Phylogeographic analyses reveal distinct lineages of the liverworts Metzgeria furcata (L.) Dumort. and Metzgeria conjugata Lindb. (Metzgeriaceae) in Europe and North America

Seed plant genera often exhibit intercontinental disjunctions where different species are found on different continents. Many morphologically circumscribed bryophyte species exhibit similar disjunctions. We used nucleotide sequences from the plastid and nuclear genomes to test hypotheses of phylogeography within representatives of the genus Metzgeria: Metzgeria furcata, Metzgeria conjugata, and Metzgeria myriopoda. The first two species have sexual and asexual populations, exhibit disjunctions between North America and Europe, and have been split into separate species, numerous subspecies or varieties. The third species occurs in eastern North America but is not reported from Europe. Phylogenetic analyses resolved three distinct lineages within the morphologically defined species, M. furcata: one in North America, and two in Europe. Similarly, three morphologically cryptic clades of M. conjugata were resolved by the molecular data: northern North America, Europe, and south‐eastern North America. For both species, molecular divergence among taxa occurred in the absence of morphological change. In the case of M. myriopoda, all plants from eastern North America were both morphologically uniform and genetically homogeneous (although not identical). The present study provides significant insight into a plant group with complex taxonomy, and indicates that these liverwort taxa with wide distributions, extreme sex ratios, and continental disjunctions harbor cryptic lineages. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 745–756.     

Systematics and macroevolution of extant and fossil scalopine moles (Mammalia,Talpidae)

Scalopini is one of the two fully fossorial mole tribes in the family Talpidae, with remarkable adaptations to subterranean lifestyles. Most living Scalopini species are distributed in North America while a sole species occurs in China. On the other hand, scalopine fossils are found in both Eurasia and North America from upper Oligocene strata onwards, implying a complex biogeographical history. The systematic relationships of both extant and fossil Scalopini across North America and Eurasia are revised by conducting phylogenetic analyses using a comprehensive morphological character matrix together with 2D geometric–morphometric analyses of the humeral shape, with a specific emphasis on Mioscalops, a genus commonly found in North America and formerly known as Scalopoides. Our phylogenetic analyses support the monophyly of the tribe Scalopini as well as a proposed two‐subtribe‐division scenario of Scalopini (i.e. Scalopina and Parascalopina), although Proscapanus could not be assigned to either subgenus. Our geometric–morphometric analyses indicate that the European Mioscalops from southern Germany should be allocated to Leptoscaptor, which in turn implies that Mioscalops may be endemic to North America and never arrived in Europe. Examination of biogeographical patterns does not unambiguously determine the geographical origin of Scalopini. Nevertheless, it does support multiple transcontinental colonization events across Asia, Europe and North America. Scapanulus oweni, distributed in central China, is the only remaining representative of one of those out‐of‐North‐America migrations, whereas scalopine moles are common in North America nowadays with up to five species.     

Molecular phylogeny of the Sphaerophorus globosus species complex

The Sphaerophorus globosus complex (Lecanorales, lichenized Ascomycota) shows a large morphological variation, and three relatively distinct morphotypes can be distinguished in parts of the distribution area. Here, we utilize a multigene‐based maximum‐parsimony approach (nITS+ LSU rDNA, mtSSU rDNA, β‐tubulin, and actin) to investigate whether these morphotypes constitute distinct species. The results show that there are at least two well‐supported monophyletic groups that we interpret as phylogenetic species within the S. globosus complex. These species do not completely correspond to the predefined morphotypes. One group, an apparently undescribed species, contains noncoralloid specimens from the North American Pacific Northwest. The other group, S. globosus, consists of two well‐supported monophyletic groups: one contains coralloid epiphytic specimens from the North American Pacific Northwest that are morphologically indistinguishable from epiphytic specimens from Europe and are presently interpreted as belonging to the same species and the other is morphologically variable and contains terrestrial specimens from Europe, North America, and southernmost South America and coralloid epiphytic and epilithic specimens from Europe. The results suggest that the population in southernmost South America originated by long‐distance dispersal from arctic populations in the Northern Hemisphere.     

