded the woody habitat, having woodboring larvae (as in extant woodwasps: the Anaxyelidae,
Siricidae, and Xiphydriidae), a condition that apparently
immediately predated the evolution of parasitism. This
scenario is corroborated by many basal parasitoid taxa (e.g.,
Orussidae, Megalyridae, Stephanidae, many Ichneumonoidea) being ectoparasitoids of woodboring insect larvae,presumably the lifestyle adopted by the ancestral parasitoid.
Many of the morphological changes occurring within thebasal Hymenoptera can be correlated with the transition to
the parasitoid lifestyle. For example, the adult mouthparts

163

Entomologische Abhandlungen 61 (2)

x
xx

x
x

x
x

x
x
xxx

A

Xyelidae
Blasticotomidae
Athaliaother Tenthredinidae
Nematus
Diprionidae
Cimbicidae
Argidae
Pergidae
Megalodontesidae
Pamphiliidae
CephidaeAnaxyelidae
Siricidae
Xiphydriidae
Orussidae
APOCRITA
B

x

x
x

x
x

x
x
x

XYELOIDEA
Xyelidae
Blasticotomidae
Argidae
Pergidae
TENTHREDINOIDEA
Athaliaother Tenthredinidae
Cimbicidae
Diprionidae
Megalodontesidae
PAMPHILIOIDEA
Pamphiliidae
Cephidae
CEPHOIDEA
Anaxyelidae
SIRICOIDEA
Siricidae
XIPHYDRIOIDEA
Xiphydriidae
ORUSSOIDEA
Orussidae
APOCRITA

st
ic
ot
Te
om
nt
id
hr
ae
ed
in
oi
de
a
s.
st
Pa
r.
m
ph
ili
M
i
da
eg
e
al
Ce odo
nt
ph
es
id
id
ae
ae
An
ax
ye
lid
Si
ae
ric
Xi ida
ph e
yd
rii
da
Or
e
us
si
da
e
AP
OC
RI
TA

Bl
a

Xy
e

lid

ae

Fig. 1. Phylogeny of the basal lineages of Hymenoptera according to A: VILHELMSEN (2001; only morphological characters); B:
SCHULMEISTER (2003; simultaneous analysis). Superfamily classification superimposed. Number of X’es below nodes indicatesdegree of clade support (the more X’es, the stronger the support).

Herbivorous

Herbivorous

Woodboring
Woodboring
Parasitoid
Parasitoid

Fig. 2. Early evolutionary history of the Hymenoptera, with thelarval lifestyle mapped onto the topology in Fig. 1B.

display posterior displacement and subsequent reduction of
the labrum (VILHELMSEN 1996); these changes permit themandibles to be employed more efficiently when chewing
an escape tunnel from the pupal chamber, which is paramount for taxa pupating in wood. Later in the evolutionary
history of the Hymenoptera, the mandibles were used as
tools in numerous other circumstances (e.g., many social
Hymenoptera; JERVIS & VILHELMSEN 2000). The ovipositor
apparatus is another instance of a character system having
undergone extensive modification throughout the order(QUICKE et al. 1999; VILHELMSEN 2000) in correlation with
oviposition in different substrates. A very elongate ovipositor was probably present in the common ancestor ofOrussidae + Apocrita; this was useful for penetrating deep
into the substrate of its presumably concealed host, but caused problems with handling when used to drill in wood. Theinternalized ovipositor of the Orussidae and the wasp-waistof the Apocrita, respectively, can be interpreted as different
solutions to these problems (VILHELMSEN et al. 2001).
Finally, the extensive reductions in larval anatomy (absenceof eyes, laterocervicalia, thoracic and abdominal legs;
reduction of antennae and mouthparts; see VILHELMSEN
2003) that can be inferred to have taken place between theancestral hymenopteran and the ancestral parasitoid wasp
can also be correlated with the transition from an externally living herbivorous lifestyle to ectoparasitism of a con-cealed host; the latter lifestyle is probably much less demanding of the sensory and locomotory apparatus.

References
JERVIS, M. & L. VILHELMSEN 2000. The occurrence and evolution of nectar extraction apparatus among Hymenoptera‘Symphyta’. – Biological Journal of the Linnean Society
70: 121–146.
QUICKE, D.L.J., A. LERALEC & L. VILHELMSEN 1999.
Ovipositor structure and function in the parasitic
Hymenoptera. – Rendiconti 47: 197–239.
RASNITSYN, A.P. 1988. An outline of the evolution of the hymenopterous insects (order Vespida). – Oriental Insects 22:
115–145.
RONQUIST, F., A.P. RASNITSYN, A. ROY, K. ERIKSON & M.
LINDGREN 1999. Phylogeny of the Hymenoptera: A cladistic reanalysis of Rasnitsyn’s (1988) data. – Zoologica
Scripta 28: 13–50.
SCHULMEISTER, S. 2003. Simultaneous analysis of basal
Hymenoptera (Insecta): introducing robust-choice sensitivity analysis. – Biological Journal of the Linnean Society
79: 245–275.
SCHULMEISTER, S., W.C. WHEELER & J.M. CARPENTER 2002.
Simultaneous analysis of the basal lineages ofHymenoptera (Insecta) using sensitivity analysis. –Cladistics 18: 455–484.
VILHELMSEN, L. 1996. The preoral cavity of lower
Hymenoptera (Insecta): comparative morphology andphylogenetic significance. – Zoologica Scripta 25:
143–170.
VILHELMSEN, L. 1997. The phylogeny of lower Hymenoptera(Insecta), with a summary of the early evolutionary
history of the order. – Journal of Zoological Systematics
and Evolutionary Research 35: 49–70.
VILHELMSEN, L. 2000. The ovipositor apparatus of basal
Hymenoptera (Insecta): phylogenetic implications andfunctional morphology. – Zoologica Scripta 29: 319–345.
VILHELMSEN, L. 2001. Phylogeny and classification of theextant basal lineages of the Hymenoptera (Insecta). –Zoological Journal of the Linnean Society 131: 393–442.
VILHELMSEN, L. 2003. Larval anatomy of Orussidae(Hymenoptera). – Journal of Hymenoptera Research 12:346–354.
VILHELMSEN, L., N. ISIDORO, R. ROMANI, H.H. BASIBUYUK &D.L.J. QUICKE 2001. Host location and oviposition in abasal group of parasitic wasps: the subgenual organ, ovipositor apparatus, and associated structures in theOrussidae (Hymenoptera, Insecta). – Zoomorphology
121: 63–84.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

