Wednesday, July 25, 2018

[Paleontology • 2018] Lingwulong shenqi • A New Middle Jurassic Diplodocoid Suggests An Earlier Dispersal and Diversification of Sauropod Dinosaurs


Lingwulong shenqi
Xu, Upchurch, Mannion, Barrett, Regalado-Fernandez, Mo, Ma & Liu, 2018


Abstract
The fragmentation of the supercontinent Pangaea has been suggested to have had a profound impact on Mesozoic terrestrial vertebrate distributions. One current paradigm is that geographic isolation produced an endemic biota in East Asia during the Jurassic, while simultaneously preventing diplodocoid sauropod dinosaurs and several other tetrapod groups from reaching this region. Here we report the discovery of the earliest diplodocoid, and the first from East Asia, to our knowledge, based on fossil material comprising multiple individuals and most parts of the skeleton of an early Middle Jurassic dicraeosaurid. The new discovery challenges conventional biogeographical ideas, and suggests that dispersal into East Asia occurred much earlier than expected. Moreover, the age of this new taxon indicates that many advanced sauropod lineages originated at least 15 million years earlier than previously realised, achieving a global distribution while Pangaea was still a coherent landmass.


Systematic paleontology
Sauropoda Marsh, 1878
Neosauropoda Bonaparte, 1986
Diplodocoidea (Marsh, 1884)

Dicraeosauridae Janensch, 1929

Lingwulong shenqi gen. et sp. nov.

Etymology: Lingwu, after the region where the specimens were found; long, the Mandarin Chinese for ‘dragon’; and shenqi, the Mandarin Chinese for ‘amazing’, reflecting the unexpected discovery of a dicraeosaurid in the Middle Jurassic of China.

Horizon and locality: Yanan Formation, late Early to early Middle Jurassic (late Toarcian–Bajocian), Lingwu Geopark, near Ciyaopu, Ningxia Hui Autonomous Region, China.

Diagnosis: Autapomorphies: prefrontal anterior process directed laterally; orbital dorsal margin strongly ornamented by deep, longitudinal grooves and tubercles; long-axes of the free tips of the basal tubera directed anteromedially; capitate process mediolaterally long (length:height ratio c. 5.0); occipital condyle articular surface wide transversely (width:height ratio c. 1.54); lateral surface of cervical prezygapophyseal process bears a ridge formed by a linear array of tubercles; subcircular facet-like region at the summit of metapophyses in middle cervical-anterior dorsal vertebrae; small process projects anterodorsally from the anterior margin of the transverse process, near its distal end, in anterior dorsal vertebrae; anterior dorsal metapophyses twisted along their length; anterior caudal neural spines bear subtriangular facet-like areas, extending from summit to spine mid-height.




Fig. 1 Cranial material of Lingwulong shenqi. Braincase in:
 left lateral (a), dorsal (b), occipital (c), and ventral (d) views. Dentary teeth in occlusal view (e). The 5th and 6th left dentary tooth crowns in labial view (f).

 Abbreviations: bpp, basipterygoid process; bt, basal tubera; f, frontal; fps, frontoparietal suture; gt, grooves and tubercles; pf, prefrontal; llp, ‘leaf’-like process; oc, occipital condyle; p, parietal; pcp, capitate process; paf, proatlantal facet; pag, proatlantal groove; pfap, prefrontal anterior process; po, postorbital; povp, postorbital ventral process; pp, paroccipital process; sc, sagittal crest; so, supraoccipital; sq, squamosal; sqhp, squamosal hook-like process; sqvp, squamosal ventral process; stf, supratemporal fenestra; wf, wear facet. Scale bars = 20 mm for a–e and 10 mm for f

Fig. 2 Skeletal reconstruction and exemplar skeletal remains of Lingwulong shenqi.
 Silhouette showing preserved elements (a); middle cervical vertebra in left lateral (b) and anterior (c) views; anterior dorsal vertebra in left lateral (d) and anterior (e) views; posterior dorsal vertebra in lateral view (f); sacrum and ilium in left lateral view (g); anterior caudal vertebra in left lateral (h) and anterior (i) views; right scapulocoracoid in lateral view (j); right humerus in anterior view (k); left pubis in lateral view (l); right ischium in lateral (m) views; right femur in posterior view (n); and right tibia in lateral view (o).

