Neglected native or undesirable alien? Resolution of a conservation dilemma concerning the pool frog Rana lessonae

Introduced species often pose serious threats to biodiversity, but occasionally confusion arises as to whether a species really is introduced or is in fact an overlooked native. A recent UK conservation dilemma has centred on the status of the pool frog Rana lessonae. This species has been the subject of documented introductions from central and southern Europe since the early 1800s, the accepted position being that all UK R. lessonae populations are descended from these introductions. However, a closer examination of early UK literature sources, and recent discoveries of isolated, native R. lessonae populations in Sweden and Norway, led some herpetologists to question whether the species was in fact present as a native at some locations prior to the introductions. Research was initiated along four major lines of enquiry: genetic, bioacoustic, archaeozoological and archival. A high degree of convergence among the genetic and bioacoustic investigations demonstrated that the potentially native UK pool frogs were closely related to Scandinavian frogs, thus ruling out introductions from further south as a potential origin. Subfossil evidence of pool frogs was found from ca. 1000 years before present, demonstrating that the species occurred in the UK prior to known introductions. Archival sources produced no historical support for introductions from northern Europe. The postglacial history inferred for these northern populations is consistent with the known climatic and geographical conditions. Taken together, the evidence for the native status of the pool frog is compelling, and furthermore the UK population appears to be part of a distinct northern clade.     

Morphology,Molecular Genetics,and Bioacoustics Support Two New Sympatric Xenophrys Toads (Amphibia: Anura: Megophryidae) in Southeast China

Given their recent worldwide declines and extinctions, characterization of species-level diversity is of critical importance for large-scale biodiversity assessments and conservation of amphibians. This task is made difficult by the existence of cryptic species complexes, species groups comprising closely related and morphologically analogous species. The combination of morphology, genetic, and bioacoustic analyses permits robust and accurate species identification. Using these methods, we discovered two undescribed Xenophrys species, namely Xenophrys lini sp. nov. and Xenophrys cheni sp. nov. from the middle range of Luoxiao Mountains, southeast China. These two new species can be reliably distinguished from other known congeners by morphological and morphometric differences, distinctness in male advertisement calls, and substantial genetic distances (>3.6%) based on the mitochondrial 16s and 12s rRNA genes. The two new species, together with X. jinggangensis, are sympatric in the middle range of Luoxiao Mountains but may be isolated altitudinally and ecologically. Our study provides a first step to help resolve previously unrecognized cryptic biodiversity and provides insights into the understanding of Xenophrys diversification in the mountain complexes of southeast China.     

Variation of wing shape in the Drosophila virilis species group (Diptera: Drosophilidae)

Wing shape variation was analysed with geometric morphometric methods in 17 laboratory strains, representing 11 closely related species (including two subspecies) of the Drosophila virilis group: D. virilis, D. lummei, D. novamexicana, D. americana americana, D. americana texana, D. montana, D. lacicola, D. flavomontana, D. borealis, D. littoralis, D. ezoana and D. kanekoi. Overall shape estimated using Procrustes coordinates of 14 landmarks was highly variable among strains and very similar in females and males. The landmarks in the distal part of the wing showed higher variation across strains than those in the proximal part. Procrustes distances between species were not consistent with phylogenetic distances previously suggested for the virilis group. Moreover, Procrustes distances between strains within species and within two major phylads (virilis and montana) were comparable with those between species and between phylads, respectively. The most different from other members of the group was the endemic D. kanekoi species, currently viewed as separate subphylad within the montana phylad. Allometric effects were found to be partly responsible for shape differences between the strains. Three most significant shape transformations were considered using the relative warp analysis and the strains were ordinated in accordance with transformation values. The pattern of relative warp scores could be easily interpreted only for the third warp explaining about 13% of shape variation. It separated the largest species, D. montana, D. ezoana and D. kanekoi, from other ones and was mainly associated with shape changes in the proximal region of the wing. The results of the present work suggest that wing shape in the virilis species group is not related to the speciation process. The observed proximal‐distal contrasts and allometric effects are in agreement with data of other studies, in which wing shape variation was analysed within Drosophila species.     

