Anthony Romilio

Reevaluation of the Lark Quarry dinosaur Tracksite (late Albian–Cenomanian Winton Formation, central-western Queensland, Australia): no longer a stampede?

ABSTRACT The Lark Quarry dinosaur tracksite has previously been recognized as recording the stampede of a mixed herd of dozens of small theropod and ornithopod dinosaurs. A reexamination of trackway material reveals that the small theropodtype tracks, previously assigned to the ichnotaxon Skartopus, can co-occur within individual trackways of the ornithopod-type tracks assigned to Wintonopus. Moreover, in singular deep tracks where the overall surface outline resembles Skartopus, the base of the track can also resemble Wintonopus. Whereas the Wintonopus holotype may reflect the pedal anatomy of a shorttoed or subunguligrade ornithopod trackmaker, the elongate ‘oe’ impressions typically associated with Skartopus (including the holotype) primarily provide information on digit movement through the sediment and, in many instances, may represent swim traces. The morphological differences between the two ichnotaxa are therefore not taxonomically significant and we formally propose that Skartopus australis should be considered a junior synonym of Wintonopus latomorum. Longitudinal depth profiles through tracks indicate that many are swim traces. The sedimentology and lithology of Lark Quarry further indicates the site represents a time-averaged assemblage formed in a fluvial-dominated floodplain under variable subaqueous conditions, with the parallel orientation of the numerous trackways formed by trackmakers under the influence of downstream current flow. This indicates that the fluvial environment may have been a preferred route for hydrophilic bipedal dinosaurs. We thus do not consider the Lark Quarry dinosaur tracksite to represent a ‘stampede.’ Instead, the tracksite may represent part of a riverine setting, where the water was shallow, in which small dinosaurs swam and/or waded.

The Dinosaurian Ichnofauna of the Lower Cretaceous (Valanginian–Barremian) Broome Sandstone of the Walmadany Area (James Price Point), Dampier Peninsula, Western Australia

