Arboreal Lifestyle Research Articles

A new darwinopteran pterosaur reveals arborealism and an opposed thumb

Pterosaurs, which lived during the Mesozoic, were the first known vertebrates to evolve powered flight.1,2 Arboreal locomotion has been proposed for some taxa,3,4 and even considered to have played a role in the origin of pterosaur flight.5,6 Even so, there is still need for comprehensive quantitative ecomorphological analyses.3,4 Furthermore, skeletal adaptations correlated to specialized lifestyles are often difficult to recognize and interpret in fossils. Here we report on a new darwinopteran pterosaur that inhabited a unique forest ecosystem from the Jurassic of China. The new species exhibits the oldest record of palmar (or true) opposition of the pollex, which is unprecedented for pterosaurs and represents a sophisticated adaptation related to arboreal locomotion. Principal-coordinate analyses suggest an arboreal lifestyle for the new species but not for other closely related species from the same locality, implying a possible case of ecological niche partitioning. The discovery adds to the known array of pterosaur adaptations and the history of arborealism in vertebrates. It also adds to the impressive early bloom of arboreal communities in the Jurassic of China, shedding light on the history of forest environments.

Hind Limb Bone Proportions Reveal Unexpected Morphofunctional Diversification in Xenarthrans

Divergence in forelimb morphology is often associated with functional habits exhibited within the Xenarthra, ranging from terrestrial-digging in armadillos to arboreal-suspension in sloths. We hypothesized that quantitative differences in hind limb form also will be predictive of the diverse lifestyles observed in this small clade. A total of 26 morphofunctional indices were calculated from 42 raw measurements of bone length/width/depth in a sample of N = 76 skeletal specimens (18 species). Index data for each species were categorized by substrate preference and use and then evaluated using a combination of stepwise Discriminant Function Analysis (DFA) and Principal Component Analysis (PCA) to determine significant osteological correlates (traits) among extant taxa. Additionally, character states of the morphometric data were inferred using a recent hypothesis of xenarthran phylogeny. DFA determined 14 distinct morphofunctional indices relating to femur robustness, hip/ankle/limb mechanical advantage, and foot and claw length as the most discriminating features. PCA clearly separated armadillos from sloths in morphospace based on overall robustness versus gracility, as well as proximal versus distal lengths of skeletal elements (including the claws), whereas these characteristics were intermediate in the hind limbs of anteaters and selected armadillos having either a larger greater trochanter or modified foot/claw proportions. Two-toed and three-toed sloths showed further separation from each other in morphospace primarily driven by proportions of their tibia and hind feet despite evidence of convergence for numerous functional traits. Moreover, the majority of the traits measured had significant phylogenetic signal and several of these indicated clear patterns of convergent and divergent evolution in xenarthrans by evaluation of their tip states. Our assessments expand functional interpretations of xenarthran limb form and identify potentially conserved and secondarily modified traits related to fossoriality across taxa, including in three-toed sloths, demonstrate possible morphological trade-offs between digging and climbing habits, and suggest derived traits adapted for arboreal lifestyle and suspensory function.

Walking on water: the unexpected evolution of arboreal lifestyle in a large top predator in the Amazon flooded forests.

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Do substrate roughness and gap distance impact gap-bridging strategies in arboreal chameleons?

Chameleons are well equipped for an arboreal lifestyle, having “zygodactylous” hands and feet as well as a fully prehensile tail. However, to what degree tail use is preferred over autopod prehension has been largely neglected. Using an indoor experimental set-up, where chameleons had to cross gaps of varying distances, we tested the effect of substrate diameter and roughness on tail use in Chamaeleo calyptratus. Our results show that when crossing greater distances, C. calyptratus is more likely to use its tail for additional stability. The animals were able to cross greater distances (up to 1.75 times the shoulder-hip length) on perches with a rougher surface. We saw that depending on the distance of the gap, chameleons would change how they use their prehensile tails when crossing. With shorter gaps the tails either do not touch, or only touch the perch without coiling around it. With larger distances the tails are fully coiled around the perch, and with the largest distances additionally they reposition the hind legs, shifting them towards the end of the perch. Males were able to cross relatively greater distances than females, likely due to their larger size and strength.

