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SUMMARY: The Phyllanthaceae family comprises a diverse range of plants with medicinal, edible, and ornamental value, extensively cultivated worldwide. Polyploid species commonly occur in Phyllanthaceae. Due to the rather complex genomes and evolutionary histories, their speciation process has been still lacking in research. In this study, we generated chromosome‐scale haplotype‐resolved genomes of two octoploid species (Phyllanthus emblica and Sauropus spatulifolius) in Phyllanthaceae family. Combined with our previously reported one tetraploid (Sauropus androgynus) and one diploid species (Phyllanthus cochinchinensis) from the same family, we explored their speciation history. The three polyploid species were all identified as allopolyploids with subgenome A/B. Each of their two distinct subgenome groups from various species was uncovered to independently share a common diploid ancestor (Ancestor‐AA and Ancestor‐BB). Via different evolutionary routes, comprising various scenarios of bifurcating divergence, allopolyploidization (hybrid polyploidization), and autopolyploidization, they finally evolved to the current tetraploid S. androgynus, and octoploid S. spatulifolius and P. emblica, respectively. We further discuss the variations in copy number of alleles and the potential impacts within the two octoploids. In addition, we also investigated the fluctuation of metabolites with medical values and identified the key factor in its biosynthesis process in octoploids species. Our study reconstructed the evolutionary history of these Phyllanthaceae species, highlighting the critical roles of polyploidization and hybridization in their speciation processes. The high‐quality genomes of the two octoploid species provide valuable genomic resources for further research of evolution and functional genomics. Significance Statement: Our study reconstructed the evolutionary history of these Phyllanthaceae species, highlighting the critical roles of polyploidization and hybridization in their speciation processes. [ABSTRACT FROM AUTHOR]

A new species of Emblica Gaertn. (Phyllanthaceae), E. chakrabartyi sp. nov. from the southern part of the Western Ghats, India is described here along with data on distribution, phenology and colour photographs. A key to the species of Emblica in India is also provided. The new species is allied to Emblica anamalayana, but differs in habit, urceolate staminate flowers with short pedicel, inner tepals of staminate flower larger than outer, short anthers which are equal in length to its column, three distinct deeply bifid styles, shape of capsule and its styles and black lineolate smooth seeds. [ABSTRACT FROM AUTHOR]

Introduction: This study describes the molecular profile and the potential antiviral activity of extracts from Phyllanthus brasiliensis, a plant widely found in the Brazilian Amazon. The research aims to shed light on the potential use of this species as a natural antiviral agent. Methods: The extracts were analysed using liquid chromatography–mass spectrometry (LC–MS) system, a potent analytical technique to discover drug candidates. In the meantime, in vitro antiviral assays were performed against Mayaro, Oropouche, Chikungunya, and Zika viruses. In addition, the antiviral activity of annotated compounds was predicted by in silico methods. Results: Overall, 44 compounds were annotated in this study. The results revealed that P. brasiliensis has a high content of fatty acids, flavones, flavan‐3‐ols, and lignans. Furthermore, in vitro assays revealed potent antiviral activity against different arboviruses, especially lignan‐rich extracts against Zika virus (ZIKV), as follows: methanolic extract from bark (MEB) [effective concentration for 50% of the cells (EC50) = 0.80 μg/mL, selectivity index (SI) = 377.59], methanolic extract from the leaf (MEL) (EC50 = 0.84 μg/mL, SI = 297.62), and hydroalcoholic extract from the leaf (HEL) (EC50 = 1.36 μg/mL, SI = 735.29). These results were supported by interesting in silico prediction, where tuberculatin (a lignan) showed a high antiviral activity score. Conclusions: Phyllanthus brasiliensis extracts contain metabolites that could be a new kick‐off point for the discovery of candidates for antiviral drug development, with lignans becoming a promising trend for further virology research. The study explores the antiviral activity of extracts from Phyllanthus brasiliensis against Mayaro, Oropouche, Chikungunya, and Zika viruses. A total of 44 compounds were annotated, including fatty acids, flavones, flavan‐3‐ols, and lignans. The lignan‐rich extracts showed potent antiviral activity, particularly against the Zika virus. In silico antiviral prediction suggested tuberculatin (a lignan) as a strong antiviral candidate. Hence, the study suggests that P. brasiliensis could be a promising source for the development of antiviral drugs, particularly those containing lignans. [ABSTRACT FROM AUTHOR]