ABSTRACT Extensive and well-preserved tracksites in the coastally exposed Lower Cretaceous (Valanginian-Barremian) Broome Sandstone of the Dampier Peninsula provide almost the entire fossil record of dinosaurs from the western half of the Australian continent. Tracks near the town of Broome were described in the late 1960s as Megalosauropus broomensis and attributed to a medium-sized theropod trackmaker. Brief reports in the early 1990s suggested the occurrence of at least another nine types of tracks, referable to theropod, sauropod, ornithopod, and thyreophoran trackmakers, at scattered tracksites spread over more than 80 km of coastline north of Broome, potentially representing one of the worlds most diverse dinosaurian ichnofaunas. More recently, it has been proposed that this number could be as high as 16 and that the sites are spread over more than 200 km. However, the only substantial research that has been published on these more recent discoveries is a preliminary study of the sauropod tracks and an account of the ways in which the heavy passage of sauropod trackmakers may have shaped the Dampier Peninsulas Early Cretaceous landscape. With the other types of dinosaurian tracks in the Broome Sandstone remaining undescribed, and the full extent and nature of the Dampier Peninsulas dinosaurian tracksites yet to be adequately addressed, the overall scientific significance of the ichnofauna has remained enigmatic. At the request of the areas Goolarabooloo Traditional Custodians, 400C hours of ichnological survey work was undertaken from 2011 to 2016 on the 25 km stretch of coastline in the Yanijarri-Lurujarri section of the Dampier Peninsula, inclusive of the coastline at Walmadany (James Price Point). Forty-eight discrete dinosaurian tracksites were identified in this area, and thousands of tracks were examined and measured in situ and using three-dimensional photogrammetry. Tracksites were concentrated in three main areas along the coast: Yanijarri in the north, Walmadany in the middle, and Kardilakan-Jajal Buru in the south. Lithofacies analysis revealed 16 repeated facies types that occurred in three distinctive lithofacies associations, indicative of an environmental transgression between the distal fluvial to deltaic portions of a large braid plain, with migrating sand bodies and periodic sheet floods. The main dinosaurian track-bearing horizons seem to have been generated between periodic sheet floods that blanketed the preexisting sand bodies within the braid plain portion of a tidally influenced delta, with much of the original, gently undulating topography now preserved over large expanses of the present day intertidal reef system. Of the tracks examined, 150 could be identified and are assignable to a least eleven and possibly as many as 21 different track types: five different types of theropod tracks, at least six types of sauropod tracks, four types of ornithopod tracks, and six types of thyreophoran tracks. Eleven of these track types can formally be assigned or compared to existing or new ichnotaxa, whereas the remaining ten represent morphotypes that, although distinct, are currently too poorly represented to confidently assign to existing or new ichnotaxa. Among the ichnotaxa that we have recognized, only two (Megalosauropus broomensis and Wintonopus latomorum) belong to existing ichnotaxa, and two compare to existing ichnotaxa but display a suite of morphological features suggesting that they may be distinct in their own right and are therefore placed in open nomenclature. Six of the ichnotaxa that we have identified are new: one theropod ichnotaxon, Yangtzepus clarkei, ichnosp. nov.; one sauropod ichnotaxon, Oobardjidama foulkesi, ichnogen. et ichnosp. nov.; two ornithopod ichnotaxa, Wintonopus middletonae, ichnosp. nov., and Walmadanyichnus hunteri, ichnogen. et ichnosp. nov.; and two thyreophoran ichnotaxa, Garbina roeorum, ichnogen. et ichnosp. nov., and Luluichnus mueckei, ichnogen. et ichnosp. nov. The level of diversity of the main track types is comparable across areas where tracksites are concentrated: Kardilakan-Jajal Buru (12), Walmadany (11), and Yanijarri (10). The overall diversity of the dinosaurian ichnofauna of the Broome Sandstone in the Yanijarri-Lurujarri section of the Dampier Peninsula is unparalleled in Australia, and even globally. In addition to being the primary record of non-avian dinosaurs in the western half of Australia, this ichnofauna provides our only detailed glimpse of Australias dinosaurian fauna during the first half of the Early Cretaceous. It indicates that the general composition of Australias mid-Cretaceous dinosaurian fauna was already in place by the Valanginian-Barremian. Both sauropods and ornithopods were diverse and abundant, and thyreophorans were the only type of quadrupedal ornithischians. Important aspects of the fauna that are not seen in the Australian mid-Cretaceous body fossil record are the presence of stegosaurians, an overall higher diversity of thyreophorans and theropods, and the presence of large-bodied hadrosauroid-like ornithopods and very large-bodied sauropods. In many respects, these differences suggest a holdover from the Late Jurassic, when the majority of dinosaurian clades had a more cosmopolitan distribution prior to the fragmentation of Pangea. Although the record for the Lower Cretaceous of Gondwana is sparse, a similar mix of taxa occurs in the Barremian-lower Aptian La Amarga Formation of Argentina and the Berriasian-Hauterivian Kirkwood Formation of South Africa. The persistence of this fauna across the Jurassic-Cretaceous boundary in South America, Africa, and Australia might be characteristic of Gondwanan dinosaurian faunas more broadly. It suggests that the extinction event that affected Laurasian dinosaurian faunas across the Jurassic-Cretaceous boundary may not have been as extreme in Gondwana, and this difference may have foreshadowed the onset of Laurasian-Eurogondwanan provincialism. The disappearance of stegosaurians and the apparent drop in diversity of theropods by the mid-Cretaceous suggests that, similar to South America, Australia passed through a period of faunal turnover between the Valanginian and Aptian.

The Evolution of Fangs, Venom, and Mimicry Systems in Blenny Fishes

Venom systems have evolved on multiple occasions across the animal kingdom, and they can act as key adaptations to protect animals from predators [1]. Consequently, venomous animals serve as models for a rich source of mimicry types, as non-venomous species benefit from reductions in predation risk by mimicking the coloration, body shape, and/or movement of toxic counterparts [2-5]. The frequent evolution of such deceitful imitations provides notable examples of phenotypic convergence and are often invoked as classic exemplars of evolution by natural selection. Here, we investigate the evolution of fangs, venom, and mimetic relationships in reef fishes from the tribe Nemophini (fangblennies). Comparative morphological analyses reveal that enlarged canine teeth (fangs) originated at the base of the Nemophini radiation and have enabled a micropredatory feeding strategy in non-venomous Plagiotremus spp. Subsequently, the evolution of deep anterior grooves and their coupling to venom secretory tissue provide Meiacanthus spp. with toxic venom that they effectively employ for defense. We find that fangblenny venom contains a number of toxic components that have been independently recruited into other animal venoms, some of which cause toxicity via interactions with opioid receptors, and result in a multifunctional biochemical phenotype that exerts potent hypotensive effects. The evolution of fangblenny venom has seemingly led to phenotypic convergence via the formation of a diverse array of mimetic relationships that provide protective (Batesian mimicry) and predatory (aggressive mimicry) benefits to other fishes [2, 6]. Our results further our understanding of how novel morphological and biochemical adaptations stimulate ecological interactions in the natural world.

