Aino Juslén

Illuminating the evolutionary history of liverworts (Marchantiophyta)—towards a natural classification

The phylogenetic relationships of liverworts were reconstructed using the sequence data of four genome regions including rbcL, rps4 and trnL‐F of the chloroplast and 26S large subunit ribosomal rRNA gene of the nucleus, and 90 characters of morphological, ultrastructural and developmental aspects. The taxa sampled consisted of 159 species including 135 liverworts (108 genera, 54 families and 29 suborders), 13 mosses, two hornworts, seven vascular plants and two charophyte algae. Analyses based on maximum parsimony using both direct optimization (POY) and static alignment (NONA), as well as Bayesian inference (MrBayes) were done. All the data sets were analyzed simultaneously. Our study confirms that liverworts compose a monophyletic group which consists of three classes. The class Treubiopsida including both Treubia and Haplomitrium is resolved as the earliest diverging liverwort lineage. Blasia and the complex thalloids are assigned to the Marchantiopsida, under which Blasiidae and Marchantiidae are divided. Marchantiidae include Sphaerocarpales and Marchantiales. The simple thalloid and leafy liverworts form the Jungermanniopsida, which is further divided to subclasses Pelliidae subclassis nov., Metzgeriidae and Jungermanniidae. Metzgeriidae here is defined to include only Metzgeriaceae, Aneuraceae and Vandiemeniaceae, and is the sister group to the leafy liverworts. The leafy liverworts Jungermanniidae include the orders Pleuroziales, Porellales and Jungermanniales. It is assumed that the Porellales and the Jungermanniales have split early, at least in the Jurassic period. In the Porellales, the diversification rate may have remained relatively constant for long periods of time but speeding up only recently within some of the families, associated with an explosive radiation of angiosperms. The Jungermanniales are most probably a recently diversified group which has attained the greatest profusion of structure and the most remarkable diversity of leaf development and protective devices for maturing sporophytes. A detailed classification scheme for liverworts is presented.

Application of the Red-List Index at a National Level for Multiple Species Groups

The International Union for Conservation of Nature (IUCN) Red List Index (RLI) is recognized as one of the key indicators of trends in the status of species. The red-list assessment done by Finnish authorities of species in Finland is taxonomically one of the most extensive national assessments. We used the Finnish Red Lists from 2000 and 2010 to calculate for the first time the national RLIs for 11 taxonomic groups at different trophic levels and with different life cycles. The red-list index is calculated on the basis of changes in red-list categories and indicates trends in the status of biological diversity of sets of species. The RLI value ranges from 0 to 1. The lower the value the faster the set of species is heading toward extinction. If the value is 1, all species in the set are least concern and if the value is 0, all species are (regionally) extinct. The overall RLI of Finnish species decreased. This means that, in Finland, these taxonomic groups were heading toward extinction faster in 2010 than in 2000. Of the analyzed groups of organisms, RLIs of 5 decreased and RLIs of 6 increased. At the national level, the RLIs and status trends varied markedly between species groups. Thus, we concluded that generalizations on the basis of RLIs of a few taxa only may yield a biased view of ongoing trends in the status of biological diversity at the species level. In addition, one overall RLI that includes many different species groups may also be misleading if variation in RLI among species groups is not considered and if RLI values are not presented separately for each group.

Why taxonomists and ecologists are not, but should be, carpooling?

Abstract Traditionally, ecologists and taxonomists have conducted mostly separated research. Ecologists have been using the well-known species groups as their model organisms for practical reasons, whereas taxonomists have focused on poorly known species groups. Extensive collaboration between these research fields has unfortunately been a rarity. We argue that increased collaboration between ecologists and taxonomists yields benefits through several pathways. Firstly, it improves the scientific quality of both disciplines through deeper insight on the study organisms and increased feasibility of the collected data. Secondly, it improves the prospects of the scientists involved and opens up potential funding sources, helping to solve the taxonomic impediment. Finally, improved collaboration could balance ecological research towards more species-rich, functionally important groups. This would increase the quality and effectiveness of nature conservation and management plans positively affecting the sustainability of environmental policy making. These obvious benefits should be swiftly acknowledged by research institutions, science leaders and funding bodies. They should actively promote for strong collaborative efforts from the present and recruited staff representing both disciplines.

