Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America

The extremes of dinosaur body size have long fascinated scientists. The smallest (<1 m length) known dinosaurs are carnivorous saurischian theropods, and similarly diminutive herbivorous or omnivorous ornithischians (the other major group of dinosaurs) are unknown. We report a new ornithischian dinosaur, Fruitadens haagarorum, from the Late Jurassic of western North America that rivals the smallest theropods in size. The largest specimens of Fruitadens represent young adults in their fifth year of development and are estimated at just 65–75 cm in total body length and 0.5–0.75 kg body mass. They are thus the smallest known ornithischians. Fruitadens is a late-surviving member of the basal dinosaur clade Heterodontosauridae, and is the first member of this clade to be described from North America. The craniodental anatomy and diminutive body size of Fruitadens suggest that this taxon was an ecological generalist with an omnivorous diet, thus providing new insights into morphological and palaeoecological diversity within Dinosauria. Late-surviving (Late Jurassic and Early Cretaceous) heterodontosaurids are smaller and less ecologically specialized than Early (Late Triassic and Early Jurassic) heterodontosaurids, and this ecological generalization may account in part for the remarkable 100-million-year-long longevity of the clade.     

A new sauropodomorph ichnogenus from the Lower Jurassic of Sichuan,China fills a gap in the track record

A dinosaur tracksite in the Lower Jurassic Ziliujing Formation of Sichuan Province, China consists of a spectacular sub-vertical exposure, with multiple track-bearing levels and trackways showing parallel and bimodal orientations. Based on well-preserved material, the new ichnogenus and ichnospecies, Liujianpus shunan ichnogen. nov. ichnosp. nov. is erected to accommodate distinctive sauropodomorph trackways occurring in this assemblage. Liujianpus has a unique combination of features, some relating to the early Jurassic basal sauropodomorph (prosauropod in traditional usage) ichnogenus Otozoum, others to the sauropod ichnogenus Brontopodus. Despite such a mix of basal sauropodomorph- and sauropod-like features, the trackmaker of Liujianpus is likely a basal sauropodomorph. This identification is consistent with the occurrence of basal sauropodomorph skeletons from geographically and chronologically close localities. The other distinct morphotype from the tracksite is linked to a sauropod trackmaker. As such, the ichnofauna consisting of two distinct foot morphotypes reflects the diversity of sauropodomorph dinosaurs in the Early Jurassic of Asia.     

The earliest fossil evidence of bone boring by terrestrial invertebrates,examples from China and South Africa

We report the oldest fossil evidence of osteophagia by terrestrial invertebrates on both the Asian and African continents. Bones attributable to the Middle Jurassic dinosaur Chuanjiesaurus (Dinosauria: Sauropoda) were found with post-mortem insect modification in the Chuanjie Formation, Yunnan Province, China. The morphology of the borings closely matches the ichnogenus Cubiculum. Based on the lack of bioglyphs observed in Cubiculum ornatus, a new ichnospecies is proposed here. The new trace fossil, Cubiculum inornatus isp. nov., is interpreted to have been constructed for pupation by an unknown taxon of insect. Additionally, we report even older borings from Early Jurassic dinosaur bones of the Elliott Formation in the Karoo Basin, which represent the second oldest occurrence of insect traces in bone from continental settings. Both trace fossils sites have palaeogeographic implications for the origins and dispersal of osteophagia amongst terrestrial invertebrates during the Mesozoic. These discoveries push back the antiquity of pupation in animal bones by more than 100 million years to the Middle Jurassic, indicating that this behaviour, and osteophagy more generally, originated early in the Mesozoic, roughly comparable with the origination of insect pupation in woody substrates (Late Triassic).     

Pisanosaurus mertii and the Triassic ornithischian crisis: could phylogeny offer a solution?

Abstract

Two recent studies have independently recovered Pisanosaurus mertii – long thought to represent the oldest known member of Ornithischia – within Silesauridae. These finds are expanded upon here, as are the implications of this hypothesis. Based upon these finds, it now appears that Ornithischia was absent in the Triassic Period entirely, which constitutes a major incongruence between the fossil record and current phylogenetic hypotheses, particularly the traditional model of dinosaur interrelationships in which Ornithischia and Saurischia are sister-taxa. It has been suggested previously that Ornithischia was simply a rare component of Late Triassic faunas, or that perhaps the clade’s ecology or geographic distribution were not conducive to producing a fossil record. Here I propose that phylogeny could hold the solution to this problem. I examine how an alternative position for Ornithischia – nested either within Theropoda or Sauropodomorpha – could be the reason behind their later appearance and relative rarity in the Early Jurassic. An Early Jurassic origin of Ornithischia would force us to consider that the anatomical similarities between ornithischians and Early Jurassic taxa might not be convergences, and to broaden the current datasets of early dinosaurs to test these ideas.     

