Marc Gottschling

• Ludwig-Maximilians-Universität in Munich

The genus Prorocentrum is diverse and comprises mostly marine dinoflagellates with a worldwide distribution. One large, common and easily recognizable planktonic species of Prorocentrum was known for a long time as P. compressum. This name, however, is incorrectly linked to a basionym of a diatom. The confusing taxonomy of the now also lectotypifie...

During evolution of Dinophyceae, size reduction of the episome has occurred in several lineages (including unarmoured Amphidiniales and armoured Prorocentrales). One such species is Amphidinium crassum, whose taxonomic identity is elusive though showing morphological similarities with Oxytoxaceae (currently placed in armoured Peridiniales). Plankto...

To determine the intraspecific variability of microscopic organisms such as dinophytes is challenging, but can be achieved using cultured material. Unusual morphologies of Peridinium tuberosum assigned to the Peridinium cinctum species group have been described as bulges on the posterior end of the cell a hundred years ago and more, but its taxonom...

In the microscopy realm, a large body of dark biodiversity still awaits to be uncovered. Unarmoured dinophytes are particularly neglected here, as they only present inconspicuous traits. In a remote German locality, we collected cells, from which a monoclonal strain was established, to study morphology using light and electron microscopy and to gai...

The purpose of naming organisms is to provide a means of reference to enable exchange of information about those organisms. A universally understood, precise and stable system of naming is therefore essential for effective scientific communication. Principle III and IV of the Shenzhen Code (“Code”: Turland & al. in Regnum Veg. 159. 2018) state that...

Among the photosynthetically active dinophytes, the Kryptoperidiniaceae are unique in having a diatom as endosymbiont instead of the widely present peridinin chloroplast. Phylogenetically, it is unresolved at present how the endosymbionts are inherited, and the taxonomic identities of two iconic dinophyte names, Kryptoperidinium foliaceum and Krypt...

Prorocentrum comprises dinophytes with several unique traits, including the presence of two large thecal plates and apical insertion of flagella. Species delimitation for many small and similar planktonic species is challenging, as SEM analyses and DNA sequence information of type material are rarely available. Based on a strain from the North Atla...

The Prorocentrales are a unique group of dinophytes based on several apomorphic traits, but species delimitation is challenging within the group. Prorocentrum triestinum was described by Josef Schiller in 1918 as an important bloom-forming species from Trieste (Mediterranean, Adriatic Sea) with a conspicuous asymmetric outline and a small, asymmetr...

Detailed anatomical studies of flowers and fruits may render traits of phylogenetic importance, but are still rare in the Boraginales. An Old World clade of Cordia comprises many dioecious species, but the floral anatomy is largely unexplored and, hence, the structural basis for sex separation. Fresh material of buds, functionally unisexual flowers...

Micracanthodinium setiferum is a very distinctive but poorly known dinophyte with characteristically delicate, unbranched, filiform and pointed setae. A recent study by Mertens et al. (2022 Mertens KN, Carbonell-Moore MC, Gardner K. 2022. A morphological comparison of two cladopyxidacean dinoflagellates: the extant Micracanthodinium setiferum and t...

The Prorocentrales are a unique group of dinophytes based on several apomorphic traits, but species delimitation is challenging within the group. Prorocentrum triestinum was described by Josef Schiller in 1918 as an important bloom-forming species from Trieste (Mediterranean, Adriatic Sea) with a conspicuous asymmetric outline and a small, asymmetr...

Character evolution in calcareous dinophytes and their relatives is poorly understood at present, particularly regarding the transformations of the epitheca. The precise taxonomic identity of Chimonodinium lomnickii var. (≡ Peridinium) wierzejskii is unclear, but two different arrangements of the epithecal plate pattern have been reported. We studi...

Dinophyte evolution is essentially inferred from the pattern of thecal plates, and two different labelling systems are used for the important subgroups Gonyaulacales and Peridiniales. The partiform hypotheca of cladopyxidoid dinophytes fits into the morphological concepts of neither group, although they are assigned to the Gonyaulacales. Here, we d...

