Introduction:

At the turn of most years, the some of the editorial board at Palaeontology [online] takes the opportunity to reflect on the past year in palaeontology. Given that we published a wonderful overview of Diploporitans in January, this year we’ve moved our look over our favourite studies from last year to February. Palaeontology and associated disciplines are fast-moving and exciting areas of science — looking back at 2018 lets us highlight just a few of the key developments that really show this. Picking just one article each is difficult, and we have been forced to miss out many of the hundreds of exciting papers published in the past 12 months. Nevertheless, we hope that our choices reflect the breadth and depth of palaeobiological research in the twenty-first century. The papers include insights into the evolution of animals, the origin of colour in dinosaur eggs, the discovery of new, unusual fossil echinoderms, and insights into the origins of plant roots. So, in alphabetical order, here are the members of our editorial board with their highlights.

Russell Garwood — Deline et al., ‘Evolution of metazoan morphological disparity’

One of my favourite papers of the past year has been the work of Deline et al., which was published in August. This work looks at the animal kingdom as a whole, and analyses the variation that we see across it. The authors have compiled a cladistic character matrix that includes members of every major animal group. In brief, they have put together a list of the characters (for example, presence of a head, eyes, legs) possessed by 212 animal species within 34 phyla. In total, they managed to code 1,767 characters. This is, to me, a really significant achievement in its own right — and a valuable resource that scientists can use in the future. The authors then ran a wide range of tests on this new data set to try to improve understanding of how animals have evolved.

A key question they addressed is whether groups reach their maximum disparity — that is, variety of forms or morphologies (Fig. 1) — shortly after they evolve, or whether they continue to become more diverse as they evolve. The answer? Well, it seems both could be the case. Many groups reach the disparity we see today quite early in their evolutionary history, but some really important ones — for example, vertebrates and their kin, and arthropods — become increasingly diverse as they evolve. In short, major episodes of innovation don’t necessarily occur at the beginning of a group’s history.

The data also suggest that phyla look so different from each other today because of the extinction of intermediate forms — that is, lineages that share characteristics of major modern groups have gone extinct over time. The authors also have a go at investigating the driving force behind the patterns they report. They find that disparity is correlated with the size of the genome, and the range in a group of molecules called microRNAs, which regulate the expression of genes. Deline et al. suggest that the evolution of this system could have influenced the disparity we see in animals, but also that factors outside the groups’ genetics — such as the environment that they live in — also have an impact.

I like this paper for a number of reasons. It is asking a wide range of interesting and important questions about evolution, using a major and really diverse group. The team has created a really useful data set, and then used it to explore a number of areas in one go: I’ve only highlighted one here! It’s a really neat way to do science, and also quite a brave one (what would have happened if the data set hadn’t shown anything?). And, like lots of exciting science, it leaves us with a whole bunch of new, but potentially more specific, questions we can ask in the future.

Stephan Lautenschlager — Wiemann et al., ‘Dinosaur egg colour had a single evolutionary origin’

My favourite paper of 2018 was published in October, although Easter would have been much more fitting. Wiemann and colleagues’ study focuses on eggs — more specifically, dinosaur eggs. This by itself is not that spectacular. We have known for more than a century that dinosaurs laid eggs like modern birds; for several decades, we have also had clear fossil evidence that birds are the descendants of dinosaurs. So what makes this study so interesting? The eggs of modern birds are unique among vertebrate eggs in showing a huge variety of colours, hues and patterns. It was generally assumed that this was a fairly modern innovation of birds, but Wiemann and colleagues’ study demonstrates that coloured eggs date back more than 150 million years and probably had a single origin, within dinosaurs. Using Raman microspectroscopy (a technique that exploits the scattering pattern of light to identify molecular structures and material composition), the authors searched for traces of pigments in fossilized dinosaur egg shells (Fig. 2). With this method, they were able to identify two pigments — one responsible for giving the egg a blue-green hue, and another resulting in a red-brown hue. However, not all dinosaur eggs had the same colour and pattern. Some theropod dinosaurs, such as Deinonychus, laid blueish-green eggs with dark speckles, whereas others laid white to beige eggs. Interestingly, many herbivorous dinosaurs, including sauropods and hadrosaurs, laid eggs without any pigments. Those were probably more similar to the eggs of modern crocodiles.

But why did different dinosaur groups lay differently coloured eggs? Probably for the same reason that modern bird eggs show such a colourful variety. Coloured and speckled eggshells probably helped to camouflage eggs in open nests, and might also have helped the breeding parents to identify their own eggs (perhaps to avoid nest parasitism from cuckoo-like dinosaurs planting their eggs in other nests). The plain eggs of sauropods and ornithischians, by contrast, were probably left in nests covered by soil and dirt, making camouflage unnecessary. Eggs of different species have in fact been found in different styles of nest, which does support these results.

This study is one of several showing how many features associated with modern birds (such as feathers) have been around for millions of years in their ancestors. At the same time, this goes to show that although we have known many of these fossils for years, if not decades, new results can be obtained with the help of new methods and technologies.

Imran Rahman — Lefebvre et al., ‘Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes’

My favourite paper was actually only officially published this month, but given that it first came online in November 2018, I think it still counts! The study in question is by Lefebvre and colleagues, who described new fossil echinoderms from 480-million-year-old rocks in Morocco. The fossils belong to an extinct class called the stylophorans, which are arguably the most controversial group of fossil echinoderms ever known. The evolutionary relationships of stylophorans are hotly contested by palaeontologists, in part owing to disagreements about what soft parts were located inside the single appendage extending from the main body. Some have suggested that this appendage was a muscular tail for moving the animal over the sea floor, whereas others thought it represented a feeding arm that housed tube feet, similar to those of modern echinoderms. This disagreement stems from the fact that soft parts had never previously been described in a fossil stylophoran, leaving the interpretation of the appendage uncertain.