Abbreviations: ap, ambiens process; ar, acromial ridge; ip, iliac peduncle; naf, notch anterior to glenoid; np, neural spine; podl, postzygodiapophyseal lamina; ppr, prezygapophyseal process ridge; prp, prezygapophysis; pvf, posteroventral fossa; slf, shallow lateral fossa; spol, spinopostzygapophyseal lamina; sprl, spinoprezygapophyseal lamina; wls, wing-like structure. Scale bars = 100 cm for a and 5 cm for b–o


Fig. 3 Time-calibrated evolutionary tree for Eusauropoda. Agreement subtree produced in TNT, with additional diplodocid taxa incorporated (see Supplementary Note 4). All macronarian taxa have been combined into a single lineage, and non-sauropod sauropodomorphs have been removed, in order to enhance clarity (see Supplementary Fig. 13 for the full version of this tree). Silhouettes of dinosaurs drawn by Scott Hartman, Mike Taylor, and Mathew Wedel, and available at Phylopic (phylopic.org) under a Creative Commons Attribution 3.0 Unported license. Global paleogeographic reconstructions from the Paleobiology Database (paleobiodb.org)
  


Xing Xu, Paul Upchurch, Philip D. Mannion, Paul M. Barrett, Omar R. Regalado-Fernandez, Jinyou Mo, Jinfu Ma and Hongan Liu. 2018. A New Middle Jurassic Diplodocoid Suggests An Earlier Dispersal and Diversification of Sauropod Dinosaurs. Nature Communications. 9, 2700.  DOI:  10.1038/s41467-018-05128-1

'Amazing Dragon' Fossil Upends Origins of World's Largest Dinosaurs on.natgeo.com/2JQPINr @NatGeoScience

  

Tuesday, July 24, 2018

[Herpetology • 2018] Cyrtodactylus chamba • A New Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from the Western Himalayas, Himachal Pradesh, India


Cyrtodactylus chamba 
Agarwal, Khandekar & Bauer, 2018


Abstract
We describe a new species of the gekkonid genus Cyrtodactylus based on a series of six specimens from the Chamba Valley in the Western Himalayas, Himachal Pradesh state, India. Cyrtodactylus chamba sp. nov. is a member of the subgenus Siwaligekko and can be diagnosed from congeners in the Western Himalayas by a combination of its small size (snout to vent length up to 63 mm), a continuous series of five precloacal pores on males, 13–15 rows of dorsal tubercles, 33–43 scales across the belly, no regular series of enlarged subcaudals, and a dorsal colour pattern of 5–7 irregular, broad, dark bands with much narrower, light interspaces. The new species is 14% divergent in ND2 sequence from the most similar sampled congener, Cyrtodactylus (Siwaligekkohimalayanus from Jammu and Kashmir, and is 0.5–1.1% divergent in nuclear sequence data from sampled Siwaligekko species. Many more undiscovered Cyrtodactylus species probably exist across the Himalayas at elevations below ~2000 m; basic field surveys for reptiles and other poorly known groups and examination of existing material should be a priority if we are to appreciate the true diversity of this spectacular mountainous landscape.

Keywords: Reptilia, Alpha taxonomy, Cyrtodactylus lawderanus, Himalayas, lizard, ND2, Siwaligekko, systematics, taxonomy

FIGURE 8. Type series of Cyrtodactylus chamba sp. nov. in life,
 BNHS 2330 (top left), BNHS 2331 (top right),
BNHS 2333 (bottom left), BNHS 2334 (bottom right).

Cyrtodactylus chamba sp. nov. 
Cyrtodactylus sp. Chamba (Agarwal et al. 2014a)

Etymology. The specific epithet is the name of the type locality of the species and is used as a noun in apposition. Cyrtodactylus chamba sp. nov. is only known from around Chamba, in the Chamba Valley, Himachal Pradesh, India. 
Suggested common name. Chamba bent-toed gecko



 Ishan Agarwal, Akshay Khandekar and Aaron M. Bauer. 2018. A New Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from the Western Himalayas, Himachal Pradesh, India. Zootaxa. 4446(4); 442–454. DOI:  10.11646/zootaxa.4446.4.2

Agarwal, I., Bauer, A.M., Jackman, T.R. & Karanth, K.P. 2014. Insights into Himalayan Biogeography from Geckos: A Molecular Phylogeny of Cyrtodactylus (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution. 80, 145–155. DOI:  10.1016/j.ympev.2014.07.018

     

[Herpetology • 2018] Phylogenomics of Montane Frogs of the Brazilian Atlantic Forest is Consistent with Isolation in Sky Islands Followed by Climatic Stability


Brachycephalus investigated in the present study. 
 
Brachycephalus brunneusB. izecksohni; B. fuscolineatus B. auroguttatus


in Pie, Faircloth, Ribeiro, et al., 2018.

