Sven Sachs

Revised Vertebral Count in the “Longest-Necked Vertebrate” Elasmosaurus platyurus Cope 1868, and Clarification of the Cervical-Dorsal Transition in Plesiosauria

Elasmosaurid plesiosaurians are renowned for their immensely long necks, and indeed, possessed the highest number of cervical vertebrae for any known vertebrate. Historically, the largest count has been attributed to the iconic Elasmosaurus platyurus from the Late Cretaceous of Kansas, but estimates for the total neck series in this taxon have varied between published reports. Accurately determining the number of vertebral centra vis-à-vis the maximum length of the neck in plesiosaurians has significant implications for phylogenetic character designations, as well as the inconsistent terminology applied to some osteological structures. With these issues in mind, we reassessed the holotype of E. platyurus as a model for standardizing the debated cervical-dorsal transition in plesiosaurians, and during this procedure, documented a “lost” cervical centrum. Our revision also advocates retention of the term “pectorals” to describe the usually three or more distinctive vertebrae close to the cranial margin of the forelimb girdle that bear a functional rib facet transected by the neurocentral suture, and thus conjointly formed by both the parapophysis on the centrum body and diapophysis from the neural arch (irrespective of rib length). This morphology is unambiguously distinguishable from standard cervicals, in which the functional rib facet is borne exclusively on the centrum, and dorsals in which the rib articulation is situated above the neurocentral suture and functionally borne only by the transverse process of the neural arch. Given these easily distinguishable definitions, the maximum number of neck vertebrae preserved in E. platyurus is 72; this is only three vertebrae shorter than the recently described Albertonectes, which together with E. platyurus constitute the “longest necked” animals ever to have lived.

Postcranium of the paradigm elasmosaurid plesiosaurian Libonectes morgani (Welles, 1949)

Elasmosauridae constitutes one of the most immediately recognizable plesiosaurian radiations. Their distinctive body plan represents the popular model for Plesiosauria, and is typified by an osteological morphology especially adapted for hyper-elongation of the neck. Nevertheless, many archetypal elasmosaurids are known only from incomplete and/or inadequately documented material, a problem that has contributed to their uncertain intra-clade relationships. A prime example of this is Libonectes morgani from the Upper Cretaceous of Texas, USA, which is frequently presented as an elasmosaurid structural proxy because of its three-dimensionally preserved holotype skull. Perplexingly though, both the taxonomic diagnosis and phylogenetic placement of L. morgani rely primarily upon the cervical vertebrae, together with the pectoral girdle and forelimb, yet most of these elements are now lost and figured only as line drawings. We therefore reviewed the remnant postcranial skeleton of L. morgani first-hand with the objective of clarifying its defining character states. Our observations showed that the existing diagnosis of L. morgani is indeed inadequate. Moreover, the only identifiable autapomorphies occurred within the axial skeleton. This concurred with an examination of character scores used in published plesiosaurian phylogenies, and highlights the persistent significance of postcranial elements for discriminating elasmosaurid taxa.

Reappraisal of Europe’s most complete Early Cretaceous plesiosaurian: Brancasaurus brancai Wegner, 1914 from the “Wealden facies” of Germany

The holotype of Brancasaurus brancai is one of the most historically famous and anatomically complete Early Cretaceous plesiosaurian fossils. It derived from the Gerdemann & Co. brickworks clay pit near Gronau (Westfalen) in North Rhine-Westphalia, northwestern Germany. Stratigraphically this locality formed part of the classic European “Wealden facies,” but is now more formally attributed to the upper-most strata of the Bückeberg Group (upper Berriasian). Since its initial description in 1914, the type skeleton of B. brancai has suffered damage both during, and after WWII. Sadly, these mishaps have resulted in the loss of substantial information, in particular many structures of the cranium and limb girdles, which are today only evidenced from published text and/or illustrations. This non-confirmable data has, however, proven crucial for determining the relationships of B. brancai within Plesiosauria: either as an early long-necked elasmosaurid, or a member of the controversial Early Cretaceous leptocleidid radiation. To evaluate these competing hypotheses and compile an updated osteological compendium, we undertook a comprehensive examination of the holotype as it is now preserved, and also assessed other Bückeberg Group plesiosaurian fossils to establish a morphological hypodigm. Phylogenetic simulations using the most species-rich datasets of Early Cretaceous plesiosaurians incorporating revised scores for B. brancai, together with a second recently named Bückeberg Group plesiosaurian Gronausaurus wegneri (Hampe, 2013), demonstrated that referral of these taxa to Leptocleididae was not unanimous, and that the topological stability of this clade is tenuous. In addition, the trait combinations manifested by B. brancai and G. wegneri were virtually identical. We therefore conclude that these monotypic individuals are ontogenetic morphs and G. wegneri is a junior synonym of B. brancai. Finally, anomalies detected in the diagnostic features for other “Wealden” plesiosaurians have prompted reconsiderations of interspecies homology versus intraspecific variability. We therefore propose that the still unresolved taxonomy of B. brancai should emphasize only those character states evident in the examinable fossil material, and specifically accommodate for growth-related modifications delimited via osteologically mature referred specimens.