Coagulating Colubrids: Evolutionary, Pathophysiological and Biodiscovery Implications of Venom Variations between Boomslang (Dispholidus typus) and Twig Snake (Thelotornis mossambicanus)

Venoms can deleteriously affect any physiological system reachable by the bloodstream, including directly interfering with the coagulation cascade. Such coagulopathic toxins may be anticoagulants or procoagulants. Snake venoms are unique in their use of procoagulant toxins for predatory purposes. The boomslang (Dispholidus typus) and the twig snakes (Thelotornis species) are iconic African snakes belonging to the family Colubridae. Both species produce strikingly similar lethal procoagulant pathologies. Despite these similarities, antivenom is only produced for treating bites by D. typus, and the mechanisms of action of both venoms have been understudied. In this study, we investigated the venom of D. typus and T. mossambicanus utilising a range of proteomic and bioactivity approaches, including determining the procoagulant properties of both venoms in relation to the human coagulation pathways. In doing so, we developed a novel procoagulant assay, utilising a Stago STA-R Max analyser, to accurately detect real time clotting in plasma at varying concentrations of venom. This approach was used to assess the clotting capabilities of the two venoms both with and without calcium and phospholipid co-factors. We found that T. mossambicanus produced a significantly stronger coagulation response compared to D. typus. Functional enzyme assays showed that T. mossambicanus also exhibited a higher metalloprotease and phospholipase activity but had a much lower serine protease activity relative to D. typus venom. The neutralising capability of the available boomslang antivenom was also investigated on both species, with it being 11.3 times more effective upon D. typus venom than T. mossambicanus. In addition to being a faster clotting venom, T. mossambicanus was revealed to be a much more complex venom composition than D. typus. This is consistent with patterns seen for other snakes with venom complexity linked to dietary complexity. Consistent with the external morphological differences in head shape between the two species, CT and MRI analyses revealed significant internal structural differences in skull architecture and venom gland anatomy. This study increases our understanding of not only the biodiscovery potential of these medically important species but also increases our knowledge of the pathological relationship between venom and the human coagulation cascade.

A multidisciplinary approach to digital mapping of dinosaurian tracksites in the Lower Cretaceous (Valanginian–Barremian) Broome Sandstone of the Dampier Peninsula, Western Australia

The abundant dinosaurian tracksites of the Lower Cretaceous (Valanginian–Barremian) Broome Sandstone of the Dampier Peninsula, Western Australia, form an important part of the West Kimberley National Heritage Place. Previous attempts to document these tracksites using traditional mapping techniques (e.g., surface overlays, transects and gridlines combined with conventional photography) have been hindered by the non-trivial challenges associated with working in this area, including, but not limited to: (1) the remoteness of many of the tracksites; (2) the occurrence of the majority of the tracksites in the intertidal zone; (3) the size and complexity of many of the tracksites, with some extending over several square kilometres. Using the historically significant and well-known dinosaurian tracksites at Minyirr (Gantheaume Point), we show how these issues can be overcome through the use of an integrated array of remote sensing tools. A combination of high-resolution aerial photography with both manned and unmanned aircraft, airborne and handheld high-resolution lidar imaging and handheld photography enabled the collection of large amounts of digital data from which 3D models of the tracksites at varying resolutions were constructed. The acquired data encompasses a very broad scale, from the sub-millimetre level that details individual tracks, to the multiple-kilometre level, which encompasses discontinuous tracksite exposures and large swathes of coastline. The former are useful for detailed ichnological work, while the latter are being employed to better understand the stratigraphic and temporal relationship between tracksites in a broader geological and palaeoecological context. These approaches and the data they can generate now provide a means through which digital conservation and temporal monitoring of the Dampier Peninsula’s dinosaurian tracksites can occur. As plans for the on-going management of the tracks in this area progress, analysis of the 3D data and 3D visualization will also likely provide an important means through which the broader public can experience these spectacular National Heritage listed landscapes.

