Citation: Baur, B.; Rusterholz, H.-P.;
Braschler, B. The Grassland
Fragmentation Experiment in the
Swiss Jura Mountains: A Synthesis.
Diversity 2023, 15, 130. https://
doi.org/10.3390/d15020130
Academic Editor: Simona Maccherini
Received: 25 November 2022
Revised: 13 January 2023
Accepted: 14 January 2023
Published: 18 January 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
diversity
Review
The Grassland Fragmentation Experiment in the Swiss Jura
Mountains: A Synthesis
Bruno Baur *, Hans-Peter Rusterholz and Brigitte Braschler
Department of Environmental Sciences, University of Basel, Bernoullistrasse 30, 4056 Basel, Switzerland
* Correspondence: bruno.baur@unibas.ch
Abstract: We synthesize findings from a 7-year fragmentation experiment in species-rich, nutrient-
poor, dry calcareous grasslands in the north-western Jura mountains, Switzerland. We used a
standardized approach with 48 fragments (0.25–20.25 m
2
) and corresponding control plots in three
sites. The 5-m-wide isolation area around the fragments was maintained by frequent mowing. Frag-
ments experienced various ecological changes, e.g., plant biomass increased along fragment edges.
We examined fragmentation effects on species richness and composition, abundance, genetic diversity,
functional diversity, species traits and species interactions (pollination, herbivory, parasitism, disease)
in a wide array of invertebrate (gastropods, spiders, woodlice, various insect groups) and plant taxa.
Responses to fragmentation differed between taxonomical groups and species. While species richness
and individual density were lower in fragments in some groups, the opposite was true for other
groups. Fragmentation effects were most pronounced on species interactions; however, some effects
only occurred with a delay. For example, fragmentation influenced foraging patterns of bumblebees,
affecting pollination, which in turn resulted in a decreased outcrossing frequency and reduced genetic
diversity in a focal plant species. We highlight key findings of the experiment and emphasize their
implications for grassland conservation.
Keywords: biodiversity; calcareous grasslands; environmental change; functional dispersion; habitat
fragmentation; invertebrates; population dynamics; time lags
1. Introduction
The current biodiversity loss due to environmental change is of great concern glob-
ally [1,2]. Habitat fragmentation is generally considered to be one of the major drivers
of this loss [1,3–6]. Fragmentation reduces the total area of original habitat, changes the
environmental conditions in remnants, creates isolated subpopulations, disrupts individual
behavior [7], interrupts the exchange of individuals and genes among populations [8], and
alters species interactions [9] and ecological processes [10]. Thus, habitat fragmentation can
influence an entire suite of processes, ranging from individual behavior through population
dynamics to ecosystem fluxes.
Habitat fragmentation occurs at different spatial scales [11,12]. It ranges from small
breaks in otherwise homogeneous habitat to widely scattered fragments in a hostile ma-
trix [13]. The relevant spatial scale is different for each species [14,15]. The response of a
species to fragmentation depends on its dispersal ability, life history, and competitiveness,
and on the type and size of the fragments, and the permeability for dispersing individuals
of the matrix surrounding the fragment [4,16].
Large-scale fragmentation on the landscape level entails many challenges unfavorable
to rigorous investigation. Habitat fragments in a landscape differ in size, shape, age (time
since isolation), distance to large homogeneous habitat, and distance to other fragments:
all of which make any proper replication of the investigation unit difficult [17]. Further-
more, corresponding control areas are frequently lacking [18]. Experimental studies with
Diversity 2023, 15, 130. https://doi.org/10.3390/d15020130 https://www.mdpi.com/journal/diversity