Nitrogen-fixing symbiosis is globally important in ecosystem functioning and agriculture, yet the evolutionary history of nodulation remains the focus of considerable debate. Recent evidence suggesting a single origin of nodulation followed by massive parallel evolutionary losses raises questions about why a few lineages in the N 2 -fixing clade retained nodulation and diversified as stable nodulators, while most did not. Within legumes, nodulation is restricted to the two most diverse subfamilies, Papilionoideae and Caesalpinioideae, which show stable retention of nodulation across their core clades.We characterize two nodule anatomy types across 128 species in 56 of the 152 genera of the legume subfamily Caesalpinioideae: fixation thread nodules (FTs), where nitrogen-fixing bacteroids are retained within the apoplast in modified infection threads, and symbiosomes, where rhizobia are symplastically internalized in the host cell cytoplasm within membranebound symbiosomes (SYMs).Using a robust phylogenomic tree based on 997 genes from 147 Caesalpinioideae genera, we show that losses of nodulation are more prevalent in lineages with FTs than those with SYMs.We propose that evolution of the symbiosome allows for a more intimate and enduring symbiosis through tighter compartmentalization of their rhizobial microsymbionts, resulting in greater evolutionary stability of nodulation across this species-rich pantropical legume clade.

We analysed thirty-five 400-m2 plots encompassing forest, savanna and intermediate vegetation types in an ecotonal area in Ghana, West Africa. Across all plots, fire frequency was over a period of 15 years relatively uniform (once in 2–4 years). Although woodlands were dominated by species typically associated with savanna-type formations, and with forest formations dominated by species usually associated with closed canopies, these associations were non-obligatory and with a discrete non-specialized species grouping also identified. Across all plots, crown area index, stem basal area and above-ground biomass were positively associated with higher soil exchangeable potassium and silt contents: this supporting recent suggestions of interplays between potassium and soil water storage potential as a significant influence on tropical vegetation structure. We also found an average NDVI cover increase of ~0.15% year−1 (1984–2011) with plots dominated by non-specialized species increasing more than those dominated by either forest- or savanna-affiliated species. Our results challenge the traditional view of a simple forest vs. savanna dichotomy controlled by fire, and with our newly identified third non-specialized species grouping also potentially important in understanding ecotonal responses to climate change.

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