Lutz Eckstein is the lead author of a paper summarizing the current knowledge on the biology of the invasive legume Lupinus polyphyllus Lindley. The paper has recently been published in the journal Perspectives in Plant Ecology, Evolution and Systematics in the series “Biological Flora of Central Europe” ( This work is a cooperation with Erik Welk (Martin-Luther-University Halle-Wittenberg and German Centre for Integrative Biodiversity Research (iDiv) Halle, Germany), Yves Klinger and Wiebke Hansen (both Justus Liebig University Giessen, Germany), Tommy Lennartsson and Jörgen Wissman (both Swedish University of Agricultural Sciences (SLU), Uppsala, Sweden), Kristin Ludewig (University of Hamburg, Germany), and Satu Ramula (University of Turku, Finland).

The paper gives a thorough review of the species’ taxonomy, presents distribution maps for North America, Europa and the world (Fig. 1), illustrates the life cycle of L. polyphyllus, and it contains a comprehensive discussion of potential management options. During the research for this review, the authors encountered some doubtful information about L. polyphyllus that uncritically reiterates in several fact sheets, reports and webpages. One such erroneous piece of information refers to the apparently very high longevity of seeds, which was taken from a modelling study on seed longevity under optimal dry and cold storage conditions. Similarly, there is some uncertainty and large variation concerning the actual lifespan of the species. Another piece of doubtful information is the deep rooting depth of L. polyphyllus, which may rather characterize a maximum than a representative average value. Finally, the species is sometimes considered a “rhizomatous perennial” although it lacks true rhizomes. These points highlight some critical knowledge gaps, which partly relate to aspects of the species’ life cycle and morphology that may be either time-consuming or labor-intensive to study.

Fig 1. Distribution of Lupinus polyphyllus s.l. (A) In North America, the native segregates in the west partly overlap in their distribution and are delimited by outlines according to the color scheme in the legend. Non-native, synanthropic occurrences are indicated by black dots. Distribution data based on digitally available herbarium specimen locations and county records (for data sources see Table 1 in the paper). (B) In Europe, numbers give the first records for the species in different countries/regions (cf. Table 6 in the paper). (C) Numbers refer to the textual descriptions (for details, see the paper) of the non-native naturalized distribution across the globe

The authors conclude that there is currently no evidence-based strategy for a cost-efficient management of L. polyphyllus. The development of such control measures is necessary because L. polyphyllus is among the most problematic non-native plant species in Europe with respect to environmental and socio-economic impacts. The species has significant negative effects on community structure, composition, species richness and diversity, especially in nutrient-poor habitats such as alpic mountain hay meadows, alpic mat-grass swards but also nutrient-poor road verges or riparian terraces.

Lutz Eckstein, professor at Karlstad University, is involved in two recently published papers, studying the effects of invasive Garden Lupine (Lupinus polyphyllus) on vegetation and seed bank of mountain meadow plant communities.

The first paper together with Wiebke Hansen (first author), Julia Wollny, Annette Otte and Kristin Ludewig, published in the journal Biological Invasions (, found that the invasion of Garden Lupine homogenizes vegetation composition. The similarity among plots increased with increasing lupine cover in three different vegetation types. L. polyphyllus affected species diversity in terms of richness and effective species number but in rather complex ways, i.e. plots with low to intermediate lupine cover had higher species diversity than control plots. Probably, the invasion though Garden Lupine is linked to significant species turnover. A very clear effect was found for community-weighted means of species trait. In all three vegetation types studied, the canopy height of the community increased with increasing lupine cover, whereas especially in the low-productive Nardus grasslands, leaf dry matter content decreased and specific leaf area increased. Thus, the Garden Lupine shifted the suite of community traits towards more competitive trait values. This may lead to overall more productive plant communities from which rare, low-growing herbs and grasses will disappear.

