Citing the taxonomic literature: what a difference a year makes

We all know that the world of scientific publishing has changed profoundly since the onset of the digital revolution. One relatively new development is the rapid publication of scientific papers online, frequently before they are copyedited and typeset, and sometimes even before being peer reviewed (Sheldon 2018). Climate of the Past is one such journal that posts manuscripts online before they have been refereed. The purpose of doing this is to allow online discussion of a manuscript while it is under review in the conventional sense. Manuscripts may thus benefit from any useful feedback from readers as well as from the formal reviews. The above developments mean that scientific articles may appear online long before being assigned to a particular volume/issue and with final page numbers. Such assignments commonly occur in the following year when the complete volumes or issues of a journal appear in print and/ or digitally. Before the digital revolution, authors had to wait perhaps 12 months or more between acceptance and final publication. Today, just a week or two may elapse before the typescript of an accepted manuscript is available online. In most respects this revolution is good, especially now that many authors aim for metricised output targets. However, such early publication of a paper may cause complications regarding its referencing, but in most cases this does not really matter so long as the reference in a bibliography leads to the retrieval of the correct publication. For example, the paper cited below as Pound and Riding (2015) was initially issued online in 2015, prior to assignment to a volume of the Journal of the Geological Society published in 2016. Before 2016 it would have also been cited as Pound and Riding (2015) but that situation would not have lasted for long and would have affected very few, if any, citations. Electronic publication of a paper prior to assignment of the volume number and final pagination can be confusing, but in most cases problems are limited to referencing. However, it has critical implications for papers with biological systematics, especially those with new nomenclatural proposals (new taxa, combinations, substitute names, etc. – so-called nomenclatural novelties). Until recently, codes of nomenclature in botany and zoology required nomenclatural novelties to be published in paper format in publicly distributed articles. However, the most recent codes permit the publication of nomenclatural novelties in a hybrid (online and paper) journal or even in a purely electronic periodical (but not in an online database or catalogue)

The Bajocian (Middle Jurassic): a key interval in the early Mesozoic phytoplankton radiation

Dinoflagellates and coccolithophores are two of the most important groups of phytoplankton in the modern oceans. These groups originated in the Triassic and radiated through the early Mesozoic, rising to ecological prominence. Within this long-term radiation, important shortterm intervals of evolutionary and ecological change can be recognised. The Bajocian (Middle Jurassic, ~170–168 Ma) was characterised by an important ecological transition within the coccolithophores, and the radiation of one of the principal families of cyst-forming dinoflagellates, the Gonyaulacaceae. During the Early Bajocian, the coccolith genus Watznaueria diversified and expanded ecologically to dominate coccolith floras, a situation which continued for the remainder of the Mesozoic. This pattern was paralleled within dinoflagellate cyst floras by the ecological dominance of the genus Dissiliodinium in the midpalaeolatitudes. These phenomena appear to be linked to a positive carbon isotope shift, and an interval of enhanced productivity driven by a shift to a more humid climate, enhanced continental weathering and nutrient flux, or by changes in ocean circulation and upwelling. The latest Early Bajocian to earliest Bathonian was then characterised by the rapid increase in diversity of dinoflagellate cysts within the family Gonyaulacaceae. Through this interval, the Gonyaulacaceae transitioned from being a relatively minor component of dinoflagellate cyst floras, to becoming one of the prominent groups of cyst-forming dinoflagellates, which has persisted to the Holocene. In Europe, the pattern of this radiation was strongly influenced by sea level, with the increase in gonyaulacacean diversity reflecting a major second-order transgression. On a finer scale, the main pulses of first appearances correlate with third-order transgressive episodes. A rise in sea level, coupled with changes in the tectonic configuration of ocean gateways, appears to have controlled the pattern of plankton diversification in Europe. These palaeoceanographic changes may have enhanced water-mass transfer between Europe, the northwest Tethys Ocean and the Hispanic Corridor, which promoted the floral interchange of dinoflagellates. Whilst sea level rise and associated large-scale palaeoenvironmental shifts appear to have controlled the pattern of dinoflagellate cyst appearances in several regions outside Europe, there is no direct correlation between dinoflagellate cyst diversity and sea level rise on a global scale. Although the Bajocian was transgressive in several regions, widespread flooded continental area was also present throughout the preceding Aalenian, an interval of low gonyaulacacean diversity. Moreover, although the Middle Jurassic was an interval of major climatic cooling, there was a ~5 myr gap between the onset of cooling and the radiation of gonyaulacaceans during the Bajocian. The Bajocian was, however, marked by a major evolutionary radiation in the pelagic realm, including ammonites, giant suspension feeding fishes and planktonic foraminifera. These phenomena may indicate an underlying ecological driver to the radiation of dinoflagellates during the Bajocian evolutionary explosion which could represent an extension of the Mesozoic Marine Revolution.This work has arisen from the PhD project of Nickolas J. Wiggan which was supported by NERC BGS DTG award reference BUFI S246, entitled The mid Jurassic plankton explosion. This was funded jointly between the British Geological Survey and the University of Cambridge. James B. Riding publishes with the approval of the Executive Director, British Geological Survey (NERC). We thank the Review Papers Coordinator, Tim Horscroft, for inviting this contribution. Nick Butterfield (Cambridge) is thanked for discussions and suggestions during NJW’s PhD project. We also thank Daniel Mantle and Fabienne Giraud, whose insightful reviews greatly improved the quality of this manuscript

