Viola stoloniflora (Violaceae), a new species from the Ryukyus

Viola stoloniflora Yokota et Higa (Violaceae) is newly described from Okinawa Island, the Ryukyus. This species is characterized by the procumbent stolons, the almost round leaf blades, the densely pilose petioles, the ebarbate white petals with violet veins, and the chromosome number of 2n=22. The present species belongs to Sect.Nomimium, Subsect.Serpentes and is closely related to theV. iwagawai group of Sect.Nomimium, Subsect.Adnatae.     

Responses of a rare (Viola elatior) and a common (Viola mirabilis) congeneric species to different management conditions in grassland — is different light competition ability responsible for different abundances?

We studied a congeneric species pair that shows very different abundances in Estonia — Viola elatior occurs in only five localities in mesic calcareous grasslands, while Viola mirabilis is abundant in mesic calcareous grasslands and forests all over the country. Both species were sown in patches of mesic calcareous grassland, in clipped and untreated plots. Both species were established successfully after sowing, indicating that they both may be dispersal limited to the same extent. V. elatior showed higher fecundity in the second year than V. mirabilis. V. elatior was more sensitive to the availability of light. In the first year, V. elatior established more successfully in clipped plots than V. mirabilis. In the second year, the number of established V. elatior individuals decreased in unmanaged plots, where competition for light was more severe. Since many calcareous grasslands in Estonia have been abandoned and the standing crop, as well as the cover of shrubs and trees, has increased, sensitivity to light competition may be one reason why V. elatior has become more rare. It cannot be the only reason for its rarity in the whole region, since many other grassland species that are vulnerable to reduced light in overgrown unmanaged grassland communities still occur in much higher numbers in the country. It was hypothesised that historical factors, e.g. relatively late arrival in the region, may also be behind the rarity of V. elatior.     

Isolation and characterization of microsatellite markers for Viola mirabilis (Violaceae) using a next‐generation sequencing platform

We isolated and characterized microsatellite loci in Viola mirabilis (Violaceae), an endangered species from South Korea. Twenty‐three polymorphic microsatellite loci were developed and tested in Korean, Chinese and Japanese populations. The number of alleles per locus varied from two to eight. The observed and expected heterozygosities within the three populations were 0.000–0.625 and 0.469–0.695, respectively. A total of six loci in the Korean population, one locus in the Chinese population and seven loci in the Japanese population deviated from Hardy–Weinberg equilibrium. We expect that these newly developed microsatellite markers will contribute to understanding the phylogeography and population genetics of V. mirabilis, which will aid in developing conservation strategies for this species.     

Phenetic analysis shows conspecificity between Hispaniolan Viola domingensis Urban and North American Viola macloskeyi sensu lato (Violaceae)

Phenetic investigations were undertaken to evaluate differences between the cloud forest endemicViola domingensis of the Dominican Republic and the morphologically similarV. macloskeyi of continental North America. Cluster, principal components (PCA), and canonical variates analyses (CVA) of leaf characters and of ratios derived from them consistently separateV. macloskeyi subsp.macloskeyi of the Pacific states fromV. domingensis and transcontinentalV. macloskeyi subsp.pallens. However, all analyses portray the latter two as extensively intergradient extremes of a continuum, withV. domingensis plants tending to be smaller. Phenetic analyses of capsule and seed features, which are often divergent among other species of subsect.Stolonosae, showV. domingensis to be nested within a broaderV. macloskeyi subsp.pallens. Similarly, floral morphology does not provide observable differences between the pair. Results support the synonymy ofV. domingensis underV. macloskeyi subsp.pallens and reveal that it represents a disjunct outlier of an otherwise continental North American violet.     

Two new species of Viola (Violaceae) from the Intermountain West,U.S.A.

Viola lithion andV. franksmithii are rock-dwelling endemics that belong to the North American perennial, caulescent, simple-leaved, blue-, purple-, or white-flowered group of species.Viola lithion, from the White Pine and Pilot mountain ranges of eastern Nevada and adjacent Utah, belongs to sectionChamaemelanium subsectionCanadenses. It most closely resembles the reniform-leaved, Olympic MountainV. flettii, together with which it possibly shares an ancestry with the widespread and allopatricV. canadensis. Viola franksmithii, restricted to the Logan Canyon drainage of the Bear River Range in northern Utah, belongs to sectionRostellatae subsectionRosulantes. It comes closest morphologically toV. howellii of the Coast Range, but likely evolved from the widespreadV. adunca or a common ancestor.     