Germination experiment from the seed bank study in the paper published in Restoration Ecology. Photo: Kristin Ludewig

The second paper with Kristin Ludewig (first author), Wiebke Hansen, Yves Klinger, and Annette Otte, published in the journal Restoration Ecology (, analyzed the effects of increasing cover of Garden Lupineon the seed bank of mountain meadows, and the potential of the seed bank of these stands for active restoration of mountain meadows in terms of species composition and number. The authors conducted a seed bank analysis on 84 plots with increasing cover of L. polyphyllus in three mountain-meadow types of the Rhön Biosphere Reserve, Germany. Seedlings from 119 species germinated from the seed bank samples, including 17 Red List species but only a few seedlings of L. polyphyllus. While the influence of L. polyphyllus on the current vegetation was visible, no effects on the seed bank were apparent. L. polyphyllus had no influence on total seed density, seed density of typical mountain-meadow species, or species numbers in the seed bank. Only the seeds of the Red List species were significantly related to the cover of L. polyphyllus. The authors conclude that the seed bank offers potential for active restoration of species-rich mountain meadows, but species absent from the seed bank have to be added by other measures.

Kristin Ludewig, Wiebke Hansen and Yves Klinger will presents these and other results from a large restoration project in the UNESCO Rhön Biosphere Reserve at the RivEM week.

Lutz Eckstein, professor at Karlstad University, together with Yves Klinger, David HorlemannAnnette Otte and Kristin Ludewig recently published the paper Germination of the invasive legume Lupinus polyphyllus depends on cutting date and seed morphology in the journal NeoBiota. The paper is accessible open access on the journal website, or you can read a summary of the paper below:

In semi-natural grasslands, mowing leads to the dispersal of species that have viable seeds at the right time. For invasive plant species in grasslands, dispersal by mowing should be avoided, and information on the effect of cutting date on the germination of invasive species is needed. We investigated the germination of seeds of the invasive legume Lupinus polyphyllus Lindl. depending on the cutting date. We measured seed traits associated with successful germination that can be assessed by managers for an improved timing of control measures. Germination patterns were highly asynchronous and differed between seeds cut at different dates. Seeds cut early, being green and soft, tended to germinate in autumn. Seeds cut late, being dark and hard, were more prone to germinate the following spring, after winter stratification. This allows the species to utilize germination niches throughout the year, indicating a bet-hedging strategy.

Lupine seeds used in the study

Seed color and the percentage of hard seeds were good predictors of germination percentage. Managers should prevent the species from producing black and hard seeds, while cutting plants carrying green and soft seeds appears less problematic. Furthermore, germination patterns differed between experiments in climate chambers and in the common garden, mainly because germination of dormant seeds was lower in climate chambers. We propose that more germination experiments under ambient weather conditions should be done, as they can give valuable information on the germination dynamics of invasive species.

Yves P. Klinger, Sarah Harvolk-Schöning, Lutz Eckstein, Wiebke Hansen, Annette Otte and Kristin Ludewig recently published the paper ”Applying landscape structure analysis to assess the spatio-temporal distribution of an invasive legume in the Rhön UNESCO Biosphere Reserve” in the journal Biological Invasions.

Lutz Eckstein, Professor in Biology at Karlstad University, writes about their work:

The legume Lupinus polyphyllus. Photo by Lutz Eckstein

“We applied a combination of aerial mapping and GIS-based landscape analysis to study the invasion of the legume, Lupinus polyphyllus, in the Rhön UNESCO Biosphere Region as a case. We assessed the changes in lupine distribution between 1998 and 2016 in a strictly protected part of the Biosphere Region by means of landscape structure analysis. The area invaded by L. polyphyllus doubled from 1998 to 2016. The number of lupine stands decreased by 25%, but average stand size increased by 300%. In 2016, large and well-connected mesic grasslands that were situated close to roads were more heavily invaded than small and remote wet grasslands. Our results show that landscape composition plays an important role for the spread of invasive plants. Specifically, invasive stand characteristics, such as stand size, form, and connectivity, are crucial for driving the invasion process. Therefore, in addition to landscape composition, invasive stand characteristics should be included in the planning of conservation measures. Overall, aerial mapping combined with landscape analysis provides a cost-effective and practical tool for landscape managers to prioritize invasive control measures.”

Access the paper here, or contact any of the authors.