A synoptic review of the genus Thaumaspis Bolívar (Orthoptera,Tettigoniidae, Meconematinae) with the description of a new genus and four new species

Two new species of the new genus Athaumaspis gen. n., Athaumaspis minutus sp. n. and Athaumaspis tibetanus sp. n. from Vietnam and China are described. The subgenus Pseudothaumaspis of Thaumaspis is elevated to generic status and another two new species Pseudothaumaspis bispinosus sp. n. and Pseudothaumaspis furcocercus sp. n. are described, the remaining species of Thaumaspis are reviewed and keyed with the four new species.     

New genus of leaf-mimicking katydids (Orthoptera: Tettigoniidae) from the Late Eocene–Early Oligocene of France and England

A new leaf-mimicking katydid (Orthoptera: Tettigoniidae: Pseudophyllinae), Archepseudophylla fossilis gen. et sp. nov. is described based on tegmina from the Early Oligocene of France. Lithymnetes laurenti Théobald, 1937 from the Late Eocene of France is transferred to Archepseudophylla comb. nov. The enigmatic ‘Poekilloptera’ melanospila Cockerell, 1921 from the Early Oligocene of the Isle of Wight (UK) is transferred to Archepseudophylla comb. nov. The present discoveries in the Palaeogene of the Isle of Wight, Céreste, and Marseilles Basins represent the first fossil records of Pseudophyllinae. In addition, these records support warm and humid forest conditions near the palaeolakes of the Isle of Wight, Camoins-les-Bains, and Céreste.     

Cytogenetics studies in Brazilian species of Pseudophyllinae (Orthoptera: Tettigoniidae): 2n(♂)=35 and fn=35 the probable basic and ancestral karyotype of the family Tettigoniidae

The karyotypes of five species of Brazilian Pseudophyllinae belonging to four tribes were here studied. The data available in the literature altogether with those obtained with species in here studied allowed us to infer that 2n(♂)=35 is the highest chromosome number found in the family Tettigoniidae and that it is present in species belonging to Pseudophyllinae, Zaprochilinae and in one species of Tettigoniinae. In spite of that all five species exhibit secondary karyotypes arisen surely by a mechanism of chromosomal rearrangement of centric fusion, tandem fusion and centric inversion types from those with 2n(♂)=35 and FN=35, they share some common traits. The X chromosome is submetacentric (FN=36), heteropicnotic during the first prophase, the largest of the set but its size is rather variable among the species and the sex chromosomal mechanism is of the XO( ♂ ), XX( ♀ ) type. The chromosomal rearrangements involved in the karyotype evolution of the Pseudophyllinae and its relationship with those of the family Tettigoniidae are discussed and we propose that the basic and the ancestral karyotype of the Tettigoniidae is formed by 2n(♂)=35, FN=35 and not by 2n(♂)=31, FN= 31, as usually accepted.     

Chromosome evolution and phylogeny in Ronderosia (Orthoptera,Acrididae, Melanoplinae): clues of survivors to the challenge of sympatry?

In an attempt to unveil the origin of neo‐sex chromosomes in Ronderosia Cigliano grasshoppers, we performed a combined phylogenetic analysis based on morphological (external morphology and male genitalia) and molecular data (COI, COII, 16S and ITS2) to explore the chromosome evolution within the genus. We also analysed the distributional patterns of the various Ronderosia species and considered the possible role of chromosome rearrangements (CRs) in speciation processes within the genus in the light of ‘suppressed‐recombination’ models. We mapped the states of three chromosomal characters on the combined tree topology. The combined evidence supported Ronderosia as a monophyletic group. The cytogenetic analyses of the genus demonstrated the importance of rearranged karyotypes with single, complex and multiples neo‐sex chromosome determination systems in all species. The chromosome character optimisation suggests X‐autosome centric fusion as the mechanism responsible for neo‐sex chromosome formation in most Ronderosia species, except in R. dubia and R. bergii. Similar autosomes were involved in fusions with the ancestral X chromosome in Ronderosia, supporting previous hypotheses on the unique origin of X‐autosome fusion for the sex chromosome in the genus. As a source of chromosome variation, autosome‐autosome centric fusion played a secondary role in Ronderosia compared with other Dichroplini. Given the homogeneity in the morphological features, the sympatric distribution of closely related species and the intrinsic property of centric fusion as suppressors of the crossing over, we suggest that CRs may have played a key role during the speciation process within Ronderosia.     

