Hans Leo Nemeschkal                                                                                  

Department of Theoretical Biology

University of Vienna

Althanstrasse 14, A-1090 Vienna (Austria)




Teaching: http://online.univie.ac.at/vlvz?titel=&zuname=Nemeschkal&vorname=&lvnr=&semester=all&match=substring

Zoological Collection/ Head of the Zoological Collection:






Topics of the current scientific interest:




Phenotypic correlation patterns in avian skeletons:




Phenotypic integration, phenotypic pattern analysis:

Quantitative morphology:

- Handschuh, S. 2017. 'Reinforcing and expanding the role of x-ray microCT as a key technique for a modern and quantitative microscopic animal morphology' (dissertation thesis, completed)


Cartesian transformations (grid transfomations):










Natural and sexual selection, assortative mating, sexual dimorphism:


 diploma theses, completed:

- Bellersen, G. 2005. Sexual selection and male genitalia: Is there a difference between mated and non-mated individuals. Investigation of Rhagonycha fulva's male genital. (in co-operation with H. Paulus)

- Parzer, H. 2006. Correlative mating: a new aspect of nonrandom mating observed in Rhagonycha fulva (Coleoptera, Cantharidae).

- Handschuh, S. 2007. Assortative mating and sexual selection in the freshwater gammarid Dikerogammarus villosus (Sowinski, 1894) (Amphipoda, Crustacea)

running master thesis:

- Nowak, H. 2013: running master thesis on 'a computer simulation of sexual selection and assortativity'

master thesis, completed:

- Laciny, A. 2014. Assortativity, sexual selection, mating success and ectoparasitic load in red soldier beetle Rhagonycha fulva (Coleoptera, Cantharidae).

dissertation thesis, completed:

- Wilhelm, G. 2015. Natural and sexual selection become visible: animal – plant interactions between the parasitic weevil Rhopalapion longirostre (Olivier, 1807) (Insecta: Coleoptera: Curculionoidea: Brentidae) and its host plant Alcea rosea (Linnaeus 1758) (Malvaceae) provide the environmental frame for the emergence of an extreme sexual dimorphism reflected in the weevil's rostrum'.

Shell morphometrics, image analysis:

diploma thesis, completed:

- Pokorny, P. 2006. Quantitative characterization of snail shells. Representation of shell aperture in Arianta arbustorum using digital image processing.

running dissertation thesis on shell morphometrics:

- Pokorny, P. 2007


Collection history:

master thesis, completed:

- Harlfinger, J. 2016. Embodiments of theory: Wax models of the Zoological Collection (ZCUV), with special regard to Berthold Hatschek's 3D publications of Trochophora and Amphioxus.

running diploma thesis:

- Rafi, F. 2017. Debating the Origin of Life: Joseph Hyrtl’s Embryological Collection.

Biometry, statistics, computer programming:

Computerintensive statistics (Mantel's test, randomization tests, bootstrap, MCMC= Markov Chain Monte Carlo)

Categorical data analyses (generalized linear models), Bayes estimation, data imputation – missing data

Periodic regression analyses

Programming in BASIC (QB64, FreeBASIC), C and JAVA (Linux, Windows, MS-DOS)



Users of FreeBASIC note please !

(A) The dislin-header for FreeBASIC has been already translated to FB bi-format (test versions only):

(a)   header for Win: prg\dislinw.zip

(b)   header for Linux/Unix: prg\dislinl.tar.gz

(for dislin and Freebasic see http://www.dislin.de and http://www.freebasic.net )

 (B) The RngStreams-header for FreeBASIC has been translated recently to FB bi-format (test versions only):

(a)   header for Win: /prg/rngw.zip

(b)   header for Linux/Unix: /prg/rngl.tar.gz

(for more information on RngStreams see http://www.iro.umontreal.ca/~simardr/indexe.html and http://statmath.wu-wien.ac.at/software/RngStreams/)

(C) simulation of natural selection – LANDE ARNOLD model (MSDOS-program code) /home/nemesch4/html/prg/selgraph.zip 

