Once you have gone through the effort of collecting a global database, you can start playing with it. And one of the most fun things to play with, I’d say, is general ecological theories – the kind that you would think hold everywhere, but that usually play out differently depending on where and how you test them. We’ve done a few of those with our MIREN plant survey database from along mountain roads, and here’s another important one, just published in the Journal of Vegetation Science (Buhaly et al. 2026).
This time, the theory on the chopping block is biotic resistance: the old and intuitive idea, going back to Elton in 1958, that a diverse, well-established native community should be harder for a newcomer to break into. More species already occupying the available niches, competing for light, water and space, should mean fewer open doors for a non-native plant trying to move in. It’s one of the most-cited explanations in invasion biology, but it rests on the assumption that competition is what dominates plant-plant interactions. And we’ve known for a while that this assumption gets shakier the harsher the environment gets: in stressful places, facilitation (plants actually helping each other, via shelter, warmth or moisture) tends to take over from competition. High-elevation, alpine environments are about as stressful as it gets. So does biotic resistance survive the trip up to the treeline and beyond?

Testing an old idea
Together with the whole consortium, and led by Meike Buhaly, we used our MIREN road survey data to find out, using 15 mountain regions on six continents. At each site, a plot runs along the roadside, and a second one starts 50 m off the road and heads into the (semi-)natural vegetation behind it, the same paired design we’ve used since 2007, and that has featured oftentimes on this blog here. On top of that, we (well, mostly the lead author team in Germany, I just collected the leaves) built a proper trait dataset, with nearly 1,500 leaf samples, run through a near-infrared spectrometer back in the lab, to calculate not just how many native species a plot holds, but how functionally different those species actually are from one another.

No resistance from richness – sometimes the opposite

For functional diversity (bottom panel), the pattern is very different
The short version: using native species richness as the classic proxy for biotic resistance, the expected resistance pattern didn’t show up anywhere in this dataset. If anything, it was reversed at low-to-mid elevations, where non-native richness was consistently higher with native richness. This was true, unexpectedly, both at the roadside and in the natural vegetation, plausibly because species-rich, structurally complex lowland communities simply offer more niches and microsites, for natives and non-natives alike, than their species-poor counterparts.
Looking at functional diversity told a different story, however. At low elevations, functionally diverse natural communities did host fewer non-natives – that’s one piece of actual biotic resistance we did find! But climb the elevation gradient, and that relationship weakens, then flips: at the highest elevations, functionally diverse native communities host more non-native species again. Exactly what you’d expect if facilitation, not competition, is running the show up there, as has been shown before for native cushion plants acting as “nurse plants” for exotic invaders in the high Andes.
So: some evidence for biotic resistance at low elevations, when using functional diversity, but overall a fairly strong case for the opposite to be happening.
Going back to Norway 2012
Then, at the end of the paper, there’s a part that I am a big fan of personally. Back in 2012, as a master’s student doing the first MIREN survey in northern Norway, I found that the ratio of non-native to native species in the roadside – compared to the untouched vegetation right behind it – increased steeply with elevation. My reading at the time: weaker resistance up high lets a disproportionate share of whatever non-natives do arrive spill off the road into the natural vegetation. That finding did get published, in my first ever peer-reviewed publication, but I always felt a bit hesitant. I trusted the pattern and process, but I felt like we just didn’t have enough data – very low invasion levels in Norway – to confidentially accept the claim as proven.
The global dataset now shows that this ratio does indeed climb toward high elevations, as we saw in Norway. However, it’s not a simple upward slope. It’s a U: the ratio of non-natives in natural vegetation relative to the roadside is highest at both ends of the elevation gradient (around 0.6 at each extreme), and dips in the middle. The high-elevation arm of that U is the one I stumbled onto in Norway — non-natives that make it up the road managing to spread into the surrounding alpine vegetation surprisingly easily, either helped along by exactly the facilitation signal described above, or by an overall reduction in interactions (my reading). The low-elevation arm is a different story, though: lowland sites are where most non-natives arrive in the first place, have had by far the longest time to build up a presence, and can draw on a far bigger regional species pool, so plenty of them eventually work their way off the road there too, just through sheer numbers and time, and regardless of the resistance levels.

Which also explains, rather neatly, why my Norwegian roads never showed that low-elevation bump: Norway’s cold climate, isolation and modest regional species pool have always made it something of an outlier when it comes to that kind of long-term, low-elevation invasion build-up. What I picked up on in 2012 was thus really just one half of a much bigger, two-sided global pattern, and it took fourteen years and a lot of collaborating mountains to see the other half. Glad!

Reference: Buhaly, M., Turner, S.C., Kreuz, N.K., Vandvik, V., Veltmann, B., Alexander, J.M., Lembrechts, J.J., et al. (2026). No signal of biotic resistance to non-native species establishment in high-elevation communities. Journal of Vegetation Science, 37, e70174. https://doi.org/10.1111/jvs.70174
































