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👩‍🎓 | Clone breeding species Japanese crucian carp, whether to acquire genetic diversity while crossing with sexual reproduction species


Clone breeding species Japanese crucian carp, will it acquire genetic diversity while crossing with sexual reproduction species?

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Evidence that cloned breeding organisms have taken up genes from sexual individuals is the first finding observed in vertebrates and is expected to provide clues to the sexual evolution of organisms.

The freshwater fish "crucian carp", which is familiar to Japanese people, includes the usual "sexual reproduction of male and female (sexual crucian carp)" and "only female ..." → Continue reading

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vertebrate(Crispy animal,Vertebrata) IsAnimal OfSortOne of theDeuterostome OfChordateBelonging to the gateMonophylyIs[1]..For convenience of animals other than vertebratesInvertebrateThat.



What is a vertebrate?mammalian,birds,reptiles,Amphibian,FishesIt is a system group consisting of.However,reptilesとFishes TheParaphylyBecause it isMonophylyFrom the standpoint of recognizing only as a phylogenetic group, as shown in the table below,Quadruped,Amniote,DiapsidsThe monophyletic group such as: will be used as a term:

TaxonimageEstimating the number of species[2]
vertebrateQuadrupedAmniotemammalianSquirrel (PSF) .png5,513
DiapsidsbirdsSecretary bird (Sagittarius serpentarius) 2.jpg10,425

(Paraphyly consisting of diapsids other than birds)

Florida Box Turtle Digon3.jpg10,711
AmphibianLithobates pipiens.jpg7,302
Fishes(Paraphyly consisting of vertebrates other than quadrupeds)Carassius wild golden fish 2013 G1.jpg32,900
Total (number of species)66,178

Listed in the table種Estimate the number ofInternational Union for Conservation of Nature(IUCN)Red list(March 2014)[2]Quoted from.This list estimates the number of modern invertebrates to be 1,305,075, which means that the number of vertebrates is less than 5% of all animals.


Positioning in system classification

Animal kingdomTo vertebrates系統 樹Is as follows[3][4]..The details of the strains far from vertebrates are omitted.For details of the omitted part, see "AnimalPlease refer to the item.


Bilateria anterior(Paraphyly)[Annotation 1]

Bilateral animal






C. elegans











Cephalochordate: Has a notochord for the entire length for the rest of his life.Cephalochordates


Tunicate: Has a notochord in the tail for a lifetime or for a period of time.Ascidian rope,Larvacea,ThaliaceaIt consists of (Hikariboya, Umitaru, Salp, etc.).


vertebrate: The spine is formed around the notochord.




Bilateral animal

There are many exceptions[129][130], Basically has the following features:

  • Complete threeGermBy sex[125], Body is bilateral (= symmetrical)[125].
  • mouthとanus, And the digestive tract that connects them, inside the bodybody cavityOr have a pseudo-cavity (nematodes, rotifers, etc.).
  • Body plans tend to distinguish between anterior (the direction in which the body travels during exercise) and posterior, ventral and dorsal, and therefore left and right.[5][130]..When exercising, it moves forward of the body, so identify what is in the direction of travelSensory organAnd food-eating mouths tend to gather forward (CephalizationCalled).
  • Many bilaterian animalsCircular muscleとLongitudinal muscleBecause I have a pair of[130], In soft-bodied animals such as earthworms, the hydrostatic skeletonPeristalsisCan move by[131].
  • For many bilateriansCiliaCan swim inLarvaThere is a time.


What is a deuterostome (new mouth animal)? Historically, the blastopore formed in the embryo is the mouth.ProtostomeIt is a taxon defined as an animal that does not have a mouth and opens a new mouth with respect to (protostome).[6]..However, when molecular phylogenetic analysis began in the 1990s, deuterostomes in this historical senseSingle systemBecause it was shown not to beChaetognatha,SiboglinidaeEtc. were removed from the deuterostome, and only those in the above phylogenetic tree were left as deuterostomes.[7].

