A new golden frog species of the genus Diasporus (Amphibia, Eleutherodactylidae) from the Cordillera Central, western Panama

We describe the frog species Diasporus citrinobapheussp. n. from the Cordillera Central of western Panama. The new species differs from all other species in its genus in coloration, disk cover and disk pad shape, skin texture, advertisement call, and size. It is most similar to Diasporus tigrillo, from which it differs in dorsal skin texture, relative tibia length, number of vomerine teeth, ventral coloration, dorsal markings, and relative tympanum size, and to Diasporus gularis, from which it can be distinguished by the lack of membranes between the toes, adult size, posterior thigh coloration, and position of the choanae. We provide data on morpho- logy, vocalization, and distribution of the new species, as well as brief information on its natural history.Describimos la especie de rana Diasporus citrinobapheus sp. n. de la Cordillera Central, occidente de Panamá. La nueva especie se distingue de otras especies del género por su coloración, su forma de la cubierta y la almohadilla de los discos, textura de la piel, canto de anúncio, y tamaño corporal. Se asemeja mas a D. tigrillo, del cual se distingue por la textura de la piel dorsal, longitud relativa de la tibia, número de dientes vomerianos, coloración ventral, patrón dorsal, y tamaño relativo del tímpano, y a D. gularis, del cual se diferencia por la ausencia de membranas entre los dedos de pie, tamaño corporal, coloración de la parte trasera del muslo, y posición de las coanas. Presentamos datos de la morfología, vocalización, y distribución de la nueva especie, así como notas concisas de su historia natural

SYSTEMATICS, TAXONOMY AND BIOGEOGRAPHY OF THE COSMOPOLITAN AND FOSSIL-RICH BUCKTHORN FAMILY (RHAMNACEAE JUSS.)

My thesis combines taxonomy, phylogenetics, divergence time estimations and ancestral area reconstructions, nomenclature, and other methods to shed light on the evolutionary processes that shaped the extant biodiversity of the buckthorn family (Rhamnaceae Juss.). In a first step, this thesis provides a major contribution to the taxonomic assessment and arrangement within the buckthorns. By describing the new genus Pseudoziziphus Hauenschild, and resurrecting four genera, this thesis highlights and compensates underestimated diversity. Simultaneously, it provides a phylogenetic framework based on the largest molecular data set in the family so far. In a second step, the taxonomic and phylogenetic framework was used to reconstruct solid divergence time estimates and ancestral area estimates for Rhamnaceae, with a focus on the ziziphoid lineage, and the genus complex Alphitonia Reissek ex Endl. sensu lato. By this, it was possible to unravel evolutionary processes within the history of Rhamnaceae and provide a biogeographic scenario for the ziziphoid lineages, including Alphitonia s.l.:General Introduction 1 Bibliography 19 Chapter 1 36 Phylogenetic relationships within the cosmopolitan buckthorn family (Rhamnaceae) support the resurrection of Sarcomphalus and the description of Pseudoziziphus gen. nov. 37 Bibliography 48 Appendix 1 50 Supplements 55 Chapter 2 56 Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s.l. supports the description of a new genus, Ventia 57 Bibliography 67 Appendix 1 68 Supplements 71 Erratum 72 Chapter 3 74 The influence of the Gondwanan break-up on the biogeographic history of the ziziphoids (Rhamnaceae) 75 Bibliography 96 Appendix 1 105 Appendix 2 115 Supplements 117 Chapter 4 118 Biogeographic analyses support an Australian origin for the IndomalesianAustralasian wet forest-adapted tropical tree and shrub genus Alphitonia and its close allies (Rhamnaceae) 119 Bibliography 144 Appendix 1 153 Appendix 2 161 Appendix 3 166 General Conclusions 172 Bibliography 182 Appendix 186 Spatio-temporal evolution of Allium L. in the Qinghai-Tibet-Plateau region: Immigration and in situ radiation 187 Curriculum Vitae 200 Declaration of Independent work 207 Author contributions 20