Сравнение способности к обучению макаков резусов ( Macaca mulatta ) и павианов гамадрилов ( Papio hamadryas ) в свете создания модели когнитивной эволюции приматов

A comparative study of the learning ability in two species of lower monkeys, rhesus monkeys (Macaca mulatta) and hamadryas baboons (Papio hamadryas), was used as a hypothetical model for the development of cognitive abilities in primates during such an evolutionary event as the transition from an arboreal lifestyle to a terrestrial one. It was found that in male rhesus monkeys the average level of learning was 68.5 ± 5.8%, in male hamadryas baboons - 78.0 ± 3.1%. The median of the distribution of learning, which characterizes the maximum number of continuous correct decisions, in rhesus monkeys is 17.0% (interquartile range, IQR = 11.0); in hamadryas baboons - 40.0% (IQR = 15.5). In the studied groups of monkeys, certain differences were found in the dynamics of skill formation. Our results illustrate the advantages of terrestrialism as a factor contributing to evolution towards rationality, and are in the best possible way with the ecological hypothesis of the development of cognitive abilities.

Revisiting the Evolution of Arboreal Life in Oribatid Mites

Though mostly soil dwelling, oribatid mites are found in all kind of habitats, with several species exclusively living on trees. Using previously published DNA sequences and eco-morphological data available from the literature, we inferred the number of transitions between soil dwelling to a truly arboreal lifestyle in oribatid mites and the shape evolution of a particular morphological structure of a sense organ (bothridial seta (= sensillus) of a trichobothrium), the shape of which was previously reported to be associated with an arboreal lifestyle. Our data suggest that a truly arboreal lifestyle evolved several times independently in oribatid mites, but much less often than previously proposed in the past. Even though all truly arboreal species indeed seem to possess a capitate sensillus, this character is not exclusive for arboreal taxa. Nonetheless, since all truly arboreal species do have a capitate sensillus, this might be considered an important (pre-)adaptation to a life on trees. We further provide guidelines on how the term “arboreal” should be applied in future mite research and emphasize the importance of exact microhabitat characterization, as this will greatly facilitate comparisons across studies.

Notes on Forefoot Characteristics of the Ryukyu Long-Furred Rat, Diplothrix legata, for Its Arboreal Lifestyle

Forefoot characteristics of the Ryukyu long-furred rat, Diplothrix legata, in the Amami-oshima, Tokunoshima, and Okinawajima Islands of the southern part of Japan (the Nansei Archipelago), were analyzed morphologically considering its arboreal lifestyle. The presence of a nail on the pollex was confirmed and seemed to be regarded as an arboreal feature, based on the presence of nails in other arboreal murids reported in previous studies. In addition, a linear palmar groove was confirmed between the interdigital pads and the thenar-hypothenar pads. We tried to allow the forefoot to take hold of a thin stick at this groove; the stick fit well in the groove, and it seemed to be possible to grip by the fingers except for pollex. Thus, the sizes and positions of palmar swollen pads forming the linear groove were considered to be useful for gripping a thin twig in an arboreal lifestyle. In addition, the entire unguis size tended to be larger in Okinawajima Island than in Amami-oshima Island as a geographical variation.

Increased terrestriality in a Neotropical primate living on islands with reduced predation risk

An arboreal lifestyle is thought to be central to primate origins, and most extant primate species still live in the trees. Nonetheless, terrestrial locomotion is a widespread adaptation that has arisen repeatedly within the primate lineage. The absence of terrestriality among the New World monkeys (Platyrrhini) is thus notable and raises questions about the ecological pressures that constrain the expansion of platyrrhines into terrestrial niches. Here, we report the results of a natural experiment, comparing patterns of terrestrial behavior in white-faced capuchin monkeys (Cebus capucinus imitator) living on two islands off the Pacific coast of Panama that lack mammalian predators (island sites) with the behavior of capuchins at three sites in central Panama with more intact predator communities (mainland sites). Surveys with camera traps revealed increased terrestriality in island vs. mainland sites. Capuchin detection rates were higher, the range of party sizes observed was larger, and individuals engaged in a wider range of terrestrial behaviors on the islands lacking mammalian predators. Furthermore, females carrying infants were frequently photographed on the ground at the island sites, but never at the mainland sites. These findings support the long-standing hypothesis that predators constrain the exploitation of terrestrial niches by primates. These results are also consistent with the hypothesis that arboreal locomotion imposes costs that primates will avoid by walking on the ground when predation risk is low.