Abstract
Despite encompassing a relatively small geographical area, montane regions harbour disproportionately high levels of species diversity and endemism. Nonetheless, relatively little is known about the evolutionary mechanisms that ultimately lead to montane diversity. In this study, we used target capture of ultraconserved elements to investigate the phylogenetic relationships and diversification patterns of Melanophryniscus (Bufonidae) and Brachycephalus (Brachycephalidae), two frog genera that occur in sky islands of the southern Atlantic Forest of Brazil. Specifically, we tested whether diversification of montane species in these genera could be explained by a single climatic shift leading to isolation in sky islands, followed by climatic stability that maintained populations in allopatry. In both genera, the topologies inferred using concatenation and coalescent-based methods were concordant and had strong nodal support, except for a few recent splits, which nevertheless tended to be supported by more informative loci. Estimation of divergence time of a combined dataset using both genera is consistent with a concordant timing of their diversification. These results support the scenario of diversification by isolation in sky islands and suggest that allopatry attributable to climatic gradients in montane regions is an important mechanism for generating species diversity and endemism in these regions.

Brachycephalus, coalescent, Melanophryniscus, target enrichment, ultraconserved elements

Figure 1. Examples of the species of Brachycephalus investigated in the present study.
E, Brachycephalus brunneus. F, Brachycephalus izecksohni. G, Brachycephalus fuscolineatus. H, Brachycephalus auroguttatus.
Photographs by L.F. Ribeiro.


Marcio R. Pie, Brant C. Faircloth, Luiz F. Ribeiro, Marcos R. Bornschein and John E Mccormack. 2018. Phylogenomics of Montane Frogs of the Brazilian Atlantic Forest is Consistent with Isolation in Sky Islands Followed by Climatic Stability. Biological Journal of the Linnean Society. bly093.   DOI: 10.1093/biolinnean/bly093   

Marcio R. Pie, Brant C Faircloth, Luiz Fernando Ribeiro, Marcos R. Bornschein and John McCormack. 2018. Phylogenomics of montane frogs of the Brazilian Atlantic Forest supports a scenario of isolation in sky islands followed by relative climatic stability. bioRxiv. 226159.  DOI: 10.1101/226159


Monday, July 23, 2018

[Ichthyology • 2018] Molecular Phylogenetic Analysis of the Catfish Species Auchenoglanis occidentalis (Valenciennes, 1840) (Pisces: Claroteidae) from Lake Turkana in East Africa: Taxonomic Iimplications.


Auchenoglanis occidentalis (Valenciennes, 1840)

 in Okwiri, Cao, Nyi & Zhang, 2018. 

Abstract
Cytochrome c oxidase subunit I (COI) gene sequences of two specimens here recognized as Auchenoglanis occidentalis from Lake Turkana in the Ethiopian section were determined. A COI gene-based phylogenetic analysis was performed for these along with sequences of African catfish species from the family Clarotidae available in GenBank. Based on results of this analysis, it is concluded that (1) the currently identified A. occidentalis is a species complex that includes several distinct species; (2) the Niger River basin harbors two distinct species of Auchenoglanis, one of which occurs in Lake Turkana, as well as A. biscutatus; and (3) A. sacchii is likely a valid species, but it is not the endemic species of Lake Turkana. It is suggested here that species diversity of Auchenoglanis requires further study based on molecular and morphological evidence.

Keywords: Pisces, Lake Turkana, Auchenoglanis occidentalis, COI, phylogenetic analysis, taxonomic implication


 Auchenoglanis occidentalis (Valenciennes, 1840)  


Brian Okwiri, Liang Cao, Dorothy Wanja Nyingi and E. Zhang. 2018. Molecular Phylogenetic Analysis of the Catfish Species Auchenoglanis occidentalis (Valenciennes, 1840) (Pisces: Claroteidae) from Lake Turkana in East Africa: Taxonomic Iimplications. Zootaxa. 4450(1); 115–124. DOI: 10.11646/zootaxa.4450.1.8

[Herpetology • 2018] Phrynopus mariellaleo • A New Species of Phrynopus (Anura: Craugastoridae) from the northeastern Andes of Peru, Its Phylogenetic Position, and Notes on the Relationships of Holoadeninae


