Favratia zoysii (Wulfen) Feer is a chasmophyte endemic to the south-eastern Alps and the sole representative of the genus Favratia Feer. Its primary occurrences are in Julian Alps and on Mt. Storžič (Kamnik-Savinja Alps) in Slovenia. This study aims at elucidating the original material utilised by Wulfen in the initial description of Campanula zoysii, later reclassified as F. zoysii by Feer, while also exploring the temporal and spatial density of the species’ herbarium collections. To this end, we examined preserved collections from six institutions, including botanical gardens and museums. A total of 127 herbarium vouchers were analysed to identify the specimens referenced by Wulfen. This comprehensive examination of historical collections, spanning over two centuries, allowed us to designate a lectotype, thereby providing a clear typification of F. zoysii. Additionally, we analysed the frequency and geographic distribution of herbarium specimens, a concept referred to as temporal and spatial herbarium density. This combined typification and spatial-temporal analysis not only strengthens the taxonomic clarity of the species, but also provides valuable insights into its botanical history, distribution patterns and conservation needs.
Chemotaxonomy is the link between the state of the art in analytical chemistry and the systematic classification and phylogenetic analysis of biota. Although the characteristic secondary metabolites from diverse biotic sources have been used in pharmacology and biological systematics since the dawn of mankind, only comparatively recently established reproducible methods have allowed the precise identification and distinction of structurally similar compounds. Reliable, rapid screening methods like TLC (Thin Layer Chromatography) can be used to investigate sufficiently large numbers of samples for chemotaxonomic purposes. Using distribution patterns of mutually exclusive naphthoquinones, it is demonstrated in this review how a simple set of chemical data from a representative sample of closely related species in the sundew family (Droseraceae, Nepenthales) provides taxonomically and phylogenetically informative signal within the investigated group and beyond.
A revision of the infrageneric (but supraspecific) names published for Pinguicula is performed, including the reference to the protologue and type for every validly published name, as well as the nomenclatural status and nomenclatural notes to explain certain cases, especially for those names that were not validly published or are illegitimate. In addition to discussing the underlying taxonomy from previous publications, we performed an updated phylogenetic analysis using representative ITS, trnK, matK, and rpl32 + rpl32-trnL sequences available in the GenBank database, as internal reference. As a result of the study, the name Brandonia Rchb. is typified with the type of Pinguicula lutea, P. sect. Brandonia is recognised due to priority as the correct name for the clade previously known as P. sect. Isoloba. Finally, some author citations that were often incorrectly cited in the literature are corrected. Pinguicula sect. Nana and P. sect. Micranthus are proposed as incertae sedis instead of forming part of P. subg. Temnoceras, and at least P. sect. Nana might be probably better included in P. subg. Pinguicula. This new placement is supported by morphological and chorological data, and partly by our molecular phylogenetic reconstructions. Chromosome numbers from the literature were also taken into account and confirmed to be useful to delineate certain infrageneric taxa.
Limb autotomy, the voluntary shedding of body parts as a strategy to escape predation or entrapment, is particularly common in insects and other arthropods that are frequently captured by the carnivorous plant genus Drosera. However, no study has previously examined the effectiveness of autotomy at facilitating escape from these passive, sessile plant predators. Using field observations of numerous Drosera species in Western Australia and Australia's Northern Territory, we present the first field evidence of limb autotomy being employed by insects and other arthropods to escape capture by carnivorous plants. Most autotomised limbs found on the Drosera traps belonged to large nematoceran flies, probably comprised primarily of Tipulidae (crane flies), which have a characteristically large, slender body with very long limbs that seems likely to allow the effective use of autotomy as a strategy to prevent fatal capture. However, autotomy was overall only rarely observed amongst the Drosera prey as most prey items were small and quickly became completely enveloped by the sticky mucilage, rendering any such escape attempts impossible. Finally, we determined that the likelihood of escape decreased markedly as multiple limbs were lost.