Incidentally,Xenacoelomorpha (Xenacoelomorpha) Is undecided as of 2016[8][9][10][11][12].


ChordateIs a phylum, including vertebrates, (at least at one time in life)NotochordHas the characteristic of having[13]..Details will be described later.

Characteristics and evolution of vertebrates in the phylum Chordate

Chordate characteristics

In order to see the characteristics and evolution of vertebrates in chordates, this section briefly describes the characteristics of chordates.Chordates have the following characteristics:

  • Between the gastrointestinal tract and the neural tubeNotochordHas a supple rod-like structure (fibrous tissue filled with large vacuoles) during the embryonic period[14]..There are also species that hold the notochord in adults[14].
  • In the embryos of other phylums, nerve cords are formed on the ventral side, whereas in the embryos of chordates, the nerve cords are circular on the dorsal side (Neural tube) Can be done[14]Develops in the central nervous system consisting of the brain and spinal cord[14].
  • Digestive tractExtends from mouth to anus[14]Groove-like structures on both sides of the pharynx (the area just behind the mouth) during the embryonic period (Pharyngeal groove) Can be done[14],therePharyngeal fissureThere is a rift[14]..The pharyngeal groove of non-vertebrates is often used as a suspension-eating organ.[14].
  • In many other phylums, the gastrointestinal tract extends to the posterior end of the body, whereas in chordates, the tail extends behind the anus.[14], Uses tail skeleton and muscles for propulsion in aquatic species[14]..However, there are many species whose tails degenerate at the stage of embryogenesis.[14].

Cephalochordate(Cephalochordate) has the following characteristics:

  • Gas exchange: Rarely done in the pharyngeal fissure, but through the body surface[14].
  • Foraging: Use cilia to take in seawater from the mouth, and the mucous membrane of the pharyngeal fissure catches the food in the seawater and sends it to the digestive tract.At that time, seawater goes out of the body from the pharyngeal fissure.[14].
  • mobile: The notochord is flexed by contracting the muscles on both sides of the notochord, and the body is shaken from side to side to move.[14].
  • Length: About XNUMX cm for adults[14]

Tunicate(Ascidians) have the following characteristics:

  • Larval period: Chordates are prominent, but some have only a few minutes of larval stage[14]
  • Adult: Sticky, significantly different from larval[14]..The tail and notochord are absorbed and the nervous system is degenerated[14].
  • Foraging: Pour seawater through the pharyngeal fissure and catch the food in the seawater with mucus.The cilia carry it to the esophagus, where water and excrement exit from the anus into the drain.[14].

Ascidians have lost 13 of the 4 Hox genes of amphioxus, and their larval body plan is formed by a mechanism different from that of other chordates.[14].

In addition, ascidians "may have lost their chordate characteristics in adults after branching from other chordates."[14].

Evolution from early chordates

Facts found from analysis of extant species

The following is known from the analysis of genes of extant species.

  • Early chordates are cephalochordatesCephalochordateIt seems to have had a mouth, notochord, dorsal neural tube, pharyngeal fissure, and tail posterior to the anus.[15].
  • "The cephalochordate's brain is not well developed, it simply has some bulging at the tip of the neural tube."[15]However, it is believed that the structure of this tip has increased in complexity and the vertebrate brain has evolved.[15]..The basis is that of vertebratesForebrain・ The homeobox gene that controls the major parts of the midbrain and retencephalon is expressed in the same pattern in amphioxus.[15].
  • 心 臓Whole-genome analysis of ascidians suggests that chordate ancestors already have genes that control vertebrate-specific structures such as and the thyroid gland.[15].
  • Cells with properties similar to the neural crest, which is a structure peculiar to vertebrates, have been found in ascidians, but ascidians do not have such cells, and ascidians may be in the middle stage of neural crest evolution.[15].