Polyploidy promotes species diversification of Allium through ecological shifts

[EN] Despite the role of polyploidy in multiple evolutionary processes, its impact on plant diversification remains controversial. An increased polyploid frequency may facilitate speciation through shifts in ecology, morphology or both. Here we used Allium to evaluate: (1) the relationship between intraspecific polyploid frequency and species diversification rate; and (2) whether this process is associated with habitat and/or trait shifts. Using eight plastid and nuclear ribosomal markers, we built a phylogeny of 448 Allium species, representing 46% of the total. We quantified intraspecific ploidy diversity, heterogeneity in diversification rates and their relationship along the phylogeny using trait-dependent diversification models. Finally, we evaluated the association between polyploidisation and habitat or trait shifts. We detected high ploidy diversity in Allium and a polyploidy-related diversification rate shift with a probability of 95% in East Asia. Allium lineages with high polyploid frequencies had higher species diversification rates than those of diploids or lineages with lower polyploidy frequencies. Shifts in speciation rates were strongly correlated with habitat shifts linked to particular soil conditions; 81.7% of edaphic variation could be explained by polyploidisation. Our study emphasises the role of intraspecific polyploid frequency combined with ecological drivers on Allium diversification, which may explain plant radiations more generally.National Natural Science Foundation of China (U1802242, 31800177); the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation)-FZT 118 to REO and ANM-R. T-SH is also supported by China Scholarship Council and Y-WX is supported by National Key R&D Program of China (2017YFC0505200) and CAS 135 programme (2017XTBG-F01)

Climate change will disproportionally affect the most genetically diverse lineages of a widespread African tree species

Global climate change is proceeding at an alarming rate with major ecological and genetic consequences for biodiversity, particularly in drylands. The response of species to climate change may difer between intraspecifc genetic groups, with major implications for conservation. We used molecular data from 10 nuclear and two chloroplast genomes to identify phylogeographic groups within 746 individuals from 29 populations of Senegalia senegal, a savannah tree species in subSaharan Africa. Three phylogroups are identifed corresponding to Sudano-Sahelian, Zambezian and Southern African biogeographic regions in West, East and Southern Africa. Genetic diversity was highest in Southern and Zambesian and lowest in the Sudano-Sahelian phylogroups. Using species distribution modeling, we infer highly divergent future distributions of the phylogroups under three climate change scenarios. Climate change will lead to severe reductions of distribution area of the genetically diverse Zambezian (−41–− 54%) and Southern (− 63–− 82%) phylogroups, but to an increase for the genetically depauperate Sudano-Sahelian (+ 7– + 26%) phylogroups. This study improves our understanding of the impact of climate change on the future distribution of this species. This knowledge is particularly useful for biodiversity management as the conservation of genetic resources needs to be considered in complementary strategies of in-situ conservation and assisted migration.The German Academic Exchange Service, the German Science Foundation, the German Federal Ministry of Education and Research and Leipzig University.https://www.nature.com/srepPlant Production and Soil Scienc

Modulation of COUP-TF Expression in a Cnidarian by Ectopic Wnt Signalling and Allorecognition

