<Emphasis Type="Italic">Elaphoglossum mickeliorum</Emphasis> (Dryopteridaceae), a new species of <Emphasis Type="Italic">Elaphoglossum</Emphasis> sect. <Emphasis Type="Italic">Polytrichia</Emphasis> from Peru

Elaphoglossum mickeliorum, a new species from the eastern slopes of the Peruvian Andes, is here described and illustrated. It belongs to E. sect. Polytrichia, which is characterized by the presence of subulate scales and absence of hydathodes on the sterile leaves of adult sporophytes. Herbarium specimens of this new species were first collected by Alwyn H. Gentry ca. 40 years ago, but these got readily confused with E. erinaceum and went undescribed since then. The new species differs from members of the E. erinaceum complex by having a nearly continuous band of planar, nonsubulate scales along the laminar margins of sterile leaves. Based on this character, E. mickeliorum resembles species such as E. glaziovii, E. ornatum, and E. scolopendrifolium. It differs from these by the presence of minute glandular hairs on petioles and costae. A distribution map and a figure with line drawings are also provided. For comparative purposes, the line drawing includes E. blepharoglottis, which is here illustrated for the first time.     

The effects of the alkaloid scopolamine on the performance and behavior of two caterpillar species

Plants have evolved many defenses against insect herbivores, including numerous chemicals that can reduce herbivore growth, performance, and fitness. One group of chemicals, the tropane alkaloids, is commonly found in the nightshade family (Solanaceae) and has been thought to reduce performance and fitness in insects. We examined the effects of the tropane alkaloid scopolamine, an alkaloid constituent of Datura wrightii, which is the most frequent host plant for the abundant and widespread insect herbivore Manduca sexta in the southwestern United States. We exposed caterpillars of two different species to scopolamine: M. sexta, which has a shared evolutionary history with Datura and other solanaceous plants, and Galleria mellonella, which does not. We showed that the addition of ecologically realistic levels of scopolamine to both the diet and the hemolymph of these two caterpillar species (M. sexta and G. mellonella) had no effect on the growth of either species. We also showed that M. sexta has no behavioral preference for or against scopolamine incorporated into an artificial diet. These results are contrary to other work showing marked differences in performance for other insect species when exposed to scopolamine, and provide evidence that scopolamine might not provide the broad-spectrum herbivore resistance typically attributed to it. It also helps to clarify the coevolutionary relationship between M. sexta and one of its main host plants, as well as the physiological mechanism of resistance against scopolamine.     

<Emphasis Type="Italic">Pseudomonas</Emphasis> spp. increases root biomass and tropane alkaloid yields in transgenic hairy roots of <Emphasis Type="Italic">Datura</Emphasis> spp.

Transgenic hairy roots of Datura spp., established using strain A4 of Agrobacterium rhizogenes, are genetically stable and produce high levels of tropane alkaloids. To increase biomass and tropane alkaloid content of this plant tissue, four Pseudomonas strains, Pseudomonas fluorescens P64, P66, C7R12, and Pseudomonas putida PP01 were assayed as biotic elicitors on transgenic hairy roots of Datura stramonium, Datura tatula, and Datura innoxia. Alkaloids were extracted from dried biomass, and hyoscyamine and scopolamine were quantified using liquid chromatography-tandem mass spectrometry analysis. D. stramonium and D. innoxia biomass production was stimulated by all Pseudomonas spp. strains after a 5-d treatment. All strains of P. fluorescens increased hyoscyamine yields compared to untreated cultures after both 5 and 10 d of treatment. Hyoscyamine yields were highest in D. tatula cultures exposed to a 5-d treatment with C7R12 (16.633 + 0.456 mg g^?1 dry weight, a 431% increase) although the highest yield increases compared to the control were observed in D. stramonium cultures exposed to strains P64 (511% increase) and C7R12 (583% increase) for 10 d. D. innoxia showed the highest scopolamine yields after elicitation with P. fluorescens strains P64 for 5 d (0.653 + 0.021 mg g^?1 dry weight, a 265% increase) and P66 for 5 and 10 d (5 d, 0.754 + 0.0.031 mg g^?1 dry weight, a 321% increase; 10 d 0.634 + 0.046 mg g^?1 dry weight, a 277% increase). These results show that the Pseudomonas strains studied here can positively and significantly affect biomass and the yields of hyoscyamine and scopolamine from transgenic roots of the three Datura species.     

