Cytological studies on the genus Holcoglossum (Orchidaceae)

The karyotypes of ten species of Holcoglossum (Orchidaceae), a highly endangered and diversified genus from China, were investigated to study the infrageneric relationships, biogeography, and speciation patterns in the Hengduan Mountains. The karyotype formulae of the studied species are as follows: 2 n  = 38 = 20m + 18sm in H. subulifolium , 2 n  = 38 = 22m + 16sm in H. amesianum , 2 n  = 38 = 26m + 12sm (6 SAT) in H. lingulatum , 2 n  = 38 = 26m + 12sm in H. wangii , 2 n  = 38 = 10m + 28sm in H. kimballianum , 2 n  = 38 = 14m + 22sm + 2st in H. flavescens , 2 n  = 38 = 24m + 12sm + 2st in H. rupestre , 2 n  = 38 = 14m + 20sm + 4st in H. sinicum , 2 n  = 38 = 16m + 14sm + 8st in H. weixiense , and 2 n  = 76 in H. tsii . The karyotypes of two tropical species, H. amesianum and H. subulifolium , are the most primitive in the genus, whereas those of four temperate alpine species, H. sinicum , H. rupestre , H. weixiense , and H. flavescens , are more advanced. H. tsii is a tetraploid and H. rupestr e may be one of its ancestors. The low frequency of polyploidy in Holcoglossum in the Hengduan Mountains region supports the conclusion that chromosome stasis during rapid speciation is common there.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 283–288.     

A study of the genus Holcoglossum of Orchidaceae

The present paper is an attempt to make a taxonomic study of the little known orchid genus Holcoglossum, as well as a comparison of the genus with its allies, such as Vanda, Papilionanthe, Ascolabium, Ascocentrum, Aěrides, Neofinetia and Saccolabium. Holcoglossum was established by Schlechter in 1919 (Orchideologiae Sino-Japonicae Prodromus) as a monotypic genus, based upon Saccolabium quasipinifolium Hayata. Five years later he published another true Holcoglossum as Aěrides flavescens, which was referred by Tang et Wang to Saccolabium in 1951. Further investigation of this genus was by Garay in 1972 who added two species, H. kimballiana and V. rupestris (synonymy of Aěrides flavescens), but considered Neofinetia, a quite different taxon, to be congeneric. It is shown that the demarcation of Holcoglossum remains cofused. During the course of our study, the species of Holcoglossum and its allied genera are carefully examined, we come to the conclusion that Ho lcoglossum is a distinct genus. It is characterized by the short stem; fleshy terete or subterete, sulcate above leaves, with their apex acute and non-lobed; thickening or keeled costa on the back of sepals, 3-lobed lip, with erect sidelobes, paralled to the column; slender and recurved spur; footless column usually with prominent wings; 2 notched pollinia attached to linear stipe which is tapered toward the base. In addition to Ascolabium, it differs from Vanda, Papilionanthe, Ascocentrum, Aěrides, Neofinetia and Saccolabium by its terete or subterete leaves on their ventral side with a furrow, from Papilionanthe by lacking footless column, from Ascolabium by sepals and spur characters, from Ascocentrum by slender and recurred spur, from Aěrides by the absence of a column-foot and the appearance of spur, from Neofinetia by stipe tapered toward the base, from Saccolabi-um by both aspects of the vegetative organs and the flowers.     

Pollination system of Holcoglossum rupestre (Orchidaceae): A special and unstable system

The pollination system of Holcoglossum rupestre was studied in northwestern Yunnan. The species is self-compatible but pollinator-dependent and offers nectar as a reward to visitors. A species of beetle, Hybovalgus bioculatus Kolbe, is identified as the pollinator among all visitors observed. The beetles carried the pollinia in two different positions, i.e. on the buttock and the foreleg, and pollinated the flowers by two different mechanisms. The analysis suggests that Holcoglossum rupestre and its pollinator may be in an unstable pollination relationship. The natural fruit set is low, which is attributed to the low visiting frequency of pollinator and the effective mechanism to prevent self-pollination.     