Inferring lifestyle for Aves and Theropoda: A model based on curvatures of extant avian ungual bones.

Claws are involved in a number of behaviours including locomotion and prey capture, and as a result animals evolve claw morphologies that enable these functions. Past authors have found geometry of the keratinous sheath of the claw to correlate with mode of life for extant birds and squamates; this relationship has frequently been cited to infer lifestyles for Mesozoic theropods including Archaeopteryx. However, many fossil claws lack keratinous sheaths and thus cannot be analysed using current methods. As the ungual phalanx within the claw is more commonly preserved in the fossil record, geometry of this bone may provide a more useful metric for paleontological analysis. In this study, ungual bones of 108 birds and 5 squamates were imaged using X-ray techniques and a relationship was found between curvatures of the ungual bone within the claw of pedal digit III and four modes of life; ground-dwelling, perching, predatory, and scansorial; using linear discriminant analysis with weighted accuracy equal to 0.79. Our model predicts arboreal lifestyles for Archaeopteryx and Microraptor and a predatory ecology for Confuciusornis. These findings demonstrate the utility of our model in answering questions of palaeoecology, the theropod-bird transition, and the evolution of avian flight. Though the metric exhibits a strong correlation with lifestyle, morphospaces for PD-III curvatures overlap and so this metric should be considered alongside additional evidence.

Is variation in tail vertebral morphology linked to habitat use in chameleons?

Chameleons (Chamaeleonidae) are known for their arboreal lifestyle, in which they make use of their prehensile tail. Yet, some species have a more terrestrial lifestyle, such as Brookesia and Rieppeleon species, as well as some chameleons of the genera Chamaeleo and Bradypodion. The main goal of this study was to identify the key anatomical features of the tail vertebral morphology associated with prehensile capacity. Both interspecific and intra-individual variation in skeletal tail morphology was investigated. For this, a 3D-shape analysis was performed on vertebral morphology using μCT-images of different species of prehensile and nonprehensile tailed chameleons. A difference in overall tail size and caudal vertebral morphology does exist between prehensile and nonprehensile taxa. Nonprehensile tailed species have a shorter tail with fewer vertebrae, a generally shorter neural spine and shorter transverse processes that are positioned more anteriorly (with respect to the vertebral center). The longer tails of prehensile species have more vertebrae as well as an increased length of the processes, likely providing a greater area for muscle attachment. At the intra-individual level, regional variation is observed with more robust proximal tail vertebrae having longer processes. The distal part has relatively longer vertebrae with shorter processes. Although longer, the small size and high number of the distal vertebrae allows the tail to coil around perches.

Jumping in arboreal salamanders: A possible tradeoff between takeoff velocity and in-air posture

Jumping performance can have important implications for an animal’s fitness by expanding its ability to evade predators and move between microhabitats. Jumping in terrestrial plethodontid salamanders is achieved through lateral bending and rapid unbending of the trunk, an action powered by axial musculature. Arboreal plethodontids, some of which are known to occupy tree crowns, tend to have more robust limbs and longer digits, which may affect their jumping kinematics and performance. We examined jumping kinematics in ten species of plethodontid salamanders, including four arboreal species of the genus Aneides, using high speed imaging and kinematic analysis. Salamanders of the genus Aneides exhibit lower takeoff velocities when compared with the terrestrial plethodontids Eurycea, Desmognathus, and Plethodon, possibly due to reduced trunk bending. Aneides also exhibit higher frequencies of two-footed takeoffs, and often orient their feet to launch from vertical surfaces when presented with the option. This suggests an alternative jumping behavior in salamanders that may reflect an arboreal lifestyle. All plethodontid species examined displayed distinctive in-air parachute postures after 45–100% of descending jumps, with Aneides showing the highest frequency of this behavior.