Application of the Red List Index as an indicator of habitat change

For the first time ever, the International Union for Conservation of Nature Red List Index for habitat types was calculated for an entire country, Finland. The RLIs were based on species threat assessments from 2000 and 2010 and included habitat definitions for all 10,131 species of 12 organism groups. The RLIs were bootstrapped to track statistically significant changes. The RLI changes of species grouped by habitats were negative for all habitat types except for forests and rural biotopes which showed a stable trend. Trends of beetles and true bugs were positive in rural and forest habitats. Other 16 observed trends of species group and habitat combinations were negative. Several trends observed were in accordance with studies focusing on particular taxa and habitats, and drivers for their change. This study demonstrates the usefulness of the RLI as a tool for observing habitat change based on species threat assessment data.

Into Africa: Molecular phylogenetics and historical biogeography of sub-Saharan African woodferns (Dryopteris)

PREMISE OF THE STUDY Our goal was to infer the phylogenetic relationships and historical biogeography of the genus Dryopteris with a focus on taxa in sub-Saharan Africa and neighboring islands. In general, little is known about the relationships between African fern species and their congeners in other geographic regions, and our aim was to determine whether the sub-Saharan African species of Dryopteris are monophyletic and evolved within Africa or arrived there via repeated dispersals into Africa from other regions. METHODS We obtained sequence data for five chloroplast markers from 214 species of Dryopteris and 18 outgroups. We performed phylogenetic and molecular dating analyses using a Bayesian relaxed clock method in BEAST with fossil and secondary calibration points and estimated ancestral ranges for the genus globally by comparing multiple models in BioGeoBEARS. KEY RESULTS We found that 22 of 27 accessions of sub-Saharan African Dryopteris belong to a large clade of 31 accessions that also includes taxa from Indian and Atlantic Ocean islands. Additional accessions of taxa from our regions of interest have Asian, Hawaiian, European, or North American species as their closest relatives. CONCLUSIONS The majority of sub-Saharan African Dryopteris species are descended from a shared common ancestor that dispersed to Africa from Asia approximately 10 Ma. There have been subsequent dispersal events from the African mainland to islands in the Atlantic and Indian Oceans, including Madagascar. Several additional species are estimated to have descended from ancestors that reached Africa via separate events over the last roughly 20 million years.

Trends of extinction risk for Lepidoptera in Finland: the first national Red List Index of butterflies and moths

The Finnish Red Lists from 2000 and 2010 were used to calculate for the first time the national Red List Index for the order Lepidoptera and different Lepidoptera families with data from 2245 species. We grouped species according to their primary habitats and analysed each habitats RLI as well. The overall RLI was significantly negative, reflecting the worsening of the group as a whole in the country. A significant negative trend was found for families Tortricidae, Gelechiidae and Hesperiidae and for coastal habitats. We also report that 76 Lepidoptera species newly established a population in Finland between 2000 and 2010, probably expanding northwards driven by climate warming.

The unknown northern green: evaluation of a national forest biodiversity research program

The taxonomic knowledge gap and lack of knowledge on species-level diversity has been a global concern within conservation biology through the recent two decades. As a national response, Finland has funded a Research Program of Poorly Known and Threatened Forest Species from the year 2003. The program is an essential part of the knowledge base applied in Finnish forest policy. The present paper evaluates the outcomes of the program, covering scientific publication, training, taxonomic coverage, increased knowledge on boreal forest biodiversity and awareness rising. The ongoing program has been funded by 6.5 million Euros to date including 59 projects. The program has produced 19 master’s theses and six doctoral theses. Due to their expertise the graduated biologists have been employed in environmental administration or become researchers. The program has produced 163 refereed scientific articles, 104 of which are recognized by the Web of Science; with an average impact factor of 1.81. The results of the program include 15 genera and 348 species new to science, and 60 genera and 1,664 species to Finland’s list of known species. Due to new knowledge gathered in the program 3,000–4,000 species could be included in the most recent National Red List assessment. The publishing of identification books on local language has proved as the most effective way of advancing general species knowledge among the public and end-users. The high number of new species to science from a biodiversity-poor boreal country describes the scale of the huge work still be done in describing the global species-level diversity. The program has achieved its goals in many ways, but the program leaves out a large part of the species that do not occur in forest biotopes. Moreover, the funding has decreased through the years despite the international goal of halting the world’s biodiversity loss has not been met.

