Ellenberg indicator values for macromycetes – a methodological approach and first applications

Ellenberg indicator values (EIVs) describe the realized niche of species and habitat parameters, and are commonly used for vascular plants, bryophytes, and lichens. We provide a methodology for EIVs and an EIV list for nearly 650 species of macromycetes. We propose a new EIV scale, namely substrate openness (O). We also give the results of two applications and compare EIV values related to the Red List classification with those related to lifestyle classification. Mycorrhizal species on average have higher demands on substrate openness and are less tolerant of high nutrient levels than saprotrophic or parasitic species. Critically endangered species have on average distinctly higher demands on openness of habitat and substrate than not threatened or less strongly threatened species, which in turn have higher demands for nutrient availability. This pattern clearly highlights the points of threat for many macromycete species.     

Lichens on dead wood: species‐substrate relationships in the epiphytic lichen floras of the Pacific Northwest and Fennoscandia

Dead wood is an important habitat feature for lichens in forest ecosystems, but little is known about how many and which lichens are dependent on dead wood. We reviewed substrate use by epiphytic lichens in the combined floras of Fennoscandia and the Pacific Northwest of North America based on literature and herbarium data and analyzed substrate affinity relative to life form, reproductive mode and major phylogenetic group within the floras. A total of 550 (43%) of the 1271 epiphytic species in the combined floras use wood, and 132 species (10%) are obligately associated with dead wood in one or both regions. Obligate and facultative wood‐dwelling guilds in the two floras were strongly similar in terms of internal guild structure in each region, but differ somewhat in species composition, while the bark‐dwelling guild differs strongly in both. Most obligate dead wood users are sexually reproducing crustose lichens. The largest numbers of species are associated with forest structural features such as logs and snags that have been greatly reduced by forest practices. Conservation of lichens inhabiting wood requires greater attention to crustose lichen species and the development of conservation strategies that look beyond numbers and volumes of dead wood and consider biologically meaningful dead wood structure types.     

Habitat selection of riparian birds at restoration sites along the Trinity River,California

Riparian habitats in the western United States are imperiled, yet they support the highest bird diversity in arid regions, making them a conservation priority. Riparian restoration efforts can be enhanced by information on species response to variation in habitat features. We examined the habitat selection of four riparian birds known as management indicators at restoration and reference sites along the Trinity River, California. We compared vegetation structure and composition at nest sites, territories, and random points to quantify used versus available habitat from 2012 to 2015. Vegetation in focal species' territories differed between site types, and from available habitat, indicating nonrandom site choice. Birds selected aspects of more structurally complex habitats, such as greater canopy cover, canopy height, and tree species richness. Yellow‐breasted Chats preferred greater shrub cover, and Yellow Warblers preferred greater cover by non‐native Himalayan blackberry. Territory preferences on restoration sites were often a subset of those on reference sites. One exception was canopy height, which was taller on restoration site territories than random points for all species, suggesting that birds preferentially used patches of remnant habitat. Few variables were significant in nest site selection. Restoration plantings along the Trinity River were only 3–10 years old during this study, and have not developed many of the characteristics of mature riparian habitat preferred by birds, but may improve in habitat value over time. Understanding habitat selection is especially important in recently human‐modified environments, where indirect cues used to assess habitat quality may become disassociated from actual habitat quality, potentially creating ecological traps.     

Movement of translocated turtles according to translocation method and habitat structure