Late Turolian Mesopithecus (Mammalia: Cercopithecidae) from Axios Valley (Macedonia, Greece): earliest presence of M. monspessulanus in Europe

The fossil mammal localities of the Axios Valley (Macedonia, Greece) yielded a rich collection of Mesopithecus remains, which were first described at the beginning of the 1990s. The late Turolian Dytiko localities include several specimens of Mesopithecus, which were originally separated in two size-related forms: the relatively larger-sized M. cf. or aff. pentelicus, and the relatively smaller-sized M. cf. monspessulanus. However, some Dytiko specimens were only partially described, or remained even undescribed because the cranium either was still in connection with the mandible or was markedly deformed. Later, careful cleaning of the materials and their revision based on larger comparative samples indicate that they represent two distinct species: the relatively larger-sized specimens belong to M. pentelicus, while the relatively smaller-sized ones to M. monspessulanus. The Dytiko M. pentelicus has slightly smaller dental dimensions compared to M. pentelicus from Pikermi, indicating a possible trend for size decrease. The Dytiko M. monspessulanus, although close to the typical form of this species, has somewhat larger dimensions. The semi-terrestrial M. pentelicus disappeared at the end of the Miocene, while some of its populations adapted to the wetter and more woody Pliocene habitats, finally giving origin to M. monspessulanus, whose elbow-joint indicates an arboreal lifestyle. M. monspessulanus was widely distributed in Europe, but its remains are very scarce and are all from the Pliocene. Some questionable indications for the presence of M. monspessulanus in the latest Miocene come from some Italian Messinian sites (Gravitelli, Baccinello V3), but this scenario still needs verification. However, even if its presence in some Italian localities was confirmed, the Dytiko M. monspessulanus nonetheless represents the earliest known occurrence of this species in Europe, as the Dytiko fauna is considered as latest Turolian (pre-Messinian, 7.0–6.0Ma).

The evolution of femoral cross-sectional properties in sciuromorph rodents: Influence of body mass and locomotor ecology.

In several groups of mammals, adaptation to differing functional demands is reflected in long bone cross-sectional properties (CSP), which relate to the resistance to compression and to bending loads in the craniocaudal and mediolateral directions. Members of the Sciuromorpha ("squirrel-like" rodents) display a diversity of locomotor ecologies and span three orders of magnitude in terms of body size. The availability of robust phylogenies is rendering them a suitable group to further substantiate the relationship of long bone CSP with locomotor ecology and body mass while taking the phylogenetic non-independence among species into account. Here, we studied 69 species of Sciuromorpha belonging to three lifestyle categories, "arboreal," "fossorial," and "aerial" (i.e., gliding). We hypothesized locomotor category specific loading regimes that act on femora during predominant or, in terms of gliding, critical locomotor behaviors of each category. High resolution computed tomography scans of the specimens' femora were obtained and cross-sections in 5% increments were analyzed. Cross-sectional area, the craniocaudal second moment of area (SMAcc ), and the mediolateral second moment of area were quantified. Further, a scaling analysis was conducted for each bone cross-section to examine how the CSP scale with body mass. Body mass accounted for variances in CSP with mainly positive allometry. The aerial sciuromorphs showed lower values of CSP compared to the arboreal and fossorial species in the distal epiphysis for all quantified parameters and along the bone for SMAcc . In contrast to previous studies on other mammalian lineages, no differences in CSP were found between the fossorial and the arboreal lifestyles.

Filling in the phylogenetic gaps: Induction, migration, and differentiation of neural crest cells in a squamate reptile, the veiled chameleon (Chamaeleo calyptratus).

Neural crest cells comprise a migratory progenitor cell population that differentiate into cell types such as neurons and glia of the peripheral nervous system, pigment cells, hormone secreting cells in glands, and skeletal and connective tissue in the head, thus making important contributions to most tissues and organs throughout the vertebrate body. The evolutionary appearance of neural crest cells is considered synonymous with the origin of vertebrates and their subsequent diversification and radiation. While the comparative biology of neural crest cells has been studied for a century and a half beginning with their discovery by Wilhelm His in 1868, most of our understanding of their development and function has come from a small number of species. Thus, critical gaps exist in our understanding of how neural crest cells mediate evolution and development. This is particularly true with respect to squamate reptiles (lizards, snakes, amphisbaenians), which account for approximately one-third of all living tetrapods. Here, we present veiled chameleons (Chamaeleo calyptratus) as a model system for studying neural crest cell development in squamates. Chameleons exhibit various morphological specializations associated with an arboreal lifestyle that may have been facilitated through neural crest cells acting as a conduit for evolutionary change.