Lefebvre and colleagues were able to address this debate through the discovery of new, exceptionally preserved fossils. The fossils come from the Fezouata biota, a deposit, or Konservat-Lagerstätten, in southeastern Morocco. Similar to other famous Lagerstätte, Fezouata preserves the soft parts of animals, as well as their hard parts, and thus offers unique insights into past life. Incredibly, some of the stylophorans described by Lefebvre et al. have soft parts preserved inside the single appendage. In at least one specimen, these soft parts consist of a canal along the length of the appendage, with tube-like extensions coming off it to the sides (Fig. 3). Chemical analyses demonstrate that the soft parts are rich in iron, clearly distinguishing them from the rest of the fossil. The organization of the soft parts is almost identical to what we see in many modern echinoderms, which possess a series of fluid-filled canals (the water vascular system). This has a number of functions in living echinoderms, but one of the most important is feeding; for example, tube feet are used to capture food particles suspended in the water. The new Moroccan fossils make it highly likely that stylophorans used their single appendage primarily as a feeding arm, although they might also have been capable of moving slowly over the sea floor.

The finding that stylophorans possessed a feeding arm does not in itself unambiguously resolve the group’s relationships with other echinoderms. Nevertheless, this is a key discovery that greatly enhances our understanding of the palaeobiology of an important fossil group. It is discoveries such as this that drive palaeontology forward, by shedding new light on old questions. Who knows what we will find next!

Alan Spencer — Hetherington and Dolan, ‘Stepwise and independent origins of roots among land plants’

One of my favorite papers of 2018 was published by Hetherington and Dolan in Nature and shed new light on one of the fundamental questions associated with early plant terrestrialization: when did plants evolve roots?

All vascular plants we know today produce roots — which have important roles in nutrient and water uptake, ground anchorage and the symbiosis, or close mutual relationship, of plants and fungi. Roots are characterized by a self-renewing structure called a root meristem. This appears at the root tip and is covered by a ‘root cap’ allowing the roots to be guided by gravity through the substance in which the plant is growing. To date, the fossil record has hinted at root-cap development with known examples from the Carboniferous period (356 million years ago to 299 million years ago) and the Permian period (299 million years ago to 252 million years ago), but poor preservation of earlier land plants has hampered our understanding of where, when and how they first developed. In their paper, Hetherington & Dolan re-investigated historical plant specimens from the 407-million-year-old Rhynie chert Lagerstätte (Scotland, UK) and discovered in the species Asteroxylon mackiei the earliest meristems without root caps associated with a terrestrial ecosystem (Fig. 4).

Through a series of observations using the exceptional preservation of other plant tissues as a basis, they ruled out the loss of root-caps through taphonomic processes. They also looked for evidence at the cellular level in the preserved promeristem, where cell-division patterns can indicate root-cap formation. They found that the pattern was inconsistent with the development of root caps. However, the cell patterns indicated that A. mackiei had a continuous surface called the epidermis. To test this hypothesis, Hetherington & Dolan constructed 3D models of the meristem surface using confocal microscopy.

This is a modern technique that uses several laser beams (at varying wavelengths) to capture 2D images at different focal depths in a sample. Their results showed that there was a continuous and smooth layer of epidermis covering the meristem, and that there was no evidence of tapering or cells breaking off. This led to them to conclude that A. mackiei  developed rooting axes from a previously unknown type of meristem, which lacked both root caps and root hairs. Furthermore, this indicates that the evolution of rooting axes in the plant group lycopsids, to which A. mackiei belongs, occurred in a stepwise fashion. Their findings point towards independent root evolution in different plant lineages at differing points throughout geological time, with them all deriving from a common rootless ancestor. This study once again proves that using historical material with new techniques can throw up unexpected but welcome discoveries!

Suggestions for further reading:

Deline, B., Greenwood, J. M., Clark, J. W., Puttick, M. N., Peterson, K. J. & Donoghue, P. C. Evolution of metazoan morphological disparity. Proceedings of the National Academy of Sciences of the United States of America 115, E8909–E8918 (2018). (DOI: 10.1073/pnas.1810575115)

Hetherington, A. J. & Dolan, L. Stepwise and independent origins of roots among land plants. Nature 561, 7722 235–238 (2018). (https://doi.org/10.1038/s41586-018-0445-z)

Lefebvre, B., Guensburg, T .E., Martin, E. L. O., Mooi, R., Nardin, E., Nohejlová, M., Saleh, F., Kouraïss, K., El Hariri, K. & David, B. Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes. Geobios 52, 27–36 (2019). (DOI: 10.1016/j.geobios.2018.11.001)

Wiemann, J., Yang, T.-R. & Norell, M. A. Dinosaur egg colour had a single evolutionary origin. Nature 563, 555–558 (2018). (DOI: 10.1038/s41586-018-0646-5)

¹ The Palaeontology [online] editorial board can be found here.

Introduction

Every now and then at Palaeontology [online], we like to take a look at the world of palaeontology and reflect on what is happening in the field. Contrary to stereotypes, we believe that palaeontology and associated disciplines represent a fast-moving and exciting area of science. To highlight this, the members of the editorial board have each chosen a favourite paper from 2017. Picking just one paper was difficult for all of us, and it means that we have highlighted just five articles out of the many hundreds published in the past 12 months. Nevertheless, we hope that our choices reflect the breadth and depth of palaeobiological research in the twenty-first century. The papers include incredibly small and ancient invertebrates, wonderfully preserved dinosaurs, important marine mammals, the application of techniques from other disciplines to help us understand ancient echinoderms, and the evolution of the royal ferns. We also couldn’t help but notice the festive season approaching — and so we have each taken the opportunity to briefly describe a favourite fossil for the holidays. So, in alphabetical order, here is our editorial board with their highlights.

Russell Garwood — A festive arachnid

One of my favourite fossils is Eophalangium sheari. Just as the end of the holiday season marks the start of a new year and all the changes this can bring, this fantastic creature is found at a turning point in evolution

It is one of the earliest land animals, a member of one of our best preserved terrestrial ecosystems from the Palaeozoic era (542 million to 251 million years ago). It is a wonderfully preserved harvestman arachnid — part of a group that is still around today — and is found alongside some of the earliest complex plant fossils and a number of other terrestrial arthropod fossils. It is so well preserved, we can even see its internal anatomy (Fig. 1). If I were to find this in my stocking, what a Christmas that would be!

Russell Garwood — Favourite invertebrate-fossil paper of the year: ‘Exceptionally preserved Cambrian loriciferans and the early animal invasion of the meiobenthos’ by Harvey and Butterfield in Nature Ecology and Evolution

There have been a lot of really exciting papers this year — in invertebrate fossils, the development of new methods and the study of evolution — which has made picking just one as a highlight something of a challenge. Nevertheless, needs must. One favourite of mine this year was the work of Harvey and Butterfield, who have described an assemblage of loriciferans from the late Cambrian period (around 497 million to 485 million years ago).

