A new species of a giant tortoise from Sandelzhausen (MN5, Burdigalian/Langhian boundary, Early/Middle Miocene, South Germany).

We describe a new species of a giant tortoise of the genus Titanochelon from the locality of Sandelzhausen in south Germany (MN5, Burdigalian/Langhian boundary, Early/Middle Miocene). The material comprises at least two different individuals, one of which is a male individual preserving large parts of the carapace and plastron and several appendicular elements. The second individual is quite fragmented, preserving parts of the bridge and the posterior rim of the carapace. The new species, Titanochelon schleichi sp. nov., is the first species of a giant tortoise named from Germany and allows reconstructing an important diversity and expansion of titanochelones in the Western Palaearctic during the earlier parts of the Neogene.

New findings on the evolution of turtles: A symposium in honor of Marcelo S. de la Fuente.

With their particular body plan within amniotes and their amazing fossil record, turtles represent a great interest for both neontologists and paleontologists with a strong anatomical background. The Turtle Evolution Symposia are regular international meetings that gather scientists working with different aspects related to the evolutionary history of turtles, from their origin and early evolution until recent times. The latest edition of the Turtle Evolution Symposium was organized in 2021 amidst the COVID-19 outbreak and held virtually from the facilities of the Museo Paleontológico Egidio Feruglio in Trelew (Patagonia, Chubut, Argentina). More than 75 scientists from 25 countries presented their latest advances on topics related to turtle evolution, some of which are published in this Special Volume of The Anatomical Record. Both the Turtle Evolution Symposium 2021 and this Special Volume are dedicated to Marcelo S. de la Fuente who was the first researcher who specialized in the study of extinct turtles in South America, and his studies have an important regional and international impact.

The latest Miocene to Pliocene small-sized fossil turtles, Testudinidae and Geoemydidae, from Makrygialos, Thermaikos Gulf, Northern Greece.

Although the existence of fossil turtles in Makrygialos and the surrounding areas, on the west side of the Thermaikos Gulf in Northern Greece, has been known for almost two decades, it has not been studied so far in detail. Over the last 5 years, we have conducted systematic fieldwork and exploration, focusing on the discovery and recovery of fossils in the coastal broader area of Makrygialos. Eleven turtle specimens have been found so far, eight of which have been collected, on the outcrops of the beaches of Makrygialos and Agiannis. Their age is estimated between the latest Miocene and Pliocene. Four of these specimens are terrestrial giant tortoises, two of them are small tortoises which we identify as the species Chersine hermanni sensu lato, and the last two are freshwater turtles which we attribute to the genus Mauremys sensu lato. Here, we present our studies on the small-sized turtle species from this region. The fossil turtle fauna of Makrygialos is among the richest and most diverse in Greece, as in only a few other Greek localities at least three species of fossil turtles have been found. The C. hermanni specimens found in Makrygialos are probably the oldest fossils of the species found in Europe. The small-turtle faunal composition during the Neogene along the east and west sides of the Thermaikos Gulf was completely different.

A new large-sized species of Chelonoidis (Testudinidae) without gibbosities from the middle Miocene of Aguada Escondida (NW Chubut, Patagonia, Argentina).

Previously, only one small-sized species of Testudinidae (Chelonoidis gringorum) was named from Lower-Middle Miocene of Central Patagonia. In this short article, we describe a new large Testudinidae, here named Chelonoidis meridiana sp. nov. This large new species (carapace up to 80 cm) differs from other large species of the southern South American fossil record by the absence of gibbosities on neural and costal carapacial bones, a short and wide entoplastron with a humeropectoral sulcus that was placed well behind its posterior end, and pectoral scutes that are medially longer, approximately one-third of the medial length of the hyoplastra. The discovery of this new species in the Middle Miocene increases the raw turtle diversity in Chubut, permitting to reduce the gap between the raw and phylogenetic diversity previously proposed. This occurrence is also important from both a temporal and geographic point of view. It permits fixing the presence of large-sized tortoises in the continent since ~15 million years ago (Middle Miocene). This occurrence predates all other large and giant tortoises from Argentina and South America and it is older than the giant Chelonoidis from the Langhian-Tortonian of Colombia. This is also the southernmost occurrence of a large-sized testudinid in the world.

Breaking the mold: telescoping drives the evolution of more integrated and heterogeneous skulls in cetaceans.