Dispersal away from the release site is among the main obstacles that reduce translocation success. Scientists should therefore test a variety of translocation methods to reduce dispersal when moving wildlife between sites. The objective of this research was to examine how translocation method (hard‐ vs. soft‐release) and habitat structure (continuous vs. patchy) affect movement of translocated turtles. A hard‐release consists of releasing individuals to their new environment without any prior acclimation, whereas a soft‐release forces animals to spend time at the release site prior to release. Our results suggest that the most effective translocation technique depends upon habitat structure. A soft‐release was effective in minimizing post‐release dispersal of translocated turtles in a continuous lotic habitat as there was no difference in the movement of soft‐released and resident turtles. However, hard‐released turtles undergo extensive movement when translocated to a continuous lotic habitat as hard‐released turtles had greater movement than resident turtles. When the release site consists of a patchy wetland complex, a hard‐release translocation may be effective as there was no difference in the movement between resident and hard‐released turtles. Our study suggests that both the habitat structure of the release site and translocation method play a role in the movement patterns of translocated wildlife. Semi‐aquatic turtles or species with poor vagility may make better candidates for hard‐release translocations in patchy habitats because these species may be less likely or unable to disperse long distances as result of their behavior, physiology, or the structure of the release site.     

When is translocation required for the population recovery of old‐growth epiphytes in a reforested landscape?

It is often assumed that species recolonization follows from the restoration of key habitat structure. Thus, forest restoration focuses on the recovery of trees into deforested landscapes, so that a multitude of associated organisms can achieve “colonization credit” and recolonize from remnant source populations into restored habitat. This opportunity for recolonization exists because species vulnerable to habitat loss may experience an “extinction debt,” during which their remnant populations decline only slowly to equilibrium with a deforested landscape. These persistent but declining populations become propagule sources for recolonization. To test limits to “colonization credit,” this study focused on old‐growth dependent lichen epiphytes, using a simulation to identify a hypothetical threshold at which: (1) the number of remnant populations, and (2) their population sizes, are too low to achieve recolonization and population recovery, despite efforts placed into forest restoration. The results show that for a landscape scenario relevant to the industrialized temperate zone, with less than 5% of old‐growth forest remaining, and ambitions for restoration to circa 10–15% forest cover, there is a failure to achieve population recovery over long timescales (i.e. within 600 years), making translocation a necessary option. This delay represents a “colonization deficit” that may be a common feature in ecological restoration more generally.     

Biocrust lichen and moss species most suitable for restoration projects

Reintroducing lichens and mosses to areas slated for restoration or rehabilitation may prove integral to project success by filling the biocrust component (niche) of arid ecosystems. In doing so, it is important to select appropriate species and genetic source material. Some bryophyte and lichen species are early pioneers and are potentially well‐suited for restoration projects. Species traits such as high reproductive rates, rapid establishment rates, and large asexual reproductive propagules can be beneficial for restoration. For instance, the large number of spores produced by some mosses are beneficial for reproductive success in arid environments. In addition to identifying the benefit of reproductive strategies, it is important to take habitat needs into consideration; lichen and moss species that are wide‐ranging both geographically and ecologically are recommended over geographically and edaphically restricted species that occur only in specific habitats, such as calcareous soils. Biocrusts used in specific restoration areas should have similar genetic source material (local provenance), and harsh environmental conditions should be ameliorated.     

Spatial and Temporal Considerations in Restoring Habitat for Wildlife

An accumulated body of theory and empirical evidence suggests that habitat selection by animals is a scale‐dependent, hierarchical process. Hierarchy theory predicts that habitat suitability is influenced by the interaction of factors at multiple spatial scales from the microsite to the landscape and that higher‐order factors impose constraints at lower levels. For instance, large‐scale factors such as landscape context may make a site unsuitable for a species even if the vegetation structure and composition are appropriate. In addition, the spatial arrangement of habitat elements at all scales must be considered when planning restoration efforts. For example, the presence of snags does not ensure that the site will be suitable for snag‐dependent species. The size, age, and spacing of snags and their juxtaposition to other habitat elements must also be considered. Finally, all habitats are dynamic, and therefore the ecological processes that contribute to those dynamics must be maintained or suitable substitutes included in the recovery plan. When considering restoring habitat for wildlife, we recommend that managers: (1) identify the wildlife species they want to target for restoration efforts, (2) consider the size and landscape context of the restoration site and whether it is appropriate for the target species, (3) identify the habitat elements that are necessary for the target species, (4) develop a strategy for restoring those elements and the ecological processes that maintain them, and (5) implement a long‐term monitoring program to gauge the success of the restoration efforts.     