Theropod and Ornithischian Footprints from the Middle Jurassic Yanan Formation of Zizhou County,Shaanxi, China

New tracksites reported from the Zizhou area elucidate the nature of the Early-Middle Jurassic dinosaurian ichnofaunas in Shaanxi Province. The assemblage is composed of footprints and trackways of medium- to large-sized theropods that show similarities with both the ichnogenera Kayentapus and Eubrontes and of small bipedal ornithischians that are referred to AS Anomoepus isp. Additionally tracks of a quadruped are present and assigned to Deltapodus isp. that may be related to a stegosaurian. Anomoepus isp. is similar to the holotype of Shensipus tungchuanensis which is, although apparently lost, re-assigned here and considered to be a subjective junior synonym of Anomoepus. It is therefore placed in new combination as Anomoepus tungchuanensis comb. nov. Identical tracks have been reported from well-preserved trackways both in the Zizhou and Shenmu areas, where they also co-occur with theropod tracks (Kayentapus and Grallator) and tracks of quadrupedal ornithischians (Shenmuichnus and Deltapodus). Thus, it appears that the carbonaceous (coal-bearing) facies of the region reveal ichnofaunas with both relatively abundant saurischian (theropod) and ornithischian tracks. This is in contrast with many areas where the ichnofaunas are heavily or exclusively saurischian (theropod) dominated.     

Review of Japanese Dinosaur Track Localities: Implications for Ichnotaxonomy,Paleogeography and Stratigraphic Correlation

Nine dinosaur ichnospecies from the Lower Jurassic to Upper Cretaceous of Japan, including two that are new, are described herein. The new ichnotaxa are Asianopodus pulvinicalx ichnogen. et ichnosp. nov. and Schizograllator otariensis ichnosp. nov. The Japanese ichnotaxa are allied to Lower Jurassic ichnospecies in South China, North America, Western Europe and South Africa, and Upper Jurassic to Lower Cretaceous ichnospecies from Southeast and East Asia. This suggests they were part of a global ichnofauna before continental drift began in the Middle Jurassic, leading to the development of a more endemic dinosaur fauna in the Cretaceous. At least two assemblages, an ornithopod-gracile-toed theropod-dominated community, in northeastern Asia, and a robust theropod- and sauropod-dominated community in the southern part of the continent, existed in the Cretaceous. This parallels North American dinosaur distribution patterns in the Cretaceous and seems to be a reflection of paleolatitudinal controls.     

Dinosaur footprints from the Phra Wihan Formation (Early Cretaceous of Thailand)

The ichnological assemblage of the Phra Wihan Formation (early Early Cretaceous of northeastern Thailand) includes the first sauropod tracks ever found in Thailand. It is reminiscent of the fauna of the underlying Jurassic Phu Kradung Formation with theropods, small ornithischians of uncertain affinities (possibly Hypsilophodon-like ornithopods) and sauropods. It suggests that, in southeastern Asia, an important faunal change occurred in the Early Cretaceous among continental vertebrates, whereas little happened at the Jurassic–Cretaceous boundary.     

Ophiomorpha irregulaire, Mesozoic trace fossil that is either well understood but rare in outcrop or poorly understood but common in core

The distinctive trace fossil, Ophiomorpha irregulaire, differs from other ichnospecies of Ophiomorpha in the so-called “meander maze” configuration of its horizontal course, and by the possession of unevenly distributed, elongated, tapering wall-lining pellets. The ichnospecies has been described, without doubt, from only two outcrop localities, both Late Cretaceous in age and both lying within the Cretaceous Western Interior Seaway. Only a single, almost convincing example of the trace fossil has been reported elsewhere, in a Lower Jurassic outcrop in Italy. Apart from this last occurrence, the distribution of O. irregulaire in outcrop suggests that the trace fossil has a restricted time span and geographical range. In contrast, O. irregulaire is identified more commonly in cores, and these occurrences show no clear geographical restrictions and are reported in sediments of Jurassic as well as Cretaceous age. Outcrop exposures of Jurassic and Cretaceous sediments have been extensively studied by ichnologists and if the large, eye-catching O. irregulaire were as common as is suggested by core material, then the trace fossil would not have escaped notice in outcrop.Identification of the trace fossil in core is generally made on the basis of two-dimensional, vertical sections. The characteristic meander maze form cannot be seen in this view, and the identification is thus based almost entirely on the special shape of the lining pellets. As is well known, the identification of large trace fossils in the restricted format of core lacks the security of extensively exposed trace fossils in outcrop. The discrepancy between age and geographic distribution of O. irregulaire in outcrop and core suggests that some misidentification is occurring in core analysis.     