The Pattern of Relationships among Superfamilies of Apocritan Hymenoptera Derived from Recent Morphological and Molecular Analyses
ANDREW AUSTIN1, MARK DOWTON2 & ANDREW DEANS3
1

Centre for Evolutionary Biology & Biodiversity, School of Earth & Environmental Sciences, The University of Adelaide, SA
5005, Australia [andrew.austin@adelaide.edu.au] – 2 Institutes of Biomolecular Science and Conservation Biology, Department of
Biology, Wollongong University, Wollongong, NSW 2522, Australia [mdowton@uow.edu.au] – 3 Department of Entomology,
University of Illinois, Urbana, IL 61801, USA [adeans@life.uiuc.edu]

Introduction
The Apocrita represent a massive radiation of insects that
may comprise 20–25 % of all insect species. This hugediversity is largely due to the evolution of several biologicalattributes: (1) parasitism where various groups have radiated to exploit other insects and spiders as hosts, (2) eusociality among the higher aculeates, and (3) various forms ofspecialised predation and phytophagy (e.g., gall-forming,pollen/nectar-feeding). The Apocrita comprise the vast
majority of Hymenoptera and they are defined by the unique character of having the first abdominal segment (propodeum) broadly fused to the posterior thorax, and a narrow
waist developed between the first and second abdominal
segments. The current classification recognises 13 superfamilies and 76 families, and of these the parasitic
Ichneumonoidea and Chalcidoidea, and the aculeate
Vespoidea (including the ants) and Apoidea constitute themost diverse groups by far.

Morphological studies
While recent studies have made significant headway inresolving the phylogeny among basal Hymenoptera (the‘Symphyta’, or sawflies), relationships within the Apocrita
remain poorly resolved, even though numerous recent studies have focused on this problem. As with sawfly phylogenetic research, RASNITSYN’s (1988) benchmark study isoften considered to be the ‘modern’ starting point for phylogenetic research on the Apocrita. The tree presented by
RASNITSYN (1988), although generated intuitively, has formed a basis for most subsequent work and intitiated substantial interest in hymenopteran phylogeny, resulting in
numerous detailed anatomical studies. Although these studies have identified a number of potentially informative
character systems, none have yet been incorporated into
broad analyses of apocritan relationships.
Overall, two studies stand out as being critically important.
RONQUIST et al. (1999) reanalysed RASNITSYN’s (1988) data
within a cladistic framework, and revealed a substantiallydifferent set of relationships compared with those predictedin the original intuitive tree. For example, this study (Fig. 1)
found no support for Ichneumonoidea + Aculeata, or
Ceraphronoidea + Evanioidea + Trigonaloidea/Megalyroidea/Stephanoidea (i.e., Rasnitsyn’s Evaniomorpha), but did
broadly support the Proctotrupomorpha (i.e., Chalcidoidea+ Platygastroidea + Cynipoidea + Proctotrupoidea, but withquite different internal relationships and the inclusion of
Ceraphronoidea) and the non-monophyly of the Proctotrupoidea. Recently, SHARKEY & ROY (2002), however, havehighlighted a number of shortcomings with RASNITSYN’s(1988) original data, and shown that much of the structurein the RONQUIST et al. (1999) tree is due to reductional characters associated with the wings. They revised all wing

1
Fig. 1. Apocritan relationships indicated by the RONQUIST et al.
(1999) reanalysis of RASNITSYN’s (1988) morphological data
(PR = Proctotrupomorpha).

2Fig. 2. Apocritan relationships developed by SHARKEY & ROY(2002) following reinterpretation of wing characters andreanalysis of the RONQUIST et al. (1999) data matrix (PR =Proctotrupomorpha).

characters, but the resulting tree (Fig. 2) was largely comblike, with the exception of a monophyletic Proctotrupomorpha (including Ceraphronoidea).

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Entomologische Abhandlungen 61 (2)given the species richness and biological diversity of manygroups; and (7) there have been general discrepancies/problems in combining and analysing independent datasets(see CARPENTER & WHEELER 1999 for discussion).

Future studies

Fig. 3. Apocritan relationships following analysis of sequence
data from multiple genes by DOWTON & AUSTIN (2001) (PR =Proctotrupomorpha), using two symphytans as the outgroup.

Where to now? In July 2003 the U.S. National Science
Foundation announced the funding of a ‘Tree of Life’ project whose primary aim is to undertake a large-scale, fully
integrated phylogenetic analysis of the Hymenoptera
employing more than 500 morphological characters andsequences from five genes for a large number of exemplar
taxa. The project will run for five years, will involve 30
research staff and graduate students and will focus on thefamily level for the whole order and at the subfamily level
for the taxon-diverse Aculeata, Chalcidoidea andIchneumonoidea. This task will require an unprecedented
level of scientific cooperation and project management, but
the outcome is likely to be a robust phylogeny that willserve as an evolutionary framework to support future research on the Hymenoptera for many years to come.