Evolutionary history

First, 5 million years ago (Cambrian explosionAt that time)(English edition)A fossil of a creature with a total length of about 3 cm, such as a lancelet, has been discovered.[16], This organism has some vertebrate characteristics[16]..Specifically, like the amphioxus, it has a mouth that seems to have eaten a suspension.[16]Has a well-developed brain like a vertebrate, small eyes, and a fish-like sarcomere structure[16].

MyllokunmingiaIs considered to be the oldest chordate that has acquired a head[16]However, the acquisition of a head with a brain and eyes has made it possible to perform complex movements and feeding behaviors.[16], I haven't acquired the spine yet[16].

The oldest vertebrate appeared about XNUMX million years ago and is one of them.ConodontKinds have only chondricular endoskeleton[16].

Vertebrates evolved further during the Ordovician and Silurian periods,Semicircular canalMaintaining a sense of balance by acquiring the inner ear with[16],Anti-finAlso won[16]..Also has a muscular pharynx[16]It is believed that this caused inhalation and eating of organisms and organic sediments living on the seabed.[16]..Around this time, there were many swimming vertebrates that protected themselves with bony shells, but all of them became extinct at the end of the Devonian period.[16].

The cartilage skeleton became bony when the instep appeared about 4 million years ago.[16]By 4 million years ago, seeds with thin bony bones covering the endoskeleton of cartilage began to appear.[16]After that, the vertebrates that acquired the jaw became ossified.[16].

Vertebrate characteristics

There is only one Hox gene in other chordates, but there are two in vertebrates, etc.[17]In vertebrates, gene duplication has occurred for important gene families that encode signal molecules and transcription factors.[17], "This may have led to innovations in the vertebrate skeletal and nervous system."[17].

Differences from other chordates

In vertebrates, the notochord and pharyngeal fissures that are characteristic of other chordates change as follows:

  • Most vertebrates develop a connected skeleton around the notochord[14], Only the remnants of the spine remain in adults[14]..In humans the notochord degenerates and becomes part of the intervertebral disc[14].
  • Pharyngeal fissure and its supportPharyngeal archIs used for gas exchange except for quadrupeds[14]..In quadrupeds, "the pharyngeal groove does not open as a pharyngeal fissure, but plays an important role in the development of head and neck structures such as the ears."[14].

Vertebrate derivative traits

In many vertebratesspineSurrounds the neural tube[17]Also on the dorsal side of the neural tube when the neural tube is closing in the embryoNeural crestHas the characteristic of being formed[17].. "The neural crest moves in the embryo and creates various structures such as teeth, parts of the skull and primordium of sensory organs such as cartilage, nerves and eyes."[17].

Phylogenetic tree from vertebrates to amniotes

The phylogenetic tree of vertebrates is as follows[18].."" Drawn in bold at the bottom rightQuadruped"" Represents "vertebrates other than fish", and "vertebrates and below excluding quadrupeds"FishesIt is.





GnathostomatachinとHard boneAcquisition)

Cartilaginous fish(shark,A,Chimaera)

Bony fish(lungOr acquisition of its derivatives)

Article fins

Lobe-finned fish(Acquisition of lobe-finned fish)

(Coelacanth eyes)



Quadruped(Acquisition of limbs with fingers)






It consists of hagfish and lamprey, and is characterized by having no jaw among vertebrates.[17]..Does not have vertebrae but has traces[17]..Hagfish were once excluded from vertebrates because they had no spine, but after that, molecular phylogenetic analysis progressed, and when traces of vertebrae were found, they were classified as vertebrates.[17].



GnathostomataIs a vertebrate with a jaw, and gnathostomata can catch and chew food with its jaw and teeth growing on it.[19]..There is a hypothesis that the jaw is an evolution of the bone that supports the anterior pharyngeal cleft (called "gill cleft" in gnathostomata).[19], The remaining gill fissures are now mainly used for gas exchange[19].