Dynamism and context‐dependency in diversification of the megadiverse plant genus Solanum (Solanaceae)

Explosive radiations—substantial increases in net species diversification—have been considered one of the most intriguing diversification patterns across the Tree of Life, but the subsequent change, movement, and extinction of the constituent lineages make radiations hard to discern or understand as geological time passes. We used the megadiverse angiosperm genus Solanum L. (Solanaceae), with ca. 1200 currently accepted species distributed worldwide in a wide array of habitats, to explore these patterns on a global scale. We synthesized phylogenetic and distributional data for this ongoing radiation to show how dispersal events and past climatic changes have interacted to shape diversification. We find that, despite the vast diversity of Solanum lineages in the Neotropics, lineages in the Old World are diversifying more rapidly. This recent increase in diversification coincides with a long‐distance dispersal event from the Neotropics to regions where major climatic changes were taking place. Two separate groups of Solanum have migrated and established in Australia, but only the arid‐adapted lineages underwent significant increases in diversification rate, as they were able to adapt to the continent's long‐term climatic trend towards seasonally dry and arid biomes (a pattern observed in the diversification of other arid‐adapted groups). Our findings provide a clear example of how successful colonization of new areas and niches can—but does not always—drive explosive diversifications.     

Diversity,systematics, and evolution of Cynodonteae inflorescences (Chloridoideae – Poaceae)

The species of the Cynodonteae tribe show great morphological diversity in their reproductive structures. Previous studies where inflorescences were comparatively analysed in the context of phylogeny have shown that although grass inflorescences seem to be excessively variable, there are certain aspects of inflorescences that store relevant information on the evolution and systematics in Poaceae. We have analysed and compared the inflorescence structures of species belonging to the Hilariinae, Monanthochloinae, Scleropogoninae, and Muhlenbergiinae subtribes. Considering the most relevant morphological characters, the most recurrent types of inflorescences in the lineage were determined by means of a principal coordinates analysis. To understand the evolution of inflorescence morphology, ancestral reconstructions of inflorescence characters were performed using the Bayesian inference method. The results obtained demonstrate that the processes of homogenization and truncation might account for the diversity observed in adult inflorescences. Five different types of inflorescences were identified out of 36 theoretical possibilities. Amongst these, inflorescence type 1 (panicle of spikelets, with a terminal spikelet, non-homogenized, and bearing third- or higher-order branches) was found to be the most frequent in the studied group. Ancestral reconstructions of morphological characters allowed us to suggest that the ancestor of the group might have had an inflorescence with the form of a raceme of spikelets, non-truncated and bearing first-order branches. More complex inflorescences bearing no terminal spikelets and having branches of higher order might have diverged this lineage.     

Diel Calling Activity and Field Survival of the Bushcricket,Sciarasaga quadrata (Orthoptera: Tettigoniidae): A Role for Sound-locating Parasitic Flies?

In this study I examine the variability in calling activity of Sciarasaga quadrata over both the short (4 d) and long term (life span), and the temporal pattern to host searching by its most significant natural enemy, the ormiine fly Homotrixa alleni. Indeed, few male S. quadrata survive the entire calling season as this acoustically orienting parasitoid fly continually ‘culls’ calling males out of the population. I found that male S. quadrata commence calling over 1 h before sunset, cease around midnight and call, on average, for over 3 h within this period (range 0–11 h 09 min). There was significant concordance and repeatability in calling activity over both the short and long term, enabling me to conclude that relative differences in calling activity among males persist throughout a male's life span. There was a distinct peak to host searching by gravid female flies, with 82 % of all flies collected at acoustic traps between sunset and midnight. A positive association between call duration (time spent calling) and fly attraction was evident from multiple collections of flies at single traps. Counter to expectation, there was no significant decline in average call duration across successive collections of males, indicating that long-call-duration males were not being selectively culled by flies from the calling population. Males at the end of the calling season, when their remaining reproductive potential is low, allocate the majority of their calling time between sunset and midnight despite the high risk of fly attack. The distribution of call durations within the population was not skewed, which if heritable, indicates a lack of history of directional selection acting on call duration. Limited evidence suggests that encounters with females are rare in S. quadrata so that males may be selected, irrespective of natural selection pressure, to call for long periods of time to optimize their chance of attracting mates. The above findings are further discussed in relation to parasitoid foraging and acoustic reproductive strategies.     