Stellaria nemorum L. andS. montana Pierrat (Caryophyllaceae) in the Forest Communities of Slovenia

Morphological and coenological differences between closely related speciesStellaria nemorum L. andS. montana Pierrat are described.Stellaria montana is frequent in Slovenia and occurs mostly in the montane belt (altitude 600–1200 m). It has been recorded in thirty two forest communities (associations or lower syntaxa). Although widely distributed in EuropeS. nemorum is more rare in Slovenia. It occurs mostly in upper-montane (altimontane) and subalpine belt (altitude 1200–1600 m) in ten forest or shrubby communities. Both,S. nemorum andS. montana are valuable diagnostic species for certain associations. They characterize sites with fresh soil, rich in nitrogen. As an example we present two syntaxa of beech forests of Slovenia, in which the species of theS. nemorum group have high constancy and cover value.     

Morphological analysis of male mating organ in the Drosophila virilis species group: a multivariate approach

The shape of the male mating organ differs among 11 closely related species of the Drosophila virilis species group. Multivariate analyses of variation of a suite of 35 morphological traits (indices) describing the phallus shape were carried out in order to characterize interspecies variability of the traits. An overwhelming majority of the traits displayed species‐specific variability. The main result of the investigation was the revelation of the differences involved in the traits studied in the evolution of the D. virilis species group. The structure of species‐specific variability of some traits was discovered to be in accordance with the generally accepted taxonomy of the species group, while that of other traits required isolation of D. virilisper se from lummei phylad (former virilis phylad), and confirmed separation of montana phylad into three subphylads: montana proper, littoralis and kanekoi. Several subsets of traits having separate variability were determined in different parts of the male mating organ which correspond to spots of evolutionarily significant variability.     

A comparison of pollinator fig wasp development in figs of Ficus montana and its hybrids with Ficus asperifolia

Figs (Moraceae) and pollinator fig wasps (Hymenoptera: Agaonidae) have a highly specific mutualistic relationship but fig wasps occasionally enter atypical hosts, and this can lead to hybrid fig trees and the potential for gene flow between species. Many fig trees are dioecious, with fig wasp offspring developing in galled ovules inside figs on male trees, whereas seeds develop only in figs on female trees. We generated experimental hybrids between the Asian Ficus montana Blume and a closely related African species Ficus asperifolia Miquel. Male F1s were sterile if entered by Kradibia tentacularis (Grandi) (Agaonidae), the pollinator of F. montana, because its offspring always failed to develop, without ovule enlargement. As with the F1s, figs on most male backcross plants [F. montana × (F. montana × F. asperifolia)] also aborted shortly after pollinator entry, resulting in a higher turnover of figs than with F. montana, although the times taken for the figs to reach receptivity were similar. Pollinator larvae nonetheless consistently managed to develop inside the figs of one backcross plant and also occasionally in a few figs from another backcross individual. In these figs, galled ovules developed as normal, whereas in figs that aborted the galled ovules failed to enlarge. The sex ratio of K. tentacularis progeny in the backcross figs was female biased and did not differ from that in F. montana figs. Sycoscapter spec. (Hymenoptera: Pteromalidae), a parasitoid of K. tentacularis, was able to lay eggs and developed normally inside male backcross figs where its host was present.     

One- and two-dimensional polyacrylamide gel analysis of the heat shock proteins of the virilis group of Drosophila

The heat shock proteins of the virilis group of Drosophila are analyzed by one- and two-dimensional polyacrylamide gel analysis. This group consists of the two closely related but distinct virilis and montana phylads. The analysis reveals that some of the heat shock proteins are highly conserved among the two phylads while others are not. The 83-, 72-, and 69-kdalton proteins comigrate in all species examined. There is, however, a noticable trend toward greater molecular weight variability in the smaller heat shock proteins. In general, the heat shock protein patterns within each phylad follow the proposed phylogenetic relationships with some exceptions. D. ezoana and D. littoralis, both members of the montana phylad, exhibit heat shock protein patterns more similar to those of the virilis phylad. The data also demonstrate that the montana phylad has almost two times the heat shock allele members that the virilis phylad has. It is also shown that F₁ and F₂ hybrid flies of crosses between Drosophila species having different patterns of heat shock proteins show Mendelian segregation of alleles. After several generations of inbred growth, however, the pattern of heat shock protein synthesis in reciprocal hybrids each resembles that of the paternal parent. The implications of these findings are discussed.This research was supported in part by Damon Runyon-Walter Winchell Grant DRG-233F to R.M.S. and NIH Grant GM 27611 to R.V.S. R.V.S. is the recipient of an NIH Research Career Development Award.     