Background: Along with the transition to the aquatic environment, cetaceans experienced profound changes in their skeletal anatomy, especially in the skull, including the posterodorsal migration of the external bony nares, the reorganization of skull bones (= telescoping) and the development of an extreme cranial asymmetry (in odontocetes). Telescoping represents an important anatomical shift in the topological organization of cranial bones and their sutural contacts; however, the impact of these changes in the connectivity pattern and integration of the skull has never been addressed. Methods: Here, we apply the novel framework provided by the Anatomical Network Analysis to quantify the organization and integration of cetacean skulls, and the impact of the telescoping process in the connectivity pattern of the skull. We built anatomical networks for 21 cetacean skulls (three stem cetaceans, three extinct and 10 extant mysticetes, and three extinct and two extant odontocetes) and estimated network parameters related to their anatomical integration, complexity, heterogeneity, and modularity. This dataset was analyzed in the context of a broader tetrapod skull sample as well (43 species of 13 taxonomic groups). Results: The skulls of crown cetaceans (Neoceti) occupy a new tetrapod skull morphospace, with better integrated, more heterogeneous and simpler skulls in comparison to other tetrapods. Telescoping adds connections and improves the integration of those bones involved in the telescoping process (e.g., maxilla, supraoccipital) as well as other ones (e.g., vomer) not directly affected by telescoping. Other underlying evolutionary processes (such as basicranial specializations linked with hearing/breathing adaptations) could also be responsible for the changes in the connectivity and integration of palatal bones. We also find prograde telescoped skulls of mysticetes distinct from odontocetes by an increased heterogeneity and modularity, whereas retrograde telescoped skulls of odontocetes are characterized by higher complexity. In mysticetes, as expected, the supraoccipital gains importance and centrality in comparison to odontocetes, increasing the heterogeneity of the skull network. In odontocetes, an increase in the number of connections and complexity is probably linked with the dominant movement of paired bones, such as the maxilla, in retrograde telescoping. Crown mysticetes (Eubalaena, Caperea, Piscobalaena, and Balaenoptera)are distinguished by having more integrated skulls in comparison to stem mysticetes (Aetiocetus and Yamatocetus), whereas crown odontocetes (Waipatia, Notocetus, Physeter, and Tursiops) have more complex skulls than stem forms (Albertocetus). Telescoping along with feeding, hearing and echolocation specializations could have driven the evolution of the different connectivity patterns of living lineages.

A modularity analysis helps improving the structure of the International Code of Zoological Nomenclature.

BACKGROUND: In a recent work I transformed a complex and integrated text like the International Code of Zoological Nomenclature into a network of interconnected parts of text. This new approach allowed understanding that a continuous body of text cannot accurately reflect the true structure of the Code, and provided a scientific methodology to identify a priori parts that could be affected by future revisions. In this next step, I investigate further the structure of the Code, seeking to use the network in order to identify the various conceptual communities grouping the various articles and other text items of the Code. METHODS: Using the first version of the network of the Code, I perform a comprehensive modularity analysis in two rounds: the first round aims to identify the fewest and largest communities or modules for the entire network, whereas the second round identifies the sub-modules within each larger module. The potential conflicts between the current structure of the Code and the module composition are evaluated with a parcellation analysis. RESULTS: The optimal modularity search identified 10 different modules in the entire network of varying size (ranging from 75 to 200 nodes). Each module can be further divided into smaller modules, that all-together allow describing the 65 conceptual groups of text items in the Code. Parcellation analysis revealed that two-thirds of the current chapters of the Code are in excellent or good accordance with the recovered conceptual modules, whereas the current composition of six chapters is in serious conflict with the conceptual structure of the Code. DISCUSSION: Judging only the composition and not the order of appearance of the Articles in the Chapters of the Code, I show that in many cases the current structure of the Code is found to correspond quite well to the concepts presented therein. The most important conflict is found on the provisions related to the various groups of names governed by the Code: family-, genus-, and species-group names. Currently, these provisions are spread out in different Articles in different Chapters, along the entire length of the Code. The modularity analysis suggests that re-organizing the Code in chapters that will deal with all aspects related to a given group (e.g., chapters including information on name formation, availability, typification, and validity for a given group), could potentially improve reader experience and, consequently, the applicability of the Code.

Fingers zipped up or baby mittens? Two main tetrapod strategies to return to the sea.

The application of network methodology in anatomical structures offers new insights on the connectivity pattern of skull bones, skeletal elements and their muscles. Anatomical networks helped to improve our understanding of the water-to-land transition and how the pectoral fins were transformed into limbs via their modular disintegration. Here, we apply the same methodology to tetrapods secondarily adapted to the marine environment. We find that these animals achieved their return to the sea with four types of morphological changes, which can be grouped into two different main strategies. In all marine mammals and the majority of the reptiles, the fin is formed by the persistence of superficial and interdigital connective tissues, like a 'baby mitten', whereas the underlying connectivity pattern of the bones does not influence the formation of the forefin. On the contrary, ichthyosaurs 'zipped up' their fingers and transformed their digits into carpal-like elements, forming a homogeneous and better-integrated forefin. These strategies led these vertebrates into three different macroevolutionary paths exploring the possible spectrum of morphological adaptations.

Radio-frequency chain selection for energy and spectral efficiency maximization in hybrid beamforming under hardware imperfections.