A diversified vertebrate ichnite fauna from the Dasheng Group (Lower Cretaceous) of southeast Shandong Province, China

Abstract The Qingquan dinosaur tracksite, from the Lower Cretaceous Dasheng Group, Shandong Province, China adds to the growing record of saurischian-dominated ichnofaunas of the region. The site reveals the presence of avian theropods (Koreanaornis) and non-avian theropods tentatively referred to Jialingpus. Sauropod tracks are referred to Brontopodus. One site shows evidence of extensive trampling attributable to sauropods and theropods that moved in the same westerly direction, though not necessarily at exactly the same time. This site is reminiscent of the famous Davenport Ranch site in Texas which has provoked much debate about the herding behavior of sauropods as a defense strategy against theropod predators.

Erratum: The Evolution of Fangs, Venom, and Mimicry Systems in Blenny Fishes (Current Biology (2017) 27(8) (1184–191) (S0960982217302695) (10.1016/j.cub.2017.02.067))

(Current Biology 27, 714–720; March 6, 2017) In this article, we unintentionally omitted to expand on a citation of previously published results. In the caption of Figure 2, we stated that ‘‘F and p values indicate the significance of a phylogenetic ANCOVA testing for intercept differences between humans and other primates (see also Smaers and Rohlf [9], Supplemental Information..., and Table S2 for more detailed results)’’ (p. 716). Wewould like to clarify that in this statement, ‘‘see also Smaers and Rohlf’’ refers, specifically and exclusively, to the phylogenetic ANCOVA of primate prefrontal cortex to primary visual cortex and frontal motor areas using the Smaers dataset in [9]. These results were depicted in a subsection of our Figure 2 (the two top left regression plots) andwere numerically presented in a subsection of our Table S2. Smaers and Rohlf presented these results as an empirical example when describing the least-squares solution of phylogenetic ANCOVA and did not discuss the wider biological implications of these results for primate brain evolution. The presentation of the previous results was discussed openly during the review process of this manuscript. The authors apologize for any confusion this oversight may have caused.

Lower Cretaceous sauropod trackways from Lishui City and an overview of dinosaur dominated track assemblages from Zhejiang Province, China

A relatively small number of tetrapod tracksites from Zhejiang Province fills out the fossil record from that region, which is also known to have yielded both saurischian (titanosauriform, coelurosaur) and ornithischian (basal ornithopod, ankylosaurian) body fossils as well as dinosaurian eggs. We use photogrammetry and 3D imaging to reinterpret the sauropod tracks from Lishui City, which revealed the existence of trackways. The track record from three documented sites in Zhejiang Province includes avian and non-avian theropod, sauropod, ornithopod and pterosaur tracks. Previous work showed that the purported new bird ichnospecies Dongyangornipes sinensis is a synonym of Uhangrichnus chuni, while we here consider Pteraichnus dongyangensis as a nomen dubium. Such “provincial ichnotaxonomy” may mask similarities in the ichnofauna across large regions. In fact, we show that the ichnofauna is similar to that found in the Early Cretaceous elsewhere in China.

Two new dinosaur tracksites from the Lower Cretaceous Jiaguan Formation of Sichuan Basin, China: specific preservation and ichnotaxonomy

Abstract Two new dinosaur tracksites are reported from the Lower Cretaceous Jiaguan Formation in the Sichuan Basin, Qijiang District of Chongqing. These are the Gaoqing-Yongsheng and the Huibu tracksites, which represent the 13th and 14th reports from this formation. The Gaoqing-Yongsheng tracksite reveals the trackway of a large biped (ornithopod) in association with isolated sauropod tracks and large indeterminate undertracks with radial cracks. These features are preserved as natural casts with pebble infillings in a coarse, cross bedded and very thick bedded sandstone sequence. The Huibu tracksite reveals isolated theropod tracks and ornithopod tracks, the latter having a quadripartite, Caririchnium-like morphology, preserved in a thin bedded sandstone sequence with intercalated mudstone.