Gonyaulacales include a considerable number of harmful algae and to understand their origin and rise, knowledge of the evolutionary relationships is necessary. Many scientific names of protists introduced prior to the availability of DNA analytics are ambiguous and impede communication about biological species and their traits in the microbial worl...

Dinophytes are widely distributed in marine‐ and fresh‐waters, but have yet to be conclusively documented in terrestrial environments. Here, we evaluated the presence of these protists from an environmental DNA metabarcoding dataset of Neotropical rainforest soils. Using a phylogenetic placement approach with a reference alignment and tree, we show...

In the current circumscription, the Thoracosphaeraceae comprise all dinophytes exhibiting calcified coccoid cells produced during their life-history. Species hitherto assigned to Ensiculifera and Pentapharsodinium are mostly based on the monadoid stage of life-history, while the link to the coccoid stage (occasionally treated taxonomically distinct...

Dinophytes are widely distributed in marine- and fresh-waters, but have yet to be conclusively documented in terrestrial environments. Here we evaluated the presence of these protists from an environmental DNA metabarcoding dataset of Neotropical rainforest soils. Using a phylogenetic placement approach with a reference alignment and tree, we showe...

Unicellular dinophytes include a considerable number of harmful algae and potent producers of toxins. The dinophyte fossil record is one of the richest among protists and indicates a geologically old origin of the group dating back to the Triassic. Besides of these records, very few molecular dating studies of dinophytes have been published to date...

Phytodinialean dinophytes are poorly known at present and their phylogenetic relationships largely elusive. Historical names of microscopic species are frequently ambiguous, and a reliable application is impeded although crucial to fully explore the biology of organisms. We collected material close to the type locality of a historical species, name...

As many other protist groups, euglenophytes are prone to false identification based solely on morphology because of a limited amount of morphological features and cryptic speciation. One of the supposedly completely asexual groups within the freshwater phototrophic representatives of euglenophytes is Trachelomonas, capable of forming an inorganic s...

Poor morphological and molecular differentiation in recently diversified lineages is a widespread phenomenon in plants. Phylogenetic relationships within such species complexes are often difficult to resolve because of the low variability in traditional molecular loci. Furthermore, biological phenomena responsible for topological incongruence such...

The taxonomy of Gymnodinium and the Gymnodiniaceae is inconsistent, because the systematic position of the type species, G. fuscum, is elusive at present. Historical names of microscopic species are frequently ambiguous, making reliable determination difficult, although it is crucial to fully explore the biology of the organisms. We collected mater...

The application of scientific names is determined by means of nomenclatural types, and every name has to be typified properly. The concept has limitations for unicellular organisms, because original material frequently consists of drawings and/or inadequately preserved physical material. Peridinium cinctum is an abundant freshwater microalga and va...

Despite their ecological importance and wide distribution, Cordiaceae have not been subject to detailed anatomical study yet. We examined flower and fruit anatomies of Cordia nodosa and Varronia bonplandii (using paraffin sectioning and light microscopy) in comparison with other woody members of Boraginales. The internal architecture of the superio...

Parvodinium elpatiewskyi, comb. nov., is a common freshwater dinophyte without intercalary plates and with various spines on hypothecal sutures. However, the taxonomy of the species has had a complex history, and its systematic placement remained unclear. The conserved type of P. elpatiewskyi, comb. nov., illustrated here for the first time using e...

Reliable determination of organisms is a prerequisite to explore their spatial and temporal occurrence and to study their evolution, ecology, and dispersal. In Europe, Bavaria (Germany) provides an excellent study system for research on the origin and diversification of freshwater organisms including dinophytes, due to the presence of extensive lak...

Dinophytes are one of few protist groups that have an extensive fossil record and are therefore appropriate for time estimations. However, insufficient sequence data and strong rate heterogeneity have been hindering to put dinophyte evolution into a time frame until now. Marine‐to‐freshwater transitions within this group are considered geologically...

Azaspiracids (AZA) are the most recently discovered group of lipophilic marine biotoxins of microalgal origin, and associated with human incidents of shellfish poisoning. They are produced by a few species of Amphidomataceae, but diversity and occurrence of the small-sized dinophytes remain poorly explored for many regions of the world. In order to...