耳唇槽舌兰——中国兰科一新种

对产于云南的兰科新种耳唇槽舌兰(Hotcoglossum auriculatum)进行了描述和绘图。该新种具有显著特征：萼片与花瓣基部有长爪;花瓣边缘具不规则齿或呈啮蚀状;唇瓣椭圆状圆形,边缘有短流苏或啮蚀状流苏,基部有爪和耳,极易区别于该属已知的种类,包括它的近缘种白唇槽舌兰(H．subulifolium)。     

Pollinators and nectar robbers cause directional selection for large spur circle in Impatiens oxyanthera (Balsaminaceae)

Nectar spurs have an important role in floral evolution and plant–pollinator coadaptation. The flowers of some species possess spurs curving into a circle. However, it is unclear whether spur circle diameter is under direct selection pressure from different sources, such as pollinators and nectar robbers. In this study, we quantified selection on some floral traits, such as spur circle diameter in Impatiens oxyanthera (Balsaminaceae) using phenotypic selection analysis and compared the relative importance of pollinators and nectar robbers as selective agents using mediation analysis. The study showed that pollinators caused significant selection on corolla length, spur curvature and spur circle diameter while nectar robbers only imposed strong selection on spur circle diameter. Pollinators favored flowers with large corolla, curly spurs and large spur circle while nectar robbers preferred flowers with small spur circle. More pollinator visits resulted in higher female reproductive success, while robbery reduced female fitness. Conflicting selection on spur traits from pollinators and nectar robbers was not found. Mediation analysis showed that selection on floral traits through nectar robbing was stronger than selection through pollination. The results suggested that pollinators and nectar robbers jointly mediated the directional selection for large spur circle, and nectar robbers caused stronger selection than pollinators on floral traits.     

Generic delimitation and a new infrageneric system in the genus Holcoglossum (Orchidaceae: Aeridinae)

Based on herbarium and field observations, we confirm the generic delimitation of Holcoglossum and present a new infrageneric system for the genus. Our work supports Holcoglossum as a natural group including H. amesianum (Rchb.f.) Christenson and H. subulifolium (Rchb.f.) Christenson. The new infrageneric system of Holcoglossum includes two subgenera: Holcoglossum subgen. Brachycentron subgen. nov. and Holcoglossum subgen. Holcoglossum , the latter comprising two new sections, Holcoglossum sect. Holcoglossum and Holcoglossum sect. Sorotylos .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 465–468.     

中国蝴蝶兰属(兰科)一新记录(英文)

报道了中国兰科植物一新记录—囊唇蝴蝶兰,并提供描述及照片。本种与麻栗坡蝴蝶兰非常相似,但它的花瓣较短而宽,花序轴呈"Z"型曲折,唇盘中央的胼胝体较小,可以与后者相区别。     

Bulbophyllum nujiangense (Bulbophyllinae,Epidendroideae, Orchidaceae), a New Species from Yunnan,China

Based on field observation and specimen examination, a new species of Orchidaceae, Bulbophyllum nujiangense, is described and illustrated. Bulbophyllum nujiangense is closely related to B. cylindraceum in sect. Brachystachya in having tufted plants, large leaf, long inflorescence, small flowers in raceme, and fleshy lip, but can be easily distinguished from the latter in having flowers deep purplish red and whorl well spaced along the rachis, lateral sepals adhere to each other slightly at base, elliptic petals with obtuse and mucronate apex, lip deep purplish red.     