 (D) lecturing-interface (various demonstration files for running lectures):














about selection theory:

http://people.oregonstate.edu/~arnoldst/pdf_files/Lande & Arnold 1983.pdf

http://people.oregonstate.edu/~arnoldst/pdf_files/Phillips & Arnold 1989.pdf

multivariate statistics course:










Scientific Networks and Contacts:


-U. Aspöck, International Research Institute for Entomology, Museum of Natural History Vienna, Austria


-R. van den Elzen, Ornithology Department, Zoologisches Forschungsinstitut und Museum Alexander Koenig (ZFMK), Bonn, Germany




-H. Keckeis, Department of Limnology, Institute of Ecology and Conservation Biology, University of Vienna, Austria




-M. Pavlicev,


-evolVienna http://www.univie.ac.at/evolvienna/

Invited lectures:

- 'Die Zoologische Sammlung der Universität Wien – Verbindungen zu Triest' im Rahmen der Eröffnungsveranstaltung zur Ausstellung 'Die k.k. Zoologische Station in Triest' 14.10.2010 – 31.1.2011, Foyer der Universitätsbibliothek, Universität Wien. http://www.univie.ac.at/zoobot/doc/Ausstellung_Triest.pdf

- 'Die Zoologische Sammlung – Gegenwärtiger Bestand, Eckpunkte der Ausrichtung und historischer Kontext.' im Rahmen des 8. Sammlungstreffens, 30.11.2010, UZA1, Althanstr.14 (Vorsitz: Mag. C. Feigl, Sammlungsprojekt der Universität Wien).

- 'Museum, Kuriositätenkabinett oder wissenschaftliche Vergleichssammlung? Aufgaben und Ziele einer universitären Sammlung.' im Rahmen der Veranstaltungsreihe 'Die Bedeutung von Sammlungen in der modernen Wissenschaft – Die Sammlungen der Zoologischen Institute der Universität Wien', ZOOBOT (Zoologisch Botanische Gesellschaft in Österreich), Wien, 16.11.2011, Hs 2, Althanstr. 14, UZA 1

- 'Measurement error in proportion data', University of Vienna, CIUS Seminar, 2017, 30th May



Computer Programming - Contributions:

Translation of the dislin-header for FreeBasic, December 2005 (http://www.freebasic.net/wiki/wikka.php?wakka=CompilerCredits).

Translation of the RngStreams-header for FreeBasic, October 2006.



 Wilhelm, G., Nemeschkal, H.L., and J. Plant. 2011. How to maximize reproduction without killing my host. Infestation balance between the parasitic weevil Rhopalapion longirostre and the Common Hollyhock Alcea rosea. 13th Congress of the European Society for Evolutionary Biology (ESEB), Tuebingen 20-25 August 2011.

Handschuh, S., Nemeschkal, H.L., Metscher, B.D., Mitteröcker, P., and G.B. Müller. 2011. Sexual selection and the evolution of sexual dimorphic appendages in amphipods. 13th Congress of the European Society for Evolutionary Biology (ESEB), Tuebingen 20-25 August 2011.

Handschuh, S., Nemeschkal, H.L., Metscher, B.D., Mitteroecker, P., and G.B. Müller. 2012. The role of post-embryonic growth and sexual selection in the evolution of sexually dimorphic appendages in Amphipods. EURO EVO DEVO 2012, Lisboa, Portugal 10-13 July 2012.

References (recently published, in press, submitted):


Laciny, A., and H.L. Nemeschkal (2015). Disassortative parasitic infestation by larval mites of the genus Trombidium (Acari: Trombidiidae) influences mate choice in a population of the cantharid beetle Rhagonycha fulva Scopoli, 1763 (Coleoptera: Cantharidae). Entomologica Austriaca 22, 1 - 15

Müller, G.B., and H.L. Nemeschkal (2015). Zoologie im Hauch der Moderne: Vom Typus zum offenen System. 355 – 369. In „Reflexive Innenansichten aus der Universität“, Fröschl, K.A., Müller, G.B., Olechowski, T., and B. Schmidt-Lauber (eds.), Vienna University Press.