Species delimitation and invasion history of the balsam woolly adelgid,Adelges (Dreyfusia) piceae (Hemiptera: Aphidoidea: Adelgidae), species complex

The Adelges (Dreyfusia) piceae (Ratzeburg) species complex is a taxonomically unstable group of six species. Three of the species are cyclically parthenogenetic [Ad. nordmannianae (Eckstein), Ad. prelli (Grossmann), and Ad. merkeri (Eichhorn)] and three are obligately asexual [Ad. piceae, Ad. schneideri (Börner), and Ad. nebrodensis (Binazzi & Covassi)]. Some species are high‐impact pests of fir (Abies) trees, so stable species names are needed to communicate effectively about management. Therefore, to refine species delimitation, guided by a reconstruction of their biogeographic history, we genotyped adelgids from Europe, North America, and the Caucasus Mountains region with 19 microsatellite loci, sequenced the COI DNA barcoding region, and compared morphology. Discriminant analysis of principal components of microsatellite genotypes revealed four distinct genetic clusters. Two clusters were morphologically consistent with Ad. nordmannianae. One of these clusters consisted of samples from the Caucasus Mountains and northern Turkey, and the other included samples from this region as well as from Europe and North America, where Ad. nordmannianae is invasive. A third cluster was morphologically consistent with Ad. piceae, and included individuals from Europe, where it is native, and North America, where it is invasive. In North America, the majority of Ad. piceae individuals were assigned to two geographically widespread clones, suggesting multiple introductions. The fourth cluster included individuals morphologically consistent with Ad. prelli or Ad. merkeri. However, based on genetic assignments, hybrid simulations, and approximate Bayesian computation, we find it likely that these are contemporary hybrids between Ad. nordmannianae and Ad. piceae that arose independently in Europe and North America, so we propose that Ad. prelli and Ad. merkeri are invalid. Finally, we synonymise Ad. schneideri (syn.n.) with Ad. nordmannianae and designate Ad. nebrodensis as subspecies Ad. piceae nebrodensis (stat.n.) . Our revised taxonomy therefore recognises two species: Ad. nordmannianae and Ad. piceae, which we estimate to have diverged recently, during one of the last two interglacial periods. Finally, we comment on this species complex being in the midst of transition between sexual and asexual reproduction, a pattern that is probably common in Adelgidae.     

The historical ecology of yellow perch (Perca flavescens[Mitchill]) and their parasites

Aim To determine the origins of the host–parasite association between among yellow perch (Perca flavescens[Mitchill]) and the parasites Crepidostomum cooperi Hopkins, Proteocephalus pearsei La Rue and Urocleidus adspectus Beverly Burton. Of secondary interest are the parasites Bunodera luciopercae (Muller) and Proteocephalus percae (Muller) predictably associated with the Eurasian perch. Location The areas considered are the Holarctic, since the upper‐Cretaceous, and contemporary North America. Methods Published and new information from host and parasite phylogenies, palaeontology, palaeogeography and plate tectonics and host biology is incorporated to assess the origins of yellow perch and several of its parasites. This information is used to determine the origins for these host–parasite associations. Results Cladistic analysis suggests a Laurasian origin for Percidae and Perca, and that Perca is sister to the other genera in the family. Parasite phylogenies support a North American origin for the three species associated with yellow perch and a Laurasian origin for B. luciopercae. Proteocephalus pearsei and P. percae are not sister taxa. The fossil record for Perca dates to the Miocene in Europe and the Pleistocene in North America. North America and Europe were connected across the North Atlantic since at least the upper Cretaceous with separation complete by the Miocene. Europe was separated from Asia by the Obik Sea from the late Cretaceous until the Oligocene. Western cordillera orogeny and its accompanying high rates of water flow and Pleistocene glaciation represent barriers to Perca dispersal. Main conclusions The origin of Perca in North America dates at least to the late Oligocene when North America and Europe were connected across the North Atlantic and Europe and Asia were separate landmasses, and does not result from Pleistocene dispersal across Beringia from Asia. The present disjunction of Perca species in North America and Europe is due to the vicariant separation of North America and Europe. Based on the available information, yellow perch and its parasites have a North America origin. The association between yellow perch and the parasites in all cases is a consequence of host switching from other sympatric host species in North America and is not explained by co‐speciation. Even the association between the host‐specific Urocleidus adspectus and yellow perch originated with a host switch and is not due to co‐speciation. The basis for this host switching is geographical and ecological sympatry, especially shared feeding habits, with other North American fish hosts.     