The genus Phyllanthus is paraphyletic as currently circumscribed, with the genera Breynia, Glochidion and Synostemon nested within it. A phylogeny based on nuclear (ITS, PHYC) and chloroplast (matK, accD‐psaI, trnS‐trnG) markers is presented, including 18/18 subgenera and 53/70 sections. Differences in habit, branching type, floral and fruit characters are discussed, and we find indications for shifts in pollination and dispersal strategies possibly underlying the convergent evolution of these characters in multiple clades. Several taxonomic issues were found in the subgeneric classification of Phyllanthus that will require new transfers and rank changes. Phyllanthus subg. Anesonemoides, subg. Conami, subg. Emblica, subg. Gomphidium, subg. Kirganelia and subg. Phyllanthus are polyphyletic, and several sections appear to be paraphyletic (e.g., P. sect. Anisonema, sect. Emblicastrum, sect. Pseudoactephila, sect. Swartziani, and sect. Xylophylla); P. subg. Phyllanthodendron is furthermore paraphyletic with the genus Glochidion nested within. To create a classification of tribe Phyllantheae that comprises exclusively monophyletic taxa, it is necessary to treat several clades at the same taxonomic rank as the genera Breynia, Glochidion and Synostemon. Since combining all genera would lead to one giant heterogeneous genus that is difficult to define, we recommend dividing Phyllanthus into several monophyletic genera, which have previously been recognized and often possess diagnostic (combinations of) morphological characters. This new classification is forthcoming. [ABSTRACT FROM AUTHOR]

Abstract: Corilagin is a component of Phyllanthus urinaria extract and has been found of possessing anti‐inflammatory, anti‐oxidative, and anti‐tumour properties in clinic treatments. However, the underlying mechanisms in anti‐cancer particularly of its induction of cell death in human breast cancer remain undefined. Our research found that corilagin‐induced apoptotic and autophagic cell death depending on reactive oxygen species (ROS) in human breast cancer cell, and it occurred in human breast cancer cell (MCF‐7) only comparing with normal cells. The expression of procaspase‐8, procaspase‐3, PARP, Bcl‐2 and procaspase‐9 was down‐regulated while caspase‐8, cleaved PARP, caspase‐9 and Bax were up‐regulated after corilagin treatment, indicating apoptosis mediated by extrinsic and mitochondrial pathways occurred in MCF‐7 cell. Meanwhile, autophagy mediated by suppressing Akt/mTOR/p70S6K pathway was detected with an increase in autophagic vacuoles and LC3‐II conversion. More significantly, inhibition of autophagy by chloroquine diphosphate salt (CQ) remarkably enhanced apoptosis, while the caspase inhibitor z‐VAD‐fmk failed in affecting autophagy, suggesting that corilagin‐induced autophagy functioned as a survival mechanism in MCF‐7 cells. In addition, corilagin induced intracellular reactive oxygen species (ROS) generation, when reduced by ROS scavenger NAC, apoptosis and autophagy were both down‐regulated. Nevertheless, in SK‐BR3 cell which expressed RIP3, necroptosis inhibitor Nec‐1 could not alleviate cell death induced by corilagin, indicating necroptosis was not triggered. Subcutaneous tumour growth in nude mice was attenuated by corilagin, consisting with the results in vitro. These results imply that corilagin inhibits cancer cell proliferation through inducing apoptosis and autophagy which regulated by ROS release. [ABSTRACT FROM AUTHOR]