Postcranial Skeleton of Henkelotherium guimarotae (Cladotheria, Mammalia) and Locomotor Adaptation

X-ray computed tomography of the cladotherian Henkelotherium guimarotae from the Kimmeridgian Guimarota coal mine (Portugal) adds new information on its postcranial skeleton. Contrary to earlier description, the humerus of Henkelotherium shows a plesiomorphic cladotherian condition similar to that seen in Dryolestes leiriensis in having individual radial and ulnar condyles on the anterior aspect of the distal joint, whereas the posterior aspect appears to be trochlear. The scapula is also more plesiomorphic than previously assumed in having a laterally flared crest on the inferior margin. In this regard, it is similar to the scapula of spalacotherioid “symmetrodontans” such as Zhangheotherium but more derived than the latter, in having a larger supraspinous fossa. The hind limb is similar to that of small extant therians, which is interpreted to be assocated with an upright limb posture. The previously proposed hypothesis of an arboreal lifestyle for Henkelotherium is corroborated by a number of skeletal features such as large humeral entepicondyles, prominent digital flexor tubercles, elongated and slender tail vertebrae, as well as a high pedal phalangeal index of 153%.

The transcriptome of the veiled chameleon (Chamaeleo calyptratus): A resource for studying the evolution and development of vertebrates.

The veiled chameleon (Chamaeleo calyptratus) is an emerging model system for studying functional morphology and evolutionary developmental biology (evo-devo). Chameleons possess body plans that are highly adapted to an arboreal life style, featuring laterally compressed bodies, split hands/ft for grasping, a projectile tongue, turreted independently moving eyes, and a prehensile tail. Despite being one of the most phenotypically divergent clades of tetrapods, genomic resources for chameleons are severely lacking. To address this lack of resources, we used RNAseq to generate 288 million raw Illumina sequence reads from four adult tissues (male and female eyes and gonads) and whole embryos at three distinct developmental stages. We used these data to assemble a largely complete de novo transcriptome consisting of only 82 952 transcripts. In addition, a majority of assembled transcripts (67%) were successfully annotated. We then demonstrated the utility of these data in the context of studying visual system evolution by examining the content of veiled chameleon opsin genes to show that chameleons possess all five ancestral tetrapod opsins. We present this de novo, annotated, multi-tissue transcriptome assembly for the Veiled Chameleon, Chamaeleo calyptratus, as a resource to address a range of evolutionary and developmental questions. The associated raw reads and final annotated transcriptome assembly are freely available for use on NCBI and Figshare, respectively.

Convergence in the functional properties of forelimb muscles in carnivorans: adaptations to an arboreal lifestyle?

The arboreal environment is complex and irregular and imposes significant constraints on the locomotor ability of vertebrates, resulting in morphological adaptations. Carnivorans are of particular interest because they radiated several times independently into the arboreal milieu, leading to convergence in long bone anatomy. We investigated whether the forelimb muscles show differences in arboreal vs. terrestrial species and tested for convergence in muscle architecture. To do so, we studied the forelimb muscles in 17 species of carnivorans with different locomotor modes (arboreal and terrestrial). Our results showed that the forelimb musculature evolved in a convergent manner in arboreal species, resulting in more forceful and heavier wrist rotators and elbow flexors, even when accounting for phylogeny. This suggests that selective forces imposed by the environment drive the evolution of the muscular system towards a convergent phenotype that provides a functional advantage for locomotion in the complex three-dimensional arboreal habitat.

Are sleeping site ecology and season linked to intestinal helminth prevalence and diversity in two sympatric, nocturnal and arboreal primate hosts (Lepilemur edwardsi and Avahi occidentalis)?