At the genetic level, gnathostomata is characterized by overlapping Hox gene clusters.[19], This is thought to have enabled the evolution of the jaw, etc.[19]..As mentioned above, Hox gene clusters are duplicated even in early vertebrates, so Hox gene clusters have increased to 2 × 2 = 4 in gnathostomata.[19].

Evolutionary history

Early fossils of gnathostomata appeared 4 million years ago[19], Which has evolved rapidly in the following 2,000 million years[19]..Cartilaginous fish, bony fish, and lobe-finned fish, which have survived to date, have all appeared by 4 million years ago.[19].

The oldest fossil gnathostomataPlacodermiAlthough it is a system with armor[19], This strain became extinct around 3 million years ago[19]..At about the same timeAcanthodiiAlthough a system called[19], Extinct 7,000 million years after the extinction of placodermi[19].

Cartilaginous fish


Characterized by having cartilage[20], Nearly 1,000 speciesElasmobranchii(shark,A) And dozens of typesHolocephali(Chimaera) Consists of[20].

Evolutionary history

As already mentioned, vertebrates have already begun to become ossified at an early stage, such as jawless fish with a shell, but cartilaginous fish are a line of secondary cartilage.[20](That is, it follows the evolutionary history of cartilage → bone → cartilage).In fact,CarboniferousThe fin skeleton of sharks has a structure similar to that of bony bones, and modern sharks also have traces of bony tissue at the base of scales and teeth.[20].

Bony fish

The name is bony "fish", but as of 2016, it reflects the phylogenetic tree.QuadrupedIs also included in teleost fish[21](If quadrupeds are not included, teleosts are paraphyly[21]).

Features / evolution

Early teleost strains had lungs that assisted in gas exchange of the gills.[21]..However, in many modern teleost fish except quadrupeds, the lungsSwim bladderEvolving into (Swim bladder)[21]..The swim bladder is clogged with air[21], Fish use swim bladders to adjust buoyancy[21]..Also, the epidermis of most modern teleost fish, except quadrupeds, is covered with scales.[21]..Mucus is secreted from the secretory glands into the epidermis, reducing resistance when swimming.[21].

Article fins

The name was given because it has a flipper that supports the fin.[21].SilurianAppeared in[21]After that, it has diversified and there are more than 27,000 species in modern life.[21].

Lobe-finned fish

"An important synapomorphy of lobe-finned fish is the presence of a rod-shaped bone surrounded by a muscular layer between the pectoral and pelvic fins."[21]..Appeared in the Silurian period as well as ray-finned fish[21].DevonianLike a coastal wetlandBrackish waterMany lobe-finned fish live in the area, and they were used not only for swimming but also for walking (as well as for modern species).[21]..By the end of the Devonian, diversity had diminished and only three strains (coelacanths, lungfish, and quadrupeds) survived.[21].



Lobe-finned fish with pectoral and pelvic fins evolved into fingered limbs[22]..Walk with limbs[22]..Adapted to land life, neck (neck) is generated and head is separated[22]..Other than completely aquatic seeds, they have no gills and instead produce ears and glandular organs during embryogenesis.[22].

Evolutionary history

It is believed that some Devonian lobe-finned fish "walked" on the mud.[22].. Lobe-finned fish 3 million years agoTiktaalikHas the characteristics of fish such as fins and gills, as well as the characteristics of quadrupeds such as ribs (which help breathing and support the body), neck, shoulders, and lungs, and the ability to move the head. Was[22].. The first limbs arose around 3 million years ago[22], Significant diversification over the next 6,000 million years[22].


Frog(カ エ ルKind, about 5420 species[23]),Ariome(SalamanderKind, about 550 species[23]),Caecilians(Caecilians, About 170 species[23]) Consists of[23]..Because amphibians exchange gas mainly in the moist epidermis[23], Its habitat is in damp places such as swamps and rainforests[23], Even species that are adapted to dryness often spend time under humid holes and moist leaves[23]..Also, amphibian eggs have no shell and lose water rapidly in the air, so amphibians lay eggs in water or on moist land.[23]..Many amphibians are in vitro fertilized[23].

