Floral morphology and phenology of Sassafras tzumu (Lauraceae).

BACKGROUND: Sassafras has been considered to belong to trib. Laureae of Lauraceae and has been assumed to have unisexual flowers. However, recent molecular phylogenetic studies have consistently suggested that Sassafras does not belong to the trib. Laureae but to Cinnamomeae and that it is nested within Cinnamomum. A recent morphological study revealed that one of the Asian species, S. randaiense, possesses bisexual flowers that are plesiomorphic in the family Lauraceae. As reports on the flower structure of the second Asian species, S. tzumu, have been contradictory, we wanted to ascertain if it has bisexual flowers or not. If the flowers were bisexual, could earlier reports that they were unisexual have been based on dichogamous flowering? RESULTS: In this study, we investigated two populations of S. tzumu. We found that this species has determinate botryoid racemes, and possesses bisexual flowers. Among the three extant species, S. tzumu is more similar to its sister species S. randaiense but markedly different from the American S. albidum: the two Asian species possess bisexual flowers while the American species has unisexual flowers. The bisexual flower of S. tzumu is protogynous, and shows two phenological phases typical of Lauraceae: 1) in a flower, the pistil functions first, the stigma is fresh and white, stamens of the outer two whorls are spreading, anthers do not open, and the staminodes secrete nectar at this stage; 2) in the second phase, the stigma becomes brown, staminodes are withered, stamens of the third whorl stand up and surround the pistil, glands of the third whorl of stamens secrete nectar, and the anthers open and release pollen. CONCLUSIONS: The similarity of racemose inflorescences between Sassafras and some members of Laureae were caused by parallel evolution; the racemose inflorescence of ancestral Sassafras originated from the thyrsoid-cymose inflorescence in Cinnamomum. The Asian species S. tzumu and S. randaiense possess bisexual flowers with two phenological phases, the American S. albidum evolved unisexual flowers independently from other clades with unisexual flowers in the Lauraceae, i.e., the Laureae, Alseodaphnopsis in the Perseeae and the unisexual clade in the Ocotea complex of the Cinnamomeae.

A comparative analysis of complete chloroplast genomes of seven Ocotea species (Lauraceae) confirms low sequence divergence within the Ocotea complex.

The genus Ocotea (Lauraceae) includes about 450 species, of which about 90% are Neotropical, while the rest is from Macaronesia, Africa and Madagascar. In this study we present the first complete chloroplast genome sequences of seven Ocotea species, six Neotropical and one from Macaronesia. Genome sizes range from 152,630 (O. porosa) to 152,685 bp (O. aciphylla). All seven plastomes contain a total of 131 (114 unique) genes, among which 87 (80 unique) encode proteins. The order of genes (if present) is the same in all Lauraceae examined so far. Two hypervariable loci were found in the LSC region (psbA-trnH, ycf2), three in the SSC region (ycf1, ndhH, trnL(UAG)-ndhF). The pairwise cp genomic alignment between the taxa showed that the LSC and SSC regions are more variable compared to the IR regions. The protein coding regions comprise 25,503-25,520 codons in the Ocotea plastomes examined. The most frequent amino acids encoded in the plastomes were leucine, isoleucine, and serine. SSRs were found to be more frequent in the two dioecious Neotropical Ocotea species than in the four bisexual species and the gynodioecious species examined (87 vs. 75-84 SSRs). A preliminary phylogenetic analysis based on 69 complete plastomes of Lauraceae species shows the seven Ocotea species as sister group to Cinnamomum sensu lato. Sequence divergence among the Ocotea species appears to be much lower than among species of the most closely related, likewise species-rich genera Cinnamomum, Lindera and Litsea.

Leaf epidermal micromorphology defining the clades in Cinnamomum (Lauraceae).