Acknowledgements
Molecular studies
Over the last ten years we have undertaken a series of studies employing DNA sequence data in an attempt to resolve relationships within Apocrita, particularly among thenine parasitic superfamilies (DOWTON & AUSTIN 1994,
2001; DOWTON et al. 1997). The most comprehensive of
these (2001) employed sequence data from three genes (16S
and 28S rDNA and COI) and 84 exemplar taxa representing
all apocritan superfamilies. Under different models of analysis a number of relationships were consistently recovered(Fig. 3), including a monophyletic Proctotrupomorpha(excluding Ceraphronoidea), the monophyly of Diapriidae+ Monomachidae + Maamingidae (a family recently described by EARLY et al. 2001), Vanhorniidae + Proctotrupidae,
and Megalyridae + Trigonalidae. Further, Chalcidoidea +Platygastroidea were recovered as sister groups except
under the simplest model, the Proctotrupoidea was always
polyphyletic, and the Ichneumonoidea was often not resolved although this is likely due to very different A-T content
between braconid and ichneumonid mitochondrial genes.
Like SHARKEY & ROY (2002), what is not evident from thisstudy is a stable backbone to the apocritan tree below theProctotrupomorpha (i.e., the basal relationships).
To date, all studies on apocritan phylogeny have been limited in some way. Most important limitations are that (1)many of Rasnitsyn’s characters (employed by RONQUIST etal. 1999 and in part by SHARKEY & ROY 2002) are not cladistic in nature (i.e., states often not properly separated as
discrete entities amenable to cladistic analysis, partly discussed by SHARKEY & ROY 2002); (2) numerous morphological characters have been coded for hypothetical groundplan taxa, not exemplars; (3) morphological nomenclature
is often not standardised and the identification of homologous structures is sometimes problematic; (4) data fromnumerous post-Rasnitsyn anatomical studies on ‘new’ character systems have not been incorporated; (5) at least 16
apocritan families are thought to be paraphyletic or polyphyletic, 14 of which are contained within just three superfamilies – Proctotrupoidea, Chalcidoidea and Apoidea; (6)taxon sampling for molecular studies has been minimal

We wish to thank Mike Sharkey, Nick Stevens and Jim
Whitfield for helpful discussions and input into this paper, andour many colleagues who have helped over the years in numerous ways.

References
CARPENTER, J.M. & W.C. WHEELER 1999. Towards simultaneous analysis of morphological and molecular data in
Hymenoptera. – Zoologica Scripta 28: 251–260.
DOWTON, M. & A.D. AUSTIN 1994. Molecular phylogeny of theinsect order Hymenoptera – apocritan relationships. –Proceedings of the National Academy of Science, USA
19: 9911–9915.
DOWTON, M., A.D. AUSTIN, N. DILLON & E. BARTOWSKY 1997.
Molecular phylogeny of the apocritan wasps with particular reference to the Proctotrupomorpha and Evaniomorpha. – Systematic Entomology 22: 245–255.
DOWTON, M. & A.D. AUSTIN 2001. Simultaneous analysis of
16S, 28S, COI and morphology in the Hymenoptera:
Apocrita – evolutionary transitions among parasitic wasps.
– Biological Journal of the Linnean Society 74: 87–111.
EARLY, J., I.D. NAUMANN, L. MASNER & A.D. AUSTIN 2001.
Maamingidae, a new family of proctotrupoid wasps fromNew Zealand. – Invertebrate Taxonomy 15: 341–352.
RASNITSYN, A.P. 1988. An outline of the evolution of the hymenopterous insects (order Vespida). – Oriental Insects 22:
115–145.
RONQUIST, F., A.P. RASNITSYN, A. ROY, K. ERIKSON & M.
LINDGREN 1999. Phylogeny of the Hymenoptera: A cladistic reanalysis of Rasnitsyn’s (1988) data. – Zoologica
Scripta 28: 13–50.
SHARKEY, M.J. & A. ROY 2002. Phylogeny of the Hymenoptera: a reanalysis of the Ronquist et al. (1999) reanalysis,
emphasizing wing venation and apocritan relationships. –Zoologica Scripta 31: 57–66.

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Phylogeny of Trichoptera
KARL M. KJER1, RALPH W. HOLZENTHAL2 & ROGER J. BLAHNIK2
1
Department of Entomology, Rutgers University, Cook College, New Brunswick, New Jersey 08901, USA [kjer@aesop.
rutgers.edu] – 2 Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, USA [holze001@ umn.edu;
blahn003@umn.edu]

Trichoptera (caddisflies) comprise the largest order of aquatic insects and occur in rivers, streams, and lakes around theworld. They are known for the remarkable cases and silken
nets built by the aquatic larvae. Larvae are integral components of aquatic ecosystems and also key species for assessing water quality in biomonitoring programs. There aresome 45 families of Trichoptera, represented by about
12,000 described species.
The monophyly of Trichoptera is well supported by a number of morphological characters such as the modified hypopharynx (= haustellum) of the adult and the apneustic aquatic larvae with reduced antennae and absence of abdominal
prolegs (except the terminal pygopods). Monophyly of
Trichoptera, as well as its sistergroup relationship to
Lepidoptera, also consistently results from molecular analyses. Despite the long interest in the biology and natural
history of the order, monophyly of the three trichopteransuborders (Annulipalpia, the fixed-retreat makers;
Integripalpia, the portable-case makers; and Spicipalpia, theclosed-cocoon makers) and their relative phylogenetic relationships was unsettled, with several quite different hypotheses having been presented.
The crux of the problem centers on the monophyly of
Spicipalpia (included families Rhyacophilidae, Hydrobiosidae, Glossosomatidae, and Hydroptilidae) and its relationship to the other suborders (each monophyletic in all
hypotheses) as either paraphyletic and allied to Integripalpia (ROSS 1967), monophyletic and allied to Annulipalpia (WEAVER 1984), monophyletic and basal within theorder (WIGGINS & WICHARD 1989), or variously polyphyletic(FRANIA & WIGGINS 1997; IVANOV 2002). MORSE (1997)
provided a summary of these hypotheses as well as a reviewof phylogenetic research within the order.
Using combined and partitioned analyses of molecular andmorphological data, we previously established the phylogenetic relationships among the three trichopteran suborders
as Annulipalpia + (Spicipalpia + Integripalpia) (KJER et al.
2001, 2002), reflecting the hypothesis originally proposed
by ROSS (1967). We tested this hypothesis with additional
data. Taxa were added to the analysis, and gene fragmentsthat had been missing in our original analysis were completed. Taxa with substantial missing data were removed, as
was the EF-1␣ data (which was missing for most taxa). The
program ‘MrBayes 3.0’ (HUELSENBECK & RONQUIST 2001)
now implements mixed models that permit more realistic
combined analyses. We analyzed our data under a combined
analysis of the different partitions, each analyzed according
to its own model, including the MK model for morphological characters (LEWIS 2001), and a doublet model for pairedsites in rRNA (SCHÖNIGER & VON HAESELER 1994). We
included data from nuclear rRNA fragments (28S, D1, D3
and 18S, V4-V5), insertions and deletions from the rRNA
data, a COI mitochondrial DNA fragment, and morphological characters previously presented by FRANIA & WIGGINS(1997). Analysis of 138 taxa revealed the same relationships
among the suborders as we previously reported.
The monophyly of Spicipalpia, while unlikely, is still
equivocal. The higher taxa among Integripalpia include
monophyletic Plenitentoria and Brevitentoria. Within
Brevitentoria, Sericostomatoidea is strongly supported, but