Variation in life-cycle between three rare and endangered floodplain violets in two regions: implications for population viability and conservation

We studied the demography of Viola elatior, V. pumila, and V. stagnina, three rare and endangered Central European floodplain species, to (i) analyse variation in life-cycles among congeners and between regions (Dyje-Morava floodplains, Czech Republic; Upper Rhine, Germany), (ii) to define sensitive stages in the life-cycles, and (iii) to identify possible threats for population viability and species conservation. Matrix models were based on the fate of marked individuals from a total of 27 populations over two years. We analysed population growth rate (λ), stage distribution, net reproductive rate (R ₀), generation time, age at first reproduction, and elasticity and calculated a life table response experiment (LTRE). Most populations were declining and λ did not differ between species or regions during the observed interval. Despite higher probabilities for survival and flowering in the Dyje populations, R ₀ was higher in the Rhine populations. Also other demographic traits showed consistent differences between regions and/or species. Complex life-cycles and large variation in λ precluded unequivocal identification of sensitive stages or vital rates for conservation. Variation between regions may be a consequence of differences in habitat quality. Our results suggest that deterministic processes such as reduced management, succession, habitat destruction, and lack of disturbance through reduced or eliminated flooding present the strongest threat for the viability and persistence of populations of the three floodplain violets as compared with stochastic processes. However, the persistent seed bank of the species may buffer populations against environmental variation and represents a reservoir for recovery after resumption of suitable land-use management.     

Two new stoloniferous species of Viola (Violaceae) from China

Two new stoloniferous species of Viola (Violaceae) from southern China are described and illustrated. Viola nitida is recognizably different from V. fargesii as the plant is evergreen and glabrous throughout, the leaf blade adaxially nitid, base cuneate or shallowly cordate, and margin serrate. Viola maoershanensis is different from V. diffusa as the leaf blade is serrate, base cordate and not decurrent to petiole, the petiole wingless, the flowers larger, the petals bluish violet or pinkish white, and the lower petal obtuse at the apex. The chromosome numbers of the two new species were counted as 2n = 24 (V. nitida) and 2n = 44 (V. maoershanensis). The taxonomic positions of the two species are discussed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 349–356.     

Effects of cleistogamy and pollen source on seed production and offspring performance in three endangered violets

In rare plants that often occur in small or isolated populations the probability of selfing between close relatives is increased as a consequence of demographic stochasticity. The mode of pollination (selfing, outcrossing) may have considerable effects on seed traits and offspring performance and hence potential viability. Since current efforts aiming at the restoration of floodplain grasslands through the transfer of plant material from species-rich source stands may lead to the establishment of initially small populations consisting of founders from different populations, the present paper experimentally investigated the effects of pollen source and floral types (i.e. chasmogamous (CH) and cleistogamous (CL) flowers) on seed traits and offspring performance in three highly endangered violet species (Viola elatior, V. pumila, V. stagnina) of these grasslands. We estimated inbreeding depression and tested the performance of selfed and outcrossed offspring in two microbial environments, i.e. in soil inoculated with (i) non-sterile substrate from the same species (‘home’-conditions) and (ii) sterilised substrate.Plants produced more CL capsules than CH flowers. Pollinator exclusion had only small effects on CH seed production. CL seeds had a significantly lower mass per seed than CH seeds. This may be related to constraints in allocation or environmental conditions. Seedling growth was reduced in plants grown under ‘home’-conditions as compared to control soils. Under ‘home’-conditions, relative fitness of selfed seedlings of V. stagnina was significantly higher than that of crossed progeny. Our results suggest that high genetic differentiation among populations as a consequence of isolation may result in outbreeding depression, e.g., through biochemical or physiological incompatibilities between genes or the breaking of coadapted gene complexes. In V. stagnina, offspring fitness differed considerably between environments, but in general we found no indications for inbreeding depression in these rare species.     

Intraspecific variation in life history traits of Viola brevistipulata (Violaceae) in Hokkaido

Intraspecific life‐history variations of Viola brevistipulata var. brevistipulata were examined with special reference to reproductive features based on quantitative and qualitative monitoring of chasmogamous (CH) and cleistogamous (CL) flowers. We selected 10 populations in southwestern Hokkaido in common deciduous forests, and marked 40–70 individual plants at each population. Marked individuals were monitored every 1–2 weeks. The expression of CH and CL flowers was distinct among the populations. That is, there were populations consisting of plants with only CH flowers, while other populations produced CL flowers after CH flowers with greater shoot longevity This phenomenon was observed even in populations located at the same site. The results suggest that V. brevistipulata var. brevistipulata shows distinct life‐history variations among the populations.     