Drosera maanyaa-gooljoo, a new annual species of Drosera section Arachnopus from the remote Buccaneer Archipelago and Yampi Peninsula in the northwest Kimberley region of Western Australia, is described and illustrated. Drosera maanyaa-gooljoo is a distinctive species that differs in morphology and indumentum from all previously known taxa of D. section Arachnopus. Comprehensive photo plates comparing all morphological details, as well as seed shape and structure, with those of the putatively related D. barrettiorum and D. hartmeyerorum are provided. Given D. maanyaa-gooljoo is only known from two small populations, a Priority One status under Conservation Codes for Western Australian Flora is recommended. Both populations occur within exclusive Native Title lands of the Dambimangari and Mayala Traditional Owners. This exciting new species discovery highlights the importance of both consultation with Indigenous people and herbarium revisions for alpha taxonomy and biodiversity research.
The carnivorous Drosera microphylla complex from southwest Western Australia comprises a group of rare, narrowly endemic species that are potentially threatened by habitat destruction and illegal collection, thus highlighting a need for accurate taxonomic classification to facilitate conservation efforts. Following extensive fieldwork over two decades, detailed studies of both Australian and European herbaria and consideration of both crucial contributions by citizen scientists and social media observations, nine species of the D. microphylla complex are here described and illustrated, including four new species: D. atrata, D. hortiorum, D. koikyennuruff, and D. reflexa. The identities of the previously described infraspecific taxa D. calycina var. minor and D. microphylla var. macropetala are clarified. Both are here lectotypified, reinstated, and elevated to species rank. A replacement name, D. rubricalyx, is provided for the former taxon. Key morphological characters distinguishing the species of this complex include the presence or absence of axillary leaves, lamina shape, petal colour, filament shape, and style length. A detailed identification key, comparison figures, and a distribution map are provided. Six of the nine species are recommended for inclusion on the Priority Flora List under the Conservation Codes for Western Australian Flora and Fauna.
Chromosome number change is a driver of speciation in eukaryotic organisms. Carnivorous sundews in the plant genus Drosera L. exhibit single chromosome number variation both among and within species, especially in the Australian Drosera subg. Ergaleium D.C., potentially linked to atypical centromeres that span much of the length of the chromosomes. We critically reviewed the literature on chromosome counts in Drosera, verified the taxonomy and quality of the original counts, and reconstructed dated phylogenies. We used the BiChrom model to test whether rates of single chromosome number increase and decrease, and chromosome number doubling differed between D. subg. Ergaleium and the other subgenera and between self-compatible and self-incompatible lineages. The best model for chromosome evolution among subgenera had equal rates of chromosome number doubling but higher rates of single chromosome number change in D. subg. Ergaleium than in the other subgenera. Contrary to expectation, self-incompatible lineages had a significantly higher rate of single chromosome loss than self-compatible lineages. We found no evidence for an association between differences in single chromosome number changes and diploidization after polyploidy or centromere type. This study presents an exemplar for critically examining published cytological data and rigorously testing factors that may impact the rates of chromosome number evolution.
Chromosome number change is a driver of speciation in eukaryotic organisms. Carnivorous sundews, the plant genus Drosera L., exhibit single chromosome number variation among and within species, especially in the Australian Drosera subg. Ergaleium D.C., potentially linked to the presence of holocentromeres. We reviewed literature, verified chromosome counts, and using an rbc L chronogram, tested alternate models where the gain, loss, and doubling rates (+1, −1, ×2) were the same or different between D . subg. Ergaleium and the other subgenera. Ancestral chromosome number estimations were performed, and the distributions of self-compatibility and genome size were visualized across the genus. The best model for chromosome evolution had equal rates of polyploidy (0.014 per million years; Myr) but higher rates of single chromosome number gain (0.19 and 0.027 per Myr) and loss (0.23 and 0.00059 per Myr) in D . subg. Ergaleium compared to the other subgenera. We found no evidence for differences in single chromosome evolution to be due to differences in diploidization after polyploidy or to holocentromeres as had been proposed. This study highlights the complexity of factors influencing rates of chromosome number evolution. ### Competing Interest Statement The authors have declared no competing interest.
Allen Lowrie was a not a university trained botanist. He was a botanist by passion. His studies and observations of Australian carnivorous plants and triggerplants for about a half-century will inevitably impact every person with an interest in those plants from the Australian flora. It is not an exaggeration to claim that he was probably the most influential person regarding our recent understanding and knowledge of the carnivorous plant flora of Australia. No other botanist – neither 20th or 21st Century nor before – discovered and described to science more new carnivorous plant species or triggerplants.