Ramler, D.., Ahnelt, H., Nemeschkal H.L.and H. Keckeis (2016). The drift of early life stages of Percidae and Gobiidae (Pisces: Teleostei) in a free-flowing section of the Austrian Danube. Hydrobiologia 781, 199 – 216 /home/nemesch4/html/papers/RamlerEtalHydrobiologia2016.pdf

Wilhelm, G., Handschuh, S., Plant, J., and H.L. Nemeschkal: (2015). Selection becomes visible: enforced sexual dimorphism caused by sexual selection in the weevil Rhopalapion longirostre (Olivier 1807) (Coleoptera: Curculionoidea: Brentidae). Biol. J. Linn. Soc. 115, 38 – 47 /home/nemesch4/html/papers/SelectionVisible.pdf




Arnaiz-Villena, A., Nemeschkal, H.L., and R. van den Elzen. 2006. Molecular modules and morphology. 23rd Int. Ornithol. Congr., August 11-17, 2002, Beijing, China; Acta Zool. Sin. 52(Suppl.), 505 – 508. http://www.actazool.org/paperdetail.asp?id=5126&volume=52&number=-101&bgpage=null&endpage=null&year=2006&month=null

Aspöck, U., and H.L. Nemeschkal. 1998. A cladistic analysis of the Berothidae (Neuroptera). Acta Zool. Fennica 209: 45 - 63.

Aspöck, U., Plant, J.D., and H.L. Nemeschkal. 2001. Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology 26: 73 -86. http://www.blackwell-synergy.com/doi/abs/10.1046/j.1365-3113.2001.00136.x

Brieschke, H., R. van den Elzen, and H.L. Nemeschkal. 1989. Morphometric extractions of functional complexes in avian skeletons: a comparison between pigeons and finches as units of different radiative levels (Aves: Columbiformes and Carduelidae). Ann. Soc. Roy. Zool. Belg. 119 (Suppl. 1):55-56.

Classen, H., R. van den Elzen, and H.L. Nemeschkal. 1987. Determining similarities in Cardueline skeletons using numerical and phenetic analysis. J. Ornithol. 128: 395-

Eberhard, S.H., Nemeschkal, H.L., and H. W. Krenn. 2009. Biometrical evidence for adaptations of the salivary glands to pollen feeding in Heliconius butterflies (Lepidoptera: Nymphalidae). Biol. J. Linn. Soc. 97, 604 – 612. /home/nemesch4/html/papers/Eberhard.pdf

Elzen, R. van den, Nemeschkal, H.L., and H. Classen. 1987. Morphological variation of skeletal characters in the bird family Carduelidae: I. General size and shape patterns in African canaries shown by principal component analyses. Bonn. zool. Beitr. 38:221-239.

Elzen, R. van den, and H. L. Nemeschkal. 1991. Radiation in African Canaries (Carduelidae) - A Comparison of Different Classificatory Approaches. Proc. XX. Congr. Int. Ornithol., Christchurch 1990:459 – 467.

Elzen, R. van den, und H. L. Nemeschkal. 2006. Variabilität im Skelettbau der Cardueliden - ökologisch, phylogenetisch, adaptiv oder konstruktionsmorphologisch dominiert ? Ökol.Vögel (Ecol. Birds) 25 (2003), 231 -247.

Elzen, R. van den, and H. L. Nemeschkal. (2007, published 2009). The impact of body mass on morphological integration in avian skeletons (Aves, Fringillidae; Carduelinae, Fringillinae). Bonn. zool. Beitr. 56, 25-35. /home/nemesch4/html/papers/BodyMass.pdf

Fürnwein, H., Nemeschkal, H.L., and H. Kratochvil. 2004. Ecological differentiation in Gambusia holbrooki Girard 1859 - a morphometric analysis of isolated populations influenced by habitat and climate. Z. Fischkunde 7, 43 – 53.