AmphibiansChytridiomycosisDue to climate change, habitat loss, environmental pollution, etc., it has been declining rapidly in the last 2016 years or so (as of 30).[23].

Phylogenetic tree of amniotes

Next, the phylogenetic tree of amniotes is placed.[24][25]..Unlike the one above, the lower phylogenetic tree also contains extinct species, and the extinct species are marked with "†" to identify them."Birds" and "mammals" are written in bold on the phylogenetic tree.Of this phylogenetic tree excluding these two, the extant species is "reptilesIt becomes.

Amniote(Amniotic eggAcquisition)

Turtle eyes


Crocodile eyes






† Saurischia other than birds










Scaled reptiles(Lizard,ヘ ビ)






It is a quadruped that is adapted to land life, and has a common derivative trait that the egg has the following four types of membranes.Amniotic eggTo say[26] :

By obtaining amniote eggs suitable for dry land, amniotes, unlike amphibians, no longer need to spend their larval stages in water.[25].

Evolutionary history

The most recent common ancestor of modern amphibians and amniotes survived 3 million years ago.[26]..Early amniotes were small lizard-like animals with sharp teeth, probably due to predation.[26].


Diapsids are animals whose common trait is to have two holes in the temporal region of the skull.[26](Right figure).

Modern diapsids other than birdskeratinHas a synapomorphic trait that is covered with scales whose main component is[27], This scale protects the skin from dryness and rubbing[27]..Also, most reptile eggs are shelled[27]..Many reptiles are externally warm animals that regulate their body temperature with extracorporeal heat such as sunlight.[27], Distinguished from endothermic animals that maintain body temperature by metabolism, such as birds and mammals[27].



A species with a thyroid bone appeared on the ventral side 2 million years ago[27], 2 million years ago, an incomplete but instep appears on the dorsal side[27].

Although turtles do not have two holes that should be synapomorphic traits of diapsids.[27], Being a diapsid is supported both from the point of phylogenetic analysis and from the fact that some extinct fossils have holes in their skulls.[27].


Crocodile ancestors date back to the Late Triassic[27]..The oldest was terrestrial and small, but later became larger and adapted to aquatic life.[27].


The first to fly with flapping wingsQuadruped[26]..A collagen membrane extends from one finger of the forelimb to the torso or hindlimb as wings.[26], This is different in structure from birds with wings extending from their forelimbs and bats with wings extending from their hands.

Appeared in the late Triassic[26]Extinct by 6600 million years ago[26].


Including birds and saurischia[26]..Ornithischian is herbivorous[26]Many species have developed defense mechanisms such as club-shaped tails and horns of the head.[26]..Saurischia include theropods (Tyrannosaurus rex, etc.) and long-necked herbivorous saurischia.[26].. It has been shown to have been agile since its discovery in the 1970s.[26]..What makes a flock[26], Nest building and parenting[26], There was something with endotherm[26]Etc. are also shown.

Non-bird dinosaurs became extinct 6600 million years before the end of the Cretaceous[26]..Birds will be described later.



Birds are equipped with wings for flight and have various adaptations for flight.First of all, to reduce the weight of the body, there is no bladder[28], The ovaries are reduced to one[28], Both males and females have small gonads except during the breeding season[28], The beak has no teeth[28]..In addition, in order to secure sufficient energy for flight, a high metabolic rate is achieved in the respiratory system with elastic lung air sacs and the circulatory system of two ventricles and two atria.[28]..Vision is also developing due to the need in flight[28], Has a relatively large brain[28], The visual cortex and motor cortex necessary for flight are particularly developed[28].

Birds are endothermic[28]Feathers and (depending on the species) fat keep metabolic heat out.[28]..Feathers are made of a protein called β-keratin, similar to reptile scales.[28]..Mating isCloacaMade by contact[28].