Premise of the Study: A novel set of EST‐SSR markers was developed for Phyllanthus emblica (Phyllanthaceae) to investigate the genetic structure and gene flow, identify novel genes of interest, and develop markers for assisted breeding. Methods and Results: Based on the transcriptome data of P. emblica, 83 EST‐SSR primer pairs were designed; 52 primer pairs were successfully amplified, with 20 showing polymorphisms in 90 individuals from three populations of P. emblica. The number of alleles per locus varied from 11 to 44. The observed and expected levels of heterozygosity for the 20 loci ranged from 0.240 to 0.868 and 0.754 to 0.933, respectively. Cross‐species amplification was successful for all 20 loci in each of the two related species, P. reticulatus and Leptopus chinensis. Conclusions: These markers will be valuable for studying the population genetics and for mining genes of P. emblica, and may be useful for studies of related species. [ABSTRACT FROM AUTHOR]

We provide a comprehensive wood anatomical survey of 11 Indian species belonging to four genera of the subfamily Antidesmatoideae (Phyllanthaceae) and evaluate their relationships and ecological adaptations in this context. Wood microstructure of the subfamily was found to be highly heterogeneous reflecting the unnatural classification of the subfamily. With some exceptions, our results do not corroborate recent morphological and molecular phylogenetic circumscriptions. The wood anatomy of Aporosa and Baccaurea was remarkably similar and they lack distinguishing wood anatomical characters. Antidesma showed adaptations to a xeric environment and Aporosa and Baccaurea showed adaptations to mesic environments. It is concluded that from a wood anatomical point of view, Antidesma and Bischofia are advanced while Aporosa and Baccaurea are primitive genera in the subfamily. [ABSTRACT FROM AUTHOR]

Aim The dated phylogeny of Breynia (including Sauropus) was used to analyse historical biogeographical patterns and to compare the speciation events with tectonic occurrences and biotic and abiotic aspects. Location Southeast Asia, Malesia and Australia. Methods Existing molecular markers ( trnS- trnG, accD- psaI, PHYC and ITS) were used in beast to make a Bayesian age estimate with an exponential relaxed molecular clock. The resulting cladograms and maximum clade credibility ( MCC) tree were used as input in statistical dispersal-vicariance analysis (S- DIVA option in the rasp package). Results Breynia originated 25-20 Ma in Thailand (western part of northern to southern Thailand) up to peninsular Malaysia. Dispersal occurred in various directions (India+Sri Lanka, China+Indochina, Malesia+Australia) establishing widespread species. Most species evolved locally in the area of origin and a relatively high number of species form limestone endemics. Main conclusions There is little correlation between tectonic movements and dispersal patterns in Breynia. Only the dispersal over Wallace's Line occurred when stepping stones were in place. Speciation, especially in the area of origin seems to be linked to soil (limestone endemics) and pollination, as indicated by the presence of deviating inflorescences in B. subgen. Sauropus (no pollinators are known yet) and the obligate moth pollination in B. sect. Breynia. Moreover, B. sect. Breynia and three widespread species of B. subgen. Sauropus have (more) fleshy fruits and all these taxa generally have wide distributions, probably because of bird dispersal. [ABSTRACT FROM AUTHOR]