紫堇科二新种

该文描述了紫堇科二新种:(1)自云南省发现的紫堇科紫堇属一新种,宽鳞紫堇。此新种与波密紫堇相近似,与后者的区别在于本新种的根状茎鳞片半透明,具三条脉,茎无叶,外花瓣无鸡冠状突起,雄蕊束具2花药,花丝条形,无脉,子房宽条形。(2)自黑龙江省发现的紫堇科紫堇属一新种,鹤岗延胡索。此种与新疆元胡相似,但本种叶的小叶多在顶端,具1~5牙齿,花序苞片有3~5小裂片,上花瓣及其瓣片和距均较小,蜜腺呈狭卵球形,弯曲,顶端尖锐而与新疆元胡相区别。     

Loss of innovative traits underlies multiple origins of Aquilegia ecalcarata

Traits are basic attributes of organisms that form the basis for speciation and diversity. The floral nectar spur is a classic example of a key innovative trait. Differences in nectar spur morphology can lead to pollinator specialization and thereby promote reproductive isolation between species. Despite its importance, the nectar spur has been lost in some members of the columbine genus (Aquilegia), resulting in a new spurless trait, and the evolutionary influence of this trait has become a topic of scientific interest. Aquilegia ecalcarata is an important representative columbine species that lacks spurs. Here, we resequenced the genomes of 324 individuals from A. ecalcarata and four related species. We found that A. ecalcarata was divided into three groups based on the phylogenetic relationships and population genetic structures. Topology weighting analysis revealed that A. ecalcarata has multiple origins, and Patterson′s D statistic showed that the spurless trait may have one origin. Floral morphological analysis revealed significant differences between A. ecalcarata and its spurred sister groups, and the floral phenotypes of the three A. ecalcarata groups have identical or similar floral phenotypes. Our results confirmed that the spurless trait not only produced the phenotype of A. ecalcarata but also contributed to the emergence of the A. rockii phenotype. Moreover, the spurless trait promoted the divergence between A. ecalcarata and its close, spurred relatives. Our research shows that the loss of key innovative traits can play a very important role in speciation and species diversity.     

陕西兰科一新记录属--套叶兰属

报道了陕西省兰科一新记录属——套叶兰属（Hippeophyllum Schltr.）及一新记录种——套叶兰（Hippeophyllum sinicwn S. C. Chen et K. Y. Lang）。     

DNA barcoding of the recently evolved genus Holcoglossum (Orchidaceae: Aeridinae): a test of DNA barcode candidates

Orchidaceae is one of the largest families of flowering plants. Many species of orchid are endangered, and all species are included in Conventions on International Trade of Endangered Species of Fauna and Flora (CITES) I and II, but it is very difficult to identify orchid species, even those with fertile parts. The genus Holcoglossum (Orchidaceae: Aeridinae) has long been problematic in taxonomy. It consists of both long-evolved and radiated species and is an excellent case to use for testing DNA barcodes for Orchidaceae. We investigated the power of a subset of proposed plant barcoding loci [rbcL, matK, atpF-atpH, psbK-psbI, trnH-psbA and internal transcribed spacer (ITS)] to discriminate between species in this genus. Our results showed that all these DNA regions, except psbK-psbI and atpF-atpH, can be amplified easily from Holcoglossum and sequenced with established primers. The DNA regions matK and ITS had the highest variability. Among the six loci, matK resolved eight of the 12 Holcoglossum species and had the highest discriminatory ability. However, the combination of matK and ITS showed a greater ability to identify species than matK alone. Single or combined DNA markers discriminated between Holcoglossum species distributed in tropical areas effectively, but had less ability to identify radiated species from the temperate Hengduan Mountains of China. In the study, matK proved to be a useful DNA barcode for the genus Holcoglossum; however, complementary DNA regions are still required to accelerate the investigation and preservation of radiated species of orchid.     