COUP transcription factors are required for the regulation of gene expression underlying development, differentiation, and homeostasis. They have an evolutionarily conserved function, being a known marker for neurogenesis from cnidarians to vertebrates. A homologue of this gene was shown previously to be a neuronal and nematocyte differentiation marker in Hydra. However, COUP-TFs had not previously been studied in a colonial cnidarian.We cloned a COUP-TF homologue from the colonial marine cnidarian Hydractinia echinata. Expression of the gene was analysed during normal development, allorecognition events and ectopic Wnt activation, using in situ hybridisation and quantitative PCR. During normal Hydractinia development, the gene was first expressed in post-gastrula stages. It was undetectable in larvae, and its mRNA was present again in putative differentiating neurons and nematocytes in post-metamorphic stages. Global activation of canonical Wnt signalling in adult animals resulted in the upregulation of COUP-TF. We also monitored a strong COUP-TF upregulation in stolons undergoing allogeneic interactions. COUP-TF mRNA was most concentrated in the tissues that contacted allogeneic, non-self tissues, and decreased in a gradient away from the contact area. Interestingly, the gene was transiently upregulated during initial contact of self stolons, but dissipated rapidly following self recognition, while in non-self contacts high expression levels were maintained.We conclude that COUP-TF is likely involved in neuronal/nematocyte differentiation in a variety of contexts. This has now been shown to include allorecognition, where COUP-TF is thought to have been co-opted to mediate allorejection by recruiting stinging cells that are the effectors of cytotoxic rejection of allogeneic tissue. Our findings that Wnt activation upregulates COUP-TF expression suggests that Wnts' role in neuronal differentiation could be mediated through COUP-TF

SYSTEMATICS, TAXONOMY AND BIOGEOGRAPHY OF THE COSMOPOLITAN AND FOSSIL-RICH BUCKTHORN FAMILY (RHAMNACEAE JUSS.)

My thesis combines taxonomy, phylogenetics, divergence time estimations and ancestral area reconstructions, nomenclature, and other methods to shed light on the evolutionary processes that shaped the extant biodiversity of the buckthorn family (Rhamnaceae Juss.). In a first step, this thesis provides a major contribution to the taxonomic assessment and arrangement within the buckthorns. By describing the new genus Pseudoziziphus Hauenschild, and resurrecting four genera, this thesis highlights and compensates underestimated diversity. Simultaneously, it provides a phylogenetic framework based on the largest molecular data set in the family so far. In a second step, the taxonomic and phylogenetic framework was used to reconstruct solid divergence time estimates and ancestral area estimates for Rhamnaceae, with a focus on the ziziphoid lineage, and the genus complex Alphitonia Reissek ex Endl. sensu lato. By this, it was possible to unravel evolutionary processes within the history of Rhamnaceae and provide a biogeographic scenario for the ziziphoid lineages, including Alphitonia s.l.:General Introduction 1 Bibliography 19 Chapter 1 36 Phylogenetic relationships within the cosmopolitan buckthorn family (Rhamnaceae) support the resurrection of Sarcomphalus and the description of Pseudoziziphus gen. nov. 37 Bibliography 48 Appendix 1 50 Supplements 55 Chapter 2 56 Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s.l. supports the description of a new genus, Ventia 57 Bibliography 67 Appendix 1 68 Supplements 71 Erratum 72 Chapter 3 74 The influence of the Gondwanan break-up on the biogeographic history of the ziziphoids (Rhamnaceae) 75 Bibliography 96 Appendix 1 105 Appendix 2 115 Supplements 117 Chapter 4 118 Biogeographic analyses support an Australian origin for the IndomalesianAustralasian wet forest-adapted tropical tree and shrub genus Alphitonia and its close allies (Rhamnaceae) 119 Bibliography 144 Appendix 1 153 Appendix 2 161 Appendix 3 166 General Conclusions 172 Bibliography 182 Appendix 186 Spatio-temporal evolution of Allium L. in the Qinghai-Tibet-Plateau region: Immigration and in situ radiation 187 Curriculum Vitae 200 Declaration of Independent work 207 Author contributions 20

SYSTEMATICS, TAXONOMY AND BIOGEOGRAPHY OF THE COSMOPOLITAN AND FOSSIL-RICH BUCKTHORN FAMILY (RHAMNACEAE JUSS.)

