出典(authority):フリー百科事典『ウィキペディア(Wikipedia)』「2012/11/23 14:45:38」(JST)
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生息年代: エディアカラ紀 - 現世
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左上から順に、イエアメガエル、モリフクロウ、トラ、ニワオニグモ、ニワノオウシュウマイマイ、アオウミガメ、en:Solitary bee、en:Asian arowana、バーバリーマカク、ノコギリエイ、en:Marbled White
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本文参照 |
動物(どうぶつ、羅: Animalia、単数: Animal)は、動物界(どうぶつかい) に分類される生物のこと、またはその総称。真核生物に含まれ、一般に運動能力と感覚を持つ多細胞生物である。
また、日常語としての「動物」は、植物の対置語として以外に、いわゆる「けもの」の意味で使われることがある。[1]
目次
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生物を動物と植物に二分する分類法は古くから存在しており、アリストテレスは感覚と運動能力の有無によりこれら二つの分類を試みている。ただし、中間的生物も存在することを認めていたようである。18世紀の生物学者リンネ (Carous Linnaeus) は、感覚をもたない植物界と、感覚と移動能力をもち従属栄養的である動物界とに、生物を二分した。
明治時代以前の日本では、生物は草、虫、魚、獣と区分する本草学が主流であり、動物という概念は、西欧の学問に親しんだ蘭学者を除き、一般的ではなかった。生物を動物と植物に二分する分類は、西欧の学問が流入した明治以降に広く普及した。
二界説の下では、動物には下記の各群以外に、原生動物を単細胞の動物と位置づけていた。生物学の進歩により、現在では、動物か植物かのみで生物を分類するのは一般的ではなく、さまざまな分類法が提案されている(参考:生物の分類)。それらに従えば、真正細菌、古細菌、原生生物、菌類など、動物にも植物にも分類されない生物も数多く存在し、動物界はそのようないくつもの系統の内の一つと見なされる。20世紀末の分子遺伝学などの流れの中で、枠組みは何度も見直され、植物界や菌界は大きくその構成が変わった。動物界に関しても、原生動物はそのような多系統の入り交じったものであることが判明している。後生動物に関しては、ほとんど変更を受けなかった。大きな変更としてはそれまで原生動物の一つと見なされていたミクソゾアがここに含められるようになった程度である。
動物の起源については、旧来から多細胞動物の起源ではないかといわれたこともある襟鞭毛虫類がそれらしいということになっている。繊毛虫やアメーバはかなり系統が遠いらしいこと、菌界が動物界に近いことなどが示されている。動物・菌類・襟鞭毛虫を含む系統はオピストコンタと呼ばれる。原生動物の各系統、あるいはその他の情報に関しては生物の分類を参照。
一般には、運動能力と感覚を持つのが大きな特徴とされるが、現在の動物界に含まれる生物すべてに当てはめることができない。以下のような特徴を持つ生物が、現在の意味での動物である。
下表は動物界を生物の分類の分類項目である「門」に分類したものである。各動物門に属する生物はそれぞれの「門」独自の基本設計(ボディプラン)を共有している。
分類法には、背骨(脊椎)をもつ動物(脊椎動物)ともたない動物(無脊椎動物)とに分ける2分法が存在する。この分類は、ヒトを含む脊椎動物をより詳しく取り上げるときなどに、あくまでも便宜的に用いられる分類であることに注意しなければならない。実際には、脊椎動物は大きな多様性を誇る動物界の1亜門に過ぎないからである(下表35門中の脊索動物門の、さらに1亜門)。
上位分類 | 門 | 胚発生等 | ||||
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海綿動物門(カイメン、カイロウドウケツ) | 無胚葉 | 器官が 明確には |
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平板動物門(センモウヒラムシ) | ||||||
真正 後生 |
放射 相称 |
刺胞動物門(クラゲ、サンゴ) | 2胚葉 | 消化管は 出入口が |
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有櫛動物門(クシクラゲ) | ||||||
左右 相称 |
旧口 動物 |
冠輪 動物 |
扁形動物門(プラナリア、キュウチュウ、サナダムシ) | 3胚葉 | ||
無腸動物門 | 器官退化? | |||||
菱形動物門(ニハイチュウ) | ||||||