RatiteConsists of exceptionally flightless birds[29], Ostrich, Rhea, Kiwi, Cassowary, Emu, etc.[29].

Evolutionary history

Feathered theropods were born 1 million years ago[29].. Among the known as of 2016ArcheopteryxIs the oldest bird[29], Urvogel has feathers on its wings[29]It has the same characteristics as modern birds, but it also has claws on its wings.[29], Beak has teeth[29]It also retains characteristics different from those of modern birds, and unlike modern birds, it did not fly from a stationary position.[29].

Clades, including the 28th modern bird, have existed for more than 6600 million years ago.[29].


Lepidosauria consist of scaled reptiles (lizards and snakes) and tuatara.[30], Squamata is the most species-rich line of reptiles[25]..Among the scaled reptiles, snakes evolved from lizards with limbs.[30].

Tuatara appeared at the latest 2 million years ago[30]Although it was prosperous until the Cretaceous[30]As of 2016, I only live on islands near New Zealand[30]..Lifespan exceeds 100 years[30].

Synapsids / mammals

Synapsids have a synapomorphic trait of having only one infratemporal window (see the figure on the right).[31], All modern synapsids belong to mammals[31]..In humans, the infratemporal flute corresponds to a hole connecting the jaw muscle and the temporal region.[31].

Mammal characteristics

Have a mammary gland and raise the child with breast milk from the female mammary gland[32]..Many species have relatively large brains[32]..It is endothermic and the hair on the surface of the body keeps the metabolic heat from escaping.[32].kidneyDue to the well-developed function of, there is little loss of water when discharging waste products.[32].

Evolutionary history

Early synapsids had no hair and were oviparous[31]..Synapsids expanded their diversity in the Permian[31]Although it was predominant among quadrupeds[31]Its diversity diminished during the mass extinction of the late Permian period[31].

The origin of mammalsTheriodontiaBelongs toCynodontsIs[33], Cynodonts survived the late Triassic mass extinctionMammaliaformesWas born[33]..Small size[31]·Nocturnality[31]・ Insect food[31]And probably had hair[31].

Mammals appeared in the Jurassic[31]..By the Cretaceous, three strains of modern mammals (Single hole,marsupial,Eutheria) Appears[31]..Mammals adaptively radiated after the extinction of many late Cretaceous dinosaurs and reptiles[31]..Adaptive radiation of mammals is closely related to continental division[33],Gondwana ContinentIn South America when it split in the late JurassicXenarthra, On the African continentAfrotheria,LaurasiaでBoreoeutheriaHas evolved[33].


[How to use footnotes]

注 釈

  1. ^ "Anterior bilaterian" is a convenient name for describing the phylum other than bilaterian, and there is no clade group called "anterior bilaterian" (Fujita (2010), p.113.)