In this study, we sampled 48 species of Asian Cinnamomum covering the species groups that were identified in recent phylogenetic studies and conducted leaf micromorphological observations using both light microscopy (LM) and scanning electron microscopy (SEM). Synapomorphies were determined by means of mapping micromorphological characters on a phylogenetic tree. The results indicate that Cinnamomum exhibits two different types of leaf upper epidermis: Type I has smooth/non-reticulate periclinal walls whereas Type II has reticulate periclinal walls and is unusual in the family Lauraceae. We found that the two types of micromorphological characters are clade-specific, sect. Camphora s.s. possesses Type I leaf upper epidermis, and sect. Cinnamomum s.l. has Type II leaf upper epidermis. Our study also reveals that C.saxatile, a member of sect. Camphora s.l. in the traditional classification, actually has Type II leaf upper epidermis, thus reinforcing the result of a recent molecular phylogeny that has this species in a clade consisting mainly of species of sect. Cinnamomum.

Floral structure and ontogeny of Syndiclis (Lauraceae).

Generic delimitation in the Beilschmiedia group of the Lauraceae remains ambiguous because flowering specimens of a few genera with confined distribution are poorly represented in herbaria, and a few floral characters important for taxonomy are still poorly known. Syndiclis is sporadically distributed in southwestern China, and is represented in the herbaria by only a few flowering specimens. We conducted field investigations to collect floral materials of four species and observed structures and ontogeny of the tiny flowers using both light microscopy (LM) and scanning electron microscopy (SEM). The results show that the genus Syndiclis possesses flowers with huge variation in both merosity and organ number. Flowers of the genus are dimerous, trimerous, or tetramerous, or have mixed merosity with monomerous and dimerous, or dimerous and trimerous, or trimerous and tetramerous whorls. The number of staminodes ranges from two to eight, depending on floral merosity, and on how many stamens of the third androecial whorl are reduced to staminodes. The staminodes of the fourth androecial whorl are comparable to the staminodes in Potameia, but the staminodes of the third androecial whorl of Syndiclis are relatively larger than the staminodes in Potameia. They are erect or curved inwards, covering the ovary. The anthers are usually two-locular, but rarely one-locular or three-locular. Each stamen of the third androecial whorl bears two conspicuous and enlarged glands at the base. The lability of floral merosity and organ number of Syndiclis may have been caused by changes of pollination system and loss of special selective pressures that are present in most Lauraceous plants with fixed floral organ number. This study furthers our understanding of variation and evolution of a few important characters of the Beilschmiedia group and provides essential data for a revised generic classification of the group.

Alseodaphnopsis: A new genus of Lauraceae based on molecular and morphological evidence.

An investigation of a questionable species of the genus Alseodaphne led to the discovery of a new genus Alseodaphnopsis H. W. Li & J. Li, gen. nov., separated from Alseodaphne Nees, and a new species Alseodaphnopsis ximengensis H. W. Li & J. Li, sp. nov., endemic to Yunnan province, China. This new species is characterized by having big, axillary, paniculate inflorescences, as well as large, subglobose fruits. Based on DNA sequence data from two gene regions (nuclear ribosomal ITS and LEAFY intron II), we investigate its phylogenetic position within the Persea group. Phylogenies using maximum parsimony (MP) and Bayesian inference (BI) support the recognition of Alseodaphnopsis as a distinct genus but do not resolve well its relationship within the Persea group. The new genus is circumscribed, eight new combinations for its species are made, and a description and illustration of the new species are provided.

Origins and evolution of cinnamon and camphor: A phylogenetic and historical biogeographical analysis of the Cinnamomum group (Lauraceae).