A partial plesiosaurian braincase from the Upper Cretaceous of Sweden

Abstract A partial exoccipital–opisthotic from the uppermost lower Campanian (Upper Cretaceous) of the Åsen locality, Kristianstad Basin, southernmost Sweden, is described and illustrated. The fossil represents the first braincase element of a plesiosaur found in Sweden. It includes the chamber for the ampulla and utriculus, openings for the caudal vertical and horizontal semicircular canals, and four foramina for cranial nerves. The incomplete braincase can be referred to an elasmosaurid plesiosaur, and closely resembles the exoccipital–opisthotic of Libonectes morgani and a referred specimen of Aristonectes parvidens. Although we discuss putative postcranial material of the elasmosaurid subfamily Aristonectinae in the uppermost lower Campanian of southernmost Sweden, the exoccipital–opisthotic from Åsen most likely belongs to a juvenile individual of a non-aristonectine elasmosaur.

Cenomanian–Turonian marine amniote remains from the Saxonian Cretaceous Basin of Germany

The Saxonian Cretaceous Basin constitutes an important source of rare Late Cretaceous marine amniote fossils from Germany. It is also historically famous, having been documented in a series of monographic works published by the distinguished German palaeontologist Hanns Bruno Geinitz in the nineteenth century. The most productive rock units include the upper Cenomanian Dolzschen Formation and upper Turonian Strehlen and Weinbohla limestones (lower Strehlen Formation). A survey of curated specimens recovered from these deposits has now identified isolated teeth of probable polycotylid and elasmosaurid plesiosaurians, as well as several humeri that are referred to protostegid marine turtles. The Saxonian Cretaceous Basin formed a continuous epeiric seaway with the Bohemian Cretaceous Basin during late Cenomanian – Turonian time. A western connection to the North Sea Basin also existed via the North German and Munsterland Cretaceous basins. The Mesozoic marine amniote remains from these regions therefore record a coeval northern European fauna that was probably homogeneous across the northern peri-Tethyan margin during Late Cretaceous time.

An ichthyosaur fragment from the Cretaceous of Northland, New Zealand

Abstract The rostrum fragment of an ichthyosaur from Northland in New Zealand is described. The specimen appears to belong to the genus Platypterygius. It is the first Mesozoic marine vertebrate fossil reported from Northland and the first cranial element of a Cretaceous ichthyosaur from New Zealand. The fragment is, unfortunately, of uncertain provenance and could have come from the widespread Northland Allochthon (Early Cretaceous‐Oligocene) or the allochthonous Houhora Complex (late Early Cretaceous) or Whatuwhiwhi Formation (latest Early to Late Cretaceous), perhaps by way of initial derivation from the Early Miocene Omapere Conglomerate.

Plesiosaurian fossils from Baltic glacial erratics: evidence of Early Jurassic marine amniotes from the southwestern margin of Fennoscandia

Abstract Early Jurassic plesiosaurian fossils are rare in the Scandinavian region, with a few isolated bones and teeth known from Bornholm, and anecdotal finds from East Greenland. The only other identifiable specimens derive from Toarcian-aged (based on ammonites) erratics deposited during Late Pleistocene glacial advances near the town of Ahrensburg, NE of Hamburg in northern Germany. The geographical source of these transported clasts is debated, but reconstructed ice-flow directions and lithofacies comparisons implicate either the offshore Baltic Sea between the Island of Bornholm and Mecklenburg–Vorpommern (Germany) or, less probably, south of the Danish Archipelago (Mecklenburg Bay). These regions collectively bordered the Fennoscandian landmass and adjacent Ringkøbing-Fyn Island in the late Early Jurassic, and were dominated by near-shore marine deltaic to basinal settings. The Ahrensburg plesiosaurian remains include postcranial elements reminiscent of both the microcleidid Seeleyosaurus and the rhomaelosaurid Meyerasaurus. These occur alongside other classic ‘Germanic province’ marine amniotes, such as the teleosaurid crocodyliform Steneosaurus and ichthyosaurian Stenopterygius cf. quadriscissus: thus, advocating faunal continuity between Scandinavia and southern Germany during the Toarcian, and a less pronounced marine reptile faunal provinciality than previously assumed.