Are solitary and gregarious Mormon crickets (Anabrus simplex, Orthoptera, Tettigoniidae) genetically distinct?

Phase polyphenisms are usually thought to reflect plastic responses of species, independent of genetic differences; however, phase differences could correlate with genetic differentiation for various reasons. Mormon crickets appear to occur in two phases that differ in morphology and behaviour. Solitary individuals are cryptic and sedentary whereas gregarious individuals form bands, migrate, and are aposematically coloured. These traits have been thought to be phenotypically plastic and induced by environmental conditions. However, there has been no previous investigation of the extent of genetic differences between solitary and gregarious populations of this widespread North American species. We sequenced two mitochondrial genes, COII and COIII, in samples of Mormon crickets from gregarious populations west of the continental divide and solitary mountain populations primarily east of the divide. Sequencing revealed two genetically distinct clades that broadly correspond with the solitary eastern populations and the mainly gregarious western populations. We used coalescent modelling to test the hypothesis that the species consists of two deep genetic clades, as opposed to a series of equally distinct populations. Results allowed us to reject the null hypothesis that a radiation independent of phase produced these clades, and molecular clock estimates indicate the time of divergence to be approximately 2 million years ago. This work establishes that the solitary populations found in the mountains on the eastern slope are part of a clade that is genetically distinct from the western populations, which are primarily gregarious, and the implications of this apparent correlation between phase and genetic differentiation are discussed.     

Origin and evolution of the Amyrel gene in the α-amylase multigene family of Diptera

Alpha-amylase genes often form multigene families in living organisms. In Diptera, a remote paralog, Amyrel, had been discovered in Drosophila, where this gene is currently used as a population and phylogenetic marker. The putative encoded protein has about 40% divergence with the classical amylases. We have searched the presence of the paralog in other families of Diptera to track its origin and understand its evolution. Amyrel was detected in a number of families of Muscomorpha (Brachycera-Cyclorrapha), suggesting an origin much older than previously thought. It has not been found elsewhere to date, and it is absent from the Anopheles gambiae genome. The intron–exon structures of the genes found so far suggest that the ancestral gene (before the duplication which gave rise to Amyrel) had two introns, and that subsequent, repeated and independent loss of one or both introns occurred in some Muscomorpha families. It seems that the Amyrel protein has experienced specific amino acid substitutions in regions generally well conserved in amylases, raising the possibility of peculiar, functional adaptations of this protein.     

Hybridization and recurrent evolution of left–right reversal in the land snail genus Schileykula (Orculidae,Pulmonata)

The land snail genus Schileykula Gittenberger, 1983 is distributed in arid limestone areas from western Turkey to north-western Iran. It comprises eight species, which display high variation in shell size and morphology. The cylindrical shells are 5–12 mm in height and the last shell whorls bear several inner lamellae and plicae. Two taxa differ in their chirality having sinistral shells, while all the others are dextrals such as the vast majority of orculids. The aim of this study was to establish a molecular genetic phylogeny of Schileykula and to test whether it conforms to the current morphology-based classification. Furthermore, we were interested in the phylogenetic position of the two sinistral forms in order to assess whether one or two reversals happened in the evolution of the genus. Nine out of ten species, including all four subspecies of Schileykula trapezensis and three of six subspecies of Schileykula scyphus, were investigated. A section of the mitochondrial cytochrome c oxidase subunit I gene was analyzed in 54 specimens of Schileykula and from a subsample, partial sequences of the mitochondrial genes for the 12S rRNA and the 16S rRNA, and a section of the nuclear H4/H3 histone gene cluster were obtained. The phylogenetic trees based on the mitochondrial sequences feature high support values for most nodes, and the species appear well differentiated from each other. The two chiral forms evolved independently and are not sister lineages. However, some groupings disagree with the present morphology-based classification and taxonomical conclusions are drawn. Schileykula trapezensis is polyphyletic in the molecular genetic trees; therefore, three of its subspecies are elevated to species level: Schileykula acampsis Hausdorf, 1996 comb. nov., Schileykula neuberti Hausdorf, 1996 comb. nov., and Schileykula contraria Neubert, 1993 comb. nov. Furthermore, Schileykula sigma is grouped within S. scyphus in the mitochondrial and nuclear trees and consequently treated as a subspecies of the latter (Schileykula scyphus sigma Hausdorf, 1996 comb. nov.). Schileykula nordsiecki, whose shell morphology is indistinguishable from that of the neighboring Schileykula scyphus lycaonica, but who differs in its genital anatomy, was confirmed to represent a distinct lineage. The phylogenies produced by the mitochondrial and nuclear data sets are to some extent conflicting. The patterns differ concerning the grouping of some specimens, suggesting at least two independent hybridization events involving S. contraria, S. scyphus and S. trapezensis. The results exemplify the importance of integrating both mitochondrial and nuclear sequence data in order to complement morphology-based taxonomy, and they provide further evidence for hybridization across distantly related lineages in land snails.     