Impacts of Channel Reconstruction on Invertebrate Assemblages in a Restored River

Ecosystem restoration often aims to recreate the physical habitat needed to support a particular life‐stage of a focal species. For example, river channel reconstruction, a common restoration practice along the Pacific coast, is typically used to enhance spawning habitat for adult Chinook salmon, a species experiencing large population declines. These restoration efforts rarely consider, however, that altering spawning habitat could have indirect effects on other life‐stages, such as juveniles, which might occur if, e.g. reconstruction alters the benthic food web. To determine how channel reconstruction impacts benthic macroinvertebrates, juvenile Chinook's primary prey, we conducted two studies at a restoration site in the Merced River, California. We asked (1) has gravel enhancement altered invertebrate assemblages in the restored reach compared with an unrestored reach? and, if so, (2) can shifts in the invertebrate community be explained by increased substrate mobility and by reduced heterogeneity that results from restoration? We show that invertebrate abundance and biomass were lower in the restored reach and that these changes were accompanied by a shift from dominance by filter‐feeding caddisflies (Hydropsyche) in the unrestored reach to grazing mayflies (Baetis) in the restored reach. Using an in situ manipulation, we demonstrated that this trend was driven by increased substrate mobility that reduces the abundance of Hydropsyche and by decreased substrate heterogeneity that reduces the abundance of Baetis. Our studies suggest that geomorphic changes typical of reconstructed rivers can alter food webs in ways that may have important implications for supporting the focal species of restoration efforts.     

The association of oribatid mites with lichens

Oribatei (Acari, Cryptostigmata) are found in a variety of terrestrial habitats, and many are associated with lichens; the relationship ranges from casual to highly dependent. Eighty-three species associated with lichens have been surveyed, and a tentative classification, based on their ecological requirements, is presented: Group A consists of species restricted to lichens as a biotope, though occasionally occurring as accidentals in other habitats; Group B consists of species which while preferring lichens as a habitat and feeding source are also adapted to existence on other plants (though in some cases their immatures may be lichen-restricted); Group C consists of species which, though frequently found on lichens, are equally common in other biotopes, particularly mosses, and must be regarded as much more generalized in their feeding habits. Certain aspects of oribatid-lichen specificity are discussed. The importance of oribatid-lichen associations from the point of view of soil fertility and energetics is emphasized.     

Patterns of biodiverse,understudied groups do not mirror those of the surrogate groups that set conservation priorities: a case study from the Mid-Atlantic Coastal Plain of eastern North America

We conducted biodiversity inventories of lichens, woody plants, and sedges at 32 sites on the Mid-Atlantic Coastal Plain of eastern North America between November 2012 and June 2015. Each site comprised a single, uniform habitat, consisting of either Coastal Plain Floodplain forest, Coastal Plain Flatwood swamp, Coastal Plain Oak-Pine forest, Maritime forest, Mixed Mesic Hardwood forest, or Tidal forest. We compared alpha diversity and community assemblages of each organismal group across the sites, and compared selected minimal reserve sets in order to visualize biodiversity patterns and assess whether specific components of vascular plants (sedges and woody plants) serve as an effective surrogate for lichens. Woody plants provide a direct substrate for lichen growth, but there is no significant correlation between the alpha diversity of these groups. For conserving maximal species richness among the studied groups, lichens outperform the sedges and woody plants as the best surrogate group for building minimum reserve sets, even though vascular plants are more commonly used as a surrogate. Likewise, sedge alpha diversity does not correlate with lichens, or with woody plants. Although no group is an effective indicator for high alpha diversity sites of other organisms, a significant correlation between the community assemblages of lichens and woody plants suggests that protecting varied types of plant communities might serve as a workable surrogate for protecting lichens. The lack of congruence between species richness patterns across organismal groups suggests that the mechanisms that shape patterns of diversity are not identical, and that identifying and incorporating specific biodiversity indicators for understudied groups in conservation policy is necessary to ensure their protection.     

Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

The Value of Animal Behavior in Evaluations of Restoration Success

Animals are key members of ecosystems, contributing to processes like pollination, seed dispersal, and herbivory. Incorporating measures of animal behavior into evaluations of restoration success will provide critical information that is not available from animal species composition and richness estimates derived from the documentation of species presence and absence. Behavioral studies will (1) allow comparisons of the habitat quality of target and reference sites based on behaviors that have fitness consequences for organisms; (2) provide valuable information about reasons for differences in habitat quality; (3) identify critical resources that make a site suitable or not for particular species; and/or (4) provide information on the mechanisms through which species contribute to ecosystem functions. When resources for the evaluation of restoration success are available, practitioners should carefully consider the costs and benefits of the different variables they could quantify. In some cases, it may be more important to compare the behavior of one or a few critical animal species that contribute to ecosystem function rather than try to document the presence or absence of all species.     

Development and application of genomic tools to the restoration of green abalone in southern California

Due to severe declines in abundance throughout southern California, the green abalone (Haliotis fulgens Philippi 1845) became protected under a state-sponsored fishery moratorium in 1997 and was declared a NOAA NMFS Species of Concern in 2004. Recently, H. fulgens was chosen for possible stock restoration via translocation of wild adults to depleted habitat and supplementation through releasing cultured individuals. Before a management plan could be developed, however, an understanding of the species’ natural population genetic structure was needed. We used a genomic technique called restriction site associated DNA sequencing (RADSeq) to address the issue. RADSeq enabled discovery of 1,209 single nucleotide polymorphisms theoretically spread genome-wide in H. fulgens. Analyses suggested the species may be panmictic throughout our sampled range, with an effective population size (N_e) of 1,100–3,600. Hence, limitations to management, such as requiring local broodstock and restricting translocation potential, might be unnecessary. Sites with larger populations may be suitable sources for restoration of depleted sites (e.g. the Palos Verdes Peninsula), although the extent of local adaptation remains unknown. Despite this potential for restoration, results gathered on a sample of cultured H. fulgens illustrated how quickly genetic diversity can be lost through captive breeding. To help mitigate a drop in N_e due to hatchery supplementation, we recommend collection and replacement of ≥100 wild abalone per generation for broodstock and close management of the proportion of cultured individuals in the wild. Successful implementation will depend on operational capacity and the resilience of the source populations to broodstock collection.     

Top‐soil translocation as a technique in the re‐creation of species‐rich meadows

Abstract. In France, most civil engineering and excavation projects are at present accompanied by compensatory measures with the aim of preserving biodiversity. In order to avoid the destruction of a habitat of high conservation interest in NE France, harbouring two legally protected plant species, an experiment of soil translocation was conducted on an area of 1 ha. The donor site was an extensively managed mesophilic meadow and the receiving site was a neighbouring arable land. The vegetation of the translocated meadow was described 8 and 17 months after soil translocation, and compared (1) with vegetation resulting from more classical restoration techniques tested on the arable land (natural regeneration and seed mixture sowing) and (2) with the soil seed bank and vegetation previously present on the donor site. Results showed that the soil translocation technique permitted the development of many meadow species, including two legally protected species, and few ruderal species despite a large area of bare ground. This technique seems effective in terms of number and abundance of meadow species compared to natural regeneration and commercial seed sowing. In the case of the two classical methods, species richness was lower and only widespread species were present. Topsoil translocation provides a good compensatory method to avoid habitat and species destruction. However, the study should be continued, with the aim of assessing the longer term development and stabilization of the vegetation of the translocated meadow.     