Ankylosaur Remains from the Early Cretaceous (Valanginian) of Northwestern Germany

A fragmentary cervico-pectoral lateral spine and partial humerus of an ankylosaur from the Early Cretaceous (early Valanginian) of Gronau in Westfalen, northwestern Germany, are described. The spine shows closest morphological similarities to the characteristic cervical and pectoral spines of Hylaeosaurus armatus from the late Valanginian of England. An extensive comparison of distal humeri among thyreophoran dinosaurs supports systematic differences in the morphology of the distal condyli between Ankylosauria and Stegosauria and a referral of the Gronau specimen to the former. The humerus fragment indicates a rather small individual, probably in the size range of H. armatus, and both specimens are determined herein as ?Hylaeosaurus sp.. A short overview of other purported ankylosaur material from the Berriasian-Valanginian of northwest Germany shows that, aside from the material described herein, only tracks can be attributed to this clade with confidence at present.

A new basal elasmosaurid (Sauropterygia: Plesiosauria) from the Lower Cretaceous of Germany

ABSTRACT Here we report on a new basal elasmosaurid plesiosaurian, Lagenanectes richterae, gen. et sp. nov., from the Lower Cretaceous (probably Upper Hauterivian) of Germany. The material includes a partial skull (cranium and mandible), the atlas-axis complex, additional cervical vertebrae, caudal vertebrae, an ilium, and limb elements. The basioccipital and atlas intercentrum are pathologically deformed, probably due to an osteomyelitic infection. Two potential autapomorphies were found in the mandible: (1) the alveolar margin at the symphysis is laterally expanded with the rostral-most alveoli being markedly procumbent and situated along the lateral margins of the dentaries; and (2) the ventral midline at the symphysis is produced into a prominent wedge-shaped platform indented by numerous irregular pits. Lagenanectes richterae, gen. et sp. nov., also shows a number of typical elasmosaurid traits, including a longitudinal lateral ridge on the cervical vertebral centra (although a ventral notch is absent) and teeth with oval cross-sections. Lagenanectes richterae, gen. et sp. nov., is one of the best-preserved plesiosaurians from the Lower Cretaceous of Europe.

First evidence of a large predatory plesiosaurian from the Lower Cretaceous non-marine ‘Wealden facies’ deposits of northwestern Germany

Sachs, S., Hornung, J.J., Lallensack, J.N. & Kear, B.P., November.2017. First evidence of a large predatory plesiosaurian from the Lower Cretaceous non-marine ‘Wealden facies’ deposits of northwestern Germany. Alcheringa 42, 501-508. ISSN 0311-5518. Here, we describe the incomplete mandible of a large-skulled ‘pliosauromorph’ plesiosaurian from the Lower Cretaceous (Berriasian) of northwestern Germany. The fossil derives from limnic–brackish ‘Wealden facies’ deposits of the Deister Formation (Bückeberg Group), and is preserved as a natural mould in fine-grained sandstone. Examination of the original remains, in conjunction with a three-dimensional photogrammetrically digitized ‘cast’, revealed a conspicuous rosette of symphyseal alveoli, which would otherwise typically characterize Early–Middle Jurassic macrophagous plesiosaurians including rhomaleosaurids and the pliosaurid Simolestes. The Deister Formation ‘pliosauromorph’ represents the first record of a large-bodied plesiosaurian macrocarnivore from the ‘Wealden-facies’ strata of Europe, and thus adds a previously unrecognized trophic level of aquatic apex predators to the Early Cretaceous non-marine ecosystems of Europe. Sven Sachs* [sachs.pal@gmail.com] Naturkunde-Museum Bielefeld, Abteilung Geowissenschaften, Adenauerplatz 2, 33602 Bielefeld, Germany and Im Hof 9, 51766 Engelskirchen, Germany; Jahn J. Hornung [jahn.hornung@yahoo.de] Niedersächsisches Landesmuseum Hannover, Willy-Brandt-Allee 5, 30169 Hannover, Germany; Jens N. Lallensack [jens.lallensack@uni-bonn.de] Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Nussallee 8, 53115 Bonn, Germany; Benjamin P. Kear [benjamin.kear@em.uu.se] Museum of Evolution, Uppsala University, Norbyvägen 16, Uppsala SE-752 36, Sweden.