Platycleis vittata (Orthoptera: Tettigoniidae) in the northwestern part of its range is close to extinction: is this the result of landscape changes?

The northwestern distributions of several steppe species of Orthoptera extend to the southeastern part of the Czech Republic (Pannonia) and occupy more or less isolated fragments of optimal habitats. Their distributional limits are not conditioned by macroclimate in most cases but reflect landscape development (physical structure, plant community type and microclimate) and the insects’ dispersal abilities. These species prefer permanent grassland, and assessment of land use records indicates that the area occupied by permanent grassland has been greatly reduced by agriculture over the last two centuries. The area occupied by permanent grasslands in Pannonia was highest in the nineteenth century, declined until the second half of the twentieth century, and slightly increased at the beginning of the twenty first century. These changes in the area of permanent grassland generally reflect economic and political processes; in particular, consolidation of land plots and expropriation of private property brought an end to grazing. The distribution of a model Orthoptera specialist, Platycleis vittata, on steppe habitats was surveyed at 48 potentially suitable locations from 2004 to 2010. The survey determined that the area with suitable habitat is currently very limited at all 48 locations. Occurrence was confirmed in just two locations that are 3 km apart. The research suggests that Platycleis vittata in the Czech Republic is on the brink of extinction because of extreme fragmentation of suitable biotopes and degradation of optimal habitats.     

Colonization history, ecological shifts and diversification in the evolution of endemic Galápagos weevils

Mitochondrial DNA sequence data were obtained for eight species of flightless Galapaganus endemic weevils and one winged close relative in order to study their colonization history and modes of diversification in the Galápagos Archipelago. Contrary to most other insular radiations, the phylogeny estimates we recovered for Galapaganus do not follow the progression rule of island biogeography. The penalized likelihood age estimates of colonization of the archipelago exceed the age of the emerged islands and underscore the potential role of now sunken seamounts for the early evolution of Galapaganus . The phylogeny proposes one intra-island origin for Galapaganus endemics, but monophyly tests suggest a larger contribution of in-situ speciation on older islands. Generalist habitat preferences were reconstructed as ancestral while shifts to highland habitats were reconstructed as having evolved independently on different islands. Magnitudes and patterns of diversification rate were found to differ between older and younger islands. Our analyses reveal that the colonization sequence of islands and timing of colonization of Galapaganus could be linked with the geological and volcanic history of the islands in a rather complex scenario. Even though most islands appear to have been colonized soon after their emergence, there are notable deviations from the pattern of sequential colonization expected under the progression rule when considering only the extant emerged islands. Patterns of diversification rate variation on older and younger islands correspond to the volcanic activity or remnants of such activity, while the pattern of independent evolution of restricted habitat preferences in different islands suggests that habitat shifts could also have contributed to species diversity in Galapaganus .     

Diversity of biting midges of the genus Culicoides Latreille (Diptera: Ceratopogonidae) in the area of the Yacyretá Dam Lake between Argentina and Paraguay

The Culicoides communities have been analyzed between 1993/1998 in the area influenced by the Yacyret Dam Lake (Paran River, Argentina-Paraguay). Adults of Culicoides were collected monthly by using CDC light traps exposed for 24 h in 9 sampling sites located at both margins of the river; 21 species were recorded. Highest values of species richness were recorded during 1993/1994, being Quiteria and Corpus the sites with the higest number of species (10 and 11, respectively). The species diversity was elevated in Quiteria, Zaim n, Candelaria, Santa Tecla, Capit n Meza and Corpus (Shannon's diversity index 1.0-1.9) while Corate , Ituzaing and Aguapey showed less richness and diversity. The more abundant species were C. insignis, C. venezuelensis, C. leopoldoi, C. limai, C. flinti, C. debilipalpis, C. paraensis and C. guttatus. C. insignis, potential vector of bluetongue virus (BTV) to domestic and wild rumiants in the Neotropical region, is the predominant species in the area and was the only species widely distributed. C. paraensis, a proven vector of Oropouche virus to humans, is a common and abundant species. C. pusillus and C. lahillei, potential vectors of BTV and a filarial parasite, respectively, were occasionally collected. The taxonomic structure of communities was constant during the study period. The occasional species were not characteristic to one particular site and their presence could be related to non-intrinsic conditions.     

