GlyCosmos Portal

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GlyCosmos Glycans

The list of validated glycans extracted from GlyTouCan, updated weekly.

Source	Last Updated
GlyTouCan	August 18, 2024

Displaying entries **1726 - 1750** of **5831** in total

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GlyTouCan ID	IUPAC Condensed	Motifs	Subsumption Level ▲	Monoisotopic Mass	Organisms
G13294EW	Gal(b1-3)[Fuc(a1-4)]GlcNAc(a1-3)Gal(b1-4)GlcNAc(b1-3)Gal(b1-4)Glc(b1-	Lea Lewis a Paragloboside	Linkage defined saccharide	1218.44	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13495RN	Neu5Ac(a2-3)Gal(b1-4)GlcNAc(b1-2)[Neu5Ac(a2-3)Gal(b1-4)GlcNAc(b1-4)]Man(a1-3)[Neu5Ac(a2-6)Gal(b1-4)GlcNAc(b1-2)Man(a1-6)]Man(b1-4)GlcNAc(b1-4)[Fuc(a1-6)]GlcNAc(?1-	N-Glycan complex Polysialic acid	Linkage defined saccharide	3025.07	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13636IT	Fuc(a1-2)Gal(b1-4)GlcNAc(b1-3)Gal(b1-4)Glc(b1-	Blood group H (type 2) Paragloboside	Linkage defined saccharide	853.31	Amniota Artiodactyla Bilateria Boreoeutheria Canidae Caniformia Canis lupus familiaris Canis lupus Canis Carnivora Catarrhini Chordata Craniata Cricetidae Cricetinae Cricetulus griseus Cricetulus Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Glires Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Laurasiatheria Mammalia Metazoa Muroidea Myomorpha Opisthokonta Primates Rodentia Sarcopterygii Simiiformes Suidae Suina Sus scrofa Sus Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13672YG	Fuc(a1-2)Gal(b1-4)GlcNAc(b1-2)Man(a1-3)[GlcNAc(b1-4)][Gal(b1-4)GlcNAc(b1-2)Man(a1-6)]Man(b1-4)GlcNAc(b1-4)[Fuc(a1-6)]GlcNAc(?1-	N-Glycan complex N-acetyllactosamine Type 2 LN	Linkage defined saccharide	2135.79	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13800FQ	Neu5Ac(a2-6)Gal(b1-4)GlcNAc(b1-2)Man(a1-3)[Man(a1-6)]Man(b1-4)GlcNAc(b1-	N-acetyllactosamine Type 2 LN	Linkage defined saccharide	1363.48	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13863XN	Neu5Ac(a2-3)Gal(b1-3)[Fuc(a1-2)Gal(b1-3)[Fuc(a1-4)]GlcNAc(b1-3)Gal(b1-4)GlcNAc(b1-3)Gal(b1-4)GlcNAc(b1-6)]GalNAc(a1-	Blood group H (type 1) I' antigen Leb Lewis b O-Glycan core 2 Sialyl T antigen Type 2 LN2	Linkage defined saccharide	2061.75	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13869TL	Man(a1-2)Man(a1-3)[Man(a1-3)[Man(a1-2)Man(a1-6)]Man(a1-6)]Man(b1-4)GlcNAc(b1-4)GlcNAc-ol		Linkage defined saccharide	1560.55	50 kb inversion clade Acalyptratae Alphainfluenzavirus influenzae Alphainfluenzavirus Amniota Archelosauria Archosauria Arthropoda Articulavirales Artiodactyla Aves Bilateria Boreoeutheria Bos taurus Bos Bovidae Bovinae Brachycera Catarrhini Chordata Coelurosauria Craniata Cyclorrhapha Deuterostomia Dicondylia Dinosauria Dipnotetrapodomorpha Diptera Discoba Drosophila melanogaster Drosophila Drosophilidae Drosophilinae Drosophilini Ecdysozoa Embryophyta Endopterygota Ephydroidea Eremoneura Euarchontoglires Euglenozoa Eukaryota Eumetazoa Euphyllophyta Euteleostomi Eutheria Fabaceae Fabales Galliformes Galloanserae Gallus gallus Gallus Glires Gnathostomata Gunneridae Haplorrhini Hexapoda Hominidae Homininae Hominoidea Homo sapiens Homo Influenza A virus Insecta