a monophyletic Leptoceroidea is still not recovered, although it cannot be strongly rejected. Relationships within
Annulipalpia, when the morphological data of FRANIA &WIGGINS (1997) are included, closely match the higher levelstructure reported by these authors, with monophyletic
Philopotamoidea and Hydropsychoidea; our results differ
only in the placement of some of the hydropsychoid families. Thus, the implementation of more realistic models,
combined with additional taxa and data, did not substantially alter our original conclusions. For the most part, support indices remained strong where they had been strong,
and weak where they had been weak. Support for more apical family groups, particularly among Limnephiloidea,‘Leptoceroidea’, and Sericostomatoidea remains low, andwill probably require additional characters, rather than amore dense taxon sampling, to resolve these relationships
with any degree of confidence.

Acknowledgements
This material is based upon work supported by the National
Science Foundation under Grant No. 9796097.

References
FRANIA, H.E. & G.B. WIGGINS 1997. Analysis of morphological and behavioral evidence for the phylogeny and higher
classification of Trichoptera (Insecta). – Royal Ontario
Museum, Life Sciences Contributions 160: 1–67.
HUELSENBECK, J.P. & F. RONQUIST 2001. MRBAYES: Bayesianinference of phylogeny. – Bioinformatics 17: 754–755.
IVANOV, V.D. 2002. Contribution to the Trichoptera phylogeny:
new family tree with consideration of TrichopteraLepidoptera relations. – Proceedings of the 10th
International Symposium on Trichoptera, Nova Supplementa Entomologica, Keltern 15(2001): 277–292.
KJER, K.M., R.J. BLAHNIK & R.W. HOLZENTHAL 2001.
Phylogeny of Trichoptera (caddisflies): characterizationsof signal and noise within multiple datasets. – Systematic
Biology 50: 781–816.
KJER, K.M., R.J. BLAHNIK & R.W. HOLZENTHAL 2002.
Phylogeny of caddisflies (Insecta, Trichoptera). –Zoologica Scripta 31: 83–91.
LEWIS, P.O. 2001. A likelihood approach to estimating phylogeny from discrete morphological character data. –Systematic Biology 50: 913–925.
MORSE, J.C. 1997. Phylogeny of Trichoptera. – Annual Review
of Entomology 42: 427–450.
ROSS, H.H. 1967. The evolution and past dispersal of the
Trichoptera. – Annual Review of Entomology 12:
169–206.
SCHÖNIGER, M. & A. VON HAESELER 1994. A stochastic modeland the evolution of autocorrelated DNA sequences. –Molecular Phylogenetics and Evolution 3: 240–247.
WEAVER, J.S., III. 1984. The evolution and classification of
Trichoptera, Part 1: the groundplan of Trichoptera. Pp.
413–419 in: J.C. MORSE (ed.), Proceedings of the 4thInternational Symposium on Trichoptera. – Dr. W. Junk
Publishers, The Hague.
WIGGINS, G.B. & W. WICHARD 1989. Phylogeny of pupation in
Trichoptera, with proposals on the origin and higher classification of the order. – Journal of the North American
Benthological Society 8: 260–276.

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Entomologische Abhandlungen 61 (2)Resolving the Basal Phylogeny of Lepidoptera: Morphological Evidence
NIELS PEDER KRISTENSEN
Zoological Museum, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen Ø, Denmark [npkristensen@zmuc.ku.dk]

Introduction
As many as some 98–99 % of the currently described (ca.
150,000) species of Lepidoptera pertain to the subordinate
clade Ditrysia, whose members are structurally overall uniform. In contrast, the lineages constituting the remaining
tiny fraction of extant lepidopterans exhibit a remarkable
morphological diversity, spanning the gap between the‘ordinal’ ground plan with overall very generalized endopterygote characters and the ditrysian ground plan with its
considerable ‘derivation load’. The non-ditrysian grade iscurrently considered a model case of a ‘Hennigian comb’,
the constituent principal extant clades of which apparently
arose in several successive splitting events (Fig. 1). Lepidopteran monophyly is strongly supported by 20+ morphological synapomorphies of the order’s basal constituent
lineages; this includes the dense scale vestiture of thewings, a protibial epiphysis for antennal grooming, aposteromedian process on the tentorial bridge, an apodemal
process issued from the mesothoracic pleural ridge below
the wing process, and non-nucleated ‘parasperm’ present inaddition to normal ‘eusperm’.