Effect of nitrogen, water and neighbor density on the growth of Hesperis matronalis and two native perennials

Disturbance is a well-recognized catalyst of exotic species invasion, depriving or releasing resources into communities and favoring the spread of some invaders. Hesperis matronalis (dame’s rocket) is widespread in North America and has potential to become a major problem in natural communities due to anthropogenic influences. We used a target-neighbor design in a greenhouse to grow H. matronalis and native seedlings (Campanula rotundifolia and Muhlenbergia montana) at various levels of neighbor density, nitrogen and water. H. matronalis clearly reduced C. rotundifolia and M. montana aboveground growth and maintained its competitive advantage across all treatments. We expected H. matronalis to maximize its growth and have the greatest negative effect on native species under high resource conditions because ruderal species are able to take advantage of excess resources. H. matronalis demonstrated a clearly negative effect on native species, but a particular ability to outcompete native species at high resource levels depended on resource and native species identity. We also expected that the native species would not exploit excess resources as well as H. matronalis, instead growing better under ambient or low resource conditions because of local adaptation. Rather, M. montana benefited from high water inputs, inconsistent with plant strategies characteristic of stress tolerators. Information on the effects of H. matronalis on native plants in a controlled setting may aid land managers to understand its potential effects in natural communities.     

Pollen morphology of Stachys (Lamiaceae) in Iran and its systematic implication

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.     

The barriers to oceanic island radiation in bryophytes: insights from the phylogeography of the moss Grimmia montana

Aim In contrast to angiosperms, bryophytes do not appear to have radiated in Macaronesia and the western Mediterranean. We evaluate if: (1) the apparent lack of radiation in bryophytes reflects our failure to recognize cryptic endemic species; (2) bryophytes are characterized by extremely low evolutionary rates; or (3) bryophytes have a high dispersal ability, which prevents genetic isolation. Location Worldwide, with a special emphasis on Macaronesia and the western Mediterranean. Methods Three chloroplast regions were sequenced from samples of the moss Grimmia montana from its entire distribution range. Network analyses, Fst and Nst statistics were used to describe and interpret the phylogeographical signal in the data. Results Despite significant phylogeographical signal in the chloroplast genome, which demonstrates limits to gene flow at the continental scale, repeated sister group relationships observed among accessions from different geographical areas suggest recurrent colonization patterns. These observations are consistent with mounting evidence that intercontinental distributions exhibited by many bryophyte species result from long‐distance dispersal rather than continental drift. Madeiran and western Mediterranean island haplotypes are either shared by, or closely related to, European and North American ones. Fst values between Madeira, western Mediterranean islands, North America and Europe are not significantly different from zero, and suggest that Madeira and the south‐western Mediterranean are subject to strong transatlantic gene flow. By contrast, haplotypes found in the Canary Islands are shared or closely related to those of populations from south‐western Europe or southern Africa. Main conclusions Multiple origins and colonization events are not consistent with the hypothesis of a relictual origin of the Macaronesian moss flora. One possible reason for the failure of taxa that experienced multiple colonization events to radiate is niche pre‐emption. We suggest that strong gene flow, coupled with the occupancy of all suitable niches, either by earlier conspecific colonizers or by other species, could be the mechanism preventing island radiation in G. montana and other cryptogams with high long‐distance dispersal abilities.     

Light acclimation of four native tree species in felling gaps within a tropical mountain rainforest

Ecuadorian mountain rainforests are declining dramatically due to deforestation. Exploitation of remaining forests has led to low abundances of native, valuable timber species. Enrichment planting of selected native tree species into forest gaps is a strategy that may increase their abundance and maintain biodiversity. However, the development of successful planting strategies requires knowledge of environmental demands on, and ecological requirements of, native species during their establishment. This knowledge is currently lacking for mid- and late-successional species in Central American forests. Two deciduous, mid-successional (Cedrela montana, Tabebuia chrysantha) and two evergreen, late-successional native tree species (Nectandra membranacea, Podocarpus sprucei) were planted into felling gaps. Photosynthetic performance and growth in height of these species were assessed along light gradients during seedling establishment to test whether species-specific light responses were related to plant successional traits. Both mid-successional species benefited from higher light levels in gaps up to 30% canopy openness_60°. In larger gaps, C. montana exhibited a significant decline in growth. As expected, growth of the late-successional species was only marginally increased at higher light levels. Nevertheless, the photosynthetic apparatus of N. membranacea displayed rapid acclimation to higher light conditions in gaps. Plant response to felling gaps may not always be predicted based on successional status. Our results suggest that the four investigated species may coexist in the same gap by occupying different niches along light gradients. This arrangement may offer an ecological basis to increase the abundance of valuable timber species through enrichment planting in Ecuador mountain rainforests.     