Insthoviricetes Kinetoplastea Laurasiatheria Magnoliopsida Mammalia Mandibulata Mesangiospermae Metakinetoplastina Metazoa Muridae Murinae Muroidea Muscomorpha Myomorpha NPAAA clade Negarnaviricota Neognathae Neoptera Opisthokonta Orthomyxoviridae Orthornavirae Panarthropoda Pancrustacea Papilionoideae Pecora Pentapetalae Phaseoleae Phaseolus vulgaris Phaseolus Phasianidae Phasianinae Polyploviricotina Primates Protostomia Pterygota Rattus Riboviria Rodentia Ruminantia Sarcopterygii Sauria Saurischia Sauropsida Schizophora Simiiformes Sophophora Spermatophyta Streptophyta Streptophytina Suidae Suina Sus scrofa Sus Teleostomi Tetrapoda Theria Theropoda Tracheophyta Trypanosoma brucei Trypanosoma Trypanosomatida Trypanosomatidae Trypanozoon Vertebrata Viridiplantae Viruses cellular organisms eudicotyledons fabids indigoferoid/millettioid clade melanogaster group melanogaster subgroup rosids unclassified Orthomyxoviridae unidentified influenza virus
G13947MP	Man(a1-6)Man(b1-4)GlcNAc(b1-4)GlcNAc-ol		Linkage defined saccharide	750.29	Aculeata Alveolata Amniota Anas platyrhynchos Anas Anatidae Anatinae Anseriformes Anthophila Apidae Apinae Apini Apis mellifera Apis Apocrita Apoidea Archelosauria Archosauria Arthropoda Artiodactyla Aves Bilateria Boreoeutheria Catarrhini Chordata Ciliophora Coelurosauria Craniata Deuterostomia Dicondylia Dinosauria Dipnotetrapodomorpha Ecdysozoa Endopterygota Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Galliformes Galloanserae Glires Gnathostomata Haplorrhini Hexapoda Hominidae Homininae Hominoidea Homo sapiens Homo Hymenoptera Hymenostomatida Insecta Intramacronucleata Laurasiatheria Mammalia Mandibulata Metazoa Muridae Murinae Muroidea Myomorpha Neognathae Neoptera Oligohymenophorea Opisthokonta Panarthropoda Pancrustacea Primates Protostomia Pterygota Rattus Rodentia Sar Sarcopterygii Sauria Saurischia Sauropsida Simiiformes Suidae Suina Sus scrofa Sus Teleostomi Tetrahymena pyriformis Tetrahymena Tetrahymenidae Tetrahymenina Tetrapoda Theria Theropoda Vertebrata cellular organisms
G13258AZ	Neu5Ac(a2-6)Gal(b1-4)GlcNAc(b1-3)Gal(b1-3)GalNAc-ol	N-acetyllactosamine Type 2 LN	Linkage defined saccharide	1041.39	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13688QP	Neu5Ac(a2-6)Gal(b1-4)GlcNAc(b1-3)Gal(b1-4)[Fuc(a1-3)]GlcNAc(b1-2)Man(a1-3)[Gal(b1-4)[Fuc(a1-3)]GlcNAc(b1-3)Gal(b1-4)[Fuc(a1-3)]GlcNAc(b1-2)Man(a1-6)]Man(b1-4)GlcNAc(b1-4)[Fuc(a1-6)]GlcNAc-ol	Lex-Lex	Linkage defined saccharide	3248.2	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13836YR	GalNAc(b1-4)GlcNAc(b1-2)Man(a1-3)[GalNAc(b1-4)GlcNAc(b1-2)[GalNAc(b1-4)GlcNAc(b1-6)]Man(a1-6)]Man(b1-4)GlcNAc(b1-4)GlcNAc(?1-	LDN LacdiNAc N-Glycan complex	Linkage defined saccharide	2128.8	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13260JU	Gal(b1-3)[Neu5Ac(a2-3)[GalNAc(b1-4)]Gal(b1-4)GlcNAc(b1-6)]GalNAc(a1-	O-Glycan core 2 Sda/CT antigen	Linkage defined saccharide	1242.45	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Glires Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Muridae Murinae Muroidea Mus musculus Mus Mus Myomorpha Opisthokonta Primates Rodentia Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13290KJ	Neu5Ac(a2-6)Gal(b1-4)GlcNAc(b1-2)Man(a1-?)