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Ao. Univ.- Prof. Dr.
Günther Pass

Department of Integrative Zoology
UZA1 Althanstraße 14, 1090 Vienna, Austria

phone: +43 1 4277 76371
fax: +43 1 4277 876371
mail: guenther.pass@univie.ac.at 


  • Mag. Alexander Böhm
  • Patrick Fahser
  • Sven Goldermann BSc
  • Judith Kregl BSc
  • Oliver Macek BSc
  • Dominik Reischl BSc


Phylogenetic relationships in arthropods

Due to the advance of molecular phylogenetic studies, classical hypotheses on the relationships of the major arthropod groups have come into question. In particular, the relationship of insects to other arthropods has attracted much attention and still remains unresolved.

Our current research focuses on the evolutionary basis of the hexapod tree. We aim to elucidate the phylogenetic relationships among the so-called apterygote insects using molecular and morphological approaches.

Current projects:

Are the Hexapoda monophyletic?
Conflicting hypotheses regarding their relationships to myriapods and crustaceans
Postdocs: Emiliano Dell´Ampio, Nikolaus Szucsich
(FWF project 20497-B17)

The most ancient splits in insect phylogeny
Relationships among primarily wingless hexapods

PhD Students: Alexander Böhm, Daniela Bartel
(FWF project 23251-B17) 

1KITE - 1000 Insect Transcriptome Evolution
This international and interdisciplinary approach aims to study the transcriptomes of 1,000 insect species encompassing representatives from all recognized insect orders. From the data we expect to infer a robust phylogenetic backbone tree of the insect clade. For more information see www.1kite.org


  • Bernhard Misof and Karen Meusemann, Alexander Koenig Museum Bonn, Germany), see DFG Priority Program „Deep Metazoan Phylogeny
  • Ryuichiro Machida, University of Tsukuba, Japan
  • Nikola Szucsich, Museum of Natural History, Vienna

Newsletter Uni Wien (PDF)

Radio Ö1 "Dimensionen"

Medienportal der Universität Wien


Radio Ö1 "Vom Lebern der Natur" 

Structure, function and evolution
of circulatory organs in insects

Many ancestral arthropods have a complex vascular system which has been reduced to various degrees during arthropod evolution. In insects, only the dorsal vessel (“heart“) has remained. To supply longer body appendages such as antennae, legs and wings, new pumps have evolved in insects, the so-called accessory pulsatile organs (“auxiliary hearts”).

These accessory pulsatile organs deserve special interest from both functional and evolutionary points of view. On the one hand, they are ideal objects for investigating the structure and function of simply organized nerve-muscle systems. On the other hand, they are good examples to study how novelties occur in evolution.

Review on the accessory pulsatile organs in insects
(pdf file);
supplementary material with an animated film for teaching purposes

Current project:

Wing circulatory organs in Drosophila
PhD Student Markus Tögel
(FWF project  19380-B17)

Newsletter Uni Wien (PDF)


  • Achim Paululat, University of Osnabrück, Germany
  • Rolf Beutel and Beny Wipfler, University of Jena, Germany
  • Christian Wirkner, University of Rostock, Germany

Selected Publications


Pass G (2018) Beyond aerodynamics: The critical roles of the circulatory and tracheal systems in maintaining insect wing functionality. Arthropod Structure & Development 47:391-407 https://doi.org/10.1016/j.asd.2018.05.004, available online

Dudley R, Pass G (2018) Wings and powered flight: Core novelties in insect evolution. Arthropod Structure & Development 47: 319-321 https://doi.org/10.1016/j.asd.2018.06.006, available online

Pass G, Tögel M, Krenn H, Paululat A (2015) The circulatory organs of insect wings: prime examples for the origin of evolutionary novelties. Zool Anz 256:82-95

Wirkner CS, Tögel M, Pass G (2013) The arthropod circulatory system. In: Minelli A, Boxshall G, Fusco G (eds) Arthropod biology and evolution – molecules, development, morphology. Springer, Heidelberg. pp. 343-391

Miller T, Pass G (2009) Chapter "Circulatory System" In: Encyclopedia of Insects, Resh V.H., Cardé RT, 2nd ed. Elsevier Science & Technology pp.169-173

Pass G (2000) Accessory pulsatile organs: evolutionary innovations in insects. Ann Rev Entomol 45: 493-516 

Pass G (1998) Accessory pulsatile organs. In: Microscopic Anatomy of Invertebrates (Harrison F, Locke M eds). Vol 11B: Insecta. Wiley: New York, pp. 621-640