This phylum of tiny animals is closely related to arthropods and to priapulid worms. Their fossil record is limited because of both their minute size and the fact that they are unlikely to fossilize. This is true of many animals of this size that live in sediments: animals that constitute something called the meiofauna. This broad collection of organisms makes significant contributions to modern ecosystems but, because of a lack of fossils, their origins and relationships are hard to untangle.

The fossils described in Harvey and Butterfield (2017) are about one-third of a millimetre across as adults, showing that early in the group’s history, the loriciferans had become part of the meiofauna  (as they have been ever since). Because they are rather complex animals, however, the authors suggest that the loriciferans evolved from a larger-bodied ancestor. That they have been recovered as fossils — the researchers dissolved mudstones in very strong acid to retrieve them — also suggests that future studies may find more meiofauna.

That could help us better understand early animal evolution. Indeed, the paper concludes by saying that these approaches “offer a direct window onto cryptic animal ecosystems through deep time”. So, as well as helping us understand the evolution of this particular group, there is also scope for exciting future discoveries with broad implications.

Stephan Lautenschlager — A festive dinosaur

Dinosaurs might not have been well equipped to deal with snow and ice, but that doesn’t mean there are no polar dinosaurs. The single known Nanuqsaurus hoglundi (the ‘polar-bear lizard’) fossil was found in northern Alaska, well beyond the Arctic Circle (Fig. 3). This small (roughly 3-metre) tyrannosaur lived during the late Cretaceous period (about 70 million years ago), when Alaska was ice-free and the temperatures were more moderate than today.

Stephan Lautenschlager — Favourite dinosaur paper of the year: ‘An exceptionally preserved three-dimensional armored dinosaur reveals insights into coloration and Cretaceous predator-prey dynamics’  by Brown et al. in Current Biology

As usual, this year has seen its fair share of dinosaur papers. Some of those have shaken the very foundations of dinosaur palaeontology by proposing a substantial reorganization of the dinosaur family tree (Baron et al., 2017). Instead of the longstanding division of dinosaurs into saurischians and ornithischians, Baron and colleagues argue for combining theropods and ornithischians in a new group, named Ornithoscelida. This has prompted a storm of public and academic discussion, but one thing became eminently clear from it: a lot more fossils — and well preserved ones — are needed to resolve such controversies.

One such fossil find this year was Borealopelta, an exceptionally well preserved ankylosaur from Alberta, Canada (Brown et al., 2017; Fig. 4). This armoured dinosaur from the early Cretaceous (around 100 million to 113 million years ago) was found in fine-grained sandstones and shales representing an offshore environment, indicating that the carcass was washed into the sea shortly after the dinosaur died. Quick sedimentation and the low oxygen levels at the sea floor made sure that the body was quickly covered, and was preserved nearly intact for the following 100 million years. As a consequence, Borealopelta is one of the best ankylosaur specimens ever found, preserving not only many details of its body still in the original position, but also remnants of skin and horn sheaths covering the body armour. The fossil therefore looks unusually lifelike and almost if it was just asleep and not dead for millions of years. But that is not all. Pigments preserved in the fossilized soft tissues further revealed that the dinosaur had a reddish-brown colour pattern, which might have had a function in camouflaging it from predators.

Rachel Racicot —  Festive fossil dolphins

Odobenocetops peruvianus and Odobenocetops leptodon are archaic dolphins that seem to be a combination of several living Arctic-specialized species, such as the narwhal (Monodon monoceros) and walrus (Odobenus rosmarus). Odobenocetops fossils come from the Pliocene Pisco Formation, a rock formation between 3 million and 4 million years old on the southern coast of Peru. They are probably closely related to or nested in the group that includes narwhals and belugas (Monodontidae). The genus has an unusual shape for a dolphin (Fig. 5), including a skull shape similar to that of the walrus and knobs at the base of the skull indicating a good amount of neck mobility. This suggests that this animal probably fed on molluscs and other invertebrates from the sea floor, similar to extant walruses.

Rachel Racicot — Favourite whale paper of the year: ‘Infrasonic and ultrasonic hearing evolved after the emergence of modern whales’ by Mourlam and Orliac in Current Biology

Many excellent papers have come out this year on marine-mammal fossils, setting the stage for renewed interest in the diversity of animals and the effects of Earth systems during critical intervals of marine-mammal evolution. Mourlam and Orliac investigated the inner-ear labyrinth of a variety of whales and their extant and extinct relatives (Fig. 6). The detailed analysis describes for the first time the anatomy of two whales from the early Eocene epoch (46 million to 43 million years ago), part of a group that was semiaquatic and could still walk on land (Protocetidae). The results show that these archaic ‘walking whales’ had an intermediate hearing range between land-dwelling artiodactyls (even-toed ungulates such as deer) and ocean-dwelling artiodactyls (whales and dolphins). The results are an important step forward in understanding the ecology and evolution of ancient whales as researchers begin to investigate further whether later-diverging, more fully aquatic-adapted whales had different hearing physiologies and ecologies from one another as they coexisted in the late Eocene epoch.

Imran Rahman — A festive starfish

In the run up to the holiday season, chances are you will see lots of Christmas trees with stars on top, but what could be more festive than your very own fossil starfish! The fossil record of starfish, or sea stars, is extensive, dating back to the Ordovician period, more than 450 million years ago. Some of the most famous examples come from a 400-million-year-old site of exceptionally preserved fossils in Germany, called the Hunsrück Slate, where starfish and other fossils are beautifully preserved as pyrite, or fool’s gold (Fig. 7).

Imran Rahman — Favourite echinoderm paper of the year:  ‘A new model of respiration in blastoid (Echinodermata) hydrospires based on computational fluid dynamic simulations of virtual 3D models’ by Waters et al. in the Journal of Paleontology

2017 was a great year for research on fossil echinoderms — the group of marine animals that includes starfish and sea urchins — including a bumper special issue in the Journal of Paleontology called ‘Progress in Echinoderm Paleobiology’ (co-edited by yours truly and currently free to access here). One of the more intriguing studies in this issue was that of Johnny Waters and colleagues, who focused on a peculiar extinct group of echinoderms called blastoids. Blastoids look superficially similar to crinoids (sea lilies), a group of echinoderms that is still alive today, but it is unclear exactly how closely related the two groups are. One particularly interesting characteristic of blastoids is their internal respiratory structures, called hydrospires. These structures are rather different from anything seen in living animals, and so working out how they functioned is a major challenge for palaeontologists.

