ABSTRACT
Syntrichia relies on external water conduction for photosynthesis, survival, and reproduction, a condition referred to as ectohydry. Capillarity spaces are abundant in Syntrichia, but the link between function and morphology is complex. The aim of this study was to provide a better understanding of species-specific morphological traits underlying the functions of water conduction and storage. We used an environmental scanning electron microscope and confocal microscopy for observing anatomical characters in the leaves of Syntrichia species. We also measured hydration/dehydration curves to understand the rate of conduction and dehydration by experimental approaches. Syntrichia is an ectohydric moss that can externally transport and store water from the base of the stem using capillary action. We propose a new framework to study ectohydric capabilities, which incorporates three morphological scales and the timing of going from completely dehydrated to fully hydrated. Characters of interest in this model include cell anatomy (papillae development, hyaline basal cells and laminar cells), architecture of the stem (concavity and orientation) and whole clump characteristics (density of stems). We report significant variations in the speed of conduction, water holding capacity and hydration associated with each species studied (11 in total). All Syntrichia species are capable of external water conduction and storage, but the relevant traits differ among species. These results help to understand potential evolutionary and ecological trade-offs among speed of water conduction, water holding capacity, ontogeny, and differing habitat requirements. An integrative view of ectohydry in Syntrichia contributes to understanding the water relationships of mosses.
ABSTRACT
PREMISE: To address the biodiversity crisis, we need to understand the evolution of all organisms and how they fill geographic and ecological space. Syntrichia is one of the most diverse and dominant genera of mosses, ranging from alpine habitats to desert biocrusts, yet its evolutionary history remains unclear. METHODS: We present a comprehensive phylogenetic analysis of Syntrichia, based on both molecular and morphological data, with most of the named species and closest outgroups represented. In addition, we provide ancestral-state reconstructions of water-related traits and a global biogeographic analysis. RESULTS: We found 10 major well-resolved subclades of Syntrichia that possess geographical or morphological coherence, in some cases representing previously accepted genera. We infer that the extant species diversity of Syntrichia likely originated in South America in the early Eocene (56.5-43.8 million years ago [Mya]), subsequently expanded its distribution to the neotropics, and finally dispersed to the northern hemisphere. There, the clade experienced a recent diversification (15-12 Mya) into a broad set of ecological niches (e.g., the S. caninervis and S. ruralis complexes). The transition from terricolous to either saxicolous or epiphytic habitats occurred more than once and was associated with changes in water-related traits. CONCLUSIONS: Our study provides a framework for understanding the evolutionary history of Syntrichia through the combination of morphological and molecular characters, revealing that migration events that shaped the current distribution of the clade have implications for morphological character evolution in relation to niche diversity.