Intraspecific independent evolution of floral spur length in response to local flower visitor size in Japanese Aquilegia in different mountain regions

Geographic differences in floral traits may reflect geographic differences in effective pollinator assemblages. Independent local adaptation to pollinator assemblages in multiple regions would be expected to cause parallel floral trait evolution, although sufficient evidence for this is still lacking. Knowing the intraspecific evolutionary history of floral traits will reveal events that occur in the early stages of trait diversification. In this study, we investigated the relationship between flower spur length and pollinator size in 16 populations of Aquilegia buergeriana var. buergeriana distributed in four mountain regions in the Japanese Alps. We also examined the genetic relationship between yellow‐ and red‐flowered individuals, to see if color differences caused genetic differentiation by pollinator isolation. Genetic relationships among 16 populations were analyzed based on genome‐wide single‐nucleotide polymorphisms. Even among populations within the same mountain region, pollinator size varied widely, and the average spur length of A. buergeriana var. buergeriana in each population was strongly related to the average visitor size of that population. Genetic relatedness between populations was not related to the similarity of spur length between populations; rather, it was related to the geographic proximity of populations in each mountain region. Our results indicate that spur length in each population evolved independently of the population genetic structure but in parallel in response to local flower visitor size in different mountain regions. Further, yellow‐ and red‐flowered individuals of A. buergeriana var. buergeriana were not genetically differentiated. Unlike other Aquilegia species in Europe and America visited by hummingbirds and hawkmoths, the Japanese Aquilegia species is consistently visited by bumblebees. As a result, genetic isolation by flower color may not have occurred.     

中国兰科植物三个新记录种

报道了中国兰科植物的三个新记录种,裂唇卷瓣兰(Bulbophyllum picturatum),长柄虾脊兰(Calan-the allizettei)和绿花毛兰(Eria lanigera)。裂唇卷瓣兰的中萼片全缘前部具一粒状附属物,唇瓣基部两侧具耳。长柄虾脊兰的中裂片非常小并具三条褶片。黄花毛兰的花黄绿色,唇瓣不裂。     

怒江石豆兰，中国兰科一新种

在野外和标本研究的基础上, 描述了中国兰科石豆兰属的一新种,怒江石豆兰, 并提供彩色图片和线条图。 怒江石豆兰与大苞石豆兰很近,但这两个种在花的颜色、侧萼片合生程度、花瓣形状等性状上区别明显。     

Pollination of Habenaria tridactylites on the Canary Islands

We investigated the pollination of Habenaria tridactylites, an endemic orchid of the Canary Islands. The entirely green, widely open flowers have a long spur containing nectar. We carried out fieldwork, a molecular clock analysis, herbarium surveys, identified pollinators by both morphology and DNA barcoding, and measured the length of floral spurs and insect tongues using a combination of traditional and innovative micro‐CT scanning methods to 1) determine the pollinator of this orchid and 2) investigate correlations between local mean spur length and age, altitude and longitude of the island. Habenaria tridactylites was found to be pollinated on Tenerife by both small and intermediate sized moth species with variable tongue lengths and mostly belonging to Geometridae and to a lesser extent Crambidae, Erebidae, Noctuidae and Tortricidae. Of the sixteen moth species identified, nine are endemic to the Canary Islands or Macaronesia. The different local populations of H. tridactylites on the islands of Gran Canaria, El Hierro, La Gomera, La Palma and Tenerife with different ages and distances from mainland Africa, did not show a significant correlation of mean spur length and altitude, but did show a significant and positive linear correlation with longitude and the geological age of the island. The latter is congruent with the evolutionary arms race theory first proposed by Darwin, suggesting that flowers gradually evolve longer spurs and pollinators longer tongues.     

Pollinator shifts and the evolution of spur length in the moth-pollinated orchid Platanthera bifolia

Background and Aims

Plant–pollinator interactions are thought to have shaped much of floral evolution. Yet the relative importance of pollinator shifts and coevolutionary interactions for among-population variation in floral traits in animal-pollinated species is poorly known. This study examined the adaptive significance of spur length in the moth-pollinated orchid Platanthera bifolia.

