Bohaiornithid
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Bohaiornithidae is a group of early
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 ...
stem-birds from the early
Cretaceous Period The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 77.1 million years, it is the ninth and longest geologi ...
of
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 ...
. All undisputed Bohaiornithidae specimens come from the northeast of present-day
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 ...
– specifically, the eastern and southern foothills of the
Greater Khingan The Greater Khingan Range or Da Hinggan Range ( zh, s=大兴安岭, t=大興安嶺, p=Dà Xīng'ān Lǐng; IPA: ) is a volcanic mountain range in the Inner Mongolia region of Northeast China. It was originally called the Xianbei Mountains, whi ...
mountains –, and date to the early
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 ...
age, roughly 125–120 million years ago. As of 2024 they are not certainly known from outside this limited region and timeframe, even though there and then they were a prominent – and probably
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
y, unlike the bulk of their relatives – element of the local fauna, namely the
Jehol Biota The Jehol Biota includes all the living organisms – the ecosystem – of northeastern China between 133 and 120 million years ago. This is the Lower Cretaceous ecosystem which left fossils in the Yixian Formation and Jiufotang Formation. These d ...
. The
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
Bohaiornithidae was established by Wang and colleagues in 2014. They defined it as the natural group formed by all descendants of the common ancestor of 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 ...
, '' Bohaiornis guoi'', and '' Shenqiornis mengi''.Wang M., Zhou Z.-H., O’Connor, J.K., and Zelenkov, N.V. (2014
A new diverse enantiornithine family (Bohaiornithidae fam. nov.) from the Lower Cretaceous of China with information from two new species
''Vertebrata PalAsiatica'', 52(1): 31-76.


Description

Similar to most enantiornitheans, bohaiornithids possessed teeth rather than a beak as in modern birds, although they could be distinguished from other enantiornitheans due to the structure of their teeth. Their teeth were large, robust and somewhat conical, but had sharply pointed tips which curved backwards. The first few teeth of the premaxillae are smaller than the rest of the teeth, but the other teeth in the front of the mouth were larger than those in the back. Bohaiornithids also had lateral trabeculae (a pair of long and thin bony projections on the rear edge of the
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
) which not only extended backwards, but also outwards. The tips of each branch of the
furcula The (Latin for "little fork"; : furculae) or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is either an interclavicle or formed by the fusion of the two clavicles. In birds, its primary function is ...
(wishbone) are wide and rounded in bohaiornithids, as opposed to the tapering tips in other enantiornitheans. Their
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
e (shoulder blades) slightly curve downwards, created a convex top edge and concave lower edge. Bohaiornithids also had gradually tapering
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 ...
s. A bohaiornithid's second (innermost) toes were thicker than their other toes, while their third (middle) toes were long and thin and all of their toe claws were very long and curved. Most adult bohaiornithids would have been rather similar in size and appearance to each other, and the family lacked the longevity and physical diversity of some other enantiornithean groups, such as the long-snouted
longipterygids Longipterygidae is a Family (biology), family of early enantiornithean avialans from the Early Cretaceous epoch of China. All known specimens come from the Jiufotang Formation and Yixian Formation, dating to the early Aptian age, 125-120 million ...
. The only specimens of bohaiornithids believed to have reached adulthood (the holotype of '' Zhouornis'' and a referred specimen of ''
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 ...
'') were also the largest specimens, and were about the size of a
pigeon Columbidae is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with small heads, relatively short necks and slender bills that in some species feature fleshy ceres. ...
, which is much larger than most other Jehol enantiornitheans with the exception of ''
Pengornis ''Pengornis'' is the largest known enantiornithean bird from the Early Cretaceous of northeast China. The name derives from "Peng", which refers to a mythological bird from Chinese folklore, and "-ornis", which means bird in Greek. ''Pengorni ...
'' and '' Xiangornis''. The smallest and youngest bohaiornithid specimens were about half the size of the largest ones. Many bohaiornithids have been found preserving
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and an exa ...
s, and a few possessed a pair of long, ribbon-like tail feathers with barbs only at the tips. These specialized feathers are known in many enantiornithean specimens (as well as ''
Confuciusornis ''Confuciusornis'' is a genus of basal crow-sized avialan from the Early Cretaceous Period of the Yixian and Jiufotang Formations of China, dating from 125 to 120 million years ago. Like modern birds, ''Confuciusornis'' had a toothless beak, bu ...
''), and may be an example of
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, with the feathers being used by male members of a species for courtship display. A subadult specimen of an indeterminate bohaiornithid described in 2017 preserved feathers on various parts of the head and body which were analyzed and determined to have been
iridescent Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear gradually to change colour as the angle of view or the angle of illumination changes. Iridescence is caused by wave interference of light in microstruc ...
in life.


Paleoecology

A referred adult specimen of ''Bohaiornis'' was preserved with several rocks concentrated in the stomach region. Rocks in the stomach of birds and other animals are often believed to be
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In ...
s swallowed to assist in the digestion of plant material. However, the rocks apparently swallowed by this specimen are much larger, coarser, and less numerous than most gastroliths, and were unlikely to have been used for the same purpose as those swallowed by herbivores. Instead, it has been suggested that they were swallowed to help clean the digestive tract, a usage which has been reported in living birds of prey as well as other carnivorous animals with high-fat diets, such as
pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammals. They comprise the extant taxon, extant families Odobenidae (whose onl ...
s. The description of this specimen claimed that it was plausible, but uncertain, that ''Bohaiornis'' was ecologically similar to birds of prey. However, these rocks were later found to be mineral concretions, probably formed from the same mineral as the fossil specimen. Thus, they are not indicative of diet. Bohaiornithids had foot proportions which were between those expected of arboreal and terrestrial birds, but their long claws make terrestrial habits unlikely. Nevertheless, bohaiornithids lacked many of the specialized foot features which allow modern birds to perch and climb tree trunks. Although long claws are known in modern birds of prey, bohaiornithids had short tarsometatarsals (ankle bones) which were dissimilar to most raptors with the exception of members of the subfamily
Perninae The raptor subfamily Perninae includes a number of medium-sized broad-winged species. These are birds of warmer climates, although the ''Pernis'' species (European honey buzzard and crested honey buzzard) have a more extensive range. Several of ...
. Among birds of prey, bohaiornithids had leg proportions most similar to
osprey The osprey (; ''Pandion haliaetus''), historically known as sea hawk, river hawk, and fish hawk, is a diurnal, fish-eating bird of prey with a cosmopolitan range. It is a large raptor, reaching more than in length and a wingspan of . It ...
s, and thus it is conceivable that they were
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
. However, their teeth suggest a diet of hard-shelled creatures.


Classification

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 was found in the phylogenetic analysis of Wang ''et al.'' 2014, which itself was an updated version of O’Connor ''et al.'' 2013.O’Connor, J.K. Zhang, Y. Chiappe, L.M. Meng, Q. Quanguo, L. & Di, L. (2013
A new enantiornithine from the Yixian Formation with the first recognized avian enamel specialization
''Journal of Vertebrate Paleontology'' 33(1):1-12.
The cladogram is based on 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. The Wang ''et al.'' phylogenetic analysis was reused during the description of '' Linyiornis'' in 2016. This study it indicated that previously known bohaiornithid taxa formed a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
with ''Linyiornis'' and '' Fortunguavis'', supporting the possibility that those two genera are also examples of the family, although ''Fortunguavis'' is notoriously difficult to place phylogenetically.


References

{{Taxonbar, from=Q15644506 Dinosaur families Aptian dinosaurs Dinosaurs of China