Development and characterization of 24 chloroplast microsatellite markers for two species of <Emphasis Type="Italic">Eranthis</Emphasis> (Ranunculaceae)

Chloroplast microsatellites for two Korean endemic species, Eranthis byunsanensis and E. pungdoensis (Ranunculaceae), were isolated to address the questions of their distributional patterns and evolutionary relationships, using next-generation sequencing. Twenty-four polymorphic chloroplast microsatellite markers for these two species were developed, and then characterized in 65 individuals (55 individuals of E. byunsanensis and 10 individuals of E. pungdoensis). The number of alleles per locus ranged from 2 to 9; the average number of alleles across all the loci scored 4.792. The unbiased diversity per locus ranged from 0.089 to 0.880; the unbiased diversity averaged over all the loci was 0.646. The developed markers were successfully amplified for three congeneric species, E. stellata, E. pinnatifida, and E. longistipitata. The markers developed in this study can provide a valuable and important tool for understanding genetic variations, population structures, evolutionary histories and phylogeography of E. byunsanensis, E. pungdoensis, and related species.     

Phytochemical and pharmacological investigation of <Emphasis Type="Italic">Spiraea chamaedryfolia</Emphasis>: a contribution to the chemotaxonomy of <Emphasis Type="Italic">Spiraea</Emphasis> genus

Objective

Diterpene alkaloids are secondary plant metabolites and chemotaxonomical markers with a strong biological activity. These compounds are characteristic for the Ranunculaceae family, while their occurrence in other taxa is rare. Several species of the Spiraea genus (Rosaceae) are examples of this rarity. Screening Spiraea species for alkaloid content is a chemotaxonomical approach to clarify the classification and phylogeny of the genus. Novel pharmacological findings make further investigations of Spiraea diterpene alkaloids promising.

Results

Seven Spiraea species were screened for diterpene alkaloids. Phytochemical and pharmacological investigations were performed on Spiraea chamaedryfolia, the species found to contain diterpene alkaloids. Its alkaloid-rich fractions were found to exert a remarkable xanthine-oxidase inhibitory activity and a moderate antibacterial activity. The alkaloid distribution within the root was clarified by microscopic techniques.

     

<Emphasis Type="BoldItalic">Barrmaelia</Emphasis> and <Emphasis Type="BoldItalic">Entosordaria</Emphasis> in Barrmaeliaceae (fam. nov., Xylariales) and critical notes on <Emphasis Type="BoldItalic">Anthostomella</Emphasis>-like genera based on multigene phylogenies

Phylogenetic analyses of a combined DNA data matrix containing ITS, LSU, rpb2 and tub2 sequences of representative Xylariales revealed that the genus Barrmaelia is a well-defined monophylum, as based on four of its described species (B. macrospora, B. moravica, B. oxyacanthae, B. rhamnicola) and the new species B. rappazii. The generic type of Entosordaria, E. perfidiosa, is revealed as the closest relative of Barrmaelia, being phylogenetically distant from the generic type of Clypeosphaeria, C. mamillana, which belongs to Xylariaceae sensu stricto. Entosordaria and Barrmaelia are highly supported and form a distinct lineage, which is recognised as the new family Barrmaeliaceae. The new species E. quercina is described. Barrmaelia macrospora, B. moravica and B. rhamnicola are epitypified and E. perfidiosa is lecto- and epitypified. Published sequences of Anthostomella and several Anthostomella-like species from the genera Alloanthostomella, Anthostomelloides, Neoanthostomella, Pseudoanthostomella and Pyriformiascoma are evaluated, demonstrating the necessity of critical inspection of published sequence data before inclusion in phylogenies. Verified isolates of several species from these genera should be re-sequenced to affirm their phylogenetic affinities. In addition, the generic type of Anthostomella should be sequenced before additional generic re-arrangements are proposed.     

Three New Species of <Emphasis Type="Italic">Eustomias</Emphasis> (Melanostomiidae) From the Southern Subtropical Waters with the Description of a New Subgenus

Eustomias diplomastiga (28° S, 84° W), E. mavka (36° S, 121° W), and E. kukuevi (23° S, 12° E) spp. n. are described from the southern subtropical waters of Atlantic and Pacific oceans. For the species E. diplomastiga sp. n. and E. crucis, a new subgenus Furcostomias subgen. n. is established. It is different from other subgenera based on the following features: stem of the chin barbel bifurcate at the middle, completely deprived of bulbs, with a pigmented shaft but without the external pigmentation of the stem and branches and without the spots associated with the photophores, the presence of a short ventral furrow, P 2, V 7. A review of the diagnostic features of the subgenera of Eustomias is given.     