Gollner, S., Zekely, J., Dover, C.L. van, Govenar, B., Le Bris, N., Nemeschkal, H.L., and M. Bright. 2006. Benthic copepod communities associated with tubeworm and mussel aggregations on the East Pacific Rise. Cah. Biol. Mar. 47, 397-402. http://www.sb-roscoff.fr/CBM/

Gollner, S., Zekely,  J., Govenar, B., Le Bris, N., Nemeschkal,  H.L., Fisher, C.R., and M. Bright. 2007. Tubeworm-associated permanent meiobenthic communities from two chemically different hydrothermal vent sites on the East Pacific Rise. Mar.Ecol.Prog.Ser. 337, 30-49 http://www.int-res.com/abstracts/meps/v337/p39-49/ (bootstrap software - Linux/ Unix: ftboot\ftboot.tar.gz and Win: ftboot\ftboot.zip )

Hirzinger, V., Keckeis, H., Nemeschkal, H.L., and F. Schiemer. 2003. The importance of inshore areas for adult fish distribution along a free-flowing section of the Danube, Austria. River Res. Applic. 19: 1 – 13. http://www3.interscience.wiley.com/cgi-bin/abstract/106570886/ABSTRACT

Keckeis, H., Bauer-Nemeschkal, E., Menshutkin, V.V., Nemeschkal, H.L., and E. Kamler. 2000. Effects of female attributes and egg properties on offspring viability in a rheophilic cyprinid, Chondrostoma nasus. Can. J. Fish. Aquat. Sci. 57: 789 - 796.

Klepal, W., and H.L. Nemeschkal. 1995. The significance of cuticular structures for the systematics of the males of Scalpellidae (Cirripedia thoracica). Amer. Zool. 35:80A.

Klepal, W., and H.L. Nemeschkal. 1995. Cuticular structures in the males of Scalpellidae (Cirripedia thoracica): A character analysis. Pp. 179 - 194 in Schram, F.R., and J.T. Hoeg, eds., Crustacean Issues, 10. New frontiers in barnacle evolution. Balkema, Rotterdam.

Klepal ,W., Gruber, D., and H.L. Nemeschkal. 1997. The use of ultrastructures to solve a classical question of zoology. European J. Cell Biol. Suppl. 45, 74: 100.

Kothbauer, H., H.L. Nemeschkal, H. Sattmann, und E. WAWRA. 1991. Über den Aussagewert von Typen und qualitativen Aufsammlungen: Eine kritische Sicht am Beispiel von Arianta arbustorum styriaca (FRAUENFELD 1868) (Pulmonata, Helicidae). Ann. Naturhist. Mus. Wien 92:229 – 240.

Lange, A., Nemeschkal, H.L., and G.B. Müller: (2014). Biased polyphenism in polydactylous cats carrying a single point mutation: The Hemingway Model for digit novelty. Evol Biol 41:262-275 (online)


Mrsic, N., H. L. Nemeschkal, F. Potocnik, G. Schwammer, und H. Schwammer. 1989. Ein Beitrag zur Herpetofauna der Quarner- Inseln (Jugoslawien - Croatien). Biol. Vestn. 37:57 - 74.

Nemeschkal, H.L. 1982. Zum Erstnachweis eines Os goniale bei Taubenvögeln (Aves - Columbae). Zool. Jb. Anat. 107: 609 - 615.

Nemeschkal, H.L. 1983. Zum Nachweis eines Os coronoideum bei Vögeln - Ein Beitrag zur Morphologie des Sauropsiden-Unterkiefers. Zool. Jb. Anat. 109: 117 - 151.

Nemeschkal, H.L. 1983. Zur Morphologie des Unterkieferskeletes: Das Sauropsidendentale - ein aus 2 Deckknochen zusammengesetztes Element (Vertebrata, Sauropsida). Zool. Jb. Anat. 109: 369 - 396.

Nemeschkal, H.L. 1983. Zur Morphologie und Ontogenie des Os goniale bei tetrapoden Wirbeltieren: Ein System homologer und homoiologer Einzelmerkmale. Morph. Jb. 129: 181 - 216.