Phrynopus mariellaleo 
Venegas, Barboza, De la Riva & Padial, 2018

 Photographs by Pablo J. Venegas.  facebook.com/CORBIDI

Abstract
We report the discovery of a geographically disjunct and morphologically distinctive species of direct-developing frog of the genus Phrynopus (Phrynopus mariellaleo sp. nov.) that changes considerably our understanding of the distribution of species in this Andean genus. The type locality lies on a subcordillera (Cerro de Campanario area) of the extreme northeastern portion of the Cordillera Central of Peru, on the headwaters of the Mayo River, Amazonas department, at 2575 m asl (6°6’42.9’’S, 77°26’24’’W). This area is situated 170 km to the NE from the northernmost record of Phrynopus known so far. Molecular phylogenetic analyses of a supermatrix (13269 aligned positions of gene sequences of four mitochondrial and ten nuclear genes) of 105 terminals (representing 93 named and 9 unnamed species of Holoadeninae) recover this new species as the sister to Phrynopus auriculatus, a species occurring more than 500 km south of the type locality of the new species. Both Phrynopus auriculatus and the new species occur at moderate elevations on the easternmost stretches of the Andean subcordilleras; their sister relationship point to a potentially broader distribution of species of Phrynopus along the poorly sampled intervening areas of the eastern hills of the Andes. The new species has a conspicuous and visibly large tympanic membrane (a trait rare in the clade), outlined by a marked bold black supratympanic fold and a black facial mask, and exhibits conspicuous dorsolateral, scapular, and middorsal Y-shaped folds. Specimens were found on the forest floor—a rocky substrate covered by a thick layer of leaf litter, moss and roots—of a primary humid montane forest (Yungas ecoregion) with scattered patches of bamboo (Chusquea spp.). Our phylogenetic analyses corroborate the monophyly of all Holoadeninae genera, including Euparkerella and Psychrophrynella, genera for which tests of monophyly were pending, and corroborates Hypodactylus nigrovittatus as part of Hypodactylus and sister to a clade that includes H. brunneus, H. elassodiscus and H. peraccai.

Keywords: Amphibia, Alto Mayo, Amazon Basin, Cordillera Central, dynamic homology, Terrarana, tree-alignment, Yungas

FIGURE 2. Dorsal (A) and ventral (B) views, and lateral view of head (C) of the female holotype (CORBIDI 11668) of Phrynopus mariellaleo sp. nov. in preservative (SVL = 39.7). Photographs by Pablo J. Venegas.

FIGURE 4. Paratypes of Phrynopus mariellaleo sp. nov. showing variation in dorsal and ventral external morphological traits.
(A, B) adult female (CORBIDI 11692), (C, D) adult female (CORBIDI 11657).

 Photographs by Pablo J. Venegas.   facebook.com/CORBIDI

Phrynopus mariellaleo sp. nov.

 Etymology. The specific name “mariellaleo” is a patronym (used as a substantive in apposition) for Mariella Leo, in recognition of her tireless efforts to preserve biological diversity in Peru. Since 1982 she has been working for the Asociación Peruana para la Conservación de la Naturaleza (APECO), one of the most important non-profit organizations dedicated to biological conservation in Peru. With APECO, Mariella continues to work for the protection of montane ecosystems in Amazonas Department, including the area where the new species was discovered.

FIGURE 4. Paratypes of Phrynopus mariellaleo sp. nov. showing variation in dorsal and ventral external morphological traits. (A, B) adult female (CORBIDI 11692), (C, D) adult female (CORBIDI 11657), and (E, F) adult male (CORBIDI 11658).
Photographs by Pablo J. Venegas.   facebook.com/CORBIDI


Pablo J. Venegas, Andy C. Barboza, Ignacio De la Riva and José M. Padial. 2018. A New Species of Phrynopus from the northeastern Andes of Peru, Its Phylogenetic Position, and Notes on the Relationships of Holoadeninae (Anura: Craugastoridae).  Zootaxa. 4446(4); 501–524.  DOI:  10.11646/zootaxa.4446.4.5


[Botany • 2018] Veronica ersin-yucelii (Plantaginaceae) • A New Veronica Species from Central Anatolia, Turkey


Veronica ersin-yucelii  

in Yaylacı, Sezer, Özgişi, Öztürk, Erkara, et al., 2018. 

Abstract
Veronica ersin-yucelii (subg. Pentasepalae, Plantaginaceae) is described and illustrated as a new species. It grows on marble rocks and screes, in one single site in Central Anatolia (Eskişehir Province), at elevations from about 1700 to 1820 metres above sea level. Diagnostic features are also given to enable comparison with the most similar species, V. caespitosa Boiss. and V. multifida L., especially regarding the indumentum, habit, inflorescence, bracts, seeds and leaves. The geographical distribution of the new species is mapped. Notes about its ecology and conservation status are also presented.