[GlcNAc(b1-2)Man(a1-?)]Man(b1-4)GlcNAc(b1-4)GlcNAc(b1-	N-Glycan complex N-acetyllactosamine Type 2 LN	Linkage defined saccharide	1769.63	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13824WQ	GalNAc(a1-3)Gal(b1-3)GlcNAc(b1-3)GalNAc(a1-	O-Glycan core 3 Type 1 LN	Linkage defined saccharide	789.3	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13515ZY	GalNAc(b1-4)GlcNAc(b1-2)[GlcNAc(b1-4)]Man(a1-3)[GalNAc(b1-4)GlcNAc(b1-2)[GlcNAc(b1-6)]Man(a1-6)]Man(b1-4)GlcNAc(b1-4)[Fuc(a1-6)]GlcNAc(?1-	LDN LacdiNAc N-Glycan complex	Linkage defined saccharide	2274.86	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13537KO	Gal(b1-4)GlcNAc(b1-2)Man(a1-3)[Gal(b1-4)GlcNAc(b1-3)Gal(b1-4)[Fuc(a1-3)]GlcNAc(b1-2)Man(a1-6)]Man(b1-4)GlcNAc(b1-4)GlcNAc(?1-	I' antigen Lewis x Lex N-Glycan complex SSEA-1 Type 2 LN2	Linkage defined saccharide	2151.78	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13583BZ	Gal(b1-4)GlcNAc(b1-2)[Gal(b1-4)GlcNAc(b1-4)]Man(a1-3)[GlcNAc(b1-2)Man(a1-6)]Man(b1-4)GlcNAc(b1-4)[Fuc(a1-6)]GlcNAc(?1-	N-Glycan complex N-acetyllactosamine Type 2 LN	Linkage defined saccharide	1989.73	Amniota Archelosauria Archosauria Articulavirales Artiodactyla Aves Bilateria Boreoeutheria Catarrhini Chordata Coelurosauria Craniata Deuterostomia Dinosauria Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Galliformes Galloanserae Gallus gallus Gallus Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Insthoviricetes Laurasiatheria Mammalia Metazoa Negarnaviricota Neognathae Opisthokonta Orthomyxoviridae Orthornavirae Phasianidae Phasianinae Polyploviricotina Primates Riboviria Sarcopterygii Sauria Saurischia Sauropsida Simiiformes Suidae Suina Sus scrofa Sus Teleostomi Tetrapoda Theria Theropoda Vertebrata Viruses cellular organisms unclassified Orthomyxoviridae unidentified influenza virus
G13894RO	Fuc(a1-2)Gal(b1-3)[Fuc(a1-2)Gal(b1-4)[Fuc(a1-3)]GlcNAc(b1-3)[Fuc(a1-2)]Gal(b1-4)GlcNAc(b1-6)]GalNAc-ol	Blood group H (type 2) Blood group H (type 2) I' antigen Leb Lewis b Ley Type 2 LN2	Linkage defined saccharide	1699.65	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13902SG			Linkage defined saccharide	254.1	Acholeplasma laidlawii Acholeplasma Acholeplasmataceae Acholeplasmatales Amniota Bacilli Bacillota Bacteria Bilateria Boreoeutheria Catarrhini Chordata Craniata Deinococcaceae Deinococcales Deinococci Deinococcota Deinococcus radiodurans Deinococcus Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Lacticaseibacillus casei Lacticaseibacillus Lactobacillaceae Lactobacillales Mammalia Metazoa Mollicutes Mycoplasmatota Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Terrabacteria group Tetrapoda Theria Vertebrata cellular organisms
