''Longusunguis'' is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of bohaiornithid

enantiornithean The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and cl ...

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

known from the

Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...

Liaoning Province ) , image_skyline = , image_alt = , image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong , image_map = Liaoning in China (+all claims hatched).svg , ...

, northeastern

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

. It contains a single species, ''Longusunguis kurochkini''.

Discovery

''Longusunguis'' was first described and named by Min Wang, Zhong-He Zhou, Jingmai K. O'Connor and Nikita V. Zelenkov in

2014 The year 2014 was marked by the surge of the Western African Ebola epidemic, West African Ebola epidemic, which began in 2013, becoming the List of Ebola outbreaks, most widespread outbreak of the Ebola, Ebola virus in human history, resul ...

and the

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

is ''Longusunguis kurochkini''. The generic name is derived from the

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

words ''longus'', meaning "long", and ''unguis'', meaning "claw", in reference to the distinctly elongated pedal unguals that this taxon shares with other bohaiornithids. The

specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...

, ''kurochkini'', honors the late Prof.

Evgeny Kurochkin Yevgeny Nikolayevich Kurochkin (Евгений Николаевич Курочкин; 12 July 1940 - 13 December 2011) was a Russian paleornithologist at the Paleontological Institute of the Russian Academy of Sciences. He served as President of ...

, a prominent

paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...

, for his contributions to the study of

fossil birds Birds evolved from certain feathered dinosaurs, feathered theropod dinosaurs, and there is no real dividing line between birds and non-avian dinosaurs except that some of the former survived the Cretaceous–Paleogene extinction event while the ...

. ''Longusunguis'' is known solely from the

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

IVPP V 17964, a complete and articulated

skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...

and

skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...

. It is preserved in a single slab and currently housed at the

Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of People's Republic of China, China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian For ...

Beijing Beijing, Chinese postal romanization, previously romanized as Peking, is the capital city of China. With more than 22 million residents, it is the world's List of national capitals by population, most populous national capital city as well as ...

. IVPP V 17964 was collected at deposits near the Lamadong Town,

Jianchang County Jianchang () is a county of Huludao City in the southwest of Liaoning province, China. It is the largest division of Huludao, with an area of , and population of 450,000 in 2020, located in mountainous terrain west of that city, serviced by Chi ...

of the

, from the

Jiufotang Formation The Jiufotang Formation () is an Early Cretaceous geological formation in Chaoyang, Liaoning which has yielded fossils of feathered dinosaurs, primitive birds, pterosaurs, and other organisms (see Jehol Biota). It is a member of the Jehol group. T ...

, dated to the

Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...

stage of the

, approximately 120.3

million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...

Description

''Longusunguis'' differs from all other known bohaiornithid

s by possessing a combination of traits, some of which are

autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...

, i.e. unique. The

maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...

of ''Longusunguis'' possesses an accessory

fenestra A fenestra (fenestration; : fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biology, biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomy, ...

on the jugal process. This trait is unique among bohaiornithids, as only ''Zhouornis'' possesses such fenestra, however on the dorsal process. Its

lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...

shows an elongate excavation on the caudal margin of its descending ramus. In ''Longusunguis'', the lateral margin of the

coracoid A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...

is more convex than in other bohaiornithids. In ''Longusunguis'', the

acromion In human anatomy, the acromion (from Greek: ''akros'', "highest", ''ōmos'', "shoulder", : acromia) or summit of the shoulder is a bony process on the scapula (shoulder blade). Together with the coracoid process, it extends laterally over the sh ...

process is strongly projecting dorsally (more than in '' Sulcavis''), whereas in '' Parabohaiornis'' this process is straight, and is parallel to the scapular shaft. Finally, its

pygostyle Pygostyle is a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main component o ...

is longer than the

tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bird bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) a ...

. The holotype of ''Longusunguis'' came from a subadult individual, as evident by the absence of fusion in some compound bones, such as the

carpometacarpus The carpometacarpus is a bone found in the hands of birds. It results from the fusion of the carpal and metacarpal bone, and is essentially a single fused bone between the wrist and the knuckles. It is a smallish bone in most birds, generally fla ...

tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These ...

and

Phylogeny

The

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

position of ''Longusunguis'' was explored by Wang ''et al.'' (2014) using a data matrix that includes 56

avialan Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative defi ...

taxa, scored based on 262 morphological traits. ''Longusunguis'' was recovered as an advanced bohaiornithid, closely related to ''

Bohaiornis ''Bohaiornis'' is a genus of enantiornithean dinosaurs. Fossils have been found from the Lower Cretaceous Jiufotang Formation of western Liaoning, China. The only known species, ''Bohaiornis guoi'', was named by Dongyu Hu, Li Li, Lianhaim Hou an ...

'' and '' Parabohaiornis'' whose specimens were collected at nearby localities, also near the Lamadong Town. The

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

below shows the phylogenetic position of ''Longusunguis'' among the Enantiornithes following this analysis.

References

{{Taxonbar, from=Q15644509 Bohaiornithidae Bird genera Aptian birds Jiufotang Formation Fossil taxa described in 2014 Birds of Asia