, 2005. Although a minor element of modern marine
fauna, agnathans were prominent among the early fish in the early
Paleozoic. Two types of Early
Cambrian animal apparently having fins,
vertebrate musculature, and gills are known from the early Cambrian
Maotianshan shales of
China:
Haikouichthys and
Myllokunmingia. They have been tentatively assigned to Agnatha by Janvier. A third possible agnathan from the same region is
Haikouella. A possible agnathan that has not been formally described was reported by Simonetti from the Middle Cambrian
Burgess Shale of
British Columbia.
Conodonts, a class of agnathans which arose in the early Cambrian, remained common enough until their extinction in the
Triassic that their teeth (the only parts of them that were usually fossilized)
are often used as
index fossils from the late Cambrian to the Triassic. Many Ordovician, Silurian, and Devonian agnathans were armored with heavy bony-spiky plates. The first armored agnathans—the
ostracoderms, precursors to the
bony fish and hence to the
tetrapods (including
humans)—are known from the middle
Ordovician, and by the Late
Silurian the agnathans had reached the high point of their evolution. Most of the ostracoderms, such as
thelodonts,
osteostracans, and
galeaspids, were more closely related to the gnathostomes than to the surviving agnathans, known as cyclostomes. Cyclostomes apparently split from other agnathans before the evolution of dentine and bone, which are present in many fossil agnathans, including
conodonts. Agnathans declined in the
Devonian and never recovered. Approximately 500 million years ago, two types of recombinatorial adaptive immune systems (AISs) arose in vertebrates. The jawed vertebrates diversify their repertoire of immunoglobulin domain-based T and B cell antigen receptors mainly through the rearrangement of V(D)J gene segments and somatic hypermutation, but none of the fundamental AIS recognition elements in jawed vertebrates have been found in jawless vertebrates. Instead, the AIS of jawless vertebrates is based on
variable lymphocyte receptors (VLRs) that are generated through recombinatorial usage of a large panel of highly diverse leucine-rich-repeat (LRR) sequences. Three VLR genes (VLRA, VLRB, and VLRC) have been identified in lampreys and hagfish, and are expressed on three distinct lymphocytes lineages. VLRA+ cells and VLRC+ cells are T-cell-like and develop in a thymus-like lympho-epithelial structure, termed thymoids. VLRB+ cells are B-cell-like, develop in hematopoietic organs, and differentiate into "VLRB antibody"-secreting plasma cells. == Classification ==