Weed suppression is a potential benefit of cover crop mixtures, as species diversity may allow for combining early and late-season competition with weeds. Here, we studied if this is possible for only-legume mixtures containing species with different growth rates, by testing two legumes, alsike clover (AC; Trifolium hybridum L.) and black medic (BM; Medicago lupulina L.) in two field trials sown in 2016 and 2017. Five AC:BM ratios (100:0, 67:33, 50:50, 33:67, and 0:100) were grown at three densities (50%, 100%, and 150% of recommended seed density). Cover crop and weed aboveground biomass (CCB and WB, respectively) were harvested three times, after establishment in spring (H1), in summer (H2), and in autumn after mulching (H3). Compared to fallow plots, all monocultures and mixtures showed early-season weed suppression in terms of biomass production and more efficiency over time with an average reduction of 42%, 52%, and 96% in 2016, and 39%, 55%, and 89% in 2017 at H1, H2, and H3, respectively. Out of 54 mixture treatments, only eight mixtures showed stronger weed suppression than monocultures. Mixtures reduced WB by 28%, as an average value, in 2017 compared to the respective monocultures, but not significantly in 2016, indicating that the crop diversity effect on weeds was dependent on the growing environment. Weed suppression was significantly higher at 100% and 150% seed density than 50%, but no significant differences were determined between 100% and 150% seed density. After mulching, no density effect was observed on CCB and WB. In conclusion, AC and BM can be used as a keystone species on weed suppression for sustainable agriculture as they possess plasticity to suppress weeds when higher biomass productivity is limited by environmental conditions. However, their diversity effects are time and condition dependent. Appropriate seed density and mulching can successfully be employed in weed management, but seed density may not have an effect after mulching.

In the face of increasingly frequent droughts threatening crop performance, ecological theory suggests that higher species diversity may help buffering productivity by making systems more resistant through resource complementarity and more resilient through higher response diversity. However, empirical evidence for these diversity effects under drought stress has remained patchy. In two pot experiments, we explored whether mixing two legume species with a contrasting response to water availability, alsike clover (AC) and black medic (BM), promotes resistance to cumulative drought stress, and resilience of aboveground crop biomass to a transient drought event. The mixture was more productive than the average of the sole crops, and this mixture effect was higher in the non-stressed than in the drought-stressed plants. However, with six levels of constant drought intensities, the mixture effect was not consistently affected by drought level. Response diversity was evident as asynchrony of growth in the two species after the drought event, with BM re-growing faster than AC. Significant resilience to drought was observed in sole AC, i.e., without response diversity. Resilience was larger in AC than in BM and increased from 44 to 72 days after sowing (DAS). The mixture was more resilient than the average resilience of the sole crops at 72 DAS, but it was never more resilient than AC, indicating that resilience is promoted by, but not dependent on response diversity. We conclude that crop diversity may contribute to drought resilience through growth asynchrony, but that species identity plays a crucial role in making systems more drought-resilient.

While intercropping is known to have positive effects on crop productivity, it is unclear whether the effects of mixing species start at the early plant stage, that is, during germination. We tested whether the germination of two legume species, alsike clover and black medic, characterized by a contrasting response to water availability and temperature is affected by mixing. We set up four experiments in each of which we compared a 1:1 mixture against the two monocultures, and combined this with various other experimental factors. These additional factors were (i) varied seed densities (50%, 100% and 150% of a reference density) in two field trials in 2016 and 2017, (ii) varied seed densities (high and low) and water availability (six levels, between 25% and 100% of water holding capacity (WHC)) in a greenhouse pot trial, (iii) varied seed spacing in a climate chamber, and (iv) varied temperatures (12 °C, 20 °C and 28 °C) and water availability (four levels between 25% and 100% of WHC) in a climate chamber. Across all experiments, the absolute mixture effects (AME) on germination ranged between −9% and +11%, with a median of +1.3%. Within experiments, significant mixture effects were observed, but the direction of these effects was inconsistent. In the field, AME on germination was significantly negative at some of the tested seed densities. A positive AME was observed in the climate chamber at 12 °C, and the mean AME decreased with increasing temperature. Higher density was associated with decreased germination in the field, indicating negative interaction through competition or allelopathy, among seedlings. Our findings indicate that interaction among seeds in species mixtures may be ongoing during germination, but that the direction of the mixture effect is affected by complex interactions with abiotic and biotic factors.