First Thyreophoran Type Tracks from the Middle Jurassic Chuanjie Formation of Yunnan Province,China

Two tridactyl footprints from the Chuanjie Formation (Middle Jurassic) of Yunnan Province, China are morphological characteristics of thyreophoran tracks. They show some similarities to Shenmuichnus, known from the Early Jurassic strata of both Shaanxi and Yunnan provinces, but are somewhat larger, thereby resembling the ichnogenus Stegopodus. Based on their general morphology and size being congruent with this ichnogenus, they are tentatively assigned here to cf. Stegopodus. This is the fourth report of large ornithischian (probably thyreophoran) tracks from the Lower-Middle Jurassic of China that indicates relatively large trackmakers that were likely to be taxonomically distinct from much smaller and gracile Anomoepus trackmakers, also of ornithischian affinity. The larger tracks indicate a hitherto unreported abundance, size range and diversity of track types attributed to this group. The parallel orientation of the two best preserved trackways may indicate gregariousness.     

Diversity variation and tempo-spatial distribution of Otozamites (Bennettitales) in the Mesozoic of China

Otozamites is a representative fossil leaf morphogenus of the extinct Bennettitales, with an extensive distribution during the Mesozoic, especially in China. Understanding the fossil diversity variation and distribution pattern of Otozamites in China will provide information on biodiversity of bennettitalean plants as well as for reconstruction of palaeogeography and palaeoclimate conditions during the Mesozoic. So far, 46 species of this genus have been described in China, excluding unspecified species. The results show that the fossils of Otozamites are extensively recorded in the Late Triassic, and then reach their maximum development in the Early Jurassic, followed by a reduction in diversity in the Middle and Late Jurassic, and finally become extinct at the end of Early Cretaceous. Geographically, they occur in both Northern and Southern Floristic Provinces in the Mesozoic of China, with a relatively higher abundance in the Southern Floristic Province. It implies that the diversity variation and distribution of Otozamites are closely related to the change of the palaeoclimatic conditions. The warm and humid climate prevailed in the Late Triassic and Early Jurassic in South China, propitious to the development of Otozamites. After the Middle Jurassic, dry and hot climate may have caused the lower diversity level and blocked the development of Otozamites; finally at the end of the Early Cretaceous, the frequent arid climate may be a major cause for the extinction of Otozamites.     

Vagorichnus,a new ichnogenus for feeding burrow systems and its occurrence as discrete and compound ichnotaxa in Jurassic lacustrine turbidites of Central China

A new ichnogenus, Vagorichnus, is figured and described from Jurassic lacustrine turbidites of the Anyao Formation, Henan Province, Central China. Vagorichnus consists of complex burrow systems composed of discontinuous curved to meandering segments in which irregular networks are formed. The ichnospecies V. anyao covers burrow systems characterized by constricted and irregularly annulated segments, and ridge‐like knobs. Although traces are preserved as hypichnial ridges on the sole of turbidite sandstones, several lines of evidence indicate that these ichnofossils record the activity of a post‐event infauna. Vagorichnus occurs as discrete and compound ichnotaxa. In the latter case, it grades into Gordia and Tuberculichnus. Vagorichnus is interpreted as invertebrate feeding structures (Fodinichnia).     

A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia

Although the group played an important role in the evolution of Late Mesozoic terrestrial ecosystems, the early evolutionary history of the ornithischian dinosaurs remains poorly understood. Here, we report on a new primitive ornithischian, Eocursor parvus gen. et sp. nov., from the Late Triassic (?Norian) Lower Elliot Formation of South Africa. Eocursor is known from a single specimen comprising substantial cranial and postcranial material and represents the most complete Triassic member of Ornithischia, providing the earliest evidence for the acquisition of many key ornithischian postcranial characters, including an opisthopubic pelvis. A new phylogenetic analysis positions this taxon near the base of Ornithischia, as the sister taxon to the important and diverse clade Genasauria. The problematic clade Heterodontosauridae is also positioned basal to Genasauria, suggesting that an enlarged grasping manus may represent a plesiomorphic ornithischian condition. This analysis provides additional phylogenetic support for limited ornithischian diversity during the Late Triassic, and suggests that several major ornithischian clades may have originated later than generally believed. There are few morphological differences between Late Triassic and Early Jurassic ornithischians, supporting previous suggestions that the Early Jurassic ornithischian radiation may simply represent the filling of vacant ecological space following Late Triassic terrestrial extinctions.     