The globally distributed family Phyllanthaceae comprises 1745 species, all of which produce unisexual flowers, and most of the species occur in the tropics. The few species in which the pollination system has been studied are pollinated by flies, wasps, or specialized moths, and at least two species are duodichogamous. Here we report on the flowering phenology and pollination of the Chinese species Bridelia stipularis and Cleistanthus sumatranus, which were studied on the tropical Hainan Island. Both species are duodichogamous and indicated to be fly-pollinated. The calliphorid species Chrysomya megacephala, Isomyia isomyia, Pierretia sp., Hemipyrellia sp., and Achoetandrus tufifacies were the most frequent flower visitors to B. stipularis; some of these species together with syrphids ( Meliscaeva cinctella, Dideopsis sp., and Eristalis sp.) were the most abundant visitors to Cl. sumatranus flowers . Fruit set in hand-pollinated flowers was higher than in open-pollinated flowers, but because of natural fruit abortion the difference was not significant, indicating that fly pollinators do not limit reproduction of these Phyllanthaceae species. This study provides an addition to the limited Asian literature on pollination by flies and the even more limited literature on duodichogamy. It could help to broaden our understanding for analysis of the evolution of duodichogamy in future research, but the specific ecological factors behind the evolution of this rare sexual system are still difficult to generalize. [ABSTRACT FROM AUTHOR]

•Obligate mutualisms involving actively pollinating seed predators are among the most remarkable insect–plant relationships known, yet almost nothing is known about the chemistry of pollinator attraction in these systems. The extreme species specificity observed in these mutualisms may be maintained by specific chemical compounds through ‘private channels’. Here, we tested this hypothesis using the monoecious Breynia vitis-idaea and its host-specific Epicephala pollinator as a model. •Headspace samples were collected from both male and female flowers of the host. Gas chromatography with electroantennographic detection (GC-EAD), coupled gas chromatography–mass spectrometry, and olfactometer bioassays were used to identify the floral compounds acting as the pollinator attractant. •Male and female flowers of B. vitis-idaea produced similar sets of general floral compounds, but in different ratios, and male flowers emitted significantly more scent than female flowers. A mixture of 2-phenylethyl alcohol and 2-phenylacetonitrile, the two most abundant compounds in male flowers, was as attractive to female moths as the male flower sample, although the individual compounds were slightly less attractive when tested separately. •Data on the floral scent signals mediating obligate mutualisms involving active pollination are still very limited. We show that system-specific chemistry is not necessary for efficient host location by exclusive pollinators in these tightly coevolved mutualisms. [ABSTRACT FROM AUTHOR]

The landmark discovery of obligate pollination mutualism between Glochidion plants and Epicephala moths has sparked increased interest in the pollination systems of Phyllantheae plants. In this paper I review current information on the natural history and evolutionary history of obligate pollination mutualism in Phyllantheae. Currently, an estimated >500 species are mutualistic with Epicephala moths that actively pollinate flowers and whose progeny feed on the resulting seeds. The Phyllantheae also includes species that are not mutualistic with Epicephala moths and are instead pollinated by bees and/or flies or ants. Phylogenetic analyses indicate that the mutualism evolved independently five times within Phyllantheae, whereas active pollination behavior, a key innovation in this mutualism, evolved once in Epicephala. Reversal of mutualism has occurred at least once in both partner lineages, involving a Breynia species that evolved an alternative pollination system and a derived clade of Epicephala that colonized ant-pollinated Phyllantheae hosts and thereby lost the pollinating habit. The plant–moth association is highly species specific, although a strict one-to-one assumption is not perfectly met. A comparison of plant and moth phylogenies suggests signs of parallel speciation, but partner switches have occurred repeatedly at a range of taxonomic levels. Overall, the remarkable species diversity and multiple originations of the mutualism provide excellent opportunities to address many important questions on mutualism and the coevolutionary process. Although research on the biology of the mutualism is still in its infancy, the Phyllantheae– Epicephala association holds promise as a new model system in ecology and evolutionary biology. [ABSTRACT FROM AUTHOR]