Nemeschkal, H.L., und H. Kothbauer. 1988. Arianta arbustorum alpicola (Ferussac, 1819) (Pulmonata, Helicidae): Über Interpretation und Realität eines Taxon. Zool. Anz. 221:343 - 354.

Nemeschkal, H.L., und H. Schwammer. 1988. Unterschiede in der Altersverteilung der Jungtiere in Brutkolonien der Weißkopfmöwe (Larus cachinnans michahellis) auf drei benachbarten Inseln des nördlichen Kvarners (Kroatien, Jugoslawien). Verh. Dtsch. Zool. Ges. 81:324.

Nemeschkal, H.L. 1989. Statistik - Grundbegriffe. Pp. 36 - 46 in H. Kratochwil, ed. Leitfaden für selbständige wissenschaftliche Arbeiten für Biowissenschaften. Universitätsverlag, Wien.

Nemeschkal, H.L. 1989. The mandibular fossa with respect to avian ancestry. Fortschritte der Zoologie 35:467 - 471.

Nemeschkal, H.L., und H. Kothbauer. 1989. Zur Spirale der Schneckenschale: Eine Varianz-Kovarianzuntersuchung bei Arianta (Gastropoda, Helicidae). Zool. Jb. Syst. 116:391 - 409.

Nemeschkal, H.L. 1990. Über die Form der Schneckenschale: Morphometrische Grundlagen und Vorbereitungen für ein statistisches Taxonmodell. Zool. Jb. Syst. 117:491 - 534.

Nemeschkal, H.L., und R. van den Elzen. 1990. Funktionskreise im Skelettsystem der Cardueliden. - Ein morphometrischer Ansatz. Proc. Int. 100. DO-G Meeting, Current Topics Avian Biol., Bonn 1988:37-41.

Nemeschkal, H.L. 1991. Character coupling for taxa discrimination: a critical appraisal of quadratic assignment procedures (QAP). Z. zool. Syst. Evolut.-forsch. 29:87-96.

Nemeschkal, H.L. 1991. Die Schalengröße der Landschnecken (Arianta, Helicidae) als ein sogenannt einfaches Merkmalssystem. - Eine Systemanalyse anhand von Klassifikation und morphologischer Integration. Zool. Jb. Syst. 118:149 - 192.

Nemeschkal, H.L., R.van den Elzen, and H. Brieschke. 1992. The morphometric extraction of character complexes accomplishing common biological roles: avian skeletons as a case study. Z. zool. Syst. Evolut.-forsch. 30:201-219.

Nemeschkal, H.L. 1993. Character Coupling in Land Snails (Arianta, Helicidae): Does Mantel's Test Detect Equality or Similarity ? Zool. Anz. 230:1-17.

Nemeschkal, H.L., H. Classen, and R. van den Elzen. 1994. Morphological Divergence, Distribution and Systematics of Canaries of the Genus Ochrospiza. Proc. VII Pan-Afr. Orn. Congr., Nairobi 1988:241 - 247.

Nemeschkal, H.L., and R.van den Elzen. 1994. Macrosystematics and Infraspecific Character Coupling - An Analysis of Morphological Integration in Avian Skeletons. Zool. Jb. Syst. 121:265-302.

Nemeschkal, H.L. 1999. Morphometric correlation patterns of adult birds (Fringillidae: Passeriformes and Columbiformes) mirror the expression of developmental control genes. Evolution 53: 899 – 918./home/nemesch4/html/papers/EVOLUTION1999.pdf

Nemeschkal, Hans L., and R. van den Elzen. 2006. Size dependence of morphological integration in the true finches, Fringillidae. 24th IOC (2006), Hamburg. J. Ornithol. 147 (Suppl. 1), 218 -

Pokorny, P., Handschuh, S., and H.L. Nemeschkal. 2008. Digital Image Analysis in Morphometrics: Automatic Measurement of Eye Area in Crustacea. 1st Int. Congr. Invertebrate Morphology, Copenhagen 2008; J. Morphol. 269, 1494-