Böhm A, Meusemann K, Misof B & Pass G (2018). Hypothesis on monochromatic vision in scorpionflies questioned by new transcriptomic data. Scientific Reports 8:9872, DOI:10.1038/s41598-018-28098-2

Antemann V, Pass G, Pflüger HJ (2018) Octopaminergic innervation and a neurohaemal release site in the antennal heart of the locust Schistocerca gregaria. J Comp Physiol A 204: 131-143 doi:10.1007/s00359-017-1213-5

Böhm A, Pass G (2016) The ocelli of Archaeognatha (Hexapoda): Functional morphology, pigment migration and chemical nature of the reflective tapetum. J Exp Biol doi:10.1242/jeb.141275, available online

Galli L, Bartel D, Capurro M, Pass G, Sará A, Shrubovych J, Szucsich N (2016) Redescription and review of the most abundant conehead in Italy: Acerentomon italicum Nosek, 1969 (Protura: Acerentomidae). Ital J Zool 83: 43-58, DOI: 10.1080/11250003.2015.1114686, available online

Shrubovych J, Bartel D, Szucsich NU, Resch MC, Pass G (2016) Morphological and genetic analysis of the Acerentomon doderoi group (Protura: Acerentomidae) with description of A. christiani sp. nov. PLoS ONE 11: e0148033, DOI:10.1371/journal.pone.0148033, April 13, 2016. available online

Misof B, Liu S, Meusemann K, Peters RS, Donath A, Mayer C, Frandsen PB, Ware J, Flouri T, Beutel RG, Niehuis O, Petersen M, Izquierdo-Carrasco F, Wappler T, Rust J, Aberer AJ, Aspöck U, Aspöck H, Bartel D, Blanke A, Berger S, Böhm A, Buckley T, Calcott B, Chen J, Friedrich F, Fukui M, Fujita M, Greve C, Grobe P, Gu S, Huang Y, Jermiin LS, Kawahara AY, Krogmann L, Kubiak M, Lanfear R, Letsch H, Li Y, Li Z, Li J, Lu H, Machida R, Mashimo Y, Kapli P, McKenna DD, Meng G, Nakagaki Y, Navarrete-Heredia JL, Ott M, Ou Y, Pass G, Podsiadlowski L, Pohl H, von Reumont BM, Schütte K, Sekiya K, Shimizu S, Slipinski A, Stamatakis A, Song W, Su X, Szucsich NU, Tan M, Tan X, Tang M, Tang J, Timelthaler G, Tomizuka S, Trautwein M, Tong X, Uchifune T, Walzl MG, Wiegmann BM, Wilbrandt J, Wipfler B, Wong TK, Wu Q, Wu G, Xie Y, Yang S, Yang Q, Yeates DK, Yoshizawa K, Zhang Q, Zhang R, Zhang W, Zhang Y, Zhao J, Zhou C, Zhou L, Ziesmann T, Zou S, Li Y, Xu X, Zhang Y, Yang H, Wang J, Wang J, Kjer KM, Zhou X (2014) Phylogenomics resolves the timing and pattern of insect evolution. Science 346: 763-767

Wipfler B, Pass G (2014) Antennal heart morphology supports relationship of Zoraptera with polyneopteran insects. Syst Ent 39: 800-805

Hustert R, Frisch M, Böhm A, Pass G (2014) A new kind of auxiliary heart in insects: functional morphology and neuronal control of the accessory pulsatile organs of the cricket ovipositor. Frontiers in Zoology 11:43

Resch C, Shrubovych J, Bartel D, Szucsich NU, Timelthaler G, Bu Y, Pass G (2014) DNA barcoding in Protura: Where subtle morphological differences are perfectly mirrored by huge genetic distances. Plos ONE 9(3): e90653.doi:10.1371/journal.pone.0090653

Dell'Ampio E, Meusemann K, Szucsich NU, Meyer B, Borner J, Petersen M, Aberer AA, Stamatakis A, Walzl MG, von Haeseler A, Ebersberger I, Pass G, Misof B (2014) Decisiveness and conflicts in phylogenomics: lessons from basal hexapods. Mol Biol Evol 31:239-249

Szucsich NU, Wirkner CS, Pass G (2013). Deconstructing morphology: reply to Scholtz (2010). Acta Zoologica 94: 215-217

Böhm A, Szucsich NU, Pass G (2012). Brain anatomy in Diplura (Hexapoda). Frontiers in Zoology 9: 26