Keywords: Endemic, Eskişehir, threatened species




Ö. Koray Yaylacı, Okan Sezer, Kurtuluş Özgişi, Derviş Öztürk, İsmühan Potoğlu Erkara, Onur Koyuncu and Atila Ocak. 2018. A New Veronica (Plantaginaceae) Species from Central Anatolia, Turkey. Phytotaxa. 362(1); 55–67. DOI: 10.11646/phytotaxa.362.1.4

[Herpetology • 2018] Computational Molecular Species Delimitation and Taxonomic Revision of the Gecko Genus Ebenavia Boettger, 1878


Ebenavia safari
Hawlitschek, Scherz, Ruthensteiner, Crottini & Glaw, 2018


Abstract
Cryptic species have been detected in many groups of organisms and must be assumed to make up a significant portion of global biodiversity. We study geckos of the Ebenavia inunguis complex from Madagascar and surrounding islands and use species delimitation algorithms (GMYC, BOLD, BPP), COI barcode divergence, diagnostic codon indels in the nuclear marker PRLR, diagnostic categorical morphological characters, and significant differences in continuous morphological characters for its taxonomic revision. BPP yielded ≥ 10 operational taxonomic units, whereas GMYC (≥ 27) and BOLD (26) suggested substantial oversplitting. In consequnce, we resurrect Ebenavia boettgeri Boulenger 1885 and describe Ebenavia tuelinae sp. nov.Ebenavia safari sp. nov., and Ebenavia robusta sp. nov., increasing the number of recognised species in Ebenavia from two to six. Further lineages of Ebenavia retrieved by BPP may warrant species or subspecies status, but further taxonomic conclusions are postponed until more data become available. Finally, we present an identification key to the genus Ebenavia, provide an updated distribution map, and discuss the diagnostic values of computational species delimitation as well as morphological and molecular diagnostic characters.

Keywords: BOLD, Operational Taxonomic Unit, Madagascar clawless gecko, Integrative taxonomy, Taxonomic inflation, Species complex 


Genus Ebenavia Boettger, 1878 
Type species: Ebenavia inunguis Boettger, 1878


Diagnosis and description: A comprehensive diagnosis and description of the genus Ebenavia are provided in Nussbaum and Raxworthy (1998). Our data on the genus leads us to agree with the findings of that study, except that we were not able to confirm the presence of claws on the pes of any of the females we examined, as reported by these authors, in agreement with Boettger (1878) and many subsequent authors. We did not examine any specimens of E. maintimainty

Content: Ebenavia inunguis Boettger, 1878; E. boettgeri Boulenger, 1885; E. maintimainty Nussbaum & Raxworthy, 1998; and three new species described below. 

Distribution: Madagascar except most of the central and western areas; all major island areas of the Comoros Archipelago; Pemba Island (Tanzania); Mauritius.


Ebenavia inunguis Boettger, 1878 

Ebenavia boettgeri Boulenger, 1885, bona species 

Ebenavia tuelinae sp. nov. 

Etymology: A matronym dedicated to the first author’s partner Tülin (alternative spelling Tuelin) for her ceaseless support of this and other works and for her excellent spotting abilities in the field.


Ebenavia safari, a newly described species from northern Madagascar, which also occurs on the Tanzanian island of Pemba


Ebenavia safari sp. nov.

Etymology: Safari’ means ‘voyage’ in the Kiswahili and Comoran (Shimaoré) languages spoken across the range of this species outside Madagascar. The name was chosen because this species dispersed over surprisingly long distances across the open ocean. It is treated as an unlatinised, invariable noun in apposition.


Ebenavia robusta sp. nov. 

Etymology: The specific name is the feminine form of the Latin adjective ‘robustus’, meaning ‘robust’ or ‘sturdy’. It was chosen because this species is the largest and most sturdily built member of this genus of small geckos.


Oliver Hawlitschek, Mark D. Scherz, Bernhard Ruthensteiner, Angelica Crottini and Frank Glaw. 2018. Computational Molecular Species Delimitation and Taxonomic Revision of the Gecko Genus Ebenavia Boettger, 1878The Science of Nature. 105:49. DOI: 10.1007/s00114-018-1574-9 

Saturday, July 21, 2018

[Herpetology • 2018] Boana icamiaba • Phylogeny of Map Tree Frogs, Boana semilineata Species Group (Anura: Hylidae), with A New Amazonian Species


Boana icamiaba
Peloso, Oliveira, Sturaro, Rodrigues, Lima-Filho, Bitar, Wheeler & Aleixo, 2018


Abstract.
Gladiator Frogs (Boana) is a Neotropical group comprised of 92 species sorted into seven species groups. Herein, we present a phylogeny of the Boana semilineata species group, including all valid nominal species currently or suspected to be assigned to it— many sequenced for the first time. Parsimony and maximum likelihood analyses of two genes (16S and COI) did not support the monophyly of the B. semilineata group as currently defined. To remedy this, B. secedens, currently assigned to the B. puchella species group, is transferred to the B. semilineata group. On the other hand, our analyses corroborate the inclusion of B. hutchinsi, B. pombali and B. wavrini in the B. semilineata group. The reinterpretation of the morphology of the lower palpebral membrane in B. pombali (previously described as absent, but actually present) corroborates that the reticulated membrane is a synapomorphy of this species group (albeit homoplastic—present in a few species not assigned to this group). A recent paper suggested (based on the analyses of 16S alone) that at least six unnamed taxa (candidate species) are hidden behind the names B. geographica and B. semilineata. Our analyses resulted in the discovery of additional unnamed lineages, one of which is described and named herein. This new species is diagnosed, among other characters, by having the flanks, anterior and posterior surfaces of thigh with dark brown or blackish stripes, and by the presence of a prepollex forming a projecting spine.