The Prorocentrales are a unique group of dinophytes based on several apomorphic traits, but species delimitation is challenging within the group. The type species of Prorocentrum, namely P. micans, cannot be determined unambiguously, as important characters are not preserved in the original material collected in the first half of the 19th century....

Poor morphological and molecular differentiation in recently diversified lineages is a widespread phenomenon in plants. Phylogenetic relationships within such species complexes are often difficult to resolve because of the low variability in traditional molecular loci, as well as various other biological phenomena responsible for topological incong...

Poor morphological and molecular differentiation in recently diversified lineages is a widespread phenomenon in plants. Phylogenetic relationships within such species complexes are often difficult to resolve because of the low variability in traditional molecular loci, as well as various other biological phenomena responsible for topological incong...

We report the complete genome sequences of Rusa timorensis papillomavirus 1 (RtimPV1) and Rusa timorensis papillomavirus 2 (RtimPV2), isolated from hair follicles of asymptomatic skin from the same Timor deer specimen. RtimPV1 and RtimPV2 are evolutionarily only distantly related. RtimPV1 lacks a canonical E2-binding site, and RtimPV2 does not carr...

Halgania (Ehretiaceae, Boraginales) comprises ~20 species of ericoid shrubs endemic to Australia. The current taxonomic concepts based on morphology are confusing and are sometimes based on misidentification or lack of information about the type material. We describe the morphological diversity and infer relationships using molecular phylogenetics....

In the course of polyphasic taxonomic work in the dinophytes, we became aware of a fundamental misapplication of the name Glenodinium triquetrum (now represented conceptually by a species of Kryptoperidinium), when Stein assigned it to Heterocapsa. Possible solutions involve a conflict between retaining Ehrenberg’s epithet in its correct applicatio...

Despite recent fruitful attempts to elucidate microbial biogeography in more detail, knowledge of distribution still lags behind for dinophytes. Evolutionary phenomena, such as cryptic speciation and modification due to the environment, hamper reliable conclusions about the distribution of this important plankton group. We combined newly collected...

Flowers that release their powdery pollen by being vibrated through buzzing are morphologically diverse and have evolved many times independently. Most prominently, buzz-pollination is known from Solanum (Solanaceae) defining the “solanoid” flower type by Knut Fægri. The flowers of buzz-pollinated, Australian Halgania resemble the actinomorphic Sol...

Peridinium cinctum is a common freshwater dinophyte with a long history of research. Erich Lindemann was the first to assess intraspecific variability in this species focusing on plate pattern variation. Since then, this issue has been neglected but with the application of DNA sequence diagnostics, a combination of morphological and molecular chara...

Miller, J.S. & M. Gottschling (2017). Generic transfers in Malagasy Boraginales. Candollea 72 : 329–332. In English, English and French abstracts. Recent molecular data have demonstrated convincingly that the Old World genus Hilsenbergia Tausch ex Meisn. (Ehretiaceae) is nested within the New World genus Bourreria P. Browne. Thus, 13 new combinatio...

Amphidoma is an old though poorly studied thecate dinophyte that has attracted attention recently as a potential producer of azaspiracids (AZA), a group of lipophilic phycotoxins. A new species, Amphidoma parvula, sp. nov. is described from the South Atlantic shelf of Argentina. With a Kofoidean thecal plate pattern Po, cp, X, 6′, 6′′, 6C, 5S, 6′′′...

Tertiary endosymbiosis is proven through dinophytes, some of which (i.e. Kryptoperidiniaceae) have engulfed diatom algae containing a secondary plastid. Chloroplasts are usually inherited together permanently with the host cell, leading to co-phylogeny. We compiled a diatom sequence data matrix of two nuclear and two chloroplast loci. Almost all en...

The contemporary occurrence of dinophytes at their type localities has not been intensely studied so far, despite the type locality's crucial importance for any reliable scientific name application. The microscopist and phycologist Ch.G. Ehrenberg described a number of dinophyte species more than 150 years ago, many of which are currently taxonomic...

Peridinialean dinophytes include a unique evolutionary group of algae harboring a diatom as endosymbiont (Kryptoperidiniaceae), whose phylogenetic origin and internal relationships are not fully resolved at present. Several interpretations of the thecal plate pattern present in Durinskia oculata currently compete and lead to considerable taxonomic...