Waters and colleagues analysed the function of blastoid hydrospires using a method called computational fluid dynamics, which is commonly used in engineering to test the performance of cars and planes. They created a virtual water tank on a computer and placed a 3D model of a fossil blastoid into it. They then ran simulations of water currents around and through the digital blastoid to test how the hydrospires affected fluid flow. Their results suggest that water entered the hydrospires through pores along the sides of the animal, travelling within these structures at low velocity to allow oxygen from the water to enter the animal’s cells, before exiting near the mouth. In life, blastoids were probably positioned with their stems bent over, so that the hydrospires were oriented horizontally (Fig. 8). Studies such as that of Waters and colleagues demonstrate the great potential of using methods from other disciplines, such as engineering, to reconstruct the lifestyles of extinct fossil animals.

Alan Spencer — A festive palaeoplant

Although plants do not celebrate Christmas, or thrive in cold, snowy conditions, there are many fossil plants that today are recovered from such environments, in particular from Antarctica.

The most famous was discovered by Robert Falcon Scott’s doomed 1911–12 expedition, and is called Glossopteris indica. This prize fossil, an extinct 250-million-year-old beech-like tree, was found next to Scott’s body. The study of the fossil and similar Glossopteris found in Australia, Africa and South America formed one of the ‘proofs’ that Earth’s land masses once made up an ancient super-continent named Gondwanaland (now just called Gondwana; Fig. 9).

Alan Spencer — Favourite palaeobotanical paper of the year: ‘The fossil Osmundales (Royal Ferns) — a phylogenetic network analysis, revised taxonomy, and evolutionary classification of anatomically preserved trunks and rhizomes’ by Bomfleur et al. in PeerJ

Papers dealing with plant evolution — whether based on morphological similarities or genetic characteristics — tend to fall into the category of ‘scientifically informative, but visually unappealing to read’. This is due in part to the nature of the information: vast data sets coding in 0 and 1, and for excitement the odd other numeral, leading to monotone evolutionary-tree diagrams. It was therefore highly pleasing to read Bomfleur et al.’s contribution dealing with the affinities of the Osmundales — also known as the royal ferns (Fig. 10) — the oldest surviving lineage of leptosporangiate ferns, which can be traced back to the Permian period  (298 million to 252 million years ago), part of the late Palaeozoic era.

The authors first tackled the thorny problem of standardization within specimen descriptions and their systematic classification (Fig. 11) — before this work, since the last family wide review in 1971 by Miller, 50 new species, 12 new genera and 4 new subfamilies have been described, all using differing terminology. The amount of work involved in standardizing them cannot be understated, and will provide a framework for the description of future fossil Osmundales specimens.

Second, in this work, each character defined for use in the ‘systematic–phylogenetic analysis’ has been illustrated in colour (Fig. 12) and the individual coding shown with symbols.

This is not only very visually pleasing, but reduces the interpretive errors of any future work by showing exactly what has been coded into the matrix. Third, this paper goes one step further, by combining the morphological data with a phylogenetic analysis based on living Osmundales. This is an interesting technique that has rarely been used in fossil-plant studies. The authors’ results have allowed a complete reclassification of the fossil Osmundales.

Conclusion

As you can see, 2017 has been a good year: methods by which we can understand past life have been developed, many exciting new fossils have been discovered, and other interesting findings have been achieved by revisiting old fossils that have been described before. Looking forward into 2018, the future is bright. Palaeontology is a vibrant and exciting discipline, and who knows what we may discover!

Suggestions for further reading

Baron, M. G., Norman, D. B. & Barrett, P. M. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543, 501–506 (2017). DOI: 10.1038/nature21700

Bomfleur, B., Grimm, G. W. & McLoughlin, S. The fossil Osmundales (Royal Ferns) — a phylogenetic network analysis, revised taxonomy, and evolutionary classification of anatomically preserved trunks and rhizomes. PeerJ 5, e3433 (2017). DOI: 10.7717/peerj.3433.

Brown, C. M., Henderson, D. M., Vinther, J., Fletcher, I., Sistiaga, A., Herrera, J. & Summons, R. E. An exceptionally preserved three-dimensional armored dinosaur reveals insights into coloration and Cretaceous predator-prey dynamics. Current Biology 27, 2514–2521 (2017). DOI: 10.1016/j.cub.2017.06.071

de Muizon, C. Walrus-like feeding adaptation in a new cetacean from the Pliocene Peru. Nature 365, 745–748 (1993). DOI: 10.1038/365745a0

de Muizon, C. & Domning, D. P. The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its palaeobiological implications. Zoological Journal of the Linnean Society 134, 423–452 (2002). DOI: 10.1046/j.1096-3642.2002.00015.x

Dunlop, J. A., Anderson, L. I. Kerp, H. & Hass, H. A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts, Aberdeenshire, Scotland. Transactions of the Royal Society of Edinburgh, Earth Science 94, 341–354 (2004). DOI: 10.1017/S0263593300000730

Fiorillo, A. R. & Tykoski, R. S. A diminutive new tyrannosaur from the top of the world. PLOS One 9, e91287 (2014). DOI: 10.1371/journal.pone.0091287

Harvey, T. H. & Butterfield, N. J. Exceptionally preserved Cambrian loriciferans and the early animal invasion of the meiobenthos. Nature Ecology & Evolution 1, 0022 (2017). DOI: 10.1038/s41559-016-0022

Miller, C. N. Evolution of the fern family Osmundaceae based on anatomical studies. Contribution from the Museum of Paleontology 23, 105–169 (1971).

Mourlam, M. J. & Orliac, M. J. Infrasonic and ultrasonic hearing evolved after the emergence of modern whales. Current Biology 27, 1776–1781 (2017). DOI: 10.1016/j.cub.2017.04.061

Waters, J. A., White L. E., Sumrall, C. D. & Nguyen, B. K. A new model of respiration in blastoid (Echinodermata) hydrospires based on computational fluid dynamic simulations of virtual 3D models. Journal of Paleontology 91, 662–671 (2017). DOI: 10.1017/jpa.2017.1

¹The Palaeontology [online] editorial board can be found here.