Keywords: Amazonia, Biodiversity, Cophomantinae, Hyla geographica, Taxonomy


  Boana icamiaba sp. nov. in life, paratype (MNRJ 90998) from Altamira, Pará, Brazil.

Photograph by Renan Oliveira.

Boana icamiaba sp. nov.

Etymology: A noun in apposition, given as homage in memory of the legendary Amazonian female warriors, the Icamiabas. These warriors, first described by friar Gaspar de Carvajal (a member of the Francisco Orellana expedition; Medina, 1894), were members of isolated tribes composed of women only. So far as we are aware this is the second taxon to honor to the icamiabas, the other one being a hooded tickspider, Cryptocellus icamiabas Tourinho and Azevedo, 2007.



Pedro L.V. Peloso, Renan M. De Oliveira, Marcelo J. Sturaro, Miguel T. Rodrigues, Geraldo R. Lima-Filho, Youszef O.C. Bitar, Ward C. Wheeler and Alexandre Aleixo. 2018. Phylogeny of Map Tree Frogs, Boana semilineata Species Group, with a New Amazonian Species (Anura: Hylidae). South American Journal of Herpetology. 13(2); 150-169. DOI:  10.2994/SAJH-D-17-00037.1

New Frog Species Named After Fabled Female Warriors 
 on.NatGeo.com/2muqL1k via @NatGeo

[Paleontology • 2018] Akainacephalus johnsoni • A New southern Laramidian Ankylosaurid from the upper Campanian Kaiparowits Formation of southern Utah, USA


Akainacephalus johnsoni 
Wiersma​ & Irmis, 2018 

 DOI: 10.7717/peerj.5016



Abstract

A partial ankylosaurid skeleton from the upper Campanian Kaiparowits Formation of southern Utah is recognized as a new taxonAkainacephalus johnsoni, gen. et sp. nov. The new taxon documents the first record of an associated ankylosaurid skull and postcranial skeleton from the Kaiparowits Formation. Preserved material includes a complete skull, much of the vertebral column, including a complete tail club, a nearly complete synsacrum, several fore- and hind limb elements, and a suite of postcranial osteoderms, making Akainacephalus johnsoni the most complete ankylosaurid from the Late Cretaceous of southern Laramidia. Arrangement and morphology of cranial ornamentation in Akainacephalus johnsoni is strikingly similar to Nodocephalosaurus kirtlandensis and some Asian ankylosaurids (e.g., Saichania chulsanensis, Pinacosaurus grangeri, and Minotaurasaurus ramachandrani); the cranium is densely ornamented with symmetrically arranged and distinctly raised ossified caputegulae which are predominantly distributed across the dorsal and dorsolateral regions of the nasals, frontals, and orbitals. Cranial caputegulae display smooth surface textures with minor pitting and possess a distinct conical to pyramidal morphology which terminates in a sharp apex. Character analysis suggests a close phylogenetic relationship with N. kirtlandensis, M. ramachandrani, Tarchia teresae, and S. chulsanensis, rather than with Late Cretaceous northern Laramidian ankylosaurids (e.g., Euoplocephalus tutus, Anodontosaurus lambei, and Ankylosaurus magniventris). These new data are consistent with evidence for distinct northern and southern biogeographic provinces in Laramidia during the late Campanian. The addition of this new ankylosaurid taxon from southern Utah enhances our understanding of ankylosaurid diversity and evolutionary relationships. Potential implications for the geographical distribution of Late Cretaceous ankylosaurid dinosaurs throughout the Western Interior suggest multiple time-transgressive biogeographic dispersal events from Asia into Laramidia.

Life reconstruction: Andrey Atuchin 


Systematic paleontology

Dinosauria Owen, 1842 sensu Padian and May, 1993
Ornithischia Seeley, 1887 sensu Padian and May, 1993
Thyreophora Nopcsa, 1915 sensu Sereno, 1986

Ankylosauria Osborn, 1923 sensu Carpenter, 1997
Ankylosauridae Brown, 1908 sensu Sereno, 1998
Ankylosaurinae Brown, 1908 sensu Sereno, 1986
Ankylosaurini Arbour and Currie, 2016

Akainacephalus, gen. nov.