A New Small-Bodied Species of <Emphasis Type="Italic">Palaeonictis</Emphasis> (Creodonta,Oxyaenidae) from the Paleocene-Eocene Thermal Maximum

Oxyaenid creodonts are extinct carnivorous mammals known from the Paleogene of North America, Europe, and Asia. The genus Palaeonictis is represented by three species that together span the late Paleocene to early Eocene of North America, and at least one species from the early Eocene of Europe. Previously, only a single trigonid of Palaeonictis was known from the interval encompassing the Paleocene-Eocene Thermal Maximum (PETM) in North America. We describe Palaeonictis wingi sp. nov. from the PETM in the Cabin Fork drainage, southeastern Bighorn Basin, Wyoming, based on associated right and left dentaries with P₂-M₂. Palaeonictis wingi sp. nov. is substantially smaller than the other North American congeners, making it similar in size to P. gigantea from the earliest Eocene of Europe and the previously described PETM specimen. We suggest that a form similar to the large-bodied late Paleocene P. peloria from North America gave rise to two smaller species in the earliest Eocene of North America (P. wingi) and Europe (P. gigantea). Palaeonictis wingi may have given rise to P. occidentalis following the PETM in North America. Dispersal of Palaeonictis to Europe coincided with rapid global warming of 5–10°C and related geographic range shifts in plants and other animals during the PETM. It has been suggested that certain mammalian lineages decreased in body size during the PETM, possibly in response to elevated temperature and/or higher CO₂ levels. Results from a dietary analysis of Palaeonictis indicate that it was an omnivore that primarily consumed meat. This suggests that the decreased nutritious quality of vegetation caused by increased CO₂ levels was not the direct contributing factor that caused body size reduction of this lineage during the PETM. Other selective pressures such as temperature, aridity, and prey size may have also contributed to the smaller body size of carnivorous mammals during this interval, although the presence of smaller species could also be explained by latitudinal range shifts of mammals during the PETM.     

Genetic variation and biogeography of the disjunct Vitis subg. Vitis (Vitaceae)

Aim Vitis subg. Vitis provides an example of a plant disjunction occurring in the Northern Hemisphere. It shows broad morphological variation but is assumed to be a species complex with limited genetic differentiation. Based on a comprehensive sampling of taxa and polymorphism in both chloroplast and nuclear DNA, we assessed genetic variation within this subgenus. Our aims were to clarify the relationships among species and to examine their historical biogeography. Location Asia, Europe, North America. Methods We analysed a total of 30 species and putative hybrids from subgenus Vitis and examined the infra‐specific variation in some species. Polymorphism in chloroplast DNA was assessed in trnL and trnH–psbA–trnK sequences (c. 2170 bp) and in 15 microsatellite loci. We also obtained nuclear data for size variation at 24 microsatellite loci. Phylogenetic inference was performed with Bayesian analyses. A maximum parsimony network was constructed to depict the evolutionary relationships among haplotypes, and microsatellite data were also subjected to hierarchical clustering analysis using the Ward distance. In addition, we assessed size homoplasy by sequencing both chloroplast and nuclear microsatellite loci. Results Chloroplast polymorphisms resolved subgenus Vitis as a monophyletic group with limited genetic variation. The ancestral haplotypes were found in Eurasia. American taxa all harboured derived haplotypes. Most of them formed a monophyletic group that did not include Vitis californica. The four main haplotypes in Vitis vinifera corresponded to two different origins. Nuclear microsatellites indicated that genetic variation was especially large in North America. Asian species exhibited a lower level of nuclear divergence and the European V. vinifera corresponded to a differentiated nuclear lineage. Main conclusions We obtained some evidence that subgenus Vitis has an Asian origin and then dispersed to Europe and North America. Geographic separation was followed by diversification, presumably during the Pleistocene, resulting in phylogeographic patterns similar to other biota. In contrast to chloroplast DNA, nuclear DNA shows a larger than expected genetic variation. Our molecular data also highlight the need to re‐examine certain aspects of the current subgeneric classification.     

Diadromus pulchellus in North America: field release against leek moth and new characters to distinguish it from Diadromus subtilicornis,a native diamondback moth parasitoid

We report successful overwintering of Diadromus pulchellus in North America (Ontario) following introduction of this species from Europe to control the leek moth, Acrolepiopsis assectella, a recently established alien species. Field rearing revealed that the native Diadromus subtilicornis emerged only from diamondback moth, Plutella xylostella, whereas D. pulchellus was reared almost exclusively from leek moth. The single D. pulchellus reared from diamondback moth was anticipated because host range studies found this species could develop on both leek moth and diamondback moth in the laboratory, although, it had not been previously reported from diamondback moth in the field in Europe. DNA barcoding of specimens of both Diadromus spp. confirmed their species status and novel morphological characters are presented to distinguish D. pulchellus from D. subtilicornis. In addition, DNA from specimens of D. subtilicornis from Europe clustered with DNA from specimens across Canada, confirming that it is a single Holarctic species. Finally, a new host association for D. subtilicornis is recorded from the dame's rocket moth: Pseudoplutella porrectella.     