BackgroundVarious factors, such as climate, body size and sociality are often linked to parasitism. This constrains the identification of other determinants driving parasite infections. Here, we investigate for the first time intestinal parasites in two sympatric arboreal primate species, which share similar activity patterns, feeding ecology, body size and sociality, and cope with the same climate conditions, but differ in sleeping site ecology: the Milne-Edward’s sportive lemur (Lepilemur edwardsi) and the Western woolly lemur (Avahi occidentalis). Comparison of these two species aimed to test whether differences in sleeping sites are related to differences in parasite infection patterns. Additionally, gender and seasonal factors were taken into account. Animals were radio-collared to record their sleeping site dynamics and to collect fecal samples to assess intestinal parasitism during both the dry and the rainy season.ResultsOnly low parasite diversity was detected, which is attributable to the strict arboreal lifestyle of these lemurs, limiting their contact with infective parasite stages. L. edwardsi, which sleeps in tree holes and repeatedly uses the same sleeping site, excreted eggs of strongyle and oxyurid nematodes, whereby strongyles always occurred in coinfection with oxyurids. In contrast, A. occidentalis, which sleeps on open branches and frequently changes sleeping sites, only excreted eggs of strongyle nematodes. This difference can be attributed to a potential favorable environment presented by tree holes for infective stages, facilitating parasitic transmission. Additionally, Strongylida in A. occidentalis were only observed in the rainy season, suggesting an arrested development during the dry season in the nematodes’ life cycle. Males and females of both lemur species showed the same frequency of parasitism. No differences in body mass of infected and non-infected individuals were observed, indicating that the animals’ body condition remains unaffected by the detected gastrointestinal parasites.ConclusionsThe comparison of two primate hosts with a very similar lifestyle suggests an influence of the sleeping site ecology on intestinal parasites. In A. occidentalis there was a clear seasonal difference in strongyle egg excretion. These results improve our understanding of the parasite ecology in these endangered primate species, which may be critical in the light of species conservation.

Early Evolution of Modern Birds Structured by Global Forest Collapse at the End-Cretaceous Mass Extinction

The fossil record and recent molecular phylogenies support an extraordinary early-Cenozoic radiation of crown birds (Neornithes) after the Cretaceous-Paleogene (K-Pg) mass extinction [1-3]. However, questions remain regarding the mechanisms underlying the survival of the deepest lineages within crown birds across the K-Pg boundary, particularly since this global catastrophe eliminated even the closest stem-group relatives of Neornithes [4]. Here, ancestral state reconstructions of neornithine ecology reveal a strong bias toward taxa exhibiting predominantly non-arboreal lifestyles across the K-Pg, with multiple convergent transitions toward predominantly arboreal ecologies later in the Paleocene and Eocene. By contrast, ecomorphological inferences indicate predominantly arboreal lifestyles among enantiornithines, the most diverse and widespread Mesozoic avialans [5-7]. Global paleobotanical and palynological data show that the K-Pg Chicxulub impact triggered widespread destruction of forests [8, 9]. We suggest that ecological filtering due to the temporary loss of significant plant cover across the K-Pg boundary selected against any flying dinosaurs (Avialae [10]) committed to arboreal ecologies, resulting in a predominantly non-arboreal post-extinction neornithine avifauna composed of total-clade Palaeognathae, Galloanserae, and terrestrial total-clade Neoaves that rapidly diversified into the broad range of avian ecologies familiar today. The explanation proposed here provides a unifying hypothesis for the K-Pg-associated mass extinction of arboreal stem birds, as well as for the post-K-Pg radiation of arboreal crown birds. It also provides a baseline hypothesis to be further refined pending the discovery of additional neornithine fossils from the Latest Cretaceous and earliest Paleogene.

First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny

A new fossil amniote from the Fossil Forest of Chemnitz (Sakmarian-Artinskian transition, Germany) is described as Ascendonanus nestleri gen. et sp. nov., based on five articulated skeletons with integumentary preservation. The slender animals exhibit a generalistic, lizard-like morphology. However, their synapsid temporal fenestration, ventrally ridged centra and enlarged iliac blades indicate a pelycosaur-grade affiliation. Using a renewed data set for certain early amniotes with a similar typology found Ascendonanus to be a basal varanopid synapsid. This is the first evidence of a varanopid from Saxony and the third from Central Europe, as well as the smallest varanopid at all. Its greatly elongated trunk, enlarged autopodia and strongly curved unguals, along with taphonomical observations, imply an arboreal lifestyle in a dense forest habitat until the whole ecosystem was buried under volcanic deposits. Ascendonanus greatly increases the knowledge on rare basal varanopids; it also reveals a so far unexpected ecotype of early synapsids. Its integumentary structures present the first detailed and soft tissue skin preservation of any Paleozoic synapsid. Further systematic results suggest a varanodontine position for Mycterosaurus, the monophyly of South African varanopids including Anningia and the distinction of a skeletal aggregation previously assigned to Heleosaurus, now renamed as Microvaranops parentis gen. et sp. nov.