Reumont BMv, Jenner RA, Wills MA, Dell’Ampio E, Pass G, Ebersberger I, Meyer B, Koenemann S, Iliffe TM, Stamatakis A, Niehuis O, Meusemann K, Misof B (2012) Pancrustacean phylogeny in the light of new phylogenomic data: support for Remipedia as the possible sister group of Hexapoda. Mol Biol Evol 29:1031-1045

Böhm A, Bartel D, Szucsich NU & G Pass (2011) Confocal imaging of the exo- and endoskeleton of Protura after nondestructive
DNA extraction Soil Organisms 83: 335–345 

Dell’Ampio E, Szucsich NU &  G Pass (2011) Protura and molecular phylogenetics: status quo of a young love. Soil Organisms 83: 347–358 

Pass G, Szucsich N (2011) 100 years of research on the Protura: many secrets still retained. Soil Organisms 83: 309-334 

Meusemann K, von Reumont BM, Simon S, Roeding F, Strauss S, Kück P, Ebersberger I, Walzl M, Pass G, Breuers S, Achter V, von Haeseler A, Burmester T, Hadrys H, Wägele JW, Misof B (2010) A phylogenomic approach to resolve the arthropod tree of life. Molecular Biology and Evolution Advance Access published online on June 9 http://mbe.oxfordjournals.org/content/27/11/2451.abstract

Eberhard MJB, Lang D, Metscher B, Pass G, Picker D, Wolf H (2010) Structure and sensory physiology of the leg scolopidial organs in Mantophasmatodea and their role in vibrational communication. Arthropod Structure & Development 39: 230-241 http://www.sciencedirect.com/science/article/pii/S1467803910000101

Dell’Ampio E, Szucsich NU., Carapelli A, Frati F, Steiner G, Steinacher A, Pass G (2009) Testing for misleading effects in the phylogenetic reconstruction of ancient lineages of hexapods: Influence of character dependence and character choice in analyses of 28S rRNA sequences. Zool Scripta 38: 155-170

Eberhard MJB, Pass G, Picker MD, Beutel R, Predel R, Gorb SN (2009) Structure and Function of the Arolium of Mantophasmatodea (Insecta). Journal of Morphology 270:1247-1261 http://onlinelibrary.wiley.com/doi/10.1002/jmor.10754/abstract

Lehmacher C, Tögel M, Pass G, Paululat A (2009) The Drosophila wing hearts consist of syncytial muscle cells that resemble adult somatic muscles. Arthropod Structure and Development 38: 101-110 http://www.sciencedirect.com/science/article/pii/S1467803908000789

von Reumont BM, Meusemann K, Szucsich NU, Dell’Ampio E, Gowri-Shankar V, Bartel D, Simon S, Letsch HO, Stocsits RR, Luan Y, Wägele JW, Pass G, Hadrys H, Misof B (2009) Can comprehensive background knowledge be incorporated into substitution models to improve phylogenetic analyses? A case study on major arthropod relationships. BMC Evol. Biol. 9:119 

Tögel M, Pass G, Paululat A (2008) The Drosophila wing hearts originate from pericardial cells and are essential for wing maturation. Developmental Biology 318:29-37 http://www.sciencedirect.com/science/article/pii/S0012160608001462

Dell’Ampio E, Szucsich NU, Carapelli A, Frati F, Steiner G, Steinacher A, Pass G (2008) Testing for misleading effects in the phylogenetic reconstructionof ancient lineages of hexapods: influence of character dependence and character choice in analyses of 28S rRNA sequences. Zoologica Scripta, 38: 155-170. 

Szucsich NU, Pass G (2008) Incongruent phylogenetic hypotheses and character conflicts in morphology: The root and early branches of the hexapodan tree. Mitteilungen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie, 16: 415-429 

Pass G, Gereben-Krenn BA, Merl M, Plant J, Szuscsich NU, Tögel M (2006) Phylogenetic relationships of the orders of Hexapoda:contributions from the circulatory organs for a morphological matrix. Arthropod Syst & Phylogeny 64: 165-203

Scheuch M., Pass G., Grünweis FM., Wrbka T. (2006) Interdisciplinary Project – “Expedition to Cultural Landscapes“ A Teacher Education Module for Nature Experience and Environmental Education. In Posch P., Kyburz-Graber, R. (Hrsg.). Reflective Practice in Teacher Education - Learning from Case Studies of Environmental Education. Peter Lang Verlag: Bern, Berlin, Bruxelles. pp. 183-193

Bruvo B, Huber B, Steinacher A, Pass G (2005) Molecular phylogeny of pholcid spiders inferred from nuclear and mitochondrial DNA sequences. Mol Phyl Evol 37:661-673