Morphology of Cell Wall Pore Channels in the Genus <Emphasis Type="Italic">Euastrum</Emphasis> Ralfs (Desmidiales)

The study of the cell wall structure was performed for ten species of Euastrum: E. ansatum (Ehrenb.) Ralfs, E. bidentatum Näg., E. binale (Turp.) Ehrenb. ex Ralfs, E. dubium Näg., E. elegans (Bréb.) Kütz. ex Ralfs, E. germanicum (Schmidle) W. Krieger, E. oblongum (Grev.) Ralfs ex Ralfs, E. pectinatum Bréb. ex Bréb. in Ralfs, E. validum West et G.S.West, E. verrucosum (Ehrenb.) ex Ralfs. The investigation of the cell wall ultrastructure has established for the first time that the pore canal in 6 of 10 species always has a coiled form to some degree. Three species (E. germanicum, E. pectinatum, and E. verrucosum) have both coiled and straight canals, and E. ansatum has only straight ones. A new type of pores (P7), typical only for representatives of the genus Euastrum, was also noted. In addition, a simple and effective method of preparing desmidium algal cells for investigation with transmission electron microscope has been developed.     

Identification of 21 novel <Emphasis Type="Italic">S-RNase</Emphasis> alleles and determination of <Emphasis Type="Italic">S</Emphasis>-genotypes in 66 loquat (<Emphasis Type="Italic">Eriobotrya</Emphasis>) accessions

As observed in other self-incompatible species in the Pyrinae subtribe, loquat (Eriobotrya japonica) demonstrates gametophytic self-incompatibility that is controlled by the S-locus, which encodes a polymorphic stylar ribonuclease (S-RNase). This allows the female reproductive organ (style) to recognize and reject the pollen from individuals with the same S-alleles, but allows the pollen from individuals with different S-alleles to effect fertilization. The S-genotype is therefore an important consideration in breeding strategies and orchard management. In an attempt to optimize the selection of parental lines in loquat production, the S-RNase alleles of 35 loquat cultivars and their 26 progeny, as well as five wild loquat species, were identified and characterized in this study. The best pollinizer cultivar combinations were also explored. A total of 28 S-alleles were detected, 21 of which constituted novel S-RNase alleles. The S-haplotypes S₂ and S₆ were the most frequent, followed by S ₂₉ , S ₃₁ , S ₅ , S ₂₄ , S ₂₈ , S ₃₃ , S ₃₄ , S ₃₂ , and S ₁₅ , while the rare alleles S ₁ , S ₉ , S ₁₄ , S ₁₆ , S ₁₇ , S ₁₈ , S ₁₉ , S ₂₀ , S ₂₁ , S ₂₂ , S ₂₃ , S ₂₇ , and S ₃₅ were only observed in one of the accessions tested. Moreover, the S-genotypes of five wild loquat species (E. prinoides, E. bengalensis, E. prinoides var. dadunensis, E. deflexa, and E. japonica) are reported here for the first time. The results will not only facilitate the selection of suitable pollinators for optimal orchard management, but could also encourage the crossbreeding of wild loquat species to enhance the genetic diversity of loquat cultivars.     

Age-based demography and reproductive biology of three <Emphasis Type="Italic">Epinephelus</Emphasis> groupers, <Emphasis Type="Italic">E. polyphekadion,E. tauvina,</Emphasis> and <Emphasis Type="Italic">E. howlandi</Emphasis> (Serranidae), inhabiting coral reefs in Okinawa

The demography and reproductive biology of three Epinephelus groupers (Serranidae), namely E. polyphekadion, E. tauvina, and E. howlandi in the Yaeyama Islands, Okinawa, were examined based on age assessment using otoliths and gonadal histology. The maximum ages for these three species were 26 year, 23 year, and 17 year. The von Bertalanffy growth functions were also determined for each species. The size and age at 50% female maturity were estimated to be 358 mm in total length (TL) and 6.0 year for E. polyphekadion, 371 mm TL and 6.7 year for E. tauvina, and 327 mm TL and 4.1 year for E. howlandi, respectively. Significant differences between the sexes in size and age frequencies were found in all three species, with males being larger and older than females, or transitional individuals. These results strongly indicated that the population of these three grouper species showed monandric protogynous hermaphroditism. The sex ratios of E. polyphekadion and E. tauvina were biased in favor of females, but that of E. howlandi was equivalent between sexes. The relative sizes of ripe testes indicated that the intensity of sperm competition varied among species suggesting different mating system of each species. Reproductive seasonality was similar among species, with active vitellogenesis coinciding with the annual rise in water temperature. The active spawning period was determined to be between April and May for E polyphekadion, in May for E. howlandi, and from March to June for E. tauvina.     