To survive in the nutrient-poor habitats, carnivorous plants capture small organisms comprising complex substances not suitable for immediate reuse. The traps of carnivorous plants, which are analogous to the digestive systems of animals, are equipped with mechanisms for the breakdown and absorption of nutrients. Such capabilities have been acquired convergently over the past tens of millions of years in multiple angiosperm lineages by modifying plant-specific organs including leaves. The epidermis of carnivorous trap leaves bears groups of specialized cells called glands, which acquire substances from their prey via digestion and absorption. The digestive glands of carnivorous plants secrete mucilage, pitcher fluids, acids, and proteins, including digestive enzymes. The same (or morphologically distinct) glands then absorb the released compounds via various membrane transport proteins or endocytosis. Thus, these glands function in a manner similar to animal cells that are physiologically important in the digestive system, such as the parietal cells of the stomach and intestinal epithelial cells. Yet, carnivorous plants are equipped with strategies that deal with or incorporate plant-specific features, such as cell walls, epidermal cuticles, and phytohormones. In this review, we provide a systematic perspective on the digestive and absorptive capacity of convergently evolved carnivorous plants, with an emphasis on the forms and functions of glands.
Prey spectra (the number and composition of captured arthropods) represent a crucial aspect of carnivorous plant ecology, yet remain poorly studied. Traditional morphology-based approaches for prey identification are time-intensive, require specialists with considerable knowledge of arthropod taxonomy, and are hampered by high numbers of unidentifiable (i.e., heavily digested) prey items. We examined prey spectra of three species of closely-related annual Drosera (Droseraceae, sundews) from tropical northern Australia using a novel DNA metabarcoding approach with in-situ macro photography as a plausibility control and to facilitate prey quantity estimations. This new method facilitated accurate analyses of carnivorous plant prey spectra (even of heavily digested prey lacking characteristic morphological features) at a taxonomic resolution and level of completeness far exceeding morphology-based methods and approaching the 100% mark at arthropod order level. Although the three studied species exhibited significant differences in detected prey spectra, little prey specialisation was observed and habitat or plant population density variations were likely the main drivers of prey spectra dissimilarity.
The complete life history of the kleptoparasitic ‘sundew flower fly’, Toxomerus basalis , is presented and illustrated. Adults of this species are photographed alive for the first time, including video recordings of larval and adult behaviour. Adult flies of both sexes visit Drosera (sundews) and show territorial behaviour around the plants, avoiding the dangerous sticky traps and demonstrating recognition of their larval host plant. Females lay eggs directly on non-sticky parts of the Drosera host plants, such as on the lower surface of the leaves and flower stalks, but apparently also on other plants growing in close proximity with the sundews.
Parasitic plants contain some of the most bizarre and fascinating organisms in the plant kingdom. Yet they are notable for their absence from botanic gardens’ plant collections and conservation strategies. Besides a handful of species, few are widespread in cultivation; indeed we estimate at least 76 per cent of species are entirely missing from collections today, and most of these have never been grown at all. Here, we place focus on the holoparasites, a group of plants long neglected due to their difficulty in cultivation. We review propagation breakthroughs in temperate and tropical botanic gardens to identify guiding principles for the cultivation of these neglected plants. We document the life cycle of a range of parasitic plants, and assess successful and failed attempts to propagate Rafflesia specifically, which has been the focus of decades of research. By uniting isolated case studies from around the world, we identify future directions for the cultivation and possible ex situ conservation of these botanical enigmas at a time when this is needed urgently. Finally, we recommend a dedicated global community of purpose as an intentional step forward: this could take the form of a Global Consortium for Conservation for parasitic plants, or a Parasitic Plant Specialist Group under the International Union for Conservation of Nature.
In a screening of 43 accessions of predominantly Australian sundew species (Drosera), naphthoquinones were detected convincingly for the first time in D. section Lasiocephala (D. petiolaris group, or ‘wooly sundews’), where these metabolites remain restricted to a minority of four closely related species. Great chemical similarity across the large geographic range confirms a close phylogenetic affinity between taxa of the D. peltata species group (of D. section Ergaleium, from tropical and Eastern Asia to New Zealand). Drosera barrettiorum (D. section Arachnopus) is chemically confirmed as a close relative of D. hartmeyerorum. The recently described species D. margaritacea in the same section is chemically different from the morphologically close D. finlaysoniana. Quinone data for African and South American sundews (D. sections Drosera, Ptycnostigma and Brasilianae) shed further light on the affinities between these taxonomically challenging plants.