Two new European species of <Emphasis Type="Italic">Docosia</Emphasis> (Diptera: Mycetophilidae)

Two new species of Docosia Winnertz, 1863 are described, Docosia chandleri sp. n. from Austria and Italy and Docosia matilei sp. n. from France and Italy. Both species resemble Docosia montana Laštovka et Ševčík, 2006 and they are characterized by bare laterotergites, yellow legs and long flagellomeres.     

First report of Culicoides boyi (Diptera: Ceratopogonidae) in France

Culicoides boyi Nielsen, Kristensen & Pape 2015 (Diptera: Ceratopogonidae), described in Denmark in 2015, is reported in France. This species is closely related to Culicoides pulicaris (Linnaeus 1758) and can only be distinguished from this species by a subtle variation in spot wing pattern and by calculation of maxillary palpal ratio.     

西藏东南部发现桐庐铁线莲

毛茛科的桐庐铁线莲(Clematis tongluensis)原知分布于尼泊尔、不丹、印度东北部和孟加拉,2019年夏天在中国西藏东南部被发现.桐庐铁线链与绣球腾(Clematis montana)在亲缘关系上接近,两者的区别在于桐庐铁线莲的萼片呈长圆形,顶端尾状渐尖和表面被毛,背面无毛.     

Contrasting needs of grassland dwellers: habitat preferences of endangered steppe beetles (Coleoptera)

Temperate grasslands are local biodiversity hotspots. In Europe, their extent was mostly reduced to isolated habitat patches, whose biota is subject to extinction debt. Knowledge on requirements of dry-grassland inhabitants is thus vital to slow down decline of grassland biodiversity. We studied habitat requirements of eight flightless steppe beetles, including some of the most endangered dry-grassland specialists of the continent. The beetles were sampled using 167 pitfall traps at a pannonian dry-grassland fragment, the Pouzdrany steppe, SE Czech Republic, from March to November 2006. The number of each species captures in each trap was related to vegetation and abiotic habitat characteristics; captures of all beetles were related to each other. Two of the studied species required relatively humid microhabitats, including tall-grass steppe with litter (Carabus hungaricus, Carabidae) and grassland of high herb cover (Meloe proscarabaeus, Meloidae). Others were associated with xeric habitats (e.g. Meloe scabriusculus) and their early-successional stages, including short-turf vegetation (Dorcadion fulvum, D. pedestre, Cerambycidae) and/or bare-ground patches (Blaps lethifera, Tenebrionidae; Meloe decorus, M. uralensis). Our findings point to key importance of early-successional vegetation for grassland biodiversity, and to the fact that locally co-occurring and closely related grassland specialists may exhibit contrasting habitat needs. Spatially and temporarily highly diversified patch management creating a fine scale mosaic of various seral stages from bare soil to tall-grass steppe is therefore the most appropriate approach for managing isolated grasslands. Prescribed burning and support of burrowing herbivores are recommended and discussed together with other measures for restoration of habitat diversity in dry-grassland fragments.     

Ecological Diversity and Biodiversity as Concepts for Conservation Planning: Comments on Ricotta

Ricotta argues against the existence of a unique measure of biodiversity by pointing out that no known measure of α-diversity satisfies all the adequacy conditions that have traditionally been set for it. While that technical claim is correct, it is not relevant in the context of defining biodiversity which is most usefully measured by β-diversity. The concept of complementarity provides a closely related family of measures of biodiversity which can be used for systematic conservation planning. Moreover, these measures cannot be replaced by summary statistics but must rely on inventories of biodiversity surrogates at candidate sites for conservation.     

