Iyuku
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''Iyuku'' (meaning "hatchling", pronounced "eye-yoo-koo") is a genus of
iguanodontian Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include ''Camptosaurus'', ''Dryosaurus'', ''Iguanodon'', ''Tenontosaurus'', and the hadrosaurids or "duck-bill ...
dinosaur from the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
Kirkwood Formation The Kirkwood Formation is a geological formation found in the Eastern and Western Cape provinces in South Africa. It is one of the four formations found within the Uitenhage Group of the Algoa Basin – its type locality – and in the neighbouri ...
of South Africa. The type species is ''Iyuku raathi''.


Discovery and naming

''Iykuku'' is known from a bonebed mostly containing remains of at least 27 (inferred from non-overlapping parts of left
femora The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with t ...
) juvenile and hatchling individuals, an occurrence never before reported. These bonebeds were excavated during three expeditions between 1995 and 1999. The remains were known as the "Kirkwood taxon" since at least 2012, until the remains were determined to represent a new genus and species in 2022, named ''Iyuku raathi''. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
specimen, AM 6067, consists of an incomplete, semi-articulated skeleton including a partial skull, vertebrae, scapulae, pelvic girdle, both legs, and ribs. The generic name, "''Iyuku''", is derived from the
Xhosa Xhosa may refer to: * Xhosa people, a nation, and ethnic group, who live in south-central and southeasterly region of South Africa * Xhosa language, one of the 11 official languages of South Africa, principally spoken by the Xhosa people See als ...
word for "hatchling", in reference to the immature status of the specimens. The specific name, "''raathi''", honors South African paleontologist Mike Raath.


Classification

The describers assigned ''Iyuku'' to the Iguanodontia. A 2015 PhD dissertation that included the then unnamed species in a
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis found it to be a member of the basal
iguanodontian Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include ''Camptosaurus'', ''Dryosaurus'', ''Iguanodon'', ''Tenontosaurus'', and the hadrosaurids or "duck-bill ...
family
Dryosauridae Dryosaurids were primitive iguanodonts. They are known from Middle Jurassic to Early Cretaceous rocks of Africa, Europe, and North America. Phylogeny Until recently many dryosaurids have been regarded as dubious ('' Callovosaurus'' and '' Kan ...
, with the parsimony analysis recovering ''Iyuku'' and ''
Dysalotosaurus ''Dysalotosaurus'' ("uncatchable lizard") is a genus of herbivorous iguanodontian dinosaur. It was a dryosaurid iguanodontian, and its fossils have been found in late Kimmeridgian-age rocks (Late Jurassic) of the Tendaguru Formation of Lindi Regi ...
'' as
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
. Forster ''et al.'' (2022) referenced this study but noted that the early ontogenetic stage of most of the material of ''Iyuku'' presents a major problem in placing it into a phylogenetic analysis, and assigned to Iguanodontia without providing an analysis. Subsequently, four phylogenetic analyses conducted by Poole (2022) recovered ''Iyuku'' as the sister taxon to ''Dysalotosaurus'' within Dryosauridae. The cladogram below displays the results of a
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
from Poole (2022):


Paleobiology

Fossil material of ''Iyuku'' represents at least four size classes, including three juvenile phases of growth (with the smallest likely representing the post-hatching stage) and the fourth, larger stage, likely a subadult. Two different estimates of total length were provided for an incomplete
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
of a subadult specimen collected from an unknown location in the Kirkwood cliffs: either ~ or at least long, i.e. four or five times the length of the largest juvenile tibia from the quarry site. Fossil material from the quarry site includes mostly isolated bones (with only five instances of articulation) of hatchling to juvenile individuals of varying size in a 20–30 cm thick horizon, preserved with no apparent preferred orientation of long bones. Forster ''et al.'' (2022) noted the similarity of this finding to the reported preservation of the juvenile remains of ''
Orodromeus ''Orodromeus'' (meaning "Mountain Runner") is a genus of herbivorous orodromine thescelosaurid dinosaur from the Late Cretaceous of North America. Only one species is known, the type species ''Orodromeus makelai''. Discovery and naming The rem ...
'', ''
Maiasaura ''Maiasaura'' (from the Greek ''μαῖα'', meaning "good mother" and ''σαύρα'', the feminine form of ''saurus'', meaning "reptile") is a large herbivorous saurolophine hadrosaurid ("duck-billed") dinosaur genus that lived in the area curre ...
'' and ''
Hypacrosaurus ''Hypacrosaurus'' (meaning "near the highest lizard" reek υπο-, ''hypo-'' = less + ακρος, ''akros'', high because it was almost but not quite as large as ''Tyrannosaurus'') was a genus of duckbill dinosaur similar in appearance to ''Co ...
'' which might represent aggradations of mortality near nesting sites. Although remains of ''Iyuku'' were not found in association with nests or eggshells, Forster ''et al.'' (2022) considered it possible that the concentrations of young individuals of this species represent seasonal attrition at or near a nesting site, with the scattering of bones likely caused by trampling prior to burial or
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a pr ...
shortly after interment. Bone
histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
of multiple young individuals reveals rest lines indicative of temporary arrests in bone deposition, hypothesized by Forster ''et al.'' (2022) as caused by repeated, possibly environmentally induced stress during the early stages of ontogeny in these individuals, such as dry and drought-like conditions and/or repeated periods of extreme heat.


Paleoenvironment

Sedimentological analysis of the holotype assemblage suggests it came from a horizon transitioning into a drier climate. Other animals from the Kirkwood Formation include the sauropod ''
Algoasaurus ''Algoasaurus'' (; "Algoa Bay reptile") is a genus of sauropod dinosaur from the Berriasian- early Valanginian-age Early Cretaceous Upper Kirkwood Formation of Cape Province, South Africa, specifically near a town called Despatch. Discovery an ...
'', the stegosaur ''
Paranthodon ''Paranthodon'' ( ) is a genus of stegosaurian dinosaur that lived in what is now South Africa during the Early Cretaceous, between 139 and 131 million years ago. Discovered in 1845, it was one of the first stegosaurians found. Its only rem ...
'', the ornithomimosaur ''
Nqwebasaurus ''Nqwebasaurus'' (; anglicized as or ) is a basal coelurosaur and is the basal-most member of the coelurosaurian clade Ornithomimosauria from the Early Cretaceous of South Africa. The name ''Nqwebasaurus'' is derived from the Xhosa word "Nqweb ...
'', and several other unnamed dinosaurs.


References

{{Ornithopoda, O. Iguanodonts Valanginian life Early Cretaceous dinosaurs of Africa Cretaceous South Africa Fossils of South Africa Fossil taxa described in 2022 Ornithischian genera