The non-glossatans
Three currently recognized extant moth families, Micropterigidae (150+ species), Agathiphagidae (2 species) andHeterobathmiidae (9 species), have retained short, generalized maxillary galeae, hence they are outside the clade
Glossata, which comprises all other Lepidoptera and forwhich the coilable proboscis formed by linked galeae is themost conspicuous groundplan autapomorphy. In these threefamilies movable mandibles are retained throughout adultlife (but it remains uncertain whether adult agathiphagids
feed at all).
Morphological evidence bearing on the basal split within
the Lepidoptera is ambiguous: either the Micropterigidae orAgathiphagidae are the sister group of the rest of the‘order’. The former solution is here preferred, because thederived character states shared by non-micropterigids include some noteworthy neoformations (including a muscleaccommodating process of the metafurcal stem, perhaps adouble-compartment spermathecal duct, and a unique larval
head-flattening muscle), while those shared by non-agathiphagids are regressive traits (loss of vein M4, loss of protibial spur and one pair of mesotibial spurs, reduced ovariole
number, small size of testes follicles) exclusively.
Since paraspermatozoa are absent in the examined
Micropterix the question has been asked whether this genus
is actually the sister group of all remaining Lepidoptera,
and the family Micropterigidae as currently delimited hence
paraphyletic. However, Micropterix does possess a suite of
apomorphies (including, e.g., antennal ‘ascoid sensilla’,
close-set radial folds around male gonopore and, particularly, a unique larval trunk histology) which seemingly provides strong support for micropterigid monophyly. The absence of parasperm in the genus is therefore in all probability
an autapomorphy.
The next split is apparently between agathiphagids and therest; the sistergroup relationship between heterobathmiids

and Glossata is supported primarily by shared noteworthy
specializations of the larval head, such as the adfrontal andhypostomal sulci and the medially opened hypostomal/postgenal bridge. The Heterobathmiidae are the first lepidopteran clade whose members consistently depend on angiosperm plants throughout the life cycle.

Basal Glossata
The restructuring of the adult mouth apparatus in the glossatan ground plan involves, besides the formation of the
galeal proboscis, also a suite of regressive traits (loss ofmandibular articulations, of the strong epistomal
sulcus/ridge, etc). Some noteworthy glossatan groundplan
neoformations are unrelated to the adult feeding mode:
incorporation of adult’s tritocerebral commissure in suboesophageal mass, and development of larval spinneret.
The first three glossatan splitting events are apparently, insequence, (1) between Eriocraniidae and the rest (characterized, i.a., by a specialized wing scale vestiture and specialized 1st thoracic spiracles); (2) between Acanthopteroctetidae and the rest (characterized, i.a., by produced meso‘basisternum’); and (3) between Lophocoronidae and therest, known as Myoglossata and characterized primarily by
the development of an inner musculature in the proboscis.
The basal split within the Myoglossata was apparently between the Neopsustidae and the rest, collectively known as
the Neolepidoptera and characterized in the ground plan by
dissociation of the metathoracic aorta from the ‘wing
heart’, by adecticous and obtect pupae, and perhaps by‘typical’ caterpillars with the crochet-bearing prolegs on
III–VI & X (the unknown neopseustid larvae may, of course, prove to be of the same type, but since the adults haveretained strong mandibular muscles, the similarly unknown
pupae are surely decticous and likely exarate). And thebasal split within the Neolepidoptera was apparently between the Exoporia (including the first lepidopteran familygroup taxon with several hundreds of species, viz., theHepialoidea) and the rest, the time-honoured taxon
Heteroneura characterized, i.a., by a simplified hindwing
radial sector.
The outlined basal phylogeny of the Lepidoptera(KRISTENSEN & SKALSKI 1998; KRISTENSEN 1998a, b) is reasonably well supported, but not without problems (homoplasies). For instance, it requires that neolepidopteran pupal
adecticousness must have been parallelled in
Lophocoronidae, and some thoracic specializations (proprecoxal bridge, prospinasternum contiguous with mesobasisternum) have been independently evolved in
Neopseustidae and stem-lineage Heteroneura.
The basal phylogeny within the Heteroneura has proved difficult to unravel. Recently accumulating morphological evidence (KRENN & KRISTENSEN 2000; KRISTENSEN 2000;
LUKHTANOV 2001) supports the monophyly of a clade
Eulepidoptera, which excludes the Nepticuloidea (andperhaps the very recently established family Andesianidae).
Moreover, this new evidence indicates that at least the
Tischeriidae are more closly related to the Ditrysia than the
Incurvarioidea are. It must be noted, though, that apomorphies in embryonic development (absence of thickened

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Proceedings of 1st Dresden Meeting on Insect Phylogeny

Fig. 1. Phylogeny of extant lepidopteran superfamilies. Width of extant superfamily lines indicate approximate numbers of described
extant species (where these numbers exceed 1000). Eulepidopteran monophyly and all proposed groupings above superfamilylevel in the Ditrysia must be considered very tentative. Redrawn (updated/corrected) from KRISTENSEN & SKALSKI (1998).

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Entomologische Abhandlungen 61 (2)serosal cells, embryo remaining immersed in yolk until
immediately before hatching) indicate the monophyly of anepticuloid/ditrysian assemblage excluding incurvarioids(KOBAYASHI 1997).
MINET’s (1991) suggestion that three major, successively
less inclusive and internally partly/largely unresolved, subordinate clades (Apoditrysia, Obtectomera andMacrolepidoptera) can be recognized within the Ditrysia
remains state-of-the-art, its tentative nature notwithstanding. Forthcoming morphological work on selected character complexes will likely add resolution to the heteroneuranfamily tree in the next few years, and obviously integration
with molecular evidence should be the ultimate goal.