Probst, R., Nemeschkal. H.L., McGrady, M., Tucakov, M., and T. Szep. 2011. Aerial hunting techniques and predation success of Hobbies Falco subbuteo on Sand Martin Riparia riparia at breeding colonies. Ardea 99, 9 – 16./home/nemesch4/html/papers/Probstetal.pdf

Sattmann, H., und H. L. Nemeschkal. 1988. Genitalmorphometrische Untersuchungen an Arianta arbustorum (Linne 1758), A. chamaeleon (L. Pfeiffer 1842) und A. schmidti (Rossmässler 1836) (Mollusca, Gastropoda, Helicidae). Ann. Naturhist. Mus. Wien 90:221 - 340.

Sattmann, H., E. Wawra, H. Kothbauer, und H.L. Nemeschkal. 1991. Eine Perle bei Arianta arbustorum (LINNAEUS, 1758) (Mollusca: Gastropoda: Pulmonata) aus der Steiermark. Ann. Naturhist. Mus. Wien 92:267 – 268.

Schiemer, F., Keckeis, H., Nemeschkal, H., Schludermann, E., Winkler, G., and I. Zweimüller. 2004. Ontogenetic patterns in thermal adaptation of fish vs. long-term temperature trends in large rivers. Intern. Congr. Ser., ITC 1275: 209 – 217.

Schindler, S., Hohmann, U., Probst, R., Nemeschkal, H.L., and G. Spitzer. (2012). Territoriality and Habitat Use of Common Buzzards (Buteo buteo) During Late Autumn in Northern Germany. Journal of Raptor Research 46, 149-157 http://www.bioone.org/doi/abs/10.3356/JRR-11-22.1

Schludermann, E., Keckeis, H., and H.L. Nemeschkal. 2009. Effect of initial size on daily growth and survival in freshwater Chondrostoma nasusus larvae: a field survey. Journal of Fish Biology 74, 939 – 955./home/nemesch4/html/papers/JFishBiol.pdf

Schwammer, H., und H. L. Nemeschkal. 1987. Bedeutung der Exposition für den Standort von Raupen-Nestern des Pinienprozessionsspinners (Thaumetopoea pityocampa Schiff) auf der Insel Krk, Dalmatien. Verh. Dtsch. Zool. Ges. 80:298.

Tadler, A., Nemeschkal, H.L., and G. Pass. 1999. Selection of male traits during and after copulation in the seedbug Lygaeus simulans (Heteroptera, Lygaeidae). Biol. J. Linn. Soc. (London) 68: 471 – 483. http://www.sciencedirect.com/science/journal/00244066

Wilhelm, G., Nemeschkal, H.L., Plant, J., and H.F. Paulus. 2010. Fitness components in the relationship between Rhopalapion longirostre (Olivier, 1807) (Insecta: Coleoptera: Apionidae) and Alcea rosea ( Linnaeus, 1758) (Malvaceae). Analysis of infestation balance of a herbivorous weevil and its host plant. Bonn zool. Bull. 57, 55 – 64. /home/nemesch4/html/papers/RhopAp.pdf

Wilhelm, G., Handschuh, S., Plant, J., and H.L. Nemeschkal. 2011. Sexual dimorphism in head structures of the weevil Rhopalapion longirostre (Olivier 1807) (Coleoptera: Curculionoidea): a response to ecological demands of egg deposition. Biol. J. Linn. Soc. 104, 642 - 660


 Zekely, J., Gollner, S., Dover, C.L. van, Govenar, B., Le Bris, N., Nemeschkal, H.L., and M. Bright. 2006. Nematode communities associated with tubeworm and mussel aggregations on the East Pacific Rise. Cah. Biol. Mar. 47, 477-482. http://www.sb-roscoff.fr/CBM/

Zekely, J., Van Dover, C.L., Nemeschkal, H.L., and M. Bright. 2006. Hydrothermal vent meiobenthos associated with Bathymodiolus aggregations from Mid-Atlantic Ridge and East Pacific Rise.  Deep Sea Research I, Vol. 53, 1363-1378




update: 13-06-2017