Novel insights into the evolutionary history of a taxonomically complex tropical plant group were gained in this study using DNA sequence data. A molecular phylogenetic analysis of the newly circumscribed and expanded tribe Poranthereae (Phyllanthaceae) is presented. Sampling included 97 accessions for 63 of c. 120 species. Largely congruent results have been obtained from nuclear ribosomal ITS and plastid matK sequences. These analyses support the recognition of Andrachne and Leptopus as distinct genera. The deceptively similar Andrachne section Phyllanthopsis, Andrachne ovalis, and Leptopus decaisnei are separate lineages to be segregated. Zimmermannia and Zimmermanniopsis are embedded in Meineckia; Oreoporanthera is embedded in Poranthera; and Archileptopus is embedded in Leptopus. Andrachne section Pseudophyllanthus is polyphyletic, the two Madagascan endemics emerging as a sister dade to Meineckia. The noncontiguous distributions of Andrachne sensu lato and Leptopus sensu lato were found to be the result of separate evolutionary histories of morphologically similar clades, whereas Andrachne sensu stricto and Meineckia remain geographically disjunct. Actephila and Leptopus are sisters with a sympatric distribution in humid Asia. Presence of petals appears to be plesiomorphic for the tribe. Petals are reduced in Actephila and Oreoporanthera and lost in the Meineckia dade. [ABSTRACT FROM AUTHOR]

Flowering plants commonly separate male and female function in time, but rarely are the two stages synchronized within and among individuals. One such temporal mating system is duodichogamy in which each plant produces two batches of male flowers that are temporally separated by a batch of female flowers, with within-individual synchrony and among-individual asynchrony to ensure mating partners. Duodichogamy is known only from a few species in four genera in unrelated families. We report on duodichogamy in the Chinese tree species Bridelia tomentosa (Phyllanthaceae), a common colonizer of disturbed habitats. In three populations monitored over 2 yr, most trees flowered in the order male → female → male, and resting periods between flowering bouts precluded selfing almost completely. Individuals flowered for several weeks, with the onset of flowering slightly asynchronous among trees. Pollination was by flies, and experimental pollen supplementation of a subset of a tree's flowers did not increase fruit set, suggesting high levels of insect visitation and possible resource limitation. Nineteen percent of the 166 trees monitored skipped the first male phase, and another 13% skipped the female phase, remaining male in both years (and also a third year). The regular presence of pure males, if genetically fixed, would make B. tomentosa androdioecious in addition to duodichogamous. Comparison of duodichogamous taxa known so far shows that all have few ovules, fitting with the hypothesis that duodichogamy may result from male competition for access to a small supply of ovules. [ABSTRACT FROM AUTHOR]

The obligate mutualisms between flowering plants and their seed-parasitic pollinators constitute fascinating examples of interspecific mutualisms, which are often characterized by high levels of species diversity and reciprocal species specificity. The diversification in these mutualisms has been thought to occur through simultaneous speciation of the partners, mediated by tight reciprocal adaptation; however, recent studies cast doubt over this general view. In this study, we examine the diversity and species specificity of Epicephala moths (Gracillariidae) that pollinate Glochidion trees (Phyllanthaceae), using analysis of mitochondrial and nuclear gene sequences. Phylogenetic analysis of Epicephala moths associated with five Glochidion species in Japan and Taiwan reveal six genetically isolated species that are also distinguishable by male genital morphology: (i) two species specific to single host species ( G. acuminatum and G. zeylanicum, respectively); (ii) two species that coexist on G. lanceolatum; and (iii) two species that share two, closely-related parapatric hosts ( G. obovatum and G. rubrum). Statistical analysis shows that the two species associated with G. lanceolatum are not sister species, indicating the colonization of novel Glochidion host in at least one lineage. Behavioural observations suggest that all six species possess the actively-pollinating habit, thus none of the studied species has become a nonmutualistic ‘cheater’ that exploits the benefit resulting from pollination by other species. Our results parallel recent findings in ecologically similar associations, namely the fig–fig wasp and yucca–yucca moth mutualisms, and contribute to a more general understanding of the factors that determine ecological and evolutionary outcomes in these mutualisms. [ABSTRACT FROM AUTHOR]