References
KOBAYASHI, Y. 1997. Embryogenesis of the fairy moth
Nemophora albiantenella issiki (Lepidoptera, Adelidae),with special emphasis on its phylogenetic implications. –International Journal of Insect Morphology andEmbryology 27: 157–166.
KRENN, H.W. & N.P. KRISTENSEN 2000. Early evolution of theproboscis of Lepidoptera (Insecta): external morphologyof the galea in basal glossatan moth lineages, withremarks on the origin of the pilifers. – Zoologischer
Anzeiger 239: 179–196.
KRISTENSEN, N.P. 1998a. The non-glossatan moths. Pp. 41–49
in: N.P. KRISTENSEN (ed.), Lepidoptera: moths and butterflies 1. Handbuch der Zoologie/Handbook of Zoology
IV/35. – Walter de Gruyter, Berlin, New York.
KRISTENSEN, N.P. 1998b. The homoneurous Glossata. Pp.
51–63 in: N.P. KRISTENSEN (ed.), Lepidoptera: moths andbutterflies 1. Handbuch der Zoologie/Handbook of
Zoology IV/35. – Walter de Gruyter, Berlin, New York.
KRISTENSEN, N.P. 2000. Inventory and cladistics of non-ditrysian moths: some recent advances. – Abstract Book 1, XXI
International Congress of Entomology, Brazil, August
20–26, 2000: 932.
KRISTENSEN, N.P. & A.W. SKALSKI 1998. Palaeontology andphylogeny. Pp. 7–25 in: N.P. KRISTENSEN (ed.),Lepidoptera: moths and butterflies 1. Handbuch der
Zoologie/Handbook of Zoology IV/35. – Walter deGruyter, Berlin, New York.
LUKHTANOV, V.A. 2000. Sex chromatin and sex chromosome
systems in nonditrysian Lepidoptera (Insecta). – Journalof Zoological Systematics and Evolutionary Research 38:
73–79.
MINET, J. 1991. Tentative reconstruction of the ditrysian phylogeny (Lepidoptera: Glossata). – Entomologica Scandinavica 22: 69–95.

A Comprehensive Phylogeny of Mecoptera and Siphonaptera
MICHAEL F. WHITING, ALISON S. WHITING & MICHAEL W. HASTRITER
Department of Integrative Biology, Brigham Young University, Provo, Utah 84602, USA [Michael_Whiting@byu.edu; mwhastriter@sprintmail.com]

Phylogenetic relationships among Mecoptera andSiphonaptera were inferred from molecular data for a wide
range of exemplar taxa and a coded morphological matrix.
Every mecopteran family and nearly every mecopterangenus was included in this analysis, for a total of ~125 species representing 1/5 of mecopteran species diversity.
Additional taxa representing 11 flea families and outgroups
were included. Genes sequenced include 18S rDNA, 28S
rDNA, cytochrome oxidase II, and cytochrome B. A morphological matrix consisting of 75 characters coded at thegeneric level was also generated. These data were analyzed
in a total evidence framework via direct optimization on an
IBM SP2 supercomputer.
These analyses support the monophyly of every mecopteranfamily. The clades Nannochoristidae, Boreidae + Siphonaptera, and the remaining Mecoptera are supported as mono-phyletic groups. However, resolution among these threeclades is still under investigation. Boreidae is placed assister group to Siphonaptera, and this result is congruent
with a number of characters associated with ovariole structures and other features. Panorpidae is sister to Panorpodidae, and relationships within Panorpidae are partiallycongruent with the designated species groups. The genera
Panorpa and Bittacus are each grossly paraphyletic. Thereis overall congruence between the morphological and molecular data partitions. These data support the paraphyly of
Mecoptera with regards to fleas, and that Boreidae should
be recognized as its own distinct order (‘Neomecoptera’). If
Nannochoristidae is supported as sister to the clade(Boreidae + Siphonaptera) + remaining Mecoptera, thenordinal designation for Nannochoristidae (‘Nannomecoptera’) is also warranted.

Preliminary Phylogeny of the Siphonaptera
KATHARINA DITTMAR DE LA CRUZ, MICHAEL F. WHITING, ALISON S. WHITING & MICHAEL W. HASTRITER
Department of Integrative Biology, Brigham Young University, Provo, Utah 84602, USA [katharinad@hotmail.com;
Michael_Whiting@byu.edu; mwhastriter@sprintmail.com]

Siphonaptera (fleas) is comprised of approximately 3000
described species and subspecies, of which 2525 are considered valid (LEWIS 1998). While it is clear that the order ismonophyletic, there has yet to be a formal quantitative analysis of flea phylogenetic relationships. LEWIS (1998) proposed a classification based on 15 families, avoiding a
superfamilial classification. MEDVEDEV (1994) performed amental analysis of morphology for 52 genera based on cha-racters associated with the head, thorax, and terminalia.
We present a robust phylogeny for the order Siphonaptera
based on a combination of molecular and morphological
data. Our taxon sampling for the molecular work is comprised of ~150 flea species which represent 11 of the 15 families, 23 of 28 subfamilies, and ~5 % of all described species. Genes sequenced include 18S rDNA, 28S rDNA, cytochrome oxidase II, and cytochrome B for a total of ~6425

170

nucleotides per taxon. The morphological matrix is comprised of 56 discrete morphological characters coded for
representatives of all families and subfamilies, and 11 ofthese were treated as multistate unordered characters. These
data were analyzed on an IBM SP2 supercomputer using
direct optimization as implemented in POY. The majority ofthe branches received high bootstrap and Bremer support
values.
These data are somewhat equivocal as to which taxon is thebasal most flea. Ceratophylloidea (= Ceratophyllidae +Ischnopsyllidae + Leptopsyllidae), Ceratophyllidae, andIschnopsyllidae are supported as monophyletic groups, but
Leptopsyllidae is paraphyletic in regards to Ischnopsyllidae
and Ceratophyllidae. Pulicidae is paraphyletic with theenigmatic Tunga placed elsewhere on the topology.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

Ctenophthalmidae is grossly non-monophyletic with majorgroups placed in at least four different positions on the
topology. In summary, these data suggest that roughly halfof the families, a third of the subfamilies, and a fifth of theflea genera which were studied in the analysis constitutenon-monophyletic groups.

References
LEWIS, R.E. 1998. Resume of the Siphonaptera (Insecta) of theworld. – Journal of Medical Entomology 35: 377–388.
MEDVEDEV, S.G. 1994. Morphological foundations of the classifications (Order Siphonaptera). – Entomological Review
73: 22–43.