Sequence-based species delineation and molecular phylogenetics of the transitional Nearctic–Neotropical grasshopper genus Taeniopoda (Orthoptera,Romaleidae)

Taeniopoda is a genus of grasshoppers currently represented by 12 species distributed from southern USA to Panama, with most of them occurring along the transitional Nearctic–Neotropical region in central and southern Mexico. Despite being a small group of conspicuous, colourful species, the systematics of Taeniopoda has been largely neglected, including its phylogenetic affinity with the morphologically similar, monotypic genus Romalea. Here, we assessed the species limits in 11 of the species of Teniopoda based on two mitochondrial (mt) markers (COI, cyt b). Phylogenetic relationships were reconstructed adding two nuclear gene markers (28S, H3). A relaxed molecular clock analysis was performed based on the mt markers. We detected nuclear mt paralogues (numts) and the probable introgression of T. tamaulipensis mtDNA in specimens of T. eques from central Mexico. Between six and 14 species of Taeniopoda were delimited by the sequence-based approaches performed (COI divergence with thresholds of 1 and 2%; General Mixed Yule-Coalescent (GMYC) model). The GMYC and 1% threshold analyses with COI were more congruent with the currently recognized morphology-based taxonomy with 10 and 11 putative species, respectively. Four of these species were regarded as ‘stable’, since they were supported by at least one of the molecular analyses and by diagnostic morphological features. The species-based phylogeny recovered Taeniopoda as paraphyletic with respect to the monotypic genus Romalea. Three morphologically and geographically congruent major clades were recovered, two with species having a considerably elevated pronotal crest and one with its members having it less elevated. The origin and subsequent diversification of Taeniopoda were estimated to occur from the mid and late Miocene to Pliocene, respectively. The current species diversity in Taeniopoda was estimated to occur during the Pleistocene, which was probably influenced by the climatic oscillations that occurred during this period and the uplift of mountain ranges in Central America.     

A new species of the genus Falconius Bolívar, 1898 (Orthoptera: Tetrigidae,Scelimeninae) from Vietnam

Falconius tschernovi sp. n. is described from Vietnam. The new species is closely related to F. dubius Günther, 1938 from Malaysia (Borneo) and F. guangxiensis Zheng et Jiang, 1997 from China (Guangxi Zhuang Autonomous Region), but differs from both in the presence of two pairs of additional lateral keels on the disc of the pronotum, in the armament of the fore femur, in the narrow hind femur, and in the coloration of the hind tibia.     

Population dynamics in the Grey Bush Cricket Platycleis albopunctata (Orthoptera: Tettigoniidae) – What causes interpopulation differences?

We found sharply differing population dynamics in Platycleis albopunctata at two one-hectare study sites differing in vegetation cover and inclination of slopes. At one site, the population remained relatively constant (about 600 individuals) over the period observed, while at the other site the population size varied during the observation period between more than 3000 and less than 100 individuals. This field observation was analysed by an experimental study of embryonic development and reproduction, both of which are expected to largely influence population dynamics. These experiments revealed a close correlation between the temperature and the duration of egg development on the one hand and fecundity on the other. We concluded that the temperature dependence of development and reproduction is responsible for the weather dependence of population dynamics. Based on our results, we developed a hypothesis to explain the striking differences in the dynamics of the two study sites which attributes them to differences in microclimate and primary production. Additionally, we applied a simulation model to investigate the weather dependence of the population dynamics. Finally, we discuss the consequences of different population dynamics for conservation: (a) the example stresses the difficulties that arise when using demographic data of any population for the prognosis of the extinction risk of a population under study; (b) the dependence of population dynamics on habitat parameters means they can be influenced by management.     