Introduction:

Geoconservation, also known as Earth Heritage Conservation, is how we protect important examples of Earth’s physical resources. Geological features can be protected for all sorts of reasons, including being important to cultural heritage, geological education and understanding, or the overall aesthetics of an area.

A great many designations, management frameworks and legal instruments have been used to govern and protect fossil-rich outcrops in the United Kingdom, but these are poorly publicized and, for example, rarely taught to palaeontologists as part of an undergraduate degree. Field work is an important part of palaeontological research, so it is a good idea for everyone who works with fossils, whether amateur or professional, to have a good understanding of the law so that they do not fall foul of it.

Here I summarize various areas of English law that apply to palaeontological research, fossil collecting and geoconservation. Historical examples have been added for context. It is important to note that this article will mainly consider the situation in England, and not the entire United Kingdom. Much legislation is similar across England, Wales, Scotland and Northern Ireland, but conservation policy is handled separately by each of these nations so there are differences. Other countries will of course have their own policies. This article does not constitute formal legal advice.

What is a fossil?

Palaeontologists may sometimes debate what is and is not a fossil, but more often than not they broadly agree on what structures represent evidence of ancient life, and what that means. The same is not true in English law, where protection of fossil specimens is complicated by a lack of clarity as to what fossils legally are. The absence of a specific definition in legislation has led to the matter being decided in case law — this is the legal precedent set by the rulings of judges in courts.

An early legal case of interest is that of Attorney General v. Tomline in 1877. Colonel George Tomline was lord of the manor of Walton-cum-Trimley in Suffolk — now the site of the Port of Felixstowe. The British War Department was a tenant on part of Tomline’s land, and had constructed a circular fort called a Martello tower to defend that part of the coastline. The land was underlain by deposits from the Pliocene epoch (5.3 million to 2.6 million years ago) containing coprolites — fossilized animal faeces — which at the time were an important source of phosphate for the fertilizer industry.

Tomline, wanting to cash in on this valuable resource, began to dig a trench on the property, removing and selling the coprolites. The War Department, not happy at the disturbance and open earthworks next to the fort, took Tomline to court to get him to stop.

The judge in the case, Justice Fry, ruled that as owner of the land, Tomline was entitled to the coprolites, but that he required the permission of the tenants to build the earthworks to remove them. Because he hadn’t sought permission, the War Department was awarded half of the profits from the coprolite sales.

As ultimate landowner, Tomline possessed the mineral rights to the property, meaning that he had the right to use any minerals found on the land. The judge cited the case of Hext v. Gill 1872, in which Lord Justice Mellish stated that the term minerals

“… includes every substance which can be got from underneath the earth for the purpose of profit, unless there is something in the context or in the nature of the transaction to induce the Court to give it a more limited meaning.”

Justice Fry ruled that coprolites were also minerals, and thus the property of Colonel Tomline. Yet, unfortunately for the colonel, the judge didn’t feel that this gave him the right to disturb the property to remove the coprolites — hence the need to pay damages to the tenant.

As a result of this ruling, it would seem that fossils still in the ground are the property of whoever owns the mineral rights, and cannot be collected without the permission of the landowner and tenant (if applicable). More-recent laws — the Theft Act, 1968, and the Criminal Damages Act, 1971 — could be used to prosecute people who remove samples without permission. Fossils no longer attached to the bedrock are a more complicated matter. In England and Wales, the prevailing opinion is that loose specimens are considered abandoned, and therefore that taking them is not stealing. However, in Scotland all abandoned property reverts to the Crown, so theoretically permission should be sought before removing loose material.

Fossil Export

Exporting fossils from the United Kingdom is also complicated. This came to light in the 1980s following the discovery of Westlothiana, possibly the earliest known reptile, from the Carboniferous period (360 million to 299 million years ago). When a German museum attempted to buy the specimen from the collector for £180,000, an export licence was applied for under the Export of Goods (Control) Order, 1987, which is required for anything valued at more than £20,000 and that is more than 50 years old. However, the Department of Trade and Industry ruled that because fossils are not manufactured, they are not subject to the Order. This therefore leaves a situation where the only legislation controlling the export of fossils is at the European Union level. As a result, there is little to prevent exports to EU countries.

Geoconservation Legislation

The various forms of legislation that cover areas where fossils can be found are complex and multifaceted. They apply to both researchers and fossil collectors. The main designation that a locality can be listed as for the purpose of geoconservation in England is that of a Site of Special Scientific Interest (SSSI), under Section 28 (amended) of the Wildlife and Countryside Act, 1981 (WC Act). In England, SSSIs are overseen by Natural England, a non-departmental public body. Land with SSSI status is still owned by whoever owned it before it was designated an SSSI, but there are restrictions on what people can do there, especially in terms of building and development. SSSIs are not normally patrolled or monitored on a regular basis by Natural England. The original WC Act made it very difficult to prosecute third parties who damaged or destroyed SSSIs, because it was necessary to prove that the person knew that they were in an SSSI. A new offence was created by Section 55 of the Natural Environment and Rural Communities Act, 2006 (NERC Act): that of intentionally or recklessly damaging or destroying the natural conservation or geological features of an SSSI. Importantly, this offence does not require prosecutors to prove that there was knowledge of being within a designated area. People who are convicted can be given a fine up to level 4 on the Standard Scale (£2,500 at time of writing). The Standard Scale is a system for setting criminal financial penalties, allowing for increases in the level of the scale when inflation requires, without the need to amend all the laws.

The management plan for each SSSI includes a list of “operations likely to damage the special interest”. Among other things, SSSIs may include controls on the “extraction of minerals, including peat, topsoil, subsoil and stone” and the “removal of geological specimens, including rock samples, minerals and fossils”. Anyone who wants to extract or remove geological specimens from an SSSI including these controls must get permission from Natural England.

Areas of geological significance are often in Areas of Outstanding Natural Beauty (AONB). This designation offers protection from development, but no extra regulation of access to or removal of specimens. The power to designate and oversee AONBs rests with Natural England.

A great deal of England’s upland areas are also open-access land where there is what is called a right to roam — meaning that anybody has the right to walk there. Natural England is able to restrict access for “the purpose of conserving flora, fauna or geological or physiographical features” under Section 26 of the Countryside and Rights of Way Act, 2000 (CROW Act). It is possible to restrict access for the purpose of geoconservation, but it is thought that this power has never been used. The CROW Act also made amendments to previous legislation about SSSIs and AONBs.