Phylogeny of True Flies (Diptera): a 250 Million Year Old Success Story
in Terrestrial Diversification
DAVID K. YEATES1, RUDOLF MEIER2 & BRIAN M. WIEGMANN3
1

CSIRO Entomology, Canberra ACT 2601, PO Box 1700, Australia [David.Yeates@csiro.au] – 2 Department of Biological
Sciences and University Scholars Programme, National University of Singapore, 14 Science Dr 4, Singapore 117543, Singapore[dbsmr@nus.edu.sg] – 3 Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613, USA[bwiegman@unity.ncsu.edu]

The insect order Diptera (true flies) is one of the most species rich, anatomically varied and ecologically innovative
groups of organisms, making up around 12 % of the known
animal species. An estimated 125,000 species of Diptera
have been described, however, the total number of extant fly
species is many times greater. The living dipteran species
have been classified into about 10,000 genera, 150 families,
22–32 superfamilies, 8–10 infraorders and 2 suborders(YEATES & WIEGMANN 1999).
The monophyly of Diptera is well established. HENNIG(1973) lists 37 autapomorphies some of which form morphologically complex structures such as the specialized
mouthparts adapted for sponging liquids. Traditionally, thebest-known autapomorphy is the transformation of the hindwings into halteres, but this character may now be in needof reinterpretation due to recent phylogenetic research suggesting a sistergroup relationship between Strepsiptera and
Diptera (WHITING et al. 1997). This work implies homology
between the Diptera halteres and Strepsiptera pseudohalteres.
The sister group of Diptera remains unknown. Morphology
suggests either Mecoptera, Siphonaptera, or a monophylum
consisting of both, but based on the aforementioned molecular evidence the Strepsiptera has to be added to the list of
candidate taxa. The first fossils attributable to Diptera are
known from the Permian, and a large number of fossil
Diptera are known from the Mesozoic (YEATES &WIEGMANN 1999).
Phylogenetic work in the strict sense on Diptera began withHENNIG (1973) and GRIFFITHS (1972). Only recently, numerical analyses have started to address the relationships
within higher-level taxa. Although molecular data has beenused increasingly to reconstruct dipteran phylogeny, mostpublished analyses to date have focussed on questions at alower level, generally within particular infraorders. An
exception is a recent detailed analysis of Brachycera relationships using over 2 kb of 28S rDNA (B. Wiegmann in
prep.). The results of the last 30 years of phylogenetic researchon the higher-level relationships of the Diptera using morphological data have been synthesized by us using supertree
techniques. This Diptera supertree (Fig. 1) forms the framework for the following discussion.

The supertree generally supports recent research and showsthat major dipteran higher categories such as Brachycera,
Eremoneura, Muscomorpha, Cyclorrhapha, Schizophora,
and Calyptrata are monophyletic. Conversely, a number of
traditional higher taxa are paraphyletic based on morphological and molecular data. These include the Nematocera,
Orthorrhapha, and Aschiza. We therefore prefer to use theinformal terms Lower Diptera, Lower Brachycera andLower Cyclorrhapha for these groups. They represent evolutionary grades at the base of major radiations of Diptera,
Brachycera, and Cyclorrhapha, respectively.
The paraphyly of the Lower Diptera has been suspected for
decades, beginning with Hennig, and demonstrated inrecent quantitative cladistic analyses using morphological
data (OOSTERBROEK & COURTNEY 1995). There have been
only a few comprehensive phylogenetic analyses of the relationships between lower dipteran families using morphological and especially molecular data. The position of thetipulids and their relatives has been very unstable; somemorphological treatments consider them the basal lineageof Diptera (HENNIG 1973), while others consider them to beclosely related to Brachycera (OOSTERBROEK & COURTNEY1995). The supertree analysis currently favors
Ptychopteromorpha + Culicomorpha as the sister group to
the remaining Diptera with Blephariceromorpha andBibionomorpha being the next lineages to emerge from thelower dipteran stem. Close to the grade transition to
Brachycera, the lower dipteran infraorders are not monophyletic, with Psychodomorpha and Tipulomorpha forminga paraphyletic grouping, the superfamily Tipuloidea being
sister to the Brachycera. The arrangement of Tipulomorpha
and Psychodomorpha represents a resolution of the incongruence between input trees.
The Brachycera is certainly a monophyletic group, with alarge number of undisputed autapomorphies. The phylogeny of the Lower Brachycera has been scrutinized intensively over the past 15 years. A recent quantitative reanalysis
of morphological characters used to define relationships
between the lower brachyceran families attempted to summarize and synthesize this research (YEATES 2002). This
study revealed weak evidence for the monophyly of a clade

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Entomologische Abhandlungen 61 (2)Ptychopteromorpha
Culicomorpha
Blephariceromorpha
Bibionomorpha
Psychodomorpha
Tipuloidea
Stratiomyomorpha
Xylophaghomorpha

Brachycera
Tabanomorpha
Nemestrinoidea

Muscomorpha

Asiloidea
Empidoidea

Eremoneura

Lower Cyclorrhapha (Part)

CyclorrhaphaPhoroidea
Syrphoidea

Calyptrata

Hippoboscoidea
Muscoidea
Oestroidea

Schizophora

Acalyptrata

Fig. 1. A phylogenetic tree for Diptera resulting from a supertree analysis of fly families summarized at the infraorder/superfamily level. The analysis was based on an MRP coded matrix of 12 primary source trees including GRIFFITHS (1972), HENNIG(1973), MCALPINE (1989), OOSTERBROEK & COURTNEY (1995) and YEATES (2002). For more details contact the senior author.