G13661YX		N-Glycan core basic	Linkage defined saccharide	2587.92	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13602QE		3-Sialyl-LN (type 2)	Linkage defined saccharide	1915.69	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G13974DV	Man(a1-3)[Man(a1-3)[Man(a1-6)]Man(a1-6)]Man(b1-4)GlcNAc(b1-4)GlcNAc-ol		Linkage defined saccharide	1236.45	Acalypheae Acalyphoideae Acalyptratae Achelata Aculeata Amniota Anthophila Apidae Apocrita Apoidea Archelosauria Archosauria Arthropoda Articulavirales Artiodactyla Aves Bilateria Boreoeutheria Bos taurus Bos Bovidae Bovinae Brachycera Catarrhini Chordata Coelurosauria Craniata Crustacea Cyclorrhapha Decapoda Deuterostomia Dicondylia Dinosauria Dipnotetrapodomorpha Diptera Drosophila melanogaster Drosophila Drosophilidae Drosophilinae Drosophilini Ecdysozoa Embryophyta Endopterygota Ephydroidea Eremoneura Euarchontoglires Eucarida Eukaryota Eumalacostraca Eumetazoa Euphorbiaceae Euphyllophyta Euteleostomi Eutheria Galliformes Galloanserae Gallus gallus Gallus Glires Gnathostomata Gunneridae Haplorrhini Hexapoda Hominidae Homininae Hominoidea Homo sapiens Homo Hymenoptera Insecta Insthoviricetes Lagomorpha Laurasiatheria Leporidae Magnoliopsida Malacostraca Malpighiales Mammalia Mandibulata Mesangiospermae Metazoa Multicrustacea Muridae Murinae Muroidea Mus musculus Mus Mus Muscomorpha Myomorpha Negarnaviricota Neognathae Neoptera Opisthokonta Orthomyxoviridae Orthornavirae Oryctolagus cuniculus Oryctolagus Palinuridae Palinuroidea Panarthropoda Pancrustacea Panulirus interruptus Panulirus Pecora Pentapetalae Phasianidae Phasianinae Pleocyemata Polyploviricotina Primates Protostomia Pterygota Rattus Riboviria Ricinus communis Ricinus Rodentia Ruminantia Sarcopterygii Sauria Saurischia Sauropsida Schizophora Simiiformes Sophophora Spermatophyta Streptophyta Streptophytina Suidae Suina Sus scrofa Sus Teleostomi Tetrapoda Theria Theropoda Tracheophyta Vertebrata Viridiplantae Viruses cellular organisms eudicotyledons fabids melanogaster group melanogaster subgroup rosids unclassified Orthomyxoviridae unidentified influenza virus
G14000ZZ	Neu5Ac(a2-3)Gal(b1-3)[Neu5Ac(a2-6)]GlcNAc(b1-	GM4 Type 1 LN	Linkage defined saccharide	965.33	Amniota Artiodactyla Bilateria Boreoeutheria Bos taurus Bos Bovidae Bovinae Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Laurasiatheria Mammalia Metazoa Opisthokonta Pecora Primates Ruminantia Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G14015IL	Gal(b1-4)GlcNAc(b1-2)[Gal(b1-4)GlcNAc(b1-4)]Man(a1-3)[Gal(b1-4)GlcNAc(b1-2)[Gal(b1-4)GlcNAc(b1-3)Gal(b1-4)GlcNAc(b1-6)]Man(a1-6)]Man(b1-4)GlcNAc(b1-4)[Fuc(a1-6)]GlcNAc(?1-	I' antigen N-Glycan complex Type 2 LN2	Linkage defined saccharide	2882.05	Amniota Bilateria Boreoeutheria Catarrhini Chordata Craniata Deuterostomia Dipnotetrapodomorpha Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Gnathostomata Haplorrhini Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Opisthokonta Primates Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms
G14023WK	GlcNAc(b1-3)Gal-ol		Linkage defined saccharide	385.16	Amniota Bacteria Bilateria Boreoeutheria Campylobacterales Campylobacterota Catarrhini Chordata Craniata Cricetidae Cricetinae Cricetulus griseus Cricetulus Deuterostomia Dipnotetrapodomorpha Epsilonproteobacteria Euarchontoglires Eukaryota Eumetazoa Euteleostomi Eutheria Glires Gnathostomata Haplorrhini Helicobacter pylori Helicobacter Helicobacteraceae Hominidae Homininae Hominoidea Homo sapiens Homo Mammalia Metazoa Muroidea Myomorpha Opisthokonta Primates Rodentia Sarcopterygii Simiiformes Teleostomi Tetrapoda Theria Vertebrata cellular organisms