A review of the Sentusidinium complex of dinoflagellate cysts

The Jurassic to Neogene (Miocene) dinoflagellate cyst genus Sentusidinium has a relatively simple overall morphology. This genus, together with Batiacasphaera, Kallosphaeridium and Pentafidia, comprises the Sentusidinium complex. This is distinct from the superficially similar laterally asymmetrical and subspheroidal/lenticular Cyclonephelium complex. The genus Sentusidinium is an acavate, subcircular, proximate to proximochorate, sexiform gonyaulacacean genus with an apical archaeopyle and typically low relief ornamentation. Since the erection of Sentusidinium in 1978, three similar genera have been established, which we consider to be taxonomic junior synonyms of that genus: Barbatacysta, Escharisphaeridia and Pilosidinium. However, we deem the Early Cretaceous to Miocene genera Batiacasphaera, Kallosphaeridium and Pentafidia are deemed to be separate from Sentusidinium. We refine the definition of the Early Cretaceous to Miocene genus Batiacasphaera to circumscribe cysts with a reticulate to rugulate autophragm and an apical archaeopyle with a free operculum. By contrast, Kallosphaeridium has a ventrally attached apical archaeopyle with five plates that can be interpreted as type (4A1I)@ or type (5A)@; it also has a small operculum relative to the overall cyst diameter. The six accepted Kallosphaeridium species are confined to the Palaeogene. The Australian genus Pentafidia is unusual in appearing to only have five precingular plates; this comprises two species from the Jurassic–Cretaceous transition of Western Australia. Therefore, we emend Sentusidinium to restrict it to acavate, proximate or proximochorate dinoflagellate cysts with an autophragm devoid of, or covered with, highly variable, non-linear ornamentation and a type (tA) apical archaeopyle. Occasionally the elements of ornamentation may be connected, but rarely is a cingulum indicated, and the tabulation is never clearly evident. A kalyptra may be occasionally present. The operculum is free. Following a comprehensive literature review, we accept 17 species in Batiacasphaera. In Kallosphaeridium we accept six species confidently and consider six species to be problematical. We list 38 (34 accepted and four problematical) species of Sentusidinium. Kallosphaeridium? helbyi is here transferred to Cyclonephelium without question. The species Batiacasphaera angularis is occasionally tabulate and hence we transfer it, with question, to Meiourgonyaulax. The Sentusidinium complex is clearly polyphyletic, and all genera considered herein belong to the order Gonyaulacales. Batiacasphaera, Kallosphaeridium and Pentafidia cannot be confidently assigned to a family, whereas Sentusidinium belongs to the Gonyaulacaceae. The number of species within the complex has been reduced from 137 to 68; furthermore, all infraspecific taxa have been eliminated

Taxonomy and nomenclature in palaeopalynology: basic principles, current challenges and future perspectives

Effective communication of taxonomic concepts is crucial to meaningful application in all biological sciences, and thus the development and following of best practices in taxonomy and the formulation of clear and practical rules of nomenclature underpin a wide range of scientific studies. The International Code of Nomenclature for algae, fungi and plants (the Code), currently the Shenzhen Code of 2018, provides these rules. Although early versions of the Code were designed mainly with extant plants in mind, the Code has been increasingly used for fossil plants and, in recent decades, for organic-walled microfossils, the study of which is called palaeopalynology, or simply palynology. However, rules embodied in the Code do not fully reflect the needs and practices of this discipline; and taxonomic practices between fossil applications, especially in palynology, have tended to diverge from practices for extant plants. Differences in these rules and practices present specific challenges. We therefore review the Shenzhen Code as it applies to palynology, clarifying procedures and recommending approaches based on best practices, for example, in the designation and use of nomenclatural types. The application of nomenclatural types leads to taxonomic stability and precise communication, and lost or degraded types are therefore problematic because they remove the basis for understanding a taxon. Such problems are addressed using examples from the older European literature in which type specimens are missing or degraded. A review of the three most important conventions for presenting palynological taxonomic information, synonymies, diagnoses/descriptions and illustrations, concludes with recommendations of best practices. Palynology continues to play an important role in biostratigraphy, palaeoenvironmental analyses, and evolutionary studies, and is contributing increasingly to our understanding of past climates and ocean systems. To contribute with full potential to such applied studies, consistent communication of taxonomic concepts, founded upon clear rules of nomenclature, is essential