Species richness correlations among primary producers in boreal forests

Close correlations in species numbers may make it possible to indirectly assess the species richness of difficult taxonomic groups by investigating indicator groups, for which data are more easily collected. We asked if species numbers correlate among the four dominating groups of primary producers in boreal forests (liverworts, macrolichens, mosses, and vascular plants) and if substrate affiliation of species (ground vs. other substrates), sample plot size (0.01–1000 m²), and stand age (young vs. old) influence correlation strength. We used three sets of study plots from northern Sweden each including wide ranges of species richness. Although there are large differences in the ecophysiology and substrate use of vascular plants and the two bryophyte groups (mosses and liverworts), we found strong positive correlations among them not previously reported from boreal forests. In contrast, no correlation in total species richness was found between macrolichens and the two bryophyte groups, despite large overlaps in their ecology. We suggest that the positive correlations among land plants (liverworts, mosses, and vascular plants) are linked to positive relationships between site moisture and species number for all three groups. In contrast, total species number of macrolichens has not been shown to be strongly associated with moisture. However, ground‐living lichens and mosses correlated negatively in old forests. This may relate to the inability of macrolichens to exploit shaded and wet old forest ground, a habitat that is used by many moss species. Furthermore, lichens and mosses of ‘other substrates’ correlated positively in old forests, probably because the amount of boulders was positively related to species richness in both groups. Generally, correlations became stronger with increasing plot size, whereas stand age had relatively little influence. We conclude that vascular plants could be used as an indicator group for species richness of mosses and liverworts in boreal landscapes.     

Use of Restored Riparian Habitat by the Endangered Least Bell's Vireo (Vireo bellii pusillus)

A primary objective of riparian restoration in California is the creation of habitat for endangered species. Four restoration sites in San Diego County were monitored between 1989 and 1993 and evaluated for their suitability as nesting habitat for Vireo bellii pusillus (Least Bell's Vireo), a state and federally endangered obligate riparian breeder. Vegetation structure at each site was quantified annually and compared to a model of canopy architecture derived from Least Bell's Vireo territories in natural habitat. Vireo use of restored habitat was documented through systematic surveys and nest monitoring. By 1993, only one site in its entirety met the habitat suitability criteria of the model, but portions of each site during all years did so. Differences between sites in the time required to develop suitable habitat—well-developed layered vegetation from the ground to under 8m in height)—were attributable largely to variation in annual rainfall. Vireos visited restoration sites to forage as early as the first growing season, but they did not establish territories or nest there until at least part of the site supported suitable habitat as determined from the model. Placement of territories and nests coincided with patches of dense vegetation characteristic of natural nesting areas. Occupation of restored sites was accelerated by the presence of adjacent mature riparian habitat, which afforded birds nest sites and/or foraging habitat lacking in the planted vegetation. Vireos nesting in restored habitat achieved success comparable to that of vireos nesting in surrounding natural habitat, and there was no evidence that productivity was reduced in created areas. These findings indicate that creating nesting habitat for this target species is feasible and suggest that the critical components of vireo nesting habitat have been captured in both the design and quantitative assessment of restoration sites.     

Evidence based conservation of butterflies

Few results of research aimed at solving questions arising from butterfly conservation are rigorously tested by manipulating populations and habitats in the field. Some factors common to successful conservation projects are analysed. In most non-migratory species, population density may vary by up to two orders of magnitude between sites or over time, and is primarily determined by the extent to which a subset of each species’ foodplant (or ant host) exists in the optimum growth-form or micro-habitat preferred by its larvae. Successful conservation projects have identified the optimum subset of each species’ larval resource before managing sites to increase its representation. In contrast, short-term fluctuations around a site’s carrying capacity or equilibrium level are mainly attributable to variation in weather, and are generally two orders of magnitude smaller than that attributable to larval habitat quality. There is little evidence that changing the abundance of adult resources, apart from shelter, influences population size or trends. The main constraint of the adult stage is the inability of many species to track the generation of new habitat patches that arise across modern landscapes. Within-patch larval habitat quality is again critical at the meta-population scale, explaining slightly more examples of patch occupancy than site isolation. This is because the higher density populations supported by optimum habitat are less likely to go extinct, and immigrants to new high-quality patches have a higher probability of founding new populations. Such patches may also generate up to a hundred times more emigrants per hectare than low-quality source patches.     