Identification of cryptic species of Miniopterus bats (Chiroptera: Miniopteridae) from Madagascar and the Comoros using bioacoustics overlaid on molecular genetic and morphological characters

The number of Miniopterus bat species on Madagascar and the nearby Comoros islands (Malagasy region) has risen from four to 11. These recently described cryptic taxa have been differentiated primarily based on molecular markers and associated a posteriori morphological characters that corroborate the different clades. Members of this Old World genus are notably conservative in morphology across their range. Several sites on Madagascar hold up to four small‐bodied taxa of this genus that are morphologically similar to one another, although they can be distinguished based on the tragus, an ear structure associated with echolocation. Miniopterus often emit species‐specific calls. In the present study, we analyze the bioacoustics of the 11 species of Miniopterus currently recognized from the Malagasy region, with an initial identification of the 87 recorded and collected individuals based on molecular markers and certain morphological characters. In most cases, bioacoustic parameters differentiate species and have taxonomic utility. Miniopterus griveaudi populations, which occur on three islands (Madagascar, Anjouan, and Grande Comore), showed no significant differences in peak echolocation frequencies. After running a discriminant function analysis based on five bioacoustic parameters, some mismatched assignments of Malagasy species were found, which include allopatric sister‐taxa and sympatric, phylogenetically not closely‐related species of similar body size. Because the peak echolocation frequencies of two species (Miniopterus sororculus and Miniopterus aelleni) were independent of body size, they were acoustically distinguishable from cryptic sympatric congeners. The small variation around the allometric relationship between body size and peak echolocation frequency of Malagasy Miniopterus species suggests that intraspecific communication rather than competition or prey detection may be the driver for the acoustic divergence of these two species. Our well‐defined echolocation data allow detailed ecological work to commence aiming to test predictions about the relative roles of competition, prey availability, and social communication on the evolution of echolocation in Malagasy Miniopterus species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 284–302.     

Composition of Leaf n‐Alkanes in Three Satureja montana L. Subspecies from the Balkan Peninsula: Ecological and Taxonomic Aspects

The composition of the epicuticular leaf n‐alkanes of eight populations of three Satureja montana subspecies (S. montana L. subsp. pisidica (Wettst.) ?ili? , S. montana L. subsp. montana, and S. montana L. subsp. variegata (Host ) P. W. Ball ), from central and western areas of the Balkan Peninsula was characterized by GC‐FID and GC/MS analyses. In the leaf waxes, 15 n‐alkane homologs with chain‐lengths ranging from C₂₁ to C₃₅ were identified. The main n‐alkane in almost all samples was n‐nonacosane (C₂₉), but differences in the contents of three other dominant n‐alkanes allowed separating the coastal from the continental populations. The diversity and variability of the epicuticular‐leaf‐n‐alkane patterns and their relation to different geographic and bioclimatic parameters were analyzed by several statistical methods (principal component, discriminant, and cluster analyses as well as the Mantel test). All tests showed a high correlation between the leaf n‐alkane pattern and the geographical distribution of the investigated populations, confirming the differentiation between S. montana subsp. pisidica and the other two subspecies. The S. montana subsp. variegata and S. montana subsp. montana populations are geographically closer and their differentiation according to the leaf‐n‐alkane patterns was not clear, even though there was some indication of discrimination between them. Moreover, most of the bioclimatic parameters related to temperature were highly correlated with the differentiation of the coastal and the continental populations.     

A biometric study of Fritillaria montana Hoppe ex W.D.J. Koch s.l. (Liliaceae) shows a single polymorphic species,with no infraspecific taxa

Abstract

The large amount of morphological variability within Fritillaria montana Hoppe ex W.D.J. Koch, a southeastern European species described from northeastern Italy, led to the creation of several further taxa: Fritillaria caussolensis Goaty & Pons ex Adoino from southeastern France, Fritillaria orsiniana Parl. from central Italy, Fritillaria intermedia N. Terracc. and Fritillaria pollinensis N. Terracc. from southern Italy. Aiming to test the taxonomic value of these taxa, a biometric study of both herbarium and living specimens of F. montana s.l. is carried out. A total of 22 morphological features were analysed and measured in 417 plants from 46 different localities, including loci classici of all the involved taxa. In addition, typical populations of F. montana, F. caussolensis, F. intermedia, F. orsiniana and F. pollinensis and also several other Italian populations resulted with 2n = 18 chromosomes. We also counted 2n = 27 chromosomes in endotriploid cells of plants from Abruzzo. However, there is no clear‐cut correlation between morphology and karyology. The biometric analysis, together with many observations on fresh material in the field, revealed that F. montana is a single polymorphic species with no infraspecific taxa.     

Correllia (Compositae: Peritylinae) : A new monotypic genus from southern Chihuahua,Mexico

A new genus,Correllia, and its single species,C. montana, are described from Sierra Mohinora in southwestern Chihuahua, Mexico. The genus belongs in subtribe Peritylinae (Compositae tribe Helenieae) and appears to be most closely related toPerityle.