containing Xylophagomorpha, Stratiomyomorpha andTabanomorpha, and these findings are reflected in the
supertree. The most basal lineage of Brachycera in the
supertree analysis contains Stratiomyomorpha +(Xylophagomorpha + Tabanomorpha), reflecting the resultsof recent quantitative cladistic analyses.
The infraorder Muscomorpha contains all brachyceranfamilies except those belonging to Stratiomyomorpha,
Xylophagomorpha and Tabanomorpha, and is a well-supported clade found on the supertree. The Nemestrinoidea,
Asiloidea and Empidoidea are monophyletic, arising fromthe main stem of the Brachycera in that sequence. Evidence
for the monophyly of nemestrinoids and asiloids is notstrong, and they appear paraphyletic in some analyses. A
number of asiloid families have received critical phylogenetic scrutiny in recent years, partly because of their proximity to Eremoneura.
Eremoneura is the muscomorphan lineage containing
Empidoidea + Cyclorrhapha, and is one of the best-supported higher-level brachyceran clades with many autapomorphies. Recent morphological work has emphasized male
genitalic characters for phylogenetic reconstruction in
Eremoneura, however some analyses of molecular data are
beginning to appear. There is strong evidence for the monophyly of the Empidoidea and its subgroups Atelestidae,
Hybotidae, Empididae and Microphoridae + Dolichopodidae
from both morphological and molecular data.
Cyclorrhaphan monophyly is well supported by characterssuch as the invagination of the larval head capsule andmodifications of the larval mouthparts, as well as pupation
within the skin of the last larval instar. These are the mostrecognizable features of this landmark in dipteran evolution. Over the last 40 years only three workers have attempted to synthesize phylogenetic evidence on cyclorrhaphan
relationships in a comprehensive fashion. All studies were
not based on explicit data sets and results differed in many
regards (HENNIG 1973; GRIFFITHS 1972; MCALPINE 1989).
Exploration of new character systems applied broadly
across cyclorrhaphan groups, for example of egg and larvalmorphology, female genitalia and internal morphology, andnucleotide sequences are urgently needed. There are a number of competing hypotheses regarding the relationships ofthe families belonging to the lower cyclorrhaphan grade,but the Syrphoidea are generally regarded the sister groupto the Schizophora.
The monophyletic Schizophora are classified into at least
80 families and comprise just over half the family-level
diversity in Diptera. Major reviews of Schizophora phylogeny are the synthetic revisions by GRIFFITHS (1972) and
MCALPINE (1989), which provided new information whilebuilding on Hennig’s earlier research. GRIFFITHS (1972)
provided detailed interpretation and scorings of male genitalic characters along with other morphological features,and MCALPINE’s (1989) fully resolved phylogenetic arrangements draw on most morphological character systems aswell as fly biology. Based on calypter morphology the
Schizophora have been traditionally subdivided into the
Acalyptratae and Calyptratae, but it has long been recognized that the calypter is too variable in both groups to be areliable phylogenetic marker. Although Acalyptratae is supported on our supertree it is generally regarded as being
paraphyletic.

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MCALPINE (1989) divided the Acalyptratae into 10 superfamilies and these are found in the supertree: Nerioidea,
Diopsoidea, Conopoidea, Tephritoidea, Lauxanioidea,
Sciomyzoidea, Opomyzoidea, Sphaeroceroidea, Carnoidea,
and Ephydroidea. Only a few superfamilies are uncontentious (e.g., Ephydroidea, Tephritoidea) while the remaining
will probably see some major rearrangements after more
intensive phylogenetic scrutiny. Generally, MCALPINE’s(1989) classification maintains HENNIG’s (1973) groupings
while GRIFFITHS (1972) proposes a more radical restructuring. In contrast to the Acalyptratae and despite the lack ofcomplex morphological autapomorphies, the Calyptratae
appear monophyletic based on molecular and morphological evidence. Of the three superfamily-level taxa, theHippoboscoidea are monophyletic, the Oestroidea may bemonophyletic, while the Muscoidea is likely paraphyletic.
Advances in understanding the relationships of flies will
accelerate with the increasing use of molecular data andquantitatively analysed data matrices. The most important
areas for future phylogenetic research in the Diptera are in
the Lower Diptera and Schizophora. In the Lower Diptera,
resolving the relationships between the infraorders and theposition of the craneflies and their relatives (Tipulidae) are
critical tasks. The Schizophora also await more focused
phylogenetic scrutiny applied to relationships between, andwithin, the superfamilies as they are currently defined.

Proceedings of 1st Dresden Meeting on Insect Phylogeny

References
GRIFFITHS, G.C.D. 1972. The phylogenetic classification ofDiptera Cyclorrhapha, with special reference to the structure of the male postabdomen. – Series Entomologica 8:
1–340.
HENNIG, W. 1973. Diptera. Pp. 1–337 in: J.G. HELMCKE, D.
STARCK & H. WERMUTH (eds.), Handbuch der Zoologie
IV/20: Arthropoda. – De Gruyter, Berlin, New York.
MCALPINE, J.F. 1989. Phylogeny and classification of theMuscomorpha. Pp. 1397–1518 in: J.F. MCALPINE (ed.),Manual of Nearctic Diptera, vol. 3. – Agriculture Canada
Monograph 32, Ottawa.
OOSTERBROEK, F.L.S. & G. COURTNEY 1995. Phylogeny of thenematocerous families of Diptera (Insecta). – Zoological
Journal of the Linnean Society 115: 267–311.
WHITING, M.F., J.C. CARPENTER, Q.D. WHEELER & W.C.
WHEELER 1997. The Strepsiptera problem: phylogeny of
the holometabolous insect orders inferred from 18S and
28S ribosomal DNA sequences and morphology. –Systematic Biology 46: 1–68.
YEATES, D.K. 2002. Relationships of the extant lower
Brachycera (Diptera): a quantitative synthesis of morphological characters. – Zoologica Scripta 31: 105–121.
YEATES, D.K. & B.M. WIEGMANN 1999. Congruence and controversy: toward a higher-level phylogeny of Diptera. –Annual Review of Entomology 44: 397–428.