Evolution, phylogeography, and taxonomy within the Viola alba complex (Violaceae)

 Taxa of the Viola alba complex were investigated using allozymes and morphometry. A taxonomic revision is presented. A wide delimitation of V. alba with only three morphological and geographical subspecies is suggested: (1) ssp. dehnhardtii distributed in the Mediterranean eastwards to Turkey; (2) ssp. alba flanking ssp. dehnhardtii in the north and east; and (3) ssp. cretica endemic to Crete. Ssp. cretica, up to now treated as a separate species, is particularly close to ssp. dehnhardtii. Viola cadevallii (NW Mediterranean) is included in the synonymy of ssp. dehnhardtii. Ssp. scotophylla (S Europe), ssp. thessala (Balkan), V. armena (Turkey), and V. besseri (Caucasus) are reduced to synonyms of V. alba ssp. alba. Viola pentelica (Greece) might represent transitional forms between ssp. alba and ssp. dehnhardtii. Glacial refugia for ssp. alba are suggested from the eastern Mediterranean via Turkey to the Caucasus, for ssp. dehnhardtii in the Mediterranean area in general, and for ssp. cretica in Crete. A key to the subspecies is provided. Taxonomic recombination: Viola alba Bess. ssp. cretica (Boiss. & Heldr.) Marcussen, comb. nov. Received June 17, 2002; accepted November 27, 2002 Published online: March 20, 2003     

云南堇菜属新种

记述了云南产堇菜属3新种,鸡足山堇菜,淡黄堇菜,宾川堇菜。     

Oviposition Site Selection by the Regal Fritillary, Speyeria idalia, as Affected by Proximity of Violet Host Plants

Selection of oviposition sites by female regal fritillary butterflies, Speyeria idalia (Drury), in relation to the location and abundance of their larval food plant, Viola pedatifida G. Don, was studied in Kansas tallgrass prairie. To identify potential cues that females use to select oviposition microsites, we compared plots in which eggs were laid with paired control plots in terms of violet abundance, distance from plot center to the nearest violet plant, plant species composition and relative abundance, plant diversity (Shannon's index), and average plant canopy height. No significant differences occurred between control and oviposition sites for any of these variables (P > 0.15). Violets were present in only 30% of the 1-m-diameter oviposition plots, compared with 24% of the corresponding controls. Furthermore, eggs were never laid directly on violets even when they were present nearby. These data imply that S. idalia females do not preferentially orient to or oviposit on their larval food plant. Nevertheless, female S. idalia did not oviposit randomly, for they invariably chose microsites near the ground that were shaded by the grass/forb overstory and usually deposited eggs on the underside of dead vegetation. We speculate that environmental stresses such as high temperatures and intense solar insolation during the egg stage or harsh overwintering conditions experienced by first-instar larvae impact survival and influence choice of oviposition sites. High fecundity (more than 2000 eggs in some individuals) may allow this species to use a conditional, sweepstakes strategy, i.e., producing and spreading many eggs over prairie that harbors host plants to compensate for low egg and larval survivorship.     

The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials

Summary Root proliferation in nutrient-rich soil patches is an important mechanism facilitating nutrient capture by plants. Although the phenomenon of root proliferation is well documented, the specific timing of this proliferation has not been investigated. We studied the timing and degree of root proliferation for three perennial species common to the Great Basin region of North America: a shrub, Artemisia tridentata, a native tussock grass, Agropyron spicatum, and an introduced tussock grass, Agropyron desertorum. One day after we applied nutrient solution to small soil patches, the mean relative growth rate of Agropyron desertorum roots in these soil patches was two to four times greater than for roots of the same plants in soil patches reated with distilled water. Most of the increased root growth came from thin, laterally branching roots within the patches. This rapid and striking root proliferation by Agropyron desertorum occurred in response to N-P-K enrichment as well as to P or N enrichment alone. A less competitive bunchgrass, Agrophyron spicatum, showed no tendency to proliferate roots in enriched soil patches during these two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in the N-P-K and P-enrichment experiments, respectively. The ability of Agropyron desertorum to proliferate roots rapidly may partly explain both its general competitive success and its superior ability to exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America.     

Characterization of newly developed microsatellite loci in the stemmed yellow violet,Viola pubescens (Violaceae)

Viola pubescens is a perennial herb with a mixed mating system involving the production of both open, chasmogamous flowers and closed, cleistogamous flowers. Using a nonradioactive protocol, 12 microsatellite primers that amplified loci with perfect and imperfect repeats were developed in the species. As examined in five populations of V. pubescens using multiplex reactions, all loci exhibited polymorphism with an average of 5.4 alleles. Most primers also amplified in 12 additional Viola species and one cultivar, revealing the possibility of wide applicability across the genus. This is the first known report of microsatellite loci developed in Viola and the Violaceae.     