The WC Act also establishes Limestone Pavement Orders to protect natural landforms made of exposed limestone. Most of these Orders are in place in the Yorkshire Dales National Park, or in the country between Kendal and Lancaster. Removal of rock from these areas, even if it is loose or lying in a field, is a criminal offence.

The National Parks & Access to the Countryside Act, 1949 (NPAC Act), allowed for the creation of national parks. These offer very limited protection to specific outcrops of rock, and are mainly concerned with controlling development. The NPAC Act also enabled the designation of National Nature Reserves. Unlike other designations, these areas of land are managed directly on behalf of the nation — usually by Natural England, but also by other bodies, such as Wildlife Trusts, the Royal Society for the Protection of Birds (RSPB) and local authorities. There may be restrictions on sampling and research in these areas. An example of a geological National Nature Reserve is the Wren’s Nest in Dudley, renowned for its coral-reef deposits from the Silurian period (444 million to 419 million years ago). These deposits contain fossils of the trilobite Calymene blumenbachi, known locally as the Dudley Bug.

There are also a number of other designations, not directly created for geoconservation, that may affect palaeontological research. These include Country Parks, Local Nature Reserves and Scheduled Ancient Monuments. Sites may also be designated as Regionally Important Geological Sites, although these are not supported by legislation and have little effect other than to recognize the importance of an area. International inscriptions, such as UNESCO World Heritage Site or Geopark, may also be used to highlight the worldwide significance of a site, but other designations would be required to control collection and research on fossiliferous outcrops.

The English conservation system has been built mainly to control of development in and around a site. SSSI or AONB status does not affect the ownership of the land, so control of access to the property is for the most part left to the landowner.

The English system has an obvious bias in favour of managing development and the actions of landowners; this is evident from the language used in the CROW Act, where “owner or occupier” is used to describe people who may apply for permission to do something that is otherwise prohibited in an SSSI. If this is refused, they may also appeal to the Secretary of State. This disregards the reality that many of those who may legitimately seek a permit to remove samples, including researchers and fossil collectors, are highly unlikely to be the landowner.

Case Study: The Long Mynd

The Long Mynd in Shropshire, England, is a heath and moorland plateau owned by the National Trust, a conservation charity operating in England, Wales and Northern Ireland. The Long Mynd is the type locality — meaning the place where certain rocks types were first defined — for the Longmyndian Supergroup, one of the thickest and most complete packages of sedimentary rock from the Precambrian (everything before 541 million years ago) in southern Britain.

The rocks here have been of interest for some time, with geologists such as John Salter and Charles Darwin writing about them and associated possible traces of life. The roughly 2,000-hectare site is protected by byelaws, which were created by the National Trust under Section 24 of the National Trust Act, 1971.

The byelaws prevent, among other things, the unauthorized removal of gravel, sands, clay or any other substance from Trust property, and the defacing of rock surfaces. Failure to comply with the byelaws can lead to a fine of up to £20, plus an extra £2 for each day the offence continues.

The Long Mynd is also in the Shropshire Hills AONB, and much of it is open-access land (Fig. 1).

As can be seen from the simplified diagram in Figure 2, a patchwork of legislation and governance affects the Long Mynd. These designations not only cover different areas, but also involve various different pieces of legislation and nongovernmental organizations. A key message of such diagrams is that understanding the constraints on palaeontological research in an area can be quite complex, and field work at a particular locality may require permissions or permits from a number of organizations.

Responsible conduct in the field — A guide

The key to conscientious field work and fossil collecting is being aware of your responsibilities. The most important of these is to understand the legal framework under which you are working. In England, the easiest way to check this is to go to magic.defra.gov.uk, which provides a detailed map with overlays for each of the various designations. If necessary, you can find further information through the Natural England website for information on restrictions in those designated areas.

You may also need permission from the landowner. Where it isn’t clear who owns the land, this information can be ordered online through the Land Registry for a small fee.

Don’t underestimate the value of fostering positive relationships with the local communities in which you work. In a conservation regime where sites are very rarely formally patrolled and monitored, it is very useful for a researcher to become friendly with members of the local community and enlist them to keep an eye on important fossil localities. As well as acting as informal year-round wardens, residents often have a sense of ownership over their local countryside, and so will be keen to be kept abreast of research in the area. This approach not only promotes stronger informal geoconservation monitoring, but also often leads to researchers getting tip-offs from the local community about prospective new sites and discoveries.

When it comes to sampling, there are a number of options. Removing specimens damage the appearance of an area, reducing its aesthetic value and possibly destroying geological features that may be of use to future researchers. Samples should therefore be taken from loose material if possible. If this is not possible, it is often best to sample in areas that are not easily seen, and are away from other specimens that may be of future importance. It is also worth considering how you remove the specimens — using a hammer and chisel along existing weaknesses in the rock tends to leave natural-looking exposures, whereas diamond disc saws would leave unsightly gashes on the outcrop. In general, the adage of ‘leave only footprints’ is a good one to follow.

Conclusions

Legislation alone cannot ensure the long-term protection of geologically significant sites; it is of little use once a rare and finite resource has been removed or destroyed. Total security can never be guaranteed, but if the most scientifically significant localities are to be preserved, it is essential to have monitored — and preferably staffed — access controls.

Fossil localities are as significant to communities through enhancement of the local cultural heritage and economy as they are to researchers and the development of knowledge. Indeed, with researchers often being temporary annual visitors, and many sites having no permanent staff or monitoring, local communities are often the best guardians of geologically significant localities.

It is therefore essential that geoconservation frameworks, both legislative and managerial, are designed in close consultation with both the scientific community and local residents. Future legislative advances should seek to simplify the current framework, and ensure that designations recognize the needs and concerns of all partners, while ensuring the long-term sustainability of these fossil resources.

How the law on geoconservation would be applied remains unknown, because few people have been prosecuted for breaking it. Significant uncertainty still exists on the definitions of certain key terms and how they would be applied in law. Moreover, awareness of geoconservation laws is very low. Sites rarely contain any signage information telling visitors of their rights and responsibilities, or what to do if they discover vandalism. Field work remains an important part of palaeontological study, and fossil collecting is common. It is therefore essential that new researchers and fossil collectors get adequate training and guidance on the legislation as it applies to them. For the former, it would seem logical to make the laws around fossil collecting a small, compulsory element of all geological undergraduate degrees. For the latter, increased engagement, discussion and more publically available resources would help improve awareness. Geological outcrops remain the foundation not only of palaeontological research, but also of teaching and outreach. Stakeholders from across society should invest more time to ensure that these finite resources are not lost forever.