My thesis combines taxonomy, phylogenetics, divergence time estimations and ancestral area reconstructions, nomenclature, and other methods to shed light on the evolutionary processes that shaped the extant biodiversity of the buckthorn family (Rhamnaceae Juss.). In a first step, this thesis provides a major contribution to the taxonomic assessment and arrangement within the buckthorns. By describing the new genus Pseudoziziphus Hauenschild, and resurrecting four genera, this thesis highlights and compensates underestimated diversity. Simultaneously, it provides a phylogenetic framework based on the largest molecular data set in the family so far. In a second step, the taxonomic and phylogenetic framework was used to reconstruct solid divergence time estimates and ancestral area estimates for Rhamnaceae, with a focus on the ziziphoid lineage, and the genus complex Alphitonia Reissek ex Endl. sensu lato. By this, it was possible to unravel evolutionary processes within the history of Rhamnaceae and provide a biogeographic scenario for the ziziphoid lineages, including Alphitonia s.l.:General Introduction 1 Bibliography 19 Chapter 1 36 Phylogenetic relationships within the cosmopolitan buckthorn family (Rhamnaceae) support the resurrection of Sarcomphalus and the description of Pseudoziziphus gen. nov. 37 Bibliography 48 Appendix 1 50 Supplements 55 Chapter 2 56 Analysis of the cosmopolitan buckthorn genera Frangula and Rhamnus s.l. supports the description of a new genus, Ventia 57 Bibliography 67 Appendix 1 68 Supplements 71 Erratum 72 Chapter 3 74 The influence of the Gondwanan break-up on the biogeographic history of the ziziphoids (Rhamnaceae) 75 Bibliography 96 Appendix 1 105 Appendix 2 115 Supplements 117 Chapter 4 118 Biogeographic analyses support an Australian origin for the IndomalesianAustralasian wet forest-adapted tropical tree and shrub genus Alphitonia and its close allies (Rhamnaceae) 119 Bibliography 144 Appendix 1 153 Appendix 2 161 Appendix 3 166 General Conclusions 172 Bibliography 182 Appendix 186 Spatio-temporal evolution of Allium L. in the Qinghai-Tibet-Plateau region: Immigration and in situ radiation 187 Curriculum Vitae 200 Declaration of Independent work 207 Author contributions 20

Cyrtandra argentii, a new species of Cyrtandra (Gesneriaceae) from the Philippines, and a review of the C. villosissima group

Cyrtandra argentii Olivar, H.J.Atkins & Muellner sp. nov., endemic to the Philippines and named after George Argent, is herein described and illustrated. Collections associated with this new species are often confused with three other species, namely C. ferruginea Merr., C. villosissima Merr., and C. hirtigera H.J.Atkins & Cronk. Distinguishing characters including keys, updated descriptions, distribution maps, and photos of live specimens are provided to aid identification of the four species. Following the International Union for the Conservation of Nature (IUCN) criteria, C. argentii sp. nov. is considered to be Near Threatened (NT) due to its distribution in a zone susceptible to anthropogenic pressure and the lack of any formal protection

Spatio-temporal evolution of Allium L. in the Qinghai–Tibet-Plateau region: Immigration and in situ radiation

A plethora of studies investigating the origin and evolution of diverse mountain taxa has assumed a causal link between geological processes (orogenesis) and a biological response (diversification). Yet, a substantial delay (up to 30 Myr) between the start of orogenesis and diversification is often observed. Evolutionary biologists should therefore identify alternative drivers of diversification and maintenance of biodiversity in mountain systems. Using phylogenetic, biogeographic, and diversification rate analyses, we could identify two independent processes that most likely explain the diversity of the widespread genus Allium in the Qinghai–Tibet Plateau (QTP) region: (1) While the QTP-related taxa of the subgenus Melanocrommyum diversified in situ, (2) QTP-related taxa of other subgenera migrated into the QTP from multiple source areas. Furthermore, shifts in diversification rates within Allium could not be attributed spatially and temporally to the uplift history of the QTP region. Instead, global cooling and climate oscillations in the Quaternary were major contributors to increased speciation rates in three clades of Allium. Our study therefore adds to the growing evidence supporting the “mountain-geo-biodiversity hypothesis”, which highlights the role of climate oscillations for the diversification of mountain organisms