The invasive shrub,<Emphasis Type="Italic"> Lonicera maackii</Emphasis> , reduces growth and fecundity of perennial forest herbs

Effects of invasive plant species on native plant species are frequently assumed or inferred from comparisons, but rarely quantified experimentally. Such quantification is important to assessing risks and impacts of invasives. We quantified the effects of Lonicera maackii, an exotic shrub invasive in many eastern North American forests, on survival, growth, and reproduction of three perennial herbs: Allium burdickii, Thalictrum thalictroides , and Viola pubescens. We predicted that the spring ephemeral, A. burdickii , would be most impacted, due to early leaf expansion of L. maackii. Field experiments were carried out in two deciduous forest stands, one (Greggs Woodlot, GW) disturbed and the other (Western Woods, WW) relatively undisturbed. In each stand, individual herbs were transplanted into a blocked design of 60 plots where L. maackii was present, absent, or removed, and monitored for 5 growing seasons. Lonicera maackii did not affect survival of transplants, but reduced growth and final size of individuals of all three species. For two of the species, A. burdickii and V. pubescens, L. maackii reduced the proportion of live plants flowering in both stands, and reduced the seed or fruit number per flowering individual in GW. For T. thalictroides the proportion flowering was not affected, but seed number per flowering plant was reduced by L. maackii in both stands. For all three species, cumulative seed production over the course of the study was reduced by L. maackii. Overall, effects on the spring ephemeral, A. burdickii, were similar to effects on the other herbs. Because mortality of these established individuals was not affected, short-term studies might conclude forest herbs are unaffected by invasive shrubs. However, the growth and reproduction impacts documented here suggest that populations are impacted in the long-term.     

中国堇菜属美丽堇菜亚属(堇菜科)植物分类学修订

对中国堇菜属美丽堇菜亚属植物进行了系统学修订,确认中国堇菜属美丽堇菜亚属植物共有6种,列出分种检索表,对《中国植物志》51卷中未记载的3种进行了详细的描述,并对塔城堇菜、深紫堇菜及隐距堇菜进行了绘图。     

Viola lilliputana sp. nov. (Viola sect. Andinium, Violaceae), one of the world's smallest violets, from the Andes of Peru

A new violet species of Viola Sect. Andinium, Viola lilliputana, is described from a single dry puna locality on an extensive intermontane plateau southeast of Cerro Palla Palla in the high Andes of Ayacucho Department in southern Peru. This diminutive rosulate violet is evidently among the smallest in the world and probably one of the smallest terrestrial dicots. It belongs to a distinctive species group with pinnatifid leaves that is endemic to central and southern Peru, including V. hillii, V. membranacea and V. weibelii. The new species is similar to V. weibelii in its large, strongly adnate stipules, elongate leaf lobes and dilated unappendaged style with ventral stigmatic orifice. It differs conspicuously from all other members of the pinnatifid-leaved group in its conduplicate leaf blades, straight, mostly nonoverlapping, oblong-lanceolate to broadly elliptical lobes with obtuse to rounded apices, and large basally fused pedicel bractlets. Despite many new collections of vascular plants from the high Andes of Peru and northern Bolivia in recent decades, this distinctive new species is still known only from its type locality, collected on the Iltis-Ugent expedition from November 1962 to January 1963.     

角堇与大花三色堇染色体核型分析

以2份角堇与4份大花三色堇自交系为试验材料,采用染色体常规压片方法,观察和分析了它们的细胞染色体数目、相对长度、平均臂比等核型指标,以明确两种植物细胞学特点,为分类以及育种提供理论依据。结果表明:(1)2份角堇自交系染色体数目均为2n=2x=26,染色体基数为x=13,染色体核型公式分别为2n=2x=26=8m+12sm+6st、2n=2x=26=4m+16sm+6st,核型不对称系数为67.20%~70.10%,核型分类均属于3B。(2)4份大花三色堇自交系均为四倍体,其中2份(EYO-1-2-1-4、DSRFY-1-1-2)染色体数目为44,核型公式为2n=4x=44=4m+16sm+6st、2n=4x=44=16m+24sm+4st;2份(G10-1-3-1-4、XXL-YB-1-1-1-1)染色体数目为48,核型公式分别为2n=4x=48=8m+20sm+20st、2n=4x=48=4m+36sm+8st,核型不对称系数为66.74%~71.77%,核型分类属于2B、3B。