¹Department of Earth Sciences, Memorial University of Newfoundland, St John’s, NL, A1B 3X5, Canada. 2Oxford University Museum of Natural History, Parks Road, Oxford, OX1 3PW, U.K. Email: jjmatthews@mun.ca.

Introduction:

We’re now into our sixth volume — and calendar year — at Palaeontology [online]. Over the years, we have introduced a lot of fossil groups, concepts from palaeontology and overviews of different parts of our field. An intention when we started was also to provide the occasional overview of happenings in the world of palaeontology: to reflect new developments and highlight some current ideas. To that end, we have chosen to start 2016 by looking back over the past year, and forward into the next. In this article, members of the Palaeontology [online] team have chosen their favourite papers from 2015, and indicated what they hope to be up to over the next 12 months. So without further ado, here is team Palaeontology [online]!

Imran Rahman:

My favourite paper of 2015 — Porro et al. (2015): One of the most important events in the history of life on Earth was the colonization of land by vertebrates, which was fundamental to the group’s subsequent success and modern diversity. Study of the fossil record has allowed palaeontologists to reconstruct this transition in some detail, using a number of fossils of early tetrapods that show the step-by-step transformation from a body adapted for life in water to one adapted for life on land. Acanthostega is one such animal: it lived during the Devonian period (about 365 million years ago) and combines a range of fish-like and tetrapod-like features. A paper published last year by Laura Porro, Emily Rayfield and Jenny Clack used computed tomography (CT) to describe the fossil in unprecedented detail. Acanthostega was originally described in the 1950s by the Swedish palaeontologist Erik Jarvik, and revised in the late 1980s by Clack, an English palaeontologist. The new study by Porro and colleagues was able to build on previous descriptions by constructing a three-dimensional (3D) computer model of Acanthostega, revealing previously hidden and poorly known aspects of the fossil’s shape and structure, or morphology (Fig. 1). This paper provides an excellent case study of the value of CT for studying fossils in great detail. It also serves as a timely reminder that detailed descriptions of fossils remain the foundation of the science of palaeontology.

My plans for 2016: One of the main goals of my research over the next year will be to investigate the evolution of symmetry. In particular, I hope to address how and why echinoderms — sea urchins, starfish and related marine creatures — evolved five-fold symmetry, a characteristic that is rare among modern animals, but common to all living species in the echinoderm group. Intriguingly, the fossil record of echinoderms shows that they were not always like this. Fossil echinoderms displayed a range of different types of symmetry, from bilateral, or mirror-plane, to three-fold and five-fold, but only the last of these remains today. I will analyse early fossil echinoderms using computer modelling to test hypotheses about why five-fold symmetry came to dominate the group.

Russell Garwood:

My favourite paper of 2015 — Brasier et al. (2015): When it comes to looking at ancient life, some of the hard questions hinge on interpretations of what you see in the rock. Does a particular structure represent something that was once living, or is it non-biological in origin? If the former, what kind of organism might it have been, and how do we tell? Answering these questions in an objective way can be very challenging, and interpretations depend in equal parts on the geological setting of a fossil (and hence its taphonomy, or preservation), and on educated speculation from biology — plus, ideally, a healthy dose of scepticism. The authors of one of my favourite papers from 2015 approach these questions using new ways of studying fossils. In this study, they analyse three putative fossil deposits to show the power of a range of cutting-edge methods. In one technique (focused-ion beam, or FIB), they use charged molecules called ions to create very thin wafers of a 3.46-billion-year-old putative fossil from an Australian rock called the Apex chert. Analysing these with very high-resolution transmission electron microscopy, they suggest that structures previously thought to be the fossils of early cells are more likely to be clay-like minerals altered by high-temperature fluids. They use related methods to create 3D reconstructions of some 3.43-billion-year-old structures from another Australian rock called the Strelley Pool Formation. The authors propose that these structures represent some of the earliest known organisms, single cells eking out an existence between sand grains. They finish by analysing structures that are widely accepted to be fossils, from a 1.88-billion-year-old rock found in Canada, the Gunflint Chert. They suggest that one particularly complex organism may be multicellular (Fig. 2) — and if so, very few organisms like it are alive today. I appreciate the authors’ approach in this paper because it demonstrates that by using new techniques and pushing boundaries, we can start to unravel the interplay between geology and biology in complex structures. As a result, interpretations of ancient fossils can be based on an increasingly wide array of evidence. On another level, I believe that to create 3D models of fossils as small as 10 micrometres across is a stunning technological achievement that deserves recognition in itself.

My plans for 2016: One goal for my research in 2016 is to publish work on specimens that document major evolutionary transitions. Over the past few years, I’ve been applying high-resolution CT to a number of fossils rather removed from my usual comfort zone of insects and arachnids. I hope that by using digital visualization to study these, my colleagues and I can provide a little extra information on fossils dating from soon after the evolution of hard parts, and after life first made its way onto land. But I am also expecting to work on a creepy-crawly or two!

Rachel Racicot:

My favourite paper of 2015 — Costidis and Rommel (2015): Vertebrate paleontology requires a detailed understanding of modern animals, particularly their skeletal and soft-tissue anatomy, so that we can make inferences about the biology of extinct animals. We have additional motivation to study the head anatomy of whales, because humans may be dramatically impacting the behaviour of these animals with naval sonar, shipping noise and other human-made sounds and activities. Beaked whales, a rarely seen group of deep-diving toothed whales that use echolocation, seem to have a particularly rough time — for example, events in which they have become stranded on land have been associated with naval use of sonar. Dissections of stranded whales have shown signs of decompression sickness, indicated by fat embolisms (similar to blood clots but with fat) in the tissues surrounding the ear and other specialized sound-reception areas in the heads of these animals. The authors of this study take a detailed anatomical look at the veins (notoriously difficult to study under most circumstances) in the heads of a number of specimens of beaked whales, with a specific focus on the sound-reception regions (Fig. 3). They find intricate connections between blood vessels and acoustic fat bodies (specialized areas important for sound reception and production). This has implications for how nitrogen gas moves between the blood and fat tissue (important for avoiding decompression sickness on deep dives), and for fat clots that may be caused by trauma induced by responses to naval sonar deployment. My favorite part of this paper was the 3D PDF with isolated anatomical regions that one can rotate in the digital version of the paper! This study really embraces modern technological advances and I look forward to seeing more from these great anatomists (and citing them in my work on fossil whales).