  1. ^ Fujita (2010), p.113.
  2. ^ a b The World Conservation Union. 2014. IUCN Red List of Threatened Species, 2014.3. Summary Statistics for Globally Threatened Species. Table 1: Numbers of threatened species by major groups of organisms (1996–2014).
  3. ^ Fujita (2010), p.113. pp.174-175.
  4. ^ Mawatari (2013), p.2.
  5. ^ Minelli, Alessandro (2009). Perspectives in Animal Phylogeny and EvolutionOxford University Press. P. 53. ISBN 978-0-19-856620-5. https://books.google.com/books?id=jIASDAAAQBAJ&pg=PA53 
  6. ^ Fujita (2010) p104
  7. ^ Fujita (2010) p108
  8. ^ Philippe, Hervé; Brinkmann, Henner; Copley, Richard R .; Moroz, Leonid L .; Nakano, Hiroaki; Poustka, Albert J .; Wallberg, Andreas; Peterson, Kevin J. et al. (2011-02). “Acoelomorph flatworms are deuterostomes related to Xenoturbella” (English). Nature 470 (7333): 255–258. two:10.1038 / nature09676. ISSN 0028-0836 . PMC PMC4025995. PMID 21307940. http://www.nature.com/articles/nature09676. 
  9. ^ Mawatari (2013), p27-p29
  10. ^ Rouse, Greg W .; Wilson, Nerida G .; Carvajal, Jose I .; Vrijenhoek, Robert C. (2016-02). “New deep-sea species of Xenoturbella and the position of Xenacoelomorpha” (English). Nature 530 (7588): 94–97. two:10.1038 / nature16545. ISSN 0028-0836 . http://www.nature.com/articles/nature16545. 
  11. ^ Cannon, Johanna Taylor; Vellutini, Bruno Cossermelli; Smith, Julian; Ronquist, Fredrik; Jondelius, Ulf; Hejnol, Andreas (2016-02). “Xenacoelomorpha is the sister group to Nephrozoa” (English). Nature 530 (7588): 89–93. two:10.1038 / nature16520. ISSN 0028-0836 . http://www.nature.com/articles/nature16520. 
  12. ^ "Taxonomy: Xenacoelomorpha is a sister group of Nephrozoa". Nature (September 2016, 2). 2018/7/20Browse.
  13. ^ Fujita (2010), pp. 174-175.
  14. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z Campbell 11th Edition pp. 826-827.
  15. ^ a b c d e f Campbell 11th Edition pp. 828-829.
  16. ^ a b c d e f g h i j k l m n o p Campbell 11th Edition pp. 831-832.
  17. ^ a b c d e f g h i Campbell 11th Edition pp. 829-830.
  18. ^ Campbell 11th Edition p.826.
  19. ^ a b c d e f g h i j k l m Campbell 11th Edition pp.832-833
  20. ^ a b c d Campbell 11th Edition pp.833-834
  21. ^ a b c d e f g h i j k l m n o Campbell 11th Edition pp.834-836
  22. ^ a b c d e f g h Campbell 11th Edition pp.837-838
  23. ^ a b c d e f g h i j Campbell 11th Edition pp.838-841
  24. ^ Campbell 11th Edition p.841
  25. ^ a b c Zoological Society of Japan 2018 pp. 98-99.
  26. ^ a b c d e f g h i j k l m n o p q r s t u Campbell 11th Edition pp.841-843
  27. ^ a b c d e f g h i j k Campbell 11th Edition pp.842-845
  28. ^ a b c d e f g h i j k l Campbell 11th Edition pp.845-846
  29. ^ a b c d e f g h i Campbell 11th Edition pp.846-847
  30. ^ a b c d e f Campbell 11th Edition pp.843-845
  31. ^ a b c d e f g h i j k l m n Campbell 11th Edition pp. 848-849.
  32. ^ a b c d Campbell 11th Edition p.848.
  33. ^ a b c d Zoological Society of Japan 2018 pp. 102-103.


  • [Campbell 11th Edition] Campbell Biology Original 11th Edition. Maruzen Publishing. (2018/3/20). ISBN 978-4621302767  
    • original:Lisa A. Urry; Michael L. Cain; Steven A. Wasserman; Peter V. Minorsky; Jane B. Reece; Neil A. Campbell (2016/10/29). Campbell Biology (11th Edition). Pearson. ISBN 978-0134093413  
  • [Fujita (2010)] Toshihiko Fujita, "Animal Phylogenetic Classification and Evolution," Jiro Ota, Koji Akasaka, Makoto Asashima, Toshiyuki Nagata, Shokabo <New Life Science Series>, April 2010, 4.ISBN 978-4785358426 .
  • The Zoological Society of Japan "Encyclopedia of Zoology" Maruzen Publishing Co., Ltd., September 2018, 9.ISBN 978-4621303092 .

For further understanding

Details of phylogenetic classification:

Sexual individuals


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