Methods

Geographical variation in the length of the floral spur of P. bifolia was documented in relation to variation in the pollinator fauna across Scandinavia, and a reciprocal translocation experiment was conducted in south-east Sweden between a long-spurred woodland population and a short-spurred grassland population.

Key Results

Spur length and pollinator fauna varied among regions and habitats, and spur length was positively correlated with the proboscis length of local pollinators. In the reciprocal translocation experiment, long-spurred woodland plants had higher pollination success than short-spurred grassland plants at the woodland site, while no significant difference was observed at the grassland site.

Conclusions

The results are consistent with the hypothesis that optimal floral phenotype varies with the morphology of the local pollinators, and that the evolution of spur length in P. bifolia has been largely driven by pollinator shifts.     

Facilitative interactions among co-flowering <Emphasis Type="Italic">Primula</Emphasis> species mediated by pollinator sharing

We investigated flowering phenology, pollinator visitation and visitor community composition in communities of self-incompatible sympatric Primula species in a high-elevation Himalayan ecosystem. Within the tight constraints imposed by short growing seasons in such ecosystems, interactions among co-occurring plants for pollinators may vary from competition to facilitation, depending on the specifics of the system. We found that pollinator community composition changed with elevation in this system: lepidopterans were the dominant visitors at lower elevations (2200–3000 masl), bees (other than bumblebees) dominated at mid-elevations (3000–3800 masl) and bumblebees dominated at higher elevations (3800–4600 masl). However, within an elevation zone, there were no significant differences in pollinators amongst co-occurring Primula species. At a focal study site where multiple Primula species co-occurred, our results showed that even while the overall flowering periods of these species broadly overlapped, the peak flowering periods of different Primula species were temporally segregated. Upon further inferring the nature of interaction amongst co-flowering Primula species, we found that plots with higher Primula diversity (≥?2 species) and density (80–100 individuals) experienced significantly higher pollinator visitation, compared with plots with single species and low flower densities (40–50 individuals). Our results suggest that in this community of sympatric, self-incompatible Primula species, a broadly aggregated, synchronous floral display of multiple species results in pollinator facilitation by attracting a greater number of pollinator visitors. Within this broadly synchronous display, the temporal segregation of peak flowering period of individual species may reduce competition for pollinators and limit heterospecific pollen transfer.     

Experimental Studies on Male and Female Reproductive Success: Effects of Variation in Spur Length and Pollinator Activity on Platanthera mandarinorum ssp. hachijoensis (Orchidaceae)

Abstract Field experiments on an orchid, Platanthera mandarinorum ssp. hachijoensis , were carried out to investigate the effects of variations in spur (nectary) length and pollinator activity on reproductive success (RS) of individual plants. Experiments comprised two kinds of treatments: spur shortening and bagging to restrict pollinator activity. Plants with a natural spur lengths range show nearly equal RS, whereas plants with shortened spurs had decreased RS proportional to spur length reduction. Pollinaria attachment (a measure of male RS) to the pollinators seems to demand stricter morphological correspondence than pollen receipt (female RS). Decrease of male and female RSs in spur-treatments was smaller in the case of high pollinator activity than low pollinator activity. Results of the bagging experiments indicated that female RS was greater than male RS in cases of high pollen carryover. These facts suggest that (1) fecundity selection operates more intensely on both male and female functions in the case of low pollinator activity and more intensely on the male function in the case of high pollen carryover, and (2) individuals with different spur lengths in natural populations have nearly equal probabilities of RS after excluding the extreme variants through the reduction of male RS.     

Oberonia swaminathanii sp. nov. (Orchidaceae) from Kerala,India

Oberonia swaminathanii, a new species of Orchidaceae from Wayanad district, Kerala, India is described and illustrated. The new species resembles Oberonia balakrishnanii, O. chandrasekharanii and O. seidenfadeniana by its 3‐lobed labellum and 2‐lobuled midlobe; but differs in having brick‐red coloured flowers, triangular labellum disc, and a subglobose column.