Akainacephalus johnsoni, sp. nov.

Figure 4: Skull of Akainacephalus johnsoni (UMNH VP 20202).
Photographs of the skull of Akainacephalus johnsoni in (A), dorsal; and (B), ventral views. Line drawings in (C), dorsal; and (D), ventral views highlight major anatomical features.


Study sites: bpt, basipterygoid; bs, basisphenoid; ch, choana; exo, exoccipital; fm, foramen magnum; fca, frontal caputegulum; ins, internarial septum; laca, lacrimal caputegulum; loca, loreal caputegulum; mx, maxilla; mxtr, maxillary tooth row; naca, nasal caputegulum; ns, nuchal shelf; oc, occipital condyle; pal, palatine; prfca, prefrontal caputegulum; pmx, premaxilla; pmxs, interpremaxillry suture with oblong depression; pop, paroccipital process; ptv, pterygoid vacuity; q, quadrate; qj, quadratojugal; qjh, quadratojuga horn; so, supra occipital; snca, supranarial caputegulum; sob, supraorbital boss; sqh, squamosal horn.

Figure 3: Skull of Akainacephalus johnsoni (UMNH VP 20202). Photographs of the skull of Akainacephalus johnsoni in (A), left lateral; and (B), right lateral views. Line drawings in (C), left lateral; and (D), right lateral views highlight major anatomical features.

Study sites: en, external naris; fca, frontal caputegulum; j, jugal; jca, jugal caputegulum; l, lacrimal; laca, lacrimal caputegulum; loca, loreal caputegulum; mx, maxilla; n, nasal; naca, nasal caputegulae; ns, nuchal shelf; orb, orbit; pmx, premaxilla; prfca, prefrontal caputegulum; snca, supranarial caputegulum; sob, supraorbital boss; q, quadrate; qjh, quadratojugal horn; sqh, squamosal horn. 

Holotype: UMNH VP 20202, a partial skeleton comprising a complete skull, both mandibles, predentary, four dorsal, four dorsosacral, three sacral, one caudosacral, and eight caudal vertebrae, dorsal ribs, a complete tail club, both scapulae, left coracoid, right humerus, right ulna, partial left ilium, left femur, left tibia, left fibula, phalanx, two partial cervical osteoderm half rings, and 17 dorsal and lateral osteoderms of various sizes and morphologies.

Type locality: UMNH VP Locality 1109 (“HMG Quarry”), Horse Mountain area, GSENM, Kane County, southern Utah, USA.

Type stratigraphic horizon and age: UMNH VP Locality 1109 is a multitaxic bonebed deposited in a crevasse splay sandstone within the lower portion of the middle unit of the upper Campanian Kaiparowits Formation (Fig. 2A). The stratigraphic position of this site is approximately 190 m from the base of the formation (Roberts et al., 2013: fig. 6.3) and within approximately one meter stratigraphic proximity of the recently dated bentonite ash bed KP-07, which has produced a U-Pb zircon age of 76.26 ± 0.10 Ma (Roberts et al., 2013), providing a precise age constraint for Akainacephalus johnsoni.

Etymology: The genus name is derived from the Greek akaina, meaning “thorn” or “spine,” referring to the thorn-like cranial caputegulae of the holotype; and “cephalus,” the Greek meaning for head. The specific epithet honors Randy Johnson, volunteer preparator at the Natural History Museum of Utah, who skillfully prepared the skull and lower jaws of UMNH VP 20202.

Diagnosis: Akainacephalus johnsoni possesses the following autapomorphies: massive supraorbital bosses in lateral view, forming a tall backswept flange extending laterally over the orbits, and enveloping the anterodorsal and posterior margins of the orbit; nearly vertical projecting triangular quadratojugal horns; frontal possesses a large, flat, and centrally positioned hexagonal-shaped caputegulum; a combination of tightly spaced, symmetrically positioned pyramidal and conical-shaped caputegulae across the frontonasal region; a distinct midline row of conical-shaped caputegulae across the nasal region, symmetrically separating caputegulae situated dorsolaterally; basioccipital foramen anterior and dorsally to the occipital condyle. A. johnsoni also possesses a unique combination of character states: shares with Nodocephalosaurus kirtlandensis the presence of a large, laterally oriented supranarial osteoderm forming the postmaxillary/lacrimal ridge dorsal to the external nares; differs from Tsagantegia longicranialis, Talarurus plicatospineus, Pinacosaurus grangeri, all northern Laramidian taxa and Ziapelta sanjuanensis but shares with Nodocephalosaurus kirtlandensis, Minotaurasaurus ramachandrani, Saichania chulsanensis, and Tarchia kielanae the presence of well-pronounced cranial ornamentation located along the nasal and frontal regions of the skull that are characterized by a dense array of well-defined caputegulae with a distinct conical (N. kirtlandensis) and pyramidal (M. ramachandrani, S. chulsanensis, T. kilanae) morphology; shares with Euoplocephalus and Zuul crurivastator a globular surface texture on the tail club knob, which differs from the smoother texture in Ankylosaurus magniventris; differs from ZPAL MgD I/113, cf. Pinacosaurus, Saichania chulsanensis, and Dyoplosaurus acutosquameus, but similar to Anodontosaurus lambei, Euoplocephalus tutus, Zuul crurivastator, and Ankylosaurus magniventris in having a wider than long tail club knob ratio; and shares with ZPAL MgD I/113, cf. Pinacosaurus, D. acutosquameus, and Zuul crurivastator triangular osteoderms along the lateral surfaces on the proximal portion of the tail.