A systematic review of the relation between species traits and extinction risk

Biodiversity is shrinking rapidly, and despite our efforts only a small part of it has been assessed for extinction risk. Identifying the traits that make species vulnerable might help us to predict the outcome for those less known. We gathered information on the relations of traits to extinction risk from 173 publications, across all taxa, spatial scales and biogeographical regions, in what we think is the most comprehensive compilation to date. The Palaearctic is the most represented biogeographical realm in extinction risk studies, but the other realms are also well sampled for all vertebrate groups. Other taxa are less well represented and there are gaps for some biogeographical realms. Many traits have been suggested to be good predictors of extinction risk. Among these, our meta-analyses were successful in identifying two as potentially useful in assessing risk for the lesser-known species: regardless of the taxon, species with small range and habitat breadth – two of the three classic dimensions of rarity – are more vulnerable to extinction. On the other hand, body size (the most studied trait) did not present a consistently positive or negative response. Large species can be more, or less, threatened than their smaller counterparts, depending on the taxon and spatial setting. In line with recent research, we hypothesize that the relationship between body size and extinction risk is shaped by different aspects, namely body size is a proxy for different phenomena depending on the taxonomic group, and there might be relevant interactions between body size and the specific threat that a species faces. With implications across all traits, a particular focus in future studies should be given to the interaction between traits and threats. Furthermore, intraspecific variability of traits should be part of future studies whenever data is available. We also propose two complementary paths to better understand extinction risk dynamics: (i) the simulation of virtual species and settings for theoretical insights, and (ii) the use of comprehensive and comparable empirical data representative of all taxa and regions under a unified study.

Tutkimus tehostaa suojelutoimia: yhteenveto Puutteellisesti tunnettujen ja uhanalaisten metsälajien tutkimusohjelman vaikuttavuudesta

Kuusela S., Anttila S., Halme P., Juslén A. (2017). Tutkimus tehostaa suojelutoimia: yhteenveto Puutteellisesti tunnettujen ja uhanalaisten metsälajien tutkimusohjelman vaikuttavuudesta. Metsätieteen aikakauskirja 2017-6987. Tieteen tori. 5 s. https://doi.org/10.14214/ma.6987 Yhteystiedot 1 Suomen ympäristökeskus (SYKE), Luontoympäristökeskus/Ekosysteemipalvelut, Helsinki; 2 Jyväskylän yliopisto, Bioja ympäristötieteiden laitos, Jyväskylä; 3 Helsingin yliopisto, Luonnontieteellinen keskusmuseo Luomus, Eläintieteen yksikkö, Helsinki Sähköposti saija.kuusela@ymparisto.fi Hyväksytty 3.3.2017

What We Do (and Don't) Know about Ferns: Dryopteris (Dryopteridaceae) as a Case Study

Abstract Ferns are the second largest group of vascular land plants after the angiosperms, but remain chronically underrepresented in studies of plant phylogeny, biogeography, physiology, and genomics. The genus Dryopteris, the woodferns, is a large group with a worldwide distribution, and recent research has made it one of the better understood fern genera and a potential model for understanding many aspects of fern biology and evolution. Here we review historical and current understanding of the genus, and outline promising avenues of future research in ferns for which Dryopteris is an ideal study system, particularly for research on polyploid complexes, biogeographic distributions, and physiological ecology.