A review of the Jurassic dinoflagellate cyst genus Gonyaulacysta Deflandre 1964 emend. nov.

The Middle–Late Jurassic gonyaulacacean dinoflagellate cyst genus Gonyaulacysta is characterised by an epicyst which is around twice the length of the hypocyst. The sulcus is L-type and may be slightly sigmoidal in shape, but is never S-type sensu stricto. The forms with slightly sigmoidal L-type sulcuses may have developed into species with S-type sulcuses. Gonyaulacysta jurassica, the holotype of which is the nomenclatural type of the genus, had a global distribution. A total of 151 species have been assigned to Gonyaulacysta although 126 of these have been transferred to other genera. Prior to this contribution, 25 species were accepted in the current Lentin and Williams Index. This is herein reduced to eight and 18 species are transferred to other genera. The accepted species are: Gonyaulacysta adecta stat. nov., emend. nov.; Gonyaulacysta australica comb. nov., emend. nov.; Gonyaulacysta ceratophora; Gonyaulacysta desmos stat. nov., emend. nov.; Gonyaulacysta dualis emend. nov.; Gonyaulacysta fenestrata emend. nov.; Gonyaulacysta jurassica emend. nov.; and Gonyaulacysta longicornis stat. nov., emend. nov. These species form a closely related plexus, and are distinguished on differences in cavation style, form of sutural crests/ridges and size of the apical horn. All except Gonyaulacysta australica are reliable marker taxa. Our main taxonomic proposals involve the elevation of all subspecies and varieties of species here retained in Gonyaulacysta to species rank, or their synonymisation. This avoids cumbersome infraspecific names. Gonyaulacysta exhibits substantial provincialism; for example Gonyaulacysta dualis is confined to the Oxfordian–Kimmeridgian of the Boreal Realm. Gonyaulacysta adecta, Gonyaulacysta desmos and Gonyaulacysta longicornis are present in the Bathonian to Oxfordian of Laurasia and surrounding areas. Three species, Gonyaulacysta australica, Gonyaulacysta ceratophora and Gonyaulacysta fenestrata, are restricted to the Oxfordian to Tithonian of Australasia. Gonyaulacysta adecta and the cosmopolitan Gonyaulacysta jurassica both exhibit overall size increases throughout the Bathonian to Kimmeridgian of Europe

Cretaceous and Cenozoic dinoflagellate cysts and other palynomorphs from the western and eastern margins of the Labrador–Baffin Seaway - Fig. 5

New palynological analysis of samples from 13 offshore wells on the Canadian Margin and six wells on the West Greenland Margin has led to a new event biostratigraphic framework for Cretaceous– Cenozoic strata of the Labrador Sea – Davis Strait – Baffin Bay (Labrador–Baffin Seaway) region. This framework is based on about 150 dinoflagellate cyst taxa and 30 acritarch, algal, fungal and plant microfossil (mostly miospore) taxa. In the systematics we include three new genera of dinocysts (Scalenodinium, Simplicidinium and Taurodinium), 16 new species of dinocysts (Chiropteridium gilbertii, Chytroeisphaeridia hadra, Cleistosphaeridium elegantulum, Cleistosphaeridium palmatum, Dapsilidinium pseudoinsertum, Deflandrea borealis, Evittosphaerula? foraminosa, Ginginodinium? flexidentatum, Hystrichosphaeridium quadratum, Hystrichostrogylon digitus, Impletosphaeridium apodastum, Scalenodinium scalenum, Surculosphaeridium convocatum, Talladinium pellis, Taurodinium granulatum and Trithyrodinium? conservatum), four emendations of dinocyst genera (Alterbidinium, Chatangiella, Chiropteridium and Surculosphaeridium), six new combinations for dinocyst species (Alterbidinium biaperturum, Deflandrea majae, Kleithriasphaeridium mantellii, Simplicidinium insolitum, Spongodinium grossum, Spongodinium obscurum), one new acritarch species (Fromea quadrangularis), one new miospore species (Baculatisporites crenulatus) and one new combination for miospores (Tiliaepollenites crassipites). Most of the taxa included provide age information, almost exclusively last occurrences (range ‘tops’), but some are useful mainly for environmental interpretations. Collectively, they provide a powerful tool for helping to establish the geological history of the Labrador–Baffin Seaway