Specific features of ecology of chars of the genus <Emphasis Type="Italic">Salvelinus</Emphasis> (Salmonidae) from the basin of Lake Kronotskoe (Kamchatka) according to parasitological data

A parasitological study was performed of chars of the genus Salvelinus inhabiting Lake Kronotskoe (Kamchatka Peninsula)—S. malma, S. albus, S. schmidtii, and S. kronocius, as well as of juvenile Salvelinus spp. Twenty-three species of parasites, including six species new for the lake, Hennequya zschokkei, Protteocephalus longicollis, Diphyllobothrium dendriticum, Crepidostomum sp., Echinorhynchus salmonis, and Paracanthobdella livanowi, were found. With consideration of published data, in chars of this water body, 28 species of parasites were recorded, including seven species (N. cf. pungitius, B. luciopercae, Crepidostomum sp., Cr. fausti, Cr. cf. cooperi, Eubothrium crassium, and Proteocephalus sp.), whose presence or species identification in the lake ecosystem need confirmation. Two species (N. rutili and Diphyllobothrium sp.) are removed from the list. Parasites common for all species of chars were revealed. They include Myxobolus arcticus, E. salvelini, D. ditretum, Crepidostotum sp., Cr. farionis, Cr. metoecus, Cystidicola farionis, Cucullanus truttae, Philonema oncorhynchi, and Salmincola carpionis. Cluster analysis of the fauna of parasites of different species of chars demonstrated considerable differences in infestation, which indicates differences between them in preference for food items and occupied biotopes and thereby supports the ecological differentiation of chars in the basin of Lake Kronotskoe. S. albus and S. kronocius are most similar in parasitofauna, which is determined by their predation; S. malma as a benthophage is infected by the same species of parasites, but considerably less intensively. Extremely high indices of population numbers of some parasite species are considered as a manifestation of the Krebs cycle in parasites under the conditions of an isolated lake.     

The complete chloroplast genome sequence of <Emphasis Type="Italic">Pseudoroegneria libanotica</Emphasis>, genomic features,and phylogenetic relationship with <Emphasis Type="Italic">Triticeae</Emphasis> species

Pseudoroegneria libanotica is an important herbage diploid species possessing the St genome. The St genome participates in the formation of nine perennial genera in Triticeae (Poaceae). The whole chloroplast (cp) genome of P. libanotica is 135 026 bp in length. The typical quadripartite structure consists of one large single copy of 80 634 bp, one small single copy of 12 766 bp and a pair of inverted regions (20 813 bp each). The cp genome contains 76 coding genes, four ribosomal RNA and 30 transfer RNA genes. Comparative sequence analysis suggested that: 1) the 737 bp deletion in the cp of P. libanotica was specific in Triticeae species and might transfer into its nuclear genome; 2) hot-spot regions, indels in intergenic regions and protein coding sequences mainly led to the length variation in Triticeae; 3) highly divergence regions combined with negative selection in rpl2, rps12, ccsA, rps8, ndhH, petD, ndhK, psbM, rps3, rps18, and ndhA were identified as effective molecular markers and could be considered in future phylogenetic studies of Triticeae species; and 4) ycf3 gene with rich cpSSRs was suitable for phylogeny analysis or could be used for DNA barcoding at low taxonomic levels. The cpSSRs distribution in the coding regions of diploid Triticeae species was shown for the first time and provided a valuable source for developing primers to study specific simple sequence repeat loci.     