Tropical and subtropical amphi-Pacific disjunction is among the most fascinating distribution patterns, but received little attention. Here we use the fossil-rich Cinnamomum group, a primarily tropical and subtropical Asian lineage with some species distributed in Neotropics, Australasia and Africa to shed light upon this disjunction pattern. Phylogenetic and biogeographic analyses were carried out using sequences of three nuclear loci from 94 Cinnamomum group and 13 outgroup samples. Results show that although there are three clades within a monophyletic Cinnamomum group, Cinnamomum and previously recognized subdivisions within this genus were all rejected as natural groups. The Cinnamomum group appears to have originated in the widespread boreotropical paleoflora of Laurasia during the early Eocene (ca. 55Ma). The formation and breakup of the boreotropics seems to have then played a key role in the formation of intercontinental disjunctions within the Cinnamomum group. The first cooling interval (50-48Ma) in the late early Eocene resulted in a floristic discontinuity between Eurasia and North America causing the tropical and subtropical amphi-Pacific disjunction. The second cooling interval in the mid-Eocene (42-38Ma) resulted in the fragmentation of the boreotropics within Eurasia, leading to an African-Asian disjunction. Multiple dispersal events from North into South America occurred from the early Eocene to late Miocene and a single migration event from Asia into Australia appears to have occurred in the early Miocene.

Molecular phylogenetic analysis of the Persea group (Lauraceae) and its biogeographic implications on the evolution of tropical and subtropical Amphi-Pacific disjunctions.

PREMISE OF THE STUDY: The Persea group (Lauraceae) has a tropical and subtropical amphi-pacific disjunct distribution with most of its members, and it includes two Macaronesian species. The relationships within the group are still controversial, and its intercontinental disjunction has not been investigated with extensive sampling and precise time dating. • METHODS: ITS and LEAFY intron II sequences of 78 Persea group species and nine other Lauraceae species were analyzed with maximum parsimony and Bayesian inference. Divergence time estimation employed Bayesian Markov chain Monte Carlo method under a relaxed clock. • KEY RESULTS: Several traditional genera or subgenera within the Persea group form well-supported monophyletic groups except Alseodaphne and Dehaasia. The divergence time of the Persea group is estimated as ∼55.3 (95% higher posterior densities [HPD] 41.4-69.9) million years ago (mya). Two major divergences within the Persea group are estimated as ∼51.9 (95% HPD 38.9-63.9) mya and ∼48.5 (95% HPD 35.9-59.9) mya. • CONCLUSIONS: Persea can be retained as a genus by the inclusion of Apollonias barbujana and exclusion a few species that do not fit into the established subgenera. A major revision is recommended for the delimitation between Alseodaphne, Dehaasia, and Nothaphoebe. We suggest that the Persea group originated from the Perseeae-Laureae radiation in early Eocene Laurasia. Its amphi-pacific disjunction results from the disruption of boreotropical flora by climatic cooling during the mid- to late Eocene. The American-Macaronesian disjunction may be explained by the long-distance dispersal.

Angiosperm phylogeny based on matK sequence information.

Plastid matK gene sequences for 374 genera representing all angiosperm orders and 12 genera of gymnosperms were analyzed using parsimony (MP) and Bayesian inference (BI) approaches. Traditionally, slowly evolving genomic regions have been preferred for deep-level phylogenetic inference in angiosperms. The matK gene evolves approximately three times faster than the widely used plastid genes rbcL and atpB. The MP and BI trees are highly congruent. The robustness of the strict consensus tree supercedes all individual gene analyses and is comparable only to multigene-based phylogenies. Of the 385 nodes resolved, 79% are supported by high jackknife values, averaging 88%. Amborella is sister to the remaining angiosperms, followed by a grade of Nymphaeaceae and Austrobaileyales. Bayesian inference resolves Amborella + Nymphaeaceae as sister to the rest, but with weak (0.42) posterior probability. The MP analysis shows a trichotomy sister to the Austrobaileyales representing eumagnoliids, monocots + Chloranthales, and Ceratophyllum + eudicots. The matK gene produces the highest internal support yet for basal eudicots and, within core eudicots, resolves a crown group comprising Berberidopsidaceae/Aextoxicaceae, Santalales, and Caryophyllales + asterids. Moreover, matK sequences provide good resolution within many angiosperm orders. Combined analyses of matK and other rapidly evolving DNA regions with available multigene data sets have strong potential to enhance resolution and internal support in deep level angiosperm phylogenetics and provide additional insights into angiosperm evolution.