Biostratigraphic correlation of the western and eastern margins of the Labrador–Baffin Seaway and implications for the regional geology. Appendix 3

New analyses of the palynological assemblages in 13 offshore wells on the Canadian margin and six on the West Greenland Margin, in conjunction with onshore data, have led to a new biostratigraphic framework for the Cretaceous–Cenozoic strata of the Labrador Sea – Davis Strait – Baffin Bay (Labrador–Baffin Seaway) region and the first broad biostratigraphic correlation of the Canadian and Greenland margins. This framework is based on 167 last occurrences and 18 local/regional peak/common-occurrence events for dinocysts, miospores, fungal spores and Azolla. Detailed biostratigraphic evidence has confirmed the following hiatuses: pre-Aptian in the Hopedale Basin; pre-Albian in the Saglek Basin; Albian–Turonian in some wells of the Hopedale Basin; Turonian–Santonian/Campanian in some areas; pre-Campanian and late Campanian – Thanetian on the Greenland Margin; late Maastrichtian and Danian in some wells of the Hopedale Basin and in the Saglek Basin; Selandian in part of the Hopedale Basin, in all the Saglek Basin wells and in two wells on the West Greenland Margin; late Ypresian and/or Lutetian on both sides; Oligocene to middle Miocene of considerable variability on both margins, with all of the Oligocene and the lower Miocene missing in all the West Greenland Margin wells; and middle to late Miocene on the western side. On the Canadian margin, the hiatuses can be partially matched with the five previously recognised regional unconformities; on the Greenland margin, however, the relationship to the five unconformities is more tenuous. Palynomorph assemblages show that most Aptian to Albian sediments were deposited in generally non-marine to marginal marine settings, interrupted by a short-lived shallow marine episode in the Aptian. A marine transgression started in the Cenomanian–Turonian and led to the most open-marine, oceanic conditions in the Campanian–Lutetian; shallowing probably started in the late Lutetian and continued into the Rupelian, when inner neritic and marginal marine palaeoenvironments predominated. Throughout the rest of the Cenozoic, inner neritic palaeoenvironments alternated with marginal marine conditions on the margins of the Labrador–Baffin Seaway. These observations broadly reflect the tectonic evolution of the seaway, with rift conditions prevailing from Aptian to Danian times, followed by drift through much of the Paleocene and Eocene, and post-drift from Oligocene to the present. Dinocysts indicate that climatic conditions in the Labrador–Baffin Seaway region were relatively temperate in the Cretaceous, but varied dramatically through the Cenozoic. The Danian was a time of increasingly warmer climate, a thermal maximum being reached around the Paleocene–Eocene boundary reflecting the global thermal event at this time. Warm to hot conditions prevailed throughout the Ypresian, but the climate began to cool in the Lutetian, a trend that accelerated through the Priabonian and Rupelian. Throughout the Neogene, temperatures generally declined, culminating in the Quaternary

Biostratigraphic correlation of the western and eastern margins of the Labrador–Baffin Seaway and implications for the regional geology. Fig. 5