One of the most common marine dinophytes is a species known as Heterocapsa triquetra. When Stein introduced the taxon Heterocapsa he formally based the type species H. triquetra on the basionym Glenodinium triquetrum. The latter was described by Ehrenberg and is most likely a species of Kryptoperidinium. In addition to that currently unresolved nom...

The Andean uplift is one of the major orographic events in the New World and has impacted considerably the diversification of numerous Neotropical lineages. Despite its importance for biogeography, the specific role of mountain ranges as a dispersal barrier between South and Central American lowland plant lineages is still poorly understood. The sw...

Kryptoperidiniaceae are a small group of dinophytes hosting a tertiary endosymbiont derived from a diatom (Tomas et al. 1973, Horiguchi & Pienaar 1994). Those so called ‘dinotoms’ (Imanian et al. 2011) include Dinothrix, Durinskia, Galeidinium, Kryptoperidinium (= Phyllodinium) and some species currently assigned to “Peridiniopsis” and “Peridinium”...

The vast majority not only of dinophytes, but also of Peridiniales, live in the marine environment, and some 350 (out of ca 2.500) species are present in freshwater habitats (Mertens et al. 2012). Many freshwater Peridiniales constitute small and only distantly related species groups embedded in predominantly marine clades (e.g., Apocalathium, Chim...

The Andean uplift is one of the major orographic events in the New World and has impacted considerably the diversification of numerous Neotropical organisms. Despite its importance for biogeography, the specific role of mountain ranges as a dispersal barrier between South and Central American lowland plant lineages is still poorly understood. The s...

Azaspiracids (AZA) are a group of food poisoning phycotoxins that are known to accumulate in shellfish. They are produced by some species of the planktonic dinophycean taxon Amphidomataceae. Azaspiracids have been first discovered in Ireland but are now reported in shellfish from numerous global sites thus showing a wide distribution. In shellfish...

Arctic regions have experienced pronounced biological and biophysical transformations as a result of global change processes over the last several decades. Current hypotheses propose an elevated impact of those environmental changes on the biodiversity, community composition and metabolic processes of species. The effects on ecosystem function and...

AimTo examine the historical biogeography of the Boraginales using molecular dating and ancestral area reconstruction. LocationWorld-wide. Methods We constructed data sets that included all major clades of Boraginales and all orders of asterids using previously published sequences of four plastid markers (trnL-trnF, rps16, ndhF, rbcL). We estimated...

The Boraginales are now universally accepted as monophyletic and firmly placed in Lamiidae. However, a consensus about familial classification has remained elusive, with some advocating recognition of a single, widely variable family, and others proposing recognition of several distinct families. A consensus classification is proposed here, based o...

Background: Rochefortia is a small taxon of woody plants in the Ehretiaceae (Boraginales) exhibiting coriaceous leaves with cystoliths, small whitish flowers and drupaceous fruits containing four pyrenes. It shares the dioecious sex distribution with its sister group Lepidocordia and can be delimited from the latter (and all other Ehretiaceae) by...

Trees or shrubs, rarely subshrubs, lianas or herbs, mostly evergreen; indumentum sericeous or hispid, plants rarely glabrous, trichomes usually simple, unicellular, eglandular, sometimes additionally glandular, rarely branched or dolabriform. Leaves mostly alternate, sometimes fasciculate, generally simple, petiolate or sessile, mostly exstipulate,...

Pollination effectiveness in Cycnoches is directly dependent on osmophores. Euglossine-bee pollinated orchids produce a bucket of fragrances acting as attractant and reward for pollinators, which collect these fragrances and use them during courtship. Despite the importance of scent glands in orchids, few studies on their anatomy and morphology hav...

Taxonomic diversity of Neotropical Rochefortia is not completely assessed at present. We report the existence of a new species: Rochefortia barloventensis sp. nov., distributed across multiple islands of the Lesser Antilles. We provide a morphological description, a molecular diagnosis and a botanical illustration. Specimens belonging to the new sp...