Pernstich A, Krenn HW, Pass G (2003) Preparation of serial sections of arthropods using 2,2-dimethoxypropane dehydration and epoxy resin embedding under vacuum. Biotechnic & Histochemistry 78: 5-9 

Hertel W, Pass G (2002) An evolutionary treatment of the morphology and physiology of circulatory organs in insects. Comp Biochem Physiol A 133: 555-575

Hertel W, Wirkner CS, Pass G (2002) Studies on the cardiac physiology of Onychophora and Chilopoda. Comp Biochem Physiol A 133: 605-609

Wirkner CS, Pass G (2002) The circulatory system in Chilopoda: functional morphology and phylogenetic aspects. Acta Zoologica (Stockholm) 83:193-202 

Pass G (2001) Die vielen Herzen der Insekten: evolutionärer Ursprung und strukturelle Diversität. Mitt Dtsch Ges Allg Angew Ent 13: 283-297

Gereben-Krenn BA, Pass G (2000) Circulatory organs of abdominal appendages in primitive insects (Hexapoda: Archaeognatha, Zygentoma and Ephemeroptera). Acta Zoologica 81: 285-292 

Micholitsch T, Krügel P, Pass G (2000) Insemination and fertilization in the seed bug Lygaeus simulans (Heteroptera: Lygaeidae). Eur J Entomol 97: 13-18

Tadler A, Nemeschkal Hl, Pass G (1999) Phenotypical selection of male characters by cryptic female choice in the seedbug Lygaeus simulans (Heteroptera, Lygaeidae). Biol J Linn Soc 68:471-483

Matus S, Pass G (1999) Antennal circulatory organ of Apis mellifera L. (Hymenoptera:Apidae) and other Hymenoptera: functional morphology and phylogenetic aspects. Int J Insect Morph & Embryol 28: 97-109

Gereben-Krenn BA, Pass G (1999) Circulatory organs of Diplura (Hexapoda): the basic design in Hexapoda? Int J Insect Morph & Embryol 28:71-79

Krenn HW, Pass G (1995) Morphological diversity and phylogenetic analysis of wing circulatory organs in insects, part II: Holometabola. Zoology 98:147-164

Krenn HW, Pass G (1994) Morphological diversity and phylogenetic analysis of wing circulatory organs in insects, part I: non-Holometabola. Zoology 98:7-22

Krenn HW, Pass G (1993) Wing-hearts in Mecoptera (Insecta). Int J Insect Morphol Embryol 22:63-76

Pass,G. (1991) Antennal circulatory organs in Onychophora, Myriapoda and Hexapoda: functional morphology and evolution. Zoomorphology 110:145-164

Hertel W, Pass G, Penzlin G (1988) The effect of the neuropeptide proctolin and of octopamine on the antennal heart of Periplaneta americana. Symp Biol Hungar 36:351-362

Pass G (1988) Functional morphology and evolutionary apsects of unusual antennal circulatory organs in Labidura riparia Pallas (Labiduridae), Forficula auricularia L. and Chelidurella acanthopygia Géné (Forficulidae) (Dermaptera: Insecta).Int J Insect Morph Embryol 17: 103-112 

Pass G, Agricola H, Birkenbeil H, Penzlin H (1988) Morphology of neurones associated with the antennal heart of Periplaneta americana (Blattodea, Insecta). Cell Tissue Res 253:319-326 

Pass G, Sperk G, Agricola H, Baumann E, Penzlin H (1988) Octopamine in a neurohaemal area within the antennal heart of the American cockroach. J exp Biol 135:495-498  

Pass G (1987) "Cercus heart" in stoneflies - a new type of accessory circulatory organ in insects. Naturwissenschaften 74:440-441 

Hertel W, Pass G, Penzlin H (1985) Electrophysiological investigation of the antennal heart of Periplaneta americana and its reactions to proctolin. J Insect Physiol 31:563-572 

Pass G (1985) Gross and fine structure of the antennal circulatory organ in cockroaches (Blattodea, Insecta). J Morph 185:255-268 

Pass G (1980) The Anatomy and Ultrastructure of the Antennal Circulatory Organs in the Cockchafer Beetle Melolontha melolontha L. (Coleoptera, Scarabaeidae). Zoomorphology 96, 77-89 

Department of
Integrative Zoology
University of Vienna
Althanstraße 14
A-1090 Vienna, Austria
University of Vienna | Universitätsring 1 | 1010 Vienna | T +43-1-4277-0