Acta Zoologica Academiae Scientiarum Hungaricae 47 (2–3), pp. 87–267, 2001

Title: The meadow and steppe vipers of Europe and Asia – the Vipera (Acridophaga) ursinii complex

Authors: Nilson, G. and C. Andrén*

Authors' address: Göteborg Natural History Museum, Box 7283, SE-40235 Göteborg, Sweden
*Reptilia Amphibia Research, Department of Zoology, Göteborg University
Box 463, SE-405 30 Göteborg, Sweden, E-mail:

Abstract: The systematic position and validity of the currently recognised subspecies in the "Vipera ursinii s. l. complex" (=Acridophaga) are evaluated from phenetic and phylogenetic perspectives. The West Asian and European mountain populations have evolved independently. A picture emerges of one group of comparatively widespread lowland taxa and three different assemblages of mountainous taxa. The lowland group covers rakosiensis; moldavica, west and east renardi, and the mountain groups consist of:

– the European ones (Italian ursinii, French ursinii, macrops and graeca)

– the Transcaucasian and Turkish (lotievi, eriwanensis, ebneri and anatolica)

– the Transcaspian and Chinese ones (tienshanica ssp.n. and parursinii ssp. n.).

To the eastern group, the lowland "Altai form of renardi" must be added. This is a taxon with "mountain morphology".

The various mountain populations ranging from France in the west to China in the east are characterised by a series of convergent adaptations (homoplasies). In addition, they are adapted to mountain and/or alpine habitats and environments.

1. Immunological and electrophoretical characteristics analysed with cladistic and phylogenetic methods support a north-south relationship between taxa (Joger et al. 1992, Nilson et al. 1993).

2. Newer observations and records revealed that several traditionally used morphological character states used in phylogenetic discussions are reversals and convergences (e.g. 19 midbody scale rows).

3. A cladistic analysis of the revised morphology (2., above) results in a unanimous phylogeny consistent with the results of the molecular analyses (1., above).

From a cladistic point of view, the most derived group are the different mountain populations, a hypothesis based on cladistic phylogenetics, and supported by two alternative approaches: molecular and morphometric.

The following taxa are monophyletic and fulfil the criteria of the phylogenetic species concept: ((((rakosiensis) (Italian ursinii) (macrops, graeca)) (French ursinii) (moldavica)) (((west-renardi) (((eriwanensis) (anatolica) (ebneri)) (lotievi)))((east- renardi) ((tienshanica) (("Altai form of renardi") (parursinii)))))). Biochemical analyses support the major pattern in this picture with the difference of placing moldavica as a sister taxon to macrops and graeca.

We consider renardi as a good species, and consequently none of its descendant populations anatolica, eriwanensis, ebneri, tienshanica, the "Altai form of renardi" and parursinii can be subspecies of ursinii s.str. To reflect the evolutionary history a nomenclature shift is performed. Earlier, several of the eastern taxa have been raised to species level, or are treated as such in the present work. The relationship of Vipera darevskii and Vipera pontica remains uncertain.