Peridinium acuminatum (Peridiniales, Dinophyceae) was described in the first half of the 19 th century, but the name has been rarely adopted since then. It was used as type of Goniodoma, Heteraulacus and Yesevius, providing various sources of nomenclatural and taxonomic confusion. Particularly, several early authors emphasised that the organisms in...

Phylogenetic relationships inferred from multilocus organellar and nuclear DNA data are often difficult to resolve because of evolutionary conflicts among gene trees. However, conflicting or 'outlier' associations (i.e., linked pairs of Operational Terminal Units in two phylogenies) among these data sets often provide valuable information on evolut...

The Gymnodiniaceae -even in a strict sense- comprise a vast diversity of dinophytes regarding morphology and ecology. Taxonomy and nomenclature of their constituent species remain problematic, although crucial to fully explore the biology of the group. Here, we present the rarely documented dinophyte Gymnodinium limneticum from its type locality at...

Peridinium acuminatum (Peridiniales, Dinophyceae) was described in the first half of the 19th century, but the name has been rarely adopted since then. It was used as type of Goniodoma, Heteraulacus and Yesevius, providing various sources of nomenclatural and taxonomic confusion. Particularly, several early authors emphasised that the organisms inv...

Species of dinophycean Amphidomataceae are producers of phycotoxins classified as azaspiracids. We provide the first records of two of their constituent species, Amphidoma languida and Azadinium dexteroporum, for the Irminger Sea off Iceland. Morphological examination and molecular characterization including uncorrected mean pair-wise distances bet...

Dioecious and thorny Rochefortia Sw. is a poorly known though distinct element of the Ehretiaceae comprising woody plants restricted to the Caribbean and the adjacent American mainland. The approximately ten species display a great morphological variability and overlapping taxonomic boundaries, which makes it difficult to differentiate them (partic...

The Scrippsiella trochoidea species complex (Thoracosphaeraceae, Peridiniales) consists of a cryptic diversity and multiple species, for which established scientific names are not available at present. Previously, the name Scrippsiella trochoidea has been taxonomically clarified, leaving a reliable determination of morphologically similar, but only...

Some species of the planktonic dinoflagellate genus Azadinium produce azaspiracids (AZAs), a group of lipophilic phycotoxins causing human poisoning after mussel consumption. We describe three new species from the North Atlantic, all of which shared the same Kofoidean plate pattern characteristic for Azadinium: Po, cp, X, 4′, 3a, 6′́, 6 C, 5S, 6′́́...

Tiquilia is very different from the other members of the Ehretiaceae (Boraginales) in many aspects of morphology and ecology. Because detailed knowledge about flower and fruit traits is necessary to reliably infer character evolution of and within Tiquilia, we investigated flower to fruit ontogeny in eight species of Tiquilia using light and electr...

Major taxa of Ehretiaceae (including parasitic Lennoaceae) have not all been included in previous molecular phylogenetic analyses. As a result, the generic limits and their circumscriptions have not been satisfactorily resolved, despite its importance for floristic studies. To clarify which monophyletic groups can be recognized within the Ehretiace...

Calcareous dinophytes produce exoskeletal calcified structures during their life history (a unique character among the alveolates) and are subsumed under the Thoracosphaeraceae as part of the Peridiniales. We provide a brief synopsis about the taxonomic history of the group, from the first descriptions of fossils in the 19th century through to the...

Multiple family-level subdivisions of Boraginales have been proposed in the past. The relationships of several constituent genera have been enigmatic, including Codon (Codonaceae), Hoplestigma (Hoplestigmataceae), Pholisma (Lennoaceae), Vahlia (Vahliaceae), and Wellstedia (Wellstediaceae), all of which are included in the present study. We present...

The Thoracosphaeraceae are dinophytes that produce calcareous shells during their life history, whose optical crystallography has been the basis for the division into subfamilies. To evaluate the validity of the classification (mainly applied by palaeontologists), living material of phylogenetic key species is necessary albeit frequently difficult...

Infection by papillomaviruses (PVs) has been linked to different types of neoplasias, in both human and non-human hosts. Knowledge about PV diversity is essential to reliably infer the evolutionary history of these pathogens and to elucidate the link between infection and disease. We cloned and sequenced the complete genome of a novel PV, EhelPV1,...