Redescription and revision of the Neotropical genus Pseudoheptascelio Szabó (Hymenoptera, Platygastridae, Scelioninae), parasitoids of eggs of short-horned grasshoppers (Orthoptera, Acrididae)

The genus Pseudoheptascelio Szabó is redescribed and its species revised. We recognize four species: Pseudoheptascelio muesebecki Szabó, Pseudoheptascelio cornopis Masner, Pseudoheptascelio ticosp. n. and Pseudoheptascelio rexsp. n. The genus is found from Guatemala south to the Brazilian state of Rio Grande do Sul. The species Pseudoheptascelio cornopis is recorded as a parasitoid of the eggs of Cornops aquaticum (Bruner) on water hyacinth, Eichhornia crassipes (Mart.) Solms.     

Effects of mating status on copulation investment by male bushcricket Gampsocleis gratiosa (Tettigoniidae, Orthoptera)

In many insect species, male will donate prey, se-cretion, and spermatophore as “nuptial gift” or “court- ship feeding” to females prior to, after or during the copulation[1,2]. The nuptial gifts represented great re-productive cost. In bushcrickets, …     

Origin and diversification of the Greater Cape flora: ancient species repository, hot-bed of recent radiation, or both?

Like island-endemic taxa, whose origins are expected to postdate the appearance of the islands on which they occur, biome-endemic taxa should be younger than the biomes to which they are endemic. Accordingly, the ages of biome-endemic lineages may offer insights into biome history. In this study, we used the ages of multiple lineages to explore the origin and diversification of two southern African biomes whose remarkable floristic richness and endemism has identified them as global biodiversity hotspots (succulent karoo and fynbos). We used parsimony optimization to identify succulent karoo- and fynbos-endemic lineages across 17 groups of plants, for which dated phylogenies had been inferred using a relaxed Bayesian (BEAST) approach. All succulent karoo-endemic lineages were less than 17.5 My old, the majority being younger than 10 My. This is largely consistent with suggestions that this biome is the product of recent radiation, probably triggered by climatic deterioration since the late Miocene. In contrast, fynbos-endemic lineages showed a broader age distribution, with some lineages originating in the Oligocene, but most being more recent. Also, in groups having both succulent karoo- and fynbos-endemic lineages, there was a tendency for the latter to be older. These patterns reflect the greater antiquity of fynbos, but also indicate considerable recent speciation, probably through a combination of climatically-induced refugium fragmentation and adaptive radiation.     

On Paphiopedilum malipoense sp. nov.—An Intermediate Form Between Paphiopedilum and Cypripedium,with a Discussion on the Origin of the genus

Paphiopedilum malipoense S. C. Chen et Tsi is a very interesting new species with its flower similar to that of Cypripedium, especially section Cypripedium. It belongs to subgenus Brachypetalum, the most primitive group of Paphiopedilum, but differs from its allied species in hgniva elliptic-lanceolate sepal with cuspidately acuminate apex, rather narrow petals and horizontal lip, which are of common occurrence in many cypripediums, but very rare in paphiopedilums. Apparently, this is an intermediate form, or a link, between Paphiopedilum and Cypripedium, but it does not seem to arise from hybridization between them, because no Cypripedium has been found wherever Paphiopedilum occurs. The new species is distributed in southeastern Yunnan of China. In this area, as well as in river valleys of western Yunnan or the Hengduan Mountains, there have been four species of the same genus reported before. As we know, the Hengduan Mountains and their adjacent areas are rich in Cypripedium. The differentiation of the genus there is remarkable. All five sections it contains occur there and three of them are quite distinctive. For example, the general appearance of the section Bracleosa is dissimilar to that of any other cypripediums, but closely resembles that of Listera. It appears that the difference between sect. Bracteosa of Cypripedium and sect. Brachypetalum of Paphiopedilum is not necessarily wider than that between sect. Bracteosa and sect. Cypripedium of the same genus. Apparently, it is reasonable to consider Paphiopedilum to be an evolutional branch of Cypripedium extending into tropical area, with its primitive group (subgenus Brachypetalum) still remaining in its northern fringe area. This primitive subgenus has eight species, distributed from western Yunnan to the Malay Peninsula. Five of them, including the intermediate and primitive form published here, are found in the hilly land of southeastern Yunnan and the river valleys of western Yunnan. All these facts suggest its area of origin: the river valleys of the Hengduan Mountains and the lower hilly land contiguous to the southof them.