Nyssa

species delineation and biogeography

Phylogenomic and biogeographic studies of Nyssa (Nyssaceae) using RAD-seq, and Fluidigm sequencing. (Advisor: Dr. Jenny Xiang)

1.Nyssa sylvatica complex consists of several woody taxa occurring in eastern North America. These taxa were recognized as two or three species including three or four varieties by different authors. Due to high morphological similarities and complexity of morphological variation, classiļ¬cation and delineation of taxa in the group have been difficult and controversial. Here we employ data from RAD-seq to elucidate the genetic structure and phylogenetic relationships within the group. Using the genetic evidence, we evaluate previous classiļ¬cations and delineate species. Our results support the recognition of two species in the complex, N. sylvatica and N. biļ¬‚ora, following the phylogenetic species concept. We found no genetic evidence supporting recognitions of intraspeciļ¬c taxa. However, we propose subsp. ursina and subsp. biļ¬‚ora within N. biļ¬‚ora due to their distinction in habits, distributions, and habitats. This work was published in MPE (DOI:)[add link]

Fig. 1. Nyssa sylvativa complex. On the left, Nyssa sylvativa leaves. Middle, Nyssa biflora leaves. Right, Nyssa biflora subsp. ursina.
Fig. 2. Results from STRUCTURE analyses showing genetic grouping of the Nyssa sylvatica complex using the M50 unlinked SNP dataset. N. biļ¬‚ora shares the same gene pool (blue region) with N. ursina, while N. sylvatica and its varieties share the second gene pool (orange region).

2.Nyssa represents a classical example of the wellā€known eastern Asianā€“eastern North American floristic disjunction. The genus consists of three species in eastern Asia, four species in eastern North America, and one species in Central America. Species of the genus are ecologically important trees in eastern North American and eastern Asian forests. The distribution of living species and a rich fossil record of the genus make it an excellent model for understanding the origin and evolution of the eastern Asianā€“eastern North American floristic disjunction. However, despite the small number of species, relationships within the genus have remained unclear and have not been elucidated using a molecular approach.

Fig. 3. Distribution of Nyssa from both eastern Asia, eastern North America, and Central America. This figure also indicate the biogeographic history of Nyssa.

Here, we integrate data from 48 nuclear genes, fossils, morphology, and ecological niche to resolve species relationships, elucidate its biogeographical history, and investigate the evolution of morphology and ecological niches, aiming at a better understanding of the wellā€known EAā€“ENA floristic disjunction. Results showed that the Central American (CAM) Nyssa talamancana was sister to the remaining species, which were divided among three, rapidly diversified subclades. Estimated divergence times and biogeographical history suggested that Nyssa had an ancestral range in Eurasia and western North America in the late Paleocene. The rapid diversification occurred in the early Eocene, followed by multiple dispersals between and within the Erasian and North American continents. The genus experienced two major episodes of extinction in the early Oligocene and end of Neogene, respectively. The Central American N. talamancana represents a relic lineage of the boreotropical flora in the Paleocene/Eocene boundary that once diversified in western North America. The results supported the importance of both the North Atlantic land bridge and the Bering land bridge (BLB) for the Paleogene dispersals of Nyssa and the Neogene dispersals, respectively, as well as the role of Central America as refugia of the Paleogene flora. The totalā€evidenceā€based dated phylogeny suggested that the pattern of macroevolution of Nyssa coincided with paleoclimatic changes.

Fig. 4. Phylogeny of Nyssa. Left, phylogenetic tree using IQ-Tree, RAxML, and MrBayes methods. Right, DensiTree plot using StarBEAST2 results.

We found a number of evolutionary changes in morphology (including wood anatomy and leaf traits) and ecological niches (precipitation and temperature) between the EAā€“ENA disjunct, supporting the ecological selection driving trait evolutions after geographic isolation. We also demonstrated challenges in phylogenomic studies of lineages with rapid diversification histories. The concatenation of gene data can lead to inference of strongly supported relationships incongruent with the species tree. However, conflicts in gene genealogies did not seem to impose a strong effect on divergence time dating in our case. Furthermore, we demonstrated that rapid diversification events may not be recovered in the divergence time dating analysis using BEAST if critical fossil constraints of the relevant nodes are not available. Our study provides an example of complex bidirectional exchanges of plants between Eurasia and North America in the Paleogene, but ā€œout of Asiaā€ migrations in the Neogene, to explain the present disjunct distribution of Nyssa in EA and ENA. This work was invited to be published in a special issue in JSE (DOI:)[add link]