Cytogenetic evidences on the evolutionary relationships between the tetraploids of the section <Emphasis Type="Italic">Rhizomatosae</Emphasis> and related diploid species (<Emphasis Type="Italic">Arachis</Emphasis>, Leguminosae)

Rhizomatosae is a taxonomic section of the South American genus Arachis, whose diagnostic character is the presence of rhizomes in all its species. This section is of particular evolutionary interest because it has three polyploid (A. pseudovillosa, A. nitida and A. glabrata, 2n?=?4x?=?40) and only one diploid (A. burkartii, 2n?=?2x?=?20) species. The phylogenetic relationships of these species as well as the polyploidy nature and the origin of the tetraploids are still controversial. The present study provides an exhaustive analysis of the karyotypes of all rhizomatous species and six closely related diploid species of the sections Erectoides and Procumbentes by cytogenetic mapping of DAPI/CMA heterochromatin bands and 5S and 18–26S rDNA loci. Chromosome banding showed variation in the DAPI heterochromatin distribution pattern, which, together with the number and distribution of rDNA loci, allowed the characterization of all species studied here. The bulk of chromosomal markers suggest that the three rhizomatous tetraploid species constitute a natural group and may have at least one common diploid ancestor. The cytogenetic data of the diploid species analyzed evidenced that the only rhizomatous diploid species—A. burkartii—has a karyotype pattern different from those of the rhizomatous tetraploids, showing that it is not likely the genome donor of the tetraploids and the non-monophyletic nature of the section Rhizomatosae. Thus, the tetraploid species should be excluded from the R genome, which should remain exclusively for A. burkartii. Instead, the karyotype features of these tetraploids are compatible with those of different species of the sections Erectoides and Procumbentes (E genome species), suggesting the hypothesis of multiple origins of these tetraploids. In addition, the polyploid nature and the group of diploid species closer to the tetraploids are discussed.     

The diversity of rhizobia nodulating the <Emphasis Type="Italic">Medicago</Emphasis>, <Emphasis Type="Italic">Melilotus</Emphasis> and <Emphasis Type="Italic">Trigonella</Emphasis> inoculation group in Egypt is marked by the dominance of two genetic types

Twenty four rhizobial strains were isolated from root nodules of Melilotus, Medicago and Trigonella plants growing wild in soils throughout Egypt. The nearly complete 16S rRNA gene sequence from each strain showed that 12 strains (50 %) were closely related to the Ensifer meliloti LMG6133^T type strain with identity values higher than 99.0 %, that 9 (37.5 %) strains were more than 99 % identical to the E. medicae WSM419^T type strain, and that 3 (12.5 %) strains showed 100 % identity with the type strain of N. huautlense S02^T. Accordingly, the diversity of rhizobial strains nodulating wild Melilotus, Medicago and Trigonella species in Egypt is marked by predominance of two genetic types, E. meliloti and E. medicae, although the frequency of isolation was slightly higher in E. meliloti. Sequencing of the symbiotic nodC gene from selected Medicago and Melilotus strains revealed that they were all similar to those of the E. meliloti LMG6133^T and E. medicae WSM419^T type strains, respectively. Similarly, nodC sequences of strains identified as members of the genus Neorhizobium were more than 99 % identical to that of N. galegae symbiovar officinalis HAMBI 114.     

New species of the buprestid genus <Emphasis Type="Italic">Endelus</Emphasis> Deyrolle (Coleoptera,Buprestidae) from China,India, and Laos

Endelus (Kubaniellus) indicus sp. n. from India, E. (K.) lao sp. n. and E. (K.) khnzoriani sp. n. from Laos, E. (s. str.) sausai sp. n. from China, and E. (s. str.) dembickyi sp. n. from India are described, the two latter species are included in the Endelus bicarinatus Théry, 1932 species-group recently established by the author. E. collinus Obenberger, 1922 is included in this group; lectotype of this species is designated. Keys to species of the subgenus Kubaniellus and of the E. collinus group are provided. E. (K.) kareni Kalashian is for the first time recorded for Shaanxi Prov., E. pacholatkoi Kalashian, E. smaragdinus Desc. et Vill., and E collinus Obenb., for Laos (the latter species, also for Myanmar).     