Evaluation of Swedish woodland key habitats using red-listed bryophytes and lichens

All Swedish forest land is at present being surveyed with the aim of mapping woodland key habitats which have an estimated number of ca. 70 000. An untested requirement of these habitats is that they should contain red-listed species. In order to investigate if they fulfill their aim, an inventory of nationally red-listed bryophytes and lichens was performed in ca. 120 randomly selected woodland key habitats distributed throughout the country. The species were recorded in line transects, covering the whole surface of the habitats. The mean number of red-listed species per habitat was 0.7 for bryophytes and 1.4 for lichens with 11 species at the richest site and 71% of the sites having at least one species. Nemoral forests and coniferous forests were the most species-rich habitat types. The habitats of northern Sweden were more species rich than the southern ones. There were weak correlations between presence of bryophytes and lichens. The study shows that a majority of the woodland key habitats contain red-listed bryophyte and lichen species. Further studies are needed in order to record more organism groups and to compare the woodland key habitats with the surrounding matrix forests.     

Evidence of weak habitat specialisation in microscopic animals

Macroecology and biogeography of microscopic organisms (any living organism smaller than 2 mm) are quickly developing into fruitful research areas. Microscopic organisms also offer the potential for testing predictions and models derived from observations on larger organisms due to the feasibility of performing lab and mesocosm experiments. However, more empirical knowledge on the similarities and differences between micro- and macro-organisms is needed to ascertain how much of the results obtained from the former can be generalised to the latter. One potential misconception, based mostly on anedoctal evidence rather than explicit tests, is that microscopic organisms may have wider ecological tolerance and a lower degree of habitat specialisation than large organisms. Here we explicitly test this hypothesis within the framework of metacommunity theory, by studying host specificify in the assemblages of bdelloid rotifers (animals about 350 μm in body length) living in different species of lichens in Sweden. Using several regression-based and ANOVA analyses and controlling for both spatial structure and the kind of substrate the lichen grow over (bark vs rock), we found evidence of significant but weak species-specific associations between bdelloids and lichens, a wide overlap in species composition between lichens, and wide ecological tolerance for most bdelloid species. This confirms that microscopic organisms such as bdelloids have a lower degree of habitat specialisation than larger organisms, although this happens in a complex scenario of ecological processes, where source-sink dynamics and geographic distances seem to have no effect on species composition at the analysed scale.     

Nesting,brood rearing,and summer habitat selection by translocated greater sage‐grouse in North Dakota,USA

Human enterprise has led to large‐scale changes in landscapes and altered wildlife population distribution and abundance, necessitating efficient and effective conservation strategies for impacted species. Greater sage‐grouse (Centrocercus urophasianus; hereafter sage‐grouse) are a widespread sagebrush (Artemisia spp.) obligate species that has experienced population declines since the mid‐1900s resulting from habitat loss and expansion of anthropogenic features into sagebrush ecosystems. Habitat loss is especially evident in North Dakota, USA, on the northeastern fringe of sage‐grouse’ distribution, where a remnant population remains despite recent development of energy‐related infrastructure. Resource managers in this region have determined a need to augment sage‐grouse populations using translocation techniques that can be important management tools for countering species decline from range contraction. Although translocations are a common tool for wildlife management, very little research has evaluated habitat following translocation, to track individual behaviors such as habitat selection and fidelity to the release site, which can help inform habitat requirements to guide selection of future release sites. We provide an example where locations from previously released radio‐marked sage‐grouse are used in a resource selection function framework to evaluate habitat selection following translocation and identify areas of seasonal habitat to inform habitat management and potential restoration needs. We also evaluated possible changes in seasonal habitat since the late 1980s using spatial data provided by the Rangeland Analysis Platform coupled with resource selection modeling results. Our results serve as critical baseline information for habitat used by translocated individuals across life stages in this study area, and will inform future evaluations of population performance and potential for long‐term recovery.