Figure 7: Variation in cranial ornamentation in selected Laramidian and Asian taxa, including Akainacephalus johnsoni.

Comparative line drawings highlighting major areas of cranial ornamentation in Akainacephalus johnsoni and closely related Laramidian and Asian taxa. Akainacephalus johnsoni (UMNH VP 20202) in (A), dorsal; (B), left lateral view compared to Nodocephalosaurus kirtlandensis (SMP VP-900) in (C), dorsal; and (D) left lateral view; Tarchia teresae (PIN 3142/250) in (E) dorsal; (F), left lateral view and Minotaurasaurus ramachandrani (INBR 21004) in (G), dorsal; and (H), left lateral view.

Study sites: acc po, accessory postorbital ossification; asob, anterior supraorbital boss; frca, frontal caputegulum; laca, lacrimal caputegulum; loca, loreal caputegulum; mso, medial supraorbital; mx, maxilla; n, external naris; naca, nasal caputegulae; nuca, nuchal caputegulae; orb, orbital; pmx, premxilla; pnca, postnarial caputegulum; pos postocular ossicles; prfca, prefrontal caputegulum; psob, posterior supraorbital boss; pt, pterygoid; q, quadrate; qjh, quadratojugal horn; snca, supranarial caputegulum; sqh, squamosal horn. 
Color scheme after Arbour & Currie (2013a). Dorsal view of N. kirtlandensis modified after Arbour et al. (2014). T. teresea (=Saichania chulsanensis in Arbour, Currie & Badamgarav, 2014) and M. ramachandrani modified after Arbour, Currie & Badamgarav (2014).


Figure 28: Preserved elements and skeletal reconstructions of Akainacephalus johnsoni.
A composite showing all holotype skeletal material of Akainacephalus johnsoni (UMNH VP 20202) anatomically arranged in dorsal view (A). Cartoon illustrating a full body reconstruction for A. johnsoni in (B), dorsal; and (C), left lateral view. Preserved material in the skeletal reconstructions is highlighted in orange.

  

Conclusion:
Akainacephalus johnsoni (UMNH VP 20202) is a new taxon of ankylosaurid dinosaur from the upper Campanian Kaiparowits Formation of southern Utah, USA. It consists of a complete cranium and significant amount of diagnostic postcranial material that can be distinguished from all other known Late Cretaceous Laramidian ankylosaurids. Akainacephalus johnsoni is closely related to its stratigraphically younger sister taxon, Nodocephalosaurus kirtlandensis from the Kirtland Formation of New Mexico. Both taxa are more closely related to Asian ankylosaurids than they are to other Laramidian ankylosaurids. This suggests multiple ankylosaurid dispersal events from Asia to Laramidia during the Late Cretaceous. Lastly, together with Dyoplosaurus acutossquameus and Scolosaurus cutleri (∼77 Ma) from northern Laramidia, Akainacephalus johnsoni represents one of the older known ankylosaurid dinosaurs (∼76.3 Ma) from the Late Cretaceous of western North America.



Jelle P. Wiersma​ and Randall B. Irmis. 2018. A New southern Laramidian Ankylosaurid, Akainacephalus johnsoni gen. et sp. nov., from the upper Campanian Kaiparowits Formation of southern Utah, USA. PeerJ. 6:e5016. DOI: 10.7717/peerj.5016

Newly discovered armored dinosaur from Utah reveals intriguing family history phys.org/news/2018-07-newly-armored-dinosaur-utah-reveals.html via @physorg_com
Mystery of Spiky-Skulled Utah Dinosaur Solved  livescience.com/63108-newfound-ankylosaur-has-asian-roots.html via @LiveScience