Species and hybrid identification of sturgeon caviar: a new molecular approach to detect illegal trade

Overexploitation of wild populations due to the high economic value of caviar has driven sturgeons to near extinction. The high prices commanded by caviar on world markets have made it a magnet for illegal and fraudulent caviar trade, often involving low‐value farmed caviar being sold as top‐quality caviar. We present a new molecular approach for the identification of pure sturgeon species and hybrids that are among the most commercialized species in Europe and North America. Our test is based on the discovery of species‐specific single nucleotide polymorphisms (SNPs) in the ribosomal protein S7, supplemented with the Vimentin gene and the mitochondrial D‐loop. Test validations performed in 702 specimens of target and nontarget sturgeon species demonstrated a 100% identification success for Acipenser naccarii, A. fulvescens, A. stellatus, A. sinensis and A. transmontanus. In addition to species identification, our approach allows the identification of Bester and AL hybrids, two of the most economically important hybrids in the world, with 80% and 100% success, respectively. Moreover, the approach has the potential to identify many other existing sturgeon hybrids. The development of a standardized sturgeon identification tool will directly benefit trade law enforcement, providing the tools to monitor and regulate the legal trade of caviar and protect sturgeon stocks from illicit producers and traders, hence contributing to safeguarding this group of heavily threatened species.     

FURTHER STUDIES IN THE GENUS DRYOPTERIS: THE ORIGIN OF D. CLINTONIANA,D. CELSA AND RELATED TAXA

Walker , S. (Liverpool Univ., England.) Further studies in the genus Dryopteris: the origin of D. clintoniana, D. celsa and related taxa. Amer. Jour. Bot. 49(5): 497–503. Illus. 1962.—Cytogenetic studies of 5 Dryopteris species, from North America, together with wild and synthesized hybrids between some of them, show that D. clintoniana and D. celsa are allohexaploid and allotetraploid respectively. Cytological and morphological evidence points to D. cristata and D. goldiana being parental to D. clintoniana, whilst D. goldiana and D. ludoviciana are parental to D. celsa. The taxon D. X australis is triploid and has been synthesized; it is the backcross between D. celsa and D. ludoviciana.     

Molecular phylogenetic analysis of Leibnitzia Cass. (Asteraceae: Mutisieae: Gerbera‐complex), an Asian–North American disjunct genus

Abstract Leibnitzia comprises six species of perennial herbs that are adapted to high elevation conditions and is one of only two Asteraceae genera known to have an exclusively disjunct distribution spanning central to eastern Asia and North America. Molecular phylogenetic analysis of Leibnitzia and other Gerbera‐complex members indicates that Leibnitzia is monophyletic, which is in contrast with our expectation that the American Leibnitzia species L. lyrata and L. occimadrensis would be more closely related to another American member of the Gerbera‐complex, namely Chaptalia. Ancestral area reconstructions show that the historical biogeography of the Gerbera‐complex mirrors that of the entire Asteraceae, with early diverging lineages located in South America that were followed by transfers to Africa and Eurasia and, most recently, to North America. Intercontinental transfer of Leibnitzia appears to have been directed from Asia to North America. Independent calibrations of nuclear (ribosomal DNA internal transcribed spacer region) and chloroplast (trnL–rpl32 intron) DNA sequence data using relaxed clock methods and either mean rate or fossil‐based priors unanimously support Miocene and younger divergence times for Gerbera‐complex taxa. The ages are not consistent with most Gondwanan vicariance episodes and, thus, the global distribution of Gerbera‐complex members must be explained in large part by long‐distance dispersal. American species of Leibnitzia are estimated to have diverged from their Asian ancestor during the Quaternary (ca. 2 mya) and either migrated overland to North America via Beringia and retreated southwards along high elevation corridors to their present location in southwestern North America or were dispersed long distance.     

European populations of Diabrotica virgifera virgifera are resistant to aldrin,but not to methyl‐parathion

The western corn rootworm, Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae), is a major pest of cultivated corn in North America and has recently begun to invade Europe. In addition to crop rotation, chemical control is an important option for D. v. virgifera management. However, resistance to chemical insecticides has evolved repeatedly in the USA. In Europe, chemical control strategies have yet to be harmonized and no surveys of insecticide resistance have been carried out. We investigated the resistance to methyl‐parathion and aldrin of samples from nine D. v. virgifera field populations originating from two European outbreaks thought to have originated from two independent introductions from North America. Diagnostic concentration bioassays revealed that all nine D. v. virgifera field populations were resistant to aldrin but susceptible to methyl‐parathion. Aldrin resistance was probably introduced independently, at least twice, from North America into Europe, as there is no evident selection pressure to account for an increase of frequency of aldrin resistance in each of the invasive outbreaks in Europe. Our results suggest that organophosphates, such as methyl‐parathion, may still provide effective control of both larval and adult D. v. virgifera in the European invasive outbreaks studied.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.     