The next-generation wireless communications require reduced energy consumption, increased data rates and better signal coverage. The millimetre-wave frequency spectrum above 30 GHz can help fulfil the performance requirements of the next-generation mobile broadband systems. Multiple-input multiple-output technology can provide performance gains to help mitigate the increased path loss experienced at millimetre-wave frequencies compared with microwave bands. Emerging hybrid beamforming architectures can reduce the energy consumption and hardware complexity with the use of fewer radio-frequency (RF) chains. Energy efficiency is identified as a key fifth-generation metric and will have a major impact on the hybrid beamforming system design. In terms of transceiver power consumption, deactivating parts of the beamformer structure to reduce power typically leads to significant loss of spectral efficiency. Our aim is to achieve the highest energy efficiency for the millimetre-wave communications system while mitigating the resulting loss in spectral efficiency. To achieve this, we propose an optimal selection framework which activates specific RF chains that amplify the digitally beamformed signals with the analogue beamforming network. Practical precoding is considered by including the effects of user interference, noise and hardware impairments in the system modelling.

Introducing a new tool to navigate, understand and use International Codes of Nomenclature.

BACKGROUND: In order to designate the various concepts of taxa in biology, evolution and paleontology, scientists have developed various rules on how to create unique names for taxa. Different Codes of Nomenclature have been developed for animals, plants, fungi, bacteria etc., with standard sets of Rules that govern the formation, publication and application of the nomina of extant and extinct species. These Codes are the result of decades of discussions, workshops, publications and revisions. The structure and complexity of these Codes have been criticized many times by zoologists. This project aims, using the International Code of Zoological Nomenclature as a case study, to show that the structure of these Codes is better reflected and understood as networks. METHODS: The majority of the text of the Code has been divided into hundreds of Nodes of different types, connected to each other with different types of Edges to form a network. The various mathematical descriptors of the entire system, as well as for the elements of the network, have been conceptually framed to help describing and understanding the Code as a network. RESULTS: The network of the Code comprises 1,379 Nodes, which are connected with 11,276 Edges. The structure of the Code can be accurately described as a network, a mathematical structure that is better suited than any kind of linear text publication to reflect its structure. DISCUSSION: Thinking of the Code as a network allows a better, in-depth understanding of the Code itself, as the user can navigate in a more efficient way, as well as to depict and analyze all the implied connections between the various parts of the Code that are not visible immediately. The network of the Code is an open access tool that could also help teaching, using and disseminating the Code. More importantly, this network is a powerful tool that allows identifying a priori the parts of the Code that could be potentially affected by upcoming amendment and revisions. This kind of analysis is not limited to nomenclature, as it could be applied to other fields that use complex textbooks with long editing history, such as Law, Medicine and Linguistics.

Total evidence analysis and body size evolution of extant and extinct tortoises (Testudines: Cryptodira: Pan-Testudinidae).

Testudinidae (tortoises) is an extant clade of terrestrial turtles of worldwide distribution and with a rich fossil record that provides an exceptional context for studying their evolutionary history. Because of the lack of global phylogenetic analyses integrating extinct taxa, our current knowledge of the relationships of the total clade of Testudinidae is rather poor. To resolve this issue, we performed the first total evidence analysis of Pan-Testudinidae. The total evidence trees are congruent with the molecular topology and agree on the dichotomy of derived Testudinidae (=Testudininae; Converted Clade Name) into two previously recognized major clades, Testudona and Geochelona (New Clade Name). The integration of extinct taxa into the analysis allowed the stratigraphic fit of the total evidence trees, indicating that crown Testudininae, Testudona and Geochelona all originated by the Late Eocene, in agreement with recent molecular estimates. Ghost lineage analysis indicates high diversification in the Late Eocene and in the Miocene. The age of crown Testudo is Late Miocene, again in accordance with some molecular dates. Phylogenetic placement of fossils demonstrates that giant body size independently evolved in multiple continental mainland taxa and confirms recent results deduced from living taxa-giantism in Testudinidae is not linked to the insular effect. An unexpected outcome is the recovery of miniaturization in Testudona (<30 cm carapace length) that emerged sometime between the Oligocene and Early Miocene. No clear correlation between body size evolution and climate is apparent, but increased taxon sampling may nevertheless demonstrate the role of cooling and warming as one of many influential variables.

The first record of a soft-shelled turtle (Testudines: Pan-Trionychidae) from southern Balkans (Pliocene, Gefira, N. Greece) and new information from bone histology.

Soft-shelled turtles (Pan-Trionychidae) are not included in the present-day chelonian fauna of Greece and have been unknown in the Greek fossil record up to now. Here, we report the first fossil occurrence of a soft-shelled turtle from Greece, originating from the Pliocene Gefira Member (Angelochori Formation), in the lower Axios valley. The corresponding specimens were discovered with several mammalian remains, most of them attributable to the mastodon of Auvergne, Anancus arvernensis. The chelonian material includes five carapacial fragments that belong to the same individual and can be attributed to Pan-Trionychidae based on the typical sculpturing on the dorsal side of the carapace. Most of these bony plates were histologically sampled and thereby provide evidence for the "plywood" structure, another characteristic of pan-trionychids. They represent the first extended sampling of trionychid plates that belong to the same individual, allowing the documentation of the variation of the relevant trionychid morphologies in the carapace. These findings expand the paleobiogeographic range of this taxon to the southern Balkans and Greece and allow a better estimation of the chelonian paleo-fauna of the area. They are also important for the temporal distribution of this clade in the Paleoarctic, as they join specimens from Italy as being the last trionychids in Europe.