中国中生代的马通蕨科植物:化石记录、多样性与时空分布特征*

现生真蕨目马通蕨科(Matoniaceae)植物仅存Matonia和Phanerosorus两属, 集中分布于马来西亚、印度尼西亚等热带地区。马通蕨科植物在中生代时期全球广布, 且主要分布于热带、亚热带地区, 有近9个属, 被作为热带、亚热带气候的标志性植物化石之一。本文梳理总结了中国中生代的马通蕨科化石记录并分析其多样性特征, 共计有2属16种, 包括异脉蕨属(Phlebopteris) 15种和准马通蕨属(Matonidium) 1种。对其化石记录和地质地理分布分析表明, 晚三叠世时期, 异脉蕨属植物广泛分布于热带—亚热带湿热气候区, 包括华南一带的四川、湖北、 云南、西藏、福建等地; 早侏罗世时期, 其分布逐渐向北方扩展, 在南、北方植物区系界线附近均有发现; 中侏罗世局限于湖北、青海等地; 早白垩世时仅在黑龙江和西藏少量发现。准马通蕨属仅在黑龙江地区的早白垩世地层中发现。整体上, 马通蕨科在中国中生代的分布范围变迁与气候带范围变化相吻合。     

The Trace Fossil Diplopodichnus from the Lower Jurassic Lacustrine Sediments of Central China and the Isopod Armadillidium vulgare (Pillbug) Lebensspuren as its Recent Analogue

The trace fossil Diplopodichnus isp. was identified in lacustrine siliciclastics of the Lower Jurassic Anyao Formation in the Henan Province, central China. This is the youngest occurrence of this ichnogenus. It is similar to some variants of recent surface traces of the isopod Armadillidium vulgare (pillbug) crawling on soft mud in temporal puddles. Therefore, isopods are suggested to be producers of the described Diplopodichnus isp. It seems that isopods should be taken into account as important component of deep lacustrine ecosystems by the Early Jurassic.     

The Boring Cunctichnus probans Fürsich,Palmer and Goodyear, 1994, from the Type Maastrichtian (Upper Cretaceous,Northeast Belgium)

The ichnogenus Cunctichnus Fürsich, Palmer and Goodyear, 1994 is monotypic, its type ichnospecies, C. probans Fürsich, Palmer and Goodyear, 1994, having hitherto been described solely from the Upper Jurassic (Tithonian) of southern England and the Paris Basin. Cunctichnus probans is recorded herein from the type Maastrichtian (Upper Cretaceous) of the province of Liège, northeast Belgium, that is, about 75 million years younger than previous published records. This specimen differs from the holotype in having a Y-shaped branch.     

Ichnotaxonomic Review of Large Ornithopod Dinosaur Tracks: Temporal and Geographic Implications

BackgroundLarge ornithopod tracks are known from the Upper Jurassic to the uppermost Cretaceous rocks of all continents but Antarctica. They include the tracks historically called Iguanodon footprints, iguanodontid footprints, hadrosaur/hadrosaurid footprints, and other large ornithopod tracks that have been used to define ichnotaxa. More than 40 ichnospecies based on large ornithopod tracks have been defined, but the validity of many of them is questionable.Conclusions/SignificanceThe monospecific ichnogenus Iguanodontipus (I. burreyi) is characterized by a small, rounded heel and elongate, narrow digit impressions. Its distribution is limited to the Berriasian-Valanginian of Europe. Caririchnium consists of four ichnospecies (C. magnificum [type ichnospecies], C. kortmeyeri, C. billsarjeanti and C. lotus) with a large, rounded heel and short, wide digit impressions. This ichnogenus ranges from the Berriasian-Hauterivian to the Aptian-Albian of South America, North America, Asia and Europe. Finally, Hadrosauropodus (three ichnospecies: H. langstoni [type ichnospecies], H. leonardii and H. kyoungsookimi) shows a large, bilobed heel and short, wide digit impressions. It is known from the Aptian-Albian to the Maastrichtian of North America, Asia and Europe. The ichnofamily Iguanodontipodidae includes large iguanodontian tracks characterized mainly by mesaxonic, tridactyl and subsymmetrical pes tracks that are as wide as (or wider than) long and have one pad impression in each digit and one in the heel. Its distribution is confidently limited to the Cretaceous of Europe, Asia, North America and South America.