Conspecificity of <Emphasis Type="Italic">Endothia eugeniae</Emphasis> and <Emphasis Type="Italic">Cryphonectria cubensis</Emphasis>: a re-evaluation based on morphology and DNA sequence data

Cryphonectria cubensis and Endothia eugeniae are fungal pathogens of Eucalyptus and clove that were reduced to synonymy on the basis of results of cross-inoculation studies, isozyme analysis, cultural studies, and morphology. A previous phylogenetic study on Cryphonectria, based on sequence variation in the ITS region of the ribosomal RNA operon, also supported the conspecificity of C. cubensis and E. eugeniae, but was based on only one E. eugeniae isolate. New collections from clove in Brazil and Indonesia have become available, providing the opportunity to reconsider the conspecificity of C. cubensis and E. eugeniae. The occurrence of C. cubensis on clove was confirmed based on morphological comparisons and phylogenetic analyses of ribosomal DNA and β-tubulin gene sequence data. In addition to C. cubensis, other fungi morphologically similar to Cryphonectria species on the basis of their orange stromata were present on some clove specimens, but no isolates were available for these fungi. Furthermore, some isolates, for which no herbarium material exists, grouped separately from the C. cubensis clade and closer to the Cryphonectria clade. The presence of more than one closely related fungus on clove raises questions relating to the legitimacy of the synonymy of E. eugeniae and C. cubensis. Based on the presence of C. cubensis on the type specimen of E. eugeniae, we recognize the synonymy of the two fungi but provide evidence that other fungi, more closely related to Cryphonectria spp. than to C. cubensis, are present on clove.     

Interactions of Algae within the Community of <Emphasis Type="Italic">Gracilaria gracilis</Emphasis> (Rhodophyta)

Interactions among the unattached red alga Gracilaria gracilis, the dominant species of an algal community, and associated algal species Chaetomorpha linum, Enteromorpha prolifera f. prolifera, and Polysiphonia sp. were studied during and after an algal bloom. It was shown that during their bloom the associated algae Enteromorpha and Polysiphonia sp. significantly decreased the photosynthetic rate of G. gracilis but did not affect its growth rate. It is suggested that the inhibition of Gracilaria gracilis photosynthesis is connected to the impact of extracellular metabolites excreted by Chaetomorpha linum, Enteromorpha prolifera f. prolifera, and Polysiphonia sp. In laboratory experiments, the photosynthetic rate of the associated species was enhanced in the presence of Gracilaria. However, no significant alterations were observed in the content of chlorophyll a, growth, and the dark respiration rates of associated algae when they were kept together with Gracilaria. It was suggested that allelopathic interactions observed among algal species during the formation of the monospecific Gracilaria community, as well as during algal blooms, are not determinative.     

Molecular analyses of <Emphasis Type="Italic">Ficus erecta</Emphasis> and its allies within the subsection <Emphasis Type="Italic">Frutescentiae</Emphasis> (Moraceae)

Ficus (Moraceae) is a keystone group in tropical and subtropical forests with remarkable diversity of species and taxonomical challenges as a consequence of fig–pollinator coevolution. Ficus subsect. Frutescentiae includes about 30 species that are predominantly shrubs or small trees with Terminalia branching. Many of these species are difficult to delimit morphologically, and the group includes a tangle of uncertain taxa and incorrectly applied names. We conducted a phylogenetic analysis with internal and external transcribed spacer data (ITS and ETS) and data from 18 polymorphic microsatellite loci to evaluate the species status of the most perplexing members of this subsection. The results confirm the monophyly of subsect. Frutescentiae, with F. pedunculosa as sister to the rest. The F. erecta complex comprises approximately 17 taxa: F. erecta, F. abelii, F. boninsimae, F. nishimurae, F. iidaiana, F. gasparriniana var. laceratifolia, F. gasparriniana var. viridescens, F. pyriformis, F. stenophylla, F. fusuiensis, F. fengkaiensis, F. sinociliata, F. tannoensis, F. vaccinioides, F. formosana, F. pandurata, and F. periptera. The last five of these were supported as good species, while the others were not well supported by the present evidence. Evidence also supported the status of the non-F. erecta complex species including. F. pedunculosa, F. ischnopoda, F. heteromorpha, and F. variolosa. Ficus filicauda and F. neriifolia are possibly conspecific. The species status of F. potingensis should be restored and it should be treated as a member of section Eriosycea. Identification of the remaining taxa (F. gasparriniana var. esquirolii, F. ruyuanensis, F. daimingshanensis, F. chapaensis, F. changii, F. trivia, and F. tuphapensis) and their relationships to the F. erecta complex were not clarified. As a whole, only ten species in this subsection are confirmed, one is excluded, one is synonymous, and the others are either unresolved or short of samples. There appears to be a consistent genetic background among these unresolved groups, which suggests that repeated hybridization (as a result of pollinator host shifts) has filled up the interspecific gaps during the fig–pollinator coevolution process.