My plans for 2016: I plan to continue my work on corals, which uses both morphological and molecular data sets to investigate how living and fossil corals are related. This work will make it easier to put fossils into groups with modern corals to form a more accurate and complete coral family tree, and may also help to plan for their conservation (which is critical given the present biodiversity crisis). Other work in the pipeline includes more projects on whale inner ears and brain evolution in both extant and fossil animals, as well as some new work on dinosaurs from the Transantarctic Mountains!

Peter Falkingham:

My favourite paper of 2015 — Molnar et al. (2015): I have quite an interest in how we can work out how much a joint would move from bones alone. If we are to reconstruct how extinct forms such as dinosaurs moved, we need to know how much their joints could move. The trouble is that you can’t just take bones and wiggle them together, because in life all the soft tissues around them — cartilage, muscles, skin, ligaments — affect the range of motion. Molnar and colleagues did a great study looking into the range of motion of vertebrae in extinct and extant crocodylomorphs, the group that includes crocodiles and their relatives. They physically measured dead specimens at various stages of dissection, and also used computer-based CT and animation techniques — a really cool combination. Their study agreed with the idea that osteoderms (bony structures in the skin) in early crocodylomorphs helped to stabilize the back.

My plans for 2016: This year should see the publication of a book on dinosaur tracks that I’ve been editing along with Annette Richter and Daniel Marty. It’s been several years in the making, but we’re finally on the home stretch, and it will be great to see it come out. I’m also hoping to make serious headway on a follow-up to my 2014 paper with Steve Gatesy. Perhaps more exciting, though, is that this year I’ll get my first PhD student. She’ll be studying plesiosaur neck hydrodynamics, and I’m looking forward to seeing that project come to life and evolve. Aside from research, I’ll be developing some new courses to teach on evolution and locomotion, as well as continuing my current teaching in those areas.

Suggestions for further reading:

Brasier, M. D., Antcliffe, J., Saunders, M. & Wacey, D. Changing the picture of Earth’s earliest fossils (3.5–1.9 Ga) with new approaches and new discoveries. Proceedings of the National Academy of Sciences 112, 4859–4864 (2015). DOI: 10.1073/pnas.1405338111
Costidis, A. M. & Rommel, S. A. The extracranial venous system in the heads of beaked whales, with implications on diving physiology and pathogenesis. Journal of Morphology 1, 34–64 (2015). DOI: 10.1002/jmor.20437
Falkingham, P. L. & Gatesy, S. M. The birth of a dinosaur footprint: Subsurface 3D motion reconstruction and discrete element simulation reveal track ontogeny. Proceedings of the National Academy of Sciences 111, 18279–18284 (2014). DOI: 10.1073/pnas.1416252111
Molnar, J., Pierce, S. E., Bhullar, B.-A. S., Turner, A. H. & Hutchinson, J. R. Morphological and functional changes in the vertebral column with increasing aquatic adaptation in crocodylomorphs. Royal Society Open Science 2, 150439 (2015). DOI: 10.1098/rsos.150439
Porro, L. B., Rayfield, E. J. & Clack, J. A. Descriptive anatomy and three-dimensional reconstruction of the skull of the early tetrapod Acanthostega gunnari Jarvik, 1952. PLoS ONE 10, e0124731 (2015). DOI: 10.1371/journal.pone.0124731
Zamora, S. & Rahman, I. A. Deciphering the early evolution of echinoderms with Cambrian fossils. Palaeontology 57, 1105–1119 (2014). DOI: 10.1111/pala.12138

Palaeontology and the history of life are topics that capture the imaginations of children and adults alike, many of whom are keen to learn all about the latest weird wonders discovered by science. Communicating cutting-edge research to the public can be difficult, however, partly because scientists tend to publish their results in technical papers aimed at other scientists, but also because these papers are generally not freely available to non-academics — although the growth of open access means that this is improving in many countries. Popular-science writers provide accurate and accessible summaries of some of the most topical work, but can cover only a fraction of the research carried out by palaeontologists, naturally focusing on current stories rather than the wealth of knowledge already out there. Moreover, misrepresentations of palaeontologists’ work are commonplace in the mainstream media, with research results often reported incorrectly, or lacking essential context. Taken together, all this hampers the public understanding of Earth history and contributes to the denial of evolution and climate change.

These issues were foremost in our thoughts when we set up Palaeontology [online] in 2011. The goal of the site is to publish monthly articles on palaeontology and related topics, written by experts, but understandable to secondary/high-school students and non-specialist adults. All of these articles are free to access, ensuring that anyone, anywhere in the world can read them whenever they want — all they need is an internet connection. The website, which is supported by the Palaeontological Association, officially launched to the public in July 2011 with an article by Alistair McGowan about biodiversity. Since then, we have published a total of 37 articles, which have been seen by more than 60,000 unique visitors from 179 countries (Fig. 1). To celebrate our three-year birthday, this month we take a quick look back at the articles that have helped to make the site the success it is today.

Of course, palaeontology isn’t just about fossils, and there are several articles dealing with large-scale patterns in Earth history (Fig. 3). These include the Paleocence–Eocene Thermal Maximum, the Cambrian explosion, the origin of life, body-size evolution and the latitudinal biodiversity gradient. In addition, there are contributions outlining the approaches and techniques used in modern palaeontology, such as the principle of parsimony, methods for collecting and analysing shape data, palaeoecology and the live–dead agreement, naming fossils and ancient DNA. Finally, there are articles about varied aspects of life as a palaeontologist, including career paths, the influence of the internet and open access, what is (and what isn’t) palaeontology and the history of the field.

Phew, I think that’s everything. As you can see, we’ve covered an awful lot in the past three years. Nevertheless, this is only the tip of the palaeontological iceberg, and with so many promising young researchers around, there will no doubt be plenty to write about for years to come. All that remains is to thank everyone who has contributed articles to the site and, in fact, everyone who has read the articles — thanks for your continued support, and here’s to many more years!