Phylogenetic relationships within tribe Phyllantheae, the largest tribe of the family Phyllanthaceae, were examined with special emphasis on the large genus Phyllanthus. Nuclear ribosomal ITS and plastid matK DNA sequence data for 95 species of tribe Phyllantheae, including representatives of all subgenera of Phyllanthus (except Cyclanthera) and several hitherto unplaced infrageneric groups, were analyzed. Results for ITS and matK are generally concordant, although some species are placed differently in the plastid and ITS trees, indicating that hybridization/paralogy is involved. Results confirm paraphyly of Phyllanthus in its traditional circumscription with embedded Breynia, Glochidion, Reverchonia, and Sauropus. We favor the inclusion of the embedded taxa in Phyllanthus over further generic segregation. Monophyletic Phyllanthus comprises an estimated 1269 species, making it one of the ‘giant’ genera. Phyllanthus maderaspatensis is sister to all other species of Phyllanthus, and the genus appears to be of paleotropical origin. Subgenera lsocladus, Kirganelia, and Phyllanthus arc polyphyletic, whereas other subgenera appear to be monophyletic. Monotypic Reverchonia is sister to P. abnormis, arborescent section Emblica to herbaceous Urinaria, free-floating aquatic P. fluitans to the weed P. caroliniensis, and the phyllocladous section Choretropsis to the delicate leafy P. clattssenii. The unique branching architecture known as ‘phyllanthoid branching’ found in most Phyllanthtls taxa has been lost (and/or has been derived) repeatedly. Taxonomic divisions within Phyllantheae based on similar pollen morphology are confirmed, and related taxa share similar distributions. We recommend recognition of six clades at generic level: Flueggea s.l. (including Richeriella), Lingelsheimia, Margaritaria, Phyllanthus s.l. (including Breynia, Glochidion, Reverchonia, and Sauropus), P. diandrus, and Savia section Heterosavia. [ABSTRACT FROM AUTHOR]

Plastid matK and a fragment of the low-copy nuclear gene PHYC were sequenced for 30 genera of Phyllanthaceae to evaluate tribal and generic delimitation. Resolution and bootstrap percentages obtained with matK are higher than that of PHYC, but both regions show nearly identical phylogenetic patterns. Phylogenetic relationships inferred from the independent and combined data are congruent and differ from previous, morphology-based classifications but are highly concordant with those of the plastid gene rbcL previously published. Phyllanthaceae is monophyletic and gives rise to two well-resolved clades (1 and F) that could be recognized as subfamilies. DNA sequence data for Keayodendron and Zimmennanniopsis are presented for the first time. Keayodendron is misplaced in tribe Phyllantheae and belongs to the Bridelia alliance. Zimmerinanniopsis is sister to Zimmennannia. Phylianthus and Cleistanthus are paraphyletic. Savia and Phyllanthus subgenus Kirganelia are not monophyletic. [ABSTRACT FROM AUTHOR]

This paper reports obligate seed-parasitic pollination mutualisms in Breynia vitis-idea and B. fruticosa (Phyllanthaceae). The genus Breynia is closely related to Glochidion and Gomphidium (a subgenus of Phyllanthus), in which pollination by species-specific, seed-parasitic Epicephala moths (Gracillariidae) have been previously reported. At night, female Epicephala moths carrying numerous pollen grains on their proboscises visited female flowers of B. vitis-idea, actively pollinated flowers, and each subsequently laid an egg. Examination of field-collected flowers indicated that pollinated flowers of B. vitis-idea and B. fruticosa almost invariably had Epicephala eggs, suggesting that these moths are the primary pollinators of the two species. Single Epicephala larvae consumed a fraction of seeds within developing fruit in B. vitis-idea and all seeds in B. fruticosa. However, some of the fruits were left untouched, and many of these had indication of moth oviposition, suggesting that egg/larval mortality of Epicephala moths is an important factor assuring seed set in these plants. The overall similarity of the specialized floral structure among Breynia species may indicate that this pollination system is fairly widespread within the genus. [ABSTRACT FROM AUTHOR]