Resurrecting extinct species is a fascinating and challenging idea for scientists and the general public. Whereas some theoretical progress has been made for animals, the resurrection of extinct plants (de-extinction sensu lato) is a relatively recently discussed topic. In this context, the term 'de-extinction' is used sensu lato to refer to the resurrection of 'extinct in the wild' species from seeds or tissues preserved in herbaria, as we acknowledge the current impossibility of knowing a priori whether a herbarium seed is alive and can germinate. In plants, this could be achieved by germinating or in vitro tissue-culturing old diaspores such as seeds or spores available in herbarium specimens. This paper reports the first list of plant de-extinction candidates based on the actual availability of seeds in herbarium specimens of globally extinct plants. We reviewed globally extinct seed plants using online resources and additional literature on national red lists, resulting in a list of 361 extinct taxa. We then proposed a method of prioritizing candidates for seed-plant de-extinction from diaspores found in herbarium specimens and complemented this with a phylogenetic approach to identify species that may maximize evolutionarily distinct features. Finally, combining data on seed storage behaviour and longevity, as well as specimen age in the novel 'best de-extinction candidate' score (DEXSCO), we identified 556 herbarium specimens belonging to 161 extinct species with available seeds. We expect that this list of de-extinction candidates and the novel approach to rank them will boost research efforts towards the first-ever plant de-extinction.
A synopsis of the Drosera (Droseraceae) species occurring in Brazil is here presented, providing tools for taxonomic identification and summarizing data as a basis for future studies on diversity and conservation of the genus in the country. Thirty-three taxa, comprising 32 species and a nothotaxon, are recognized for the Brazilian territory. We present an identification key, and, for each taxon, we provide a succint description, a differential diagnosis, notes on distribution, habitat, phenology, conservation status, and a list of representative specimens. Drosera hirtella var. lutescens is lectotypified and raised to species status, lectotypes are further designated for other 18 names, and a neotype is designated for D. intermedia var. tenuis. Drosera species are recorded from all Brazilian states, Minas Gerais being the richest with a total of 21 species. The campos rupestres of eastern Brazil are recognized as the main center of Drosera species diversity in the Neotropics, with 23 species, 13 of which are endemic. Twelve out of the 32 recognized species are considered threatened with extinction (37.5%), reinforcing the Brazilian responsibility for the conservation of global Droseraceae diversity.
In a commentary on our paper (Renner et al., Oecologia 195:825-831, 2021), Harder and Miksha lay out why they think that our finding of higher honeybee abundances reducing wild bee abundances in an urban botanical garden is not statistically supported. Here, we explain the statistical test provided in our paper, which took advantage of a natural experiment offered by 2019 being a poorer year for bee keeping than 2020.
SummaryPhilcoxia rhizomatosa is a highly specialised carnivorous perennial herb endemic to the deep, white quartzite sand pans of the campos rupestres vegetation of eastern Brazil's Espinhaço Range, specifically within the northern portion of the range in the state of Minas Gerais. Philcoxia is among the most threatened genera of carnivorous plants and thus far all attempts to cultivate them have failed.
A detailed study of 44 accessions representing 38 taxa (76% of the diversity known at present) of pygmy sundews (Drosera sect. Bryastrum) reveals the first naphthoquinone patterns in this lineage, in which previous studies have not yielded reliable evidence for naphthoquinones. While most samples do not display detectable amounts of naphthoquinones as previously reported for the group, ramentaceone is detected in three mutually related taxa, and both ramentaceone and its regio-isomer plumbagin is present in all accessions investigated of D. pulchella.
Utricularia multifida is carnivorous bladderwort from Western Australia and belongs to a phylogenetically early-diverging lineage of the genus. We present a prey spectrum analysis resulting from a snapshot sampling of 17 traps-the first of this species to our knowledge. The most abundant prey groups were Ostracoda, Copepoda, and Cladocera. The genus cf. Cypretta (Cyprididae, Ostracoda) was the predominant prey. However, a high variety of other prey organisms with different taxonomic backgrounds was also detected. Our results indicate that U. multifida may potentially be specialized in capturing substrate-bound prey. Future approaches should sample plants from different localities to allow for robust comparative analyses.