Population genetics of the metabolically related Adh,Gpdh and Tpi polymorphisms in Drosophila melanogaster I. Geographic variation in Gpdh and Tpi allele frequencies in different continents

Among Australasian populations from above 32.5° latitude there is a significant negative relationship between Gpdh ^F frequency and distance from the equator which is not explained by gametic disequilibrium with the linked inversion In(2L)t. This is consistent with the associations reported earlier for Gpdh ^F among populations covering comparable latitudes in North America and Europe/Asia. By contrast, Tpi allele frequencies are found to be significantly associated with distance from the equator in Australasia but not North America or Europe/Asia. The Tpi pattern in the different zones is essentially the same as that reported earlier for the Acph polymorphism, which maps only 0.2 cM away from the Tpi locus.There are now ten enzyme polymorphisms in D. melanogaster which have been screened for latitudinal associations in Australasia, North America and Europe/Asia. Allele frequencies at six of these loci show significant relationships with distance from the equator which are consistent across all three zones. These latitudinal associations are more prevalent for Group II than Group I enzymes. Values of genic heterozygosity averaged over the ten polymorphic loci and eleven other monomorphic systems do not vary with latitude but differ substantially between zones. Values of Nei's genetic distance between North American and European/Asian populations calculated from all 21 systems are equivalent to subspecific differences elsewhere in the genus.     

Population Genetics and Identity of an Introduced Terrestrial Slug: Arion subfuscus s.l. in the North-east USA (Gastropoda, Pulmonata, Arionidae)

Several European species of the terrestrial slug genus Arion have been introduced into North America. A case in point is the species complex A. subfuscus s.l. which has become one of the most abundant slug taxa in North America. In Europe this complex consists of at least two cryptic species, viz. A. fuscus and A. subfuscus s.s., the latter of which is further subdivided in five strongly divergent mtDNA lineages (A. subfuscus S1–S5). In order to determine which of these A. subfsucus s.l. taxa are present in the NE USA and in order to assess their population genetic structure, we compared mtDNA, nDNA and allozyme variation between populations from the NE USA and Europe. Our results show that (1) at least A. subfuscus S1 has become successfully established in the NE USA, (2) founder effects are the most likely explanation for the loss of a large amount of molecular genetic variation in populations from the NE USA (i.e. a loss of 96% of the 16S rDNA haplotypes, 67% of the ITS1 alleles and 46% of the alleles at polymorphic allozyme loci), and (3) part of the remaining genetic variation in NE USA populations was probably due to multiple introductions from the British Isles and the European mainland, and the hybrid structure of most of these source populations. Apparently, the extreme loss of molecular genetic variation in this introduced species has not prevented it from successfully establishing and spreading in novel environments.     

A comparison of Dichodontium flavescens (Dicks.) Lindb. and D. pellucidum (Hedw.) Lindb. (Bryopsida)

Abstract

Fifty-three herbarium samples of the genus Dichodontium, originating from 15 countries in Europe and North America, were examined to critically assess new distinguishing characters between D. flavescens and D. pellucidum. The examination, including SEM, revealed some new characteristics in the gametophyte, notably leaf shape, leaf length: width ratio, and width of the nerve, permitting accurate determinations even on sterile material. D. flavescens is recognized here as a distinct species.     

Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (±)-catechin

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the “novel weapons” hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the “evolution of increased competitive ability” hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (±)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

NewLeptographium species from Indonesia and Eastern North America

Leptographium species have predominantly been described from North America, Canada and Europe. These fungi generally occur on conifers and many cause blue-stain of lumber. MostLeptographium species are also associated with insects and in particular, bark beetles (Coleoptera: Scolytidae). Recently, an unknown species ofLeptographium was isolated from pine infested with anIps sp. in Indonesia. In addition, two unknown species have been collected from red spruce (Picea rubra) and balsam fir (Abies balsamea) roots from high elevation sites in Eastern North America. The latter isolates are unusual in that they are associated with the feeding wounds made by the conifer swift mothKorscheltellus gracilus (Lepidoptera: Hepialidae), which is a habitat unique for species ofLeptographium. Comparison with knownLeptographium species has revealed that the isolates from Indonesia and those from Eastern North America represent three previously undescribed taxa. They are, therefore, described in this study asL. pineti sp. nov,L. abieticolens sp. nov. andL. peucophilum sp. nov.