New analyses of the palynological assemblages in 13 offshore wells on the Canadian margin and six on the West Greenland Margin, in conjunction with onshore data, have led to a new biostratigraphic framework for the Cretaceous–Cenozoic strata of the Labrador Sea – Davis Strait – Baffin Bay (Labrador–Baffin Seaway) region and the first broad biostratigraphic correlation of the Canadian and Greenland margins. This framework is based on 167 last occurrences and 18 local/regional peak/common-occurrence events for dinocysts, miospores, fungal spores and Azolla. Detailed biostratigraphic evidence has confirmed the following hiatuses: pre-Aptian in the Hopedale Basin; pre-Albian in the Saglek Basin; Albian–Turonian in some wells of the Hopedale Basin; Turonian–Santonian/Campanian in some areas; pre-Campanian and late Campanian – Thanetian on the Greenland Margin; late Maastrichtian and Danian in some wells of the Hopedale Basin and in the Saglek Basin; Selandian in part of the Hopedale Basin, in all the Saglek Basin wells and in two wells on the West Greenland Margin; late Ypresian and/or Lutetian on both sides; Oligocene to middle Miocene of considerable variability on both margins, with all of the Oligocene and the lower Miocene missing in all the West Greenland Margin wells; and middle to late Miocene on the western side. On the Canadian margin, the hiatuses can be partially matched with the five previously recognised regional unconformities; on the Greenland margin, however, the relationship to the five unconformities is more tenuous. Palynomorph assemblages show that most Aptian to Albian sediments were deposited in generally non-marine to marginal marine settings, interrupted by a short-lived shallow marine episode in the Aptian. A marine transgression started in the Cenomanian–Turonian and led to the most open-marine, oceanic conditions in the Campanian–Lutetian; shallowing probably started in the late Lutetian and continued into the Rupelian, when inner neritic and marginal marine palaeoenvironments predominated. Throughout the rest of the Cenozoic, inner neritic palaeoenvironments alternated with marginal marine conditions on the margins of the Labrador–Baffin Seaway. These observations broadly reflect the tectonic evolution of the seaway, with rift conditions prevailing from Aptian to Danian times, followed by drift through much of the Paleocene and Eocene, and post-drift from Oligocene to the present. Dinocysts indicate that climatic conditions in the Labrador–Baffin Seaway region were relatively temperate in the Cretaceous, but varied dramatically through the Cenozoic. The Danian was a time of increasingly warmer climate, a thermal maximum being reached around the Paleocene–Eocene boundary reflecting the global thermal event at this time. Warm to hot conditions prevailed throughout the Ypresian, but the climate began to cool in the Lutetian, a trend that accelerated through the Priabonian and Rupelian. Throughout the Neogene, temperatures generally declined, culminating in the Quaternary

Biostratigraphic correlation of the western and eastern margins of the Labrador-Baffin Seaway and implications for the regional geology

New analyses of the palynological assemblages in 13 offshore wells on the Canadian margin and six on the West Greenland Margin, in conjunction with onshore data, have led to a new biostratigraphic framework for the Cretaceous–Cenozoic strata of the Labrador Sea – Davis Strait – Baffin Bay (Labrador–Baffin Seaway) region and the first broad biostratigraphic correlation of the Canadian and Greenland margins. This framework is based on 167 last occurrences and 18 local/regional peak/common-occurrence events for dinocysts, miospores, fungal spores and Azolla. Detailed biostratigraphic evidence has confirmed the following hiatuses: pre-Aptian in the Hopedale Basin; pre-Albian in the Saglek Basin; Albian–Turonian in some wells of the Hopedale Basin; Turonian–Santonian/Campanian in some areas; pre-Campanian and late Campanian – Thanetian on the Greenland Margin; late Maastrichtian and Danian in some wells of the Hopedale Basin and in the Saglek Basin; Selandian in part of the Hopedale Basin, in all the Saglek Basin wells and in two wells on the West Greenland Margin; late Ypresian and/or Lutetian on both sides; Oligocene to middle Miocene of considerable variability on both margins, with all of the Oligocene and the lower Miocene missing in all the West Greenland Margin wells; and middle to late Miocene on the western side. On the Canadian margin, the hiatuses can be partially matched with the five previously recognised regional unconformities; on the Greenland margin, however, the relationship to the five unconformities is more tenuous. Palynomorph assemblages show that most Aptian to Albian sediments were deposited in generally non-marine to marginal marine settings, interrupted by a short-lived shallow marine episode in the Aptian. A marine transgression started in the Cenomanian–Turonian and led to the most open-marine, oceanic conditions in the Campanian–Lutetian; shallowing probably started in the late Lutetian and continued into the Rupelian, when inner neritic and marginal marine palaeoenvironments predominated. Throughout the rest of the Cenozoic, inner neritic palaeoenvironments alternated with marginal marine conditions on the margins of the Labrador–Baffin Seaway. These observations broadly reflect the tectonic evolution of the seaway, with rift conditions prevailing from Aptian to Danian times, followed by drift through much of the Paleocene and Eocene, and post-drift from Oligocene to the present. Dinocysts indicate that climatic conditions in the Labrador–Baffin Seaway region were relatively temperate in the Cretaceous, but varied dramatically through the Cenozoic. The Danian was a time of increasingly warmer climate, a thermal maximum being reached around the Paleocene–Eocene boundary reflecting the global thermal event at this time. Warm to hot conditions prevailed throughout the Ypresian, but the climate began to cool in the Lutetian, a trend that accelerated through the Priabonian and Rupelian. Throughout the Neogene, temperatures generally declined, culminating in the Quaternary