直泳動物門(キリオキンクタ) | ||||||
紐形動物門(ヒモムシ) | 消化管は 出口と |
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顎口動物門 (ハプログナチア) | ||||||
腹毛動物門(イタチムシ、オビムシ) | ||||||
輪形動物門(ワムシ) | ||||||
内肛動物門(スズコケムシ) | ||||||
外肛動物門(チゴケムシ、コケムシ) | ||||||
箒虫動物門(ホウキムシ) | ||||||
腕足動物門(ホオズキガイ、シャミセンガイ) | ||||||
星口動物門(ホシムシ) | ||||||
ユムシ動物門(ユムシ) | ||||||
毛顎動物門(ヤムシ) | ||||||
有輪動物門(シンビオン) | ||||||
微顎動物門 (リムノグナシア) | ||||||
環形動物門(ミミズ、ゴカイ) | ||||||
軟体動物門(貝類、イカ、タコ) | ||||||
脱皮 動物 |
線形動物門(回虫) | |||||
類線形動物門(ハリガネムシ) | ||||||
鰓曳動物門(エラヒキムシ) | ||||||
胴甲動物門(コウラムシ) | ||||||
動吻動物門(トゲカワ) | ||||||
緩歩動物門(クマムシ) | ||||||
有爪動物門(カギムシ) | ||||||
節足動物門(昆虫類、甲殻類) | ||||||
新口動物 | 珍渦虫動物門 (チンウズムシ) | |||||
棘皮動物門(ヒトデ、クモヒトデ、ナマコ) | ||||||
半索動物門(ギボシムシ) | ||||||
脊索動物門(ホヤ、脊椎動物) |
現在、分子系統解析が進展中ということもあり、後生動物の内部分類にも多少の振れ幅がある。上表は今後も若干の修正が加えられていくものと思われる。
菱形動物と直泳動物はまとめて中生動物とすることもある。ほかに胞胚様動物門(一胚葉動物門)(Monoblastozoa) がサリネラという単一種によってたてられているが、この動物は存在が疑問視されている。
ミクソゾア、舌形動物、鉤頭動物は、独立の動物門とする場合もあるが、それぞれ、刺胞動物、節足動物、輪形動物に含まれるとする見方もある。上表では独立門としていない。
ユムシ動物、星口動物は、環形動物門に含まれるという見方もある。また、従来の腕足動物と箒虫動物は同系統であるとして、腕動物門あるいは腕足動物門としてひとつにまとめる立場もある。また最近では、無腸類と珍渦虫が同系統であるとして、珍無腸動物門という新門が設けられる場合もある。
現在の分子系統学では、左右相称動物(三胚葉動物)の単系統性は支持されているものの、その他の、海綿動物、平板動物、刺胞動物、有櫛動物を含めた5者間の系統関係はまったく分かっていない状況である。従来言われてきた真正後生動物というくくりも学説のうちのひとつとして理解するべきである。
平板動物門(板状動物門)と海綿動物門の2門(側生動物亜界)は器官が分化しておらず、不定形であるが、その他の動物(後生動物、真正後生動物亜界)は器官系が分化している。これらの器官をもつ後生動物は、規則的な形状をしている。放射相称(刺胞動物門、有櫛動物門)または左右対称(その他の動物)のいずれかの形状を有しているのである。
すべての動物は、受精卵が卵割していくと、細胞でできた中空のボールである胚胞を形成する。後生動物では胚胞の一部が陥入し、開口部が1つある嚢胚を形成する。嚢胚形成後、細胞は2層(2胚葉)または3層(3胚葉)の組織に分化する。3層の場合、各組織層は外胚葉、中胚葉、内胚葉とよばれる。外胚葉は主に表皮、神経系に、中胚葉は主に筋肉に、内胚葉は主に消化管になる。
嚢胚形成時の陥入箇所、原口が後に消化管のどちらになるかは重要で、口になる旧口動物と、同じ陥入箇所が後に肛門になる新口動物の2つに分けられる。刺胞動物、有櫛動物と扁形動物では、原口から続く消化管の反対側に、新しい口が開かず、消化管は口以外の出入り口を持たない。
また、外胚葉と内胚葉の間には接合していない部分が存在し、この空所を体腔と呼ぶ。この体腔は伝統的にその発達の度合いが進化の度合いを反映しているとして動物の門分類等で重要視されてきた。空所はあるが中胚葉の裏打ちがない場合を偽体腔と呼び、裏打ちのあるものを真体腔と呼ぶ。
発生からは分類や進化に関する知見が多く得られる。幼生の形も、分類群やそれらの間の類縁を示す場合があり、重要である。フジツボが甲殻類に含まれることがわかったときの決め手は、幼生がノープリウスであったことである。複数の動物群に共通の幼生がある場合、それらは類縁であると判断される。その典型がトロコフォアである。このことを拡張したのがエルンスト・ヘッケルの反復説である。
消化器系・呼吸器系・循環系・神経系・排出系などの各器官がどのような構造で、どのような配置であるかは、門によってほぼ決まっている。
これらの形質を元に、動物の系統関係が論じられてきた。最近は、分岐分類学や分子遺伝学的情報に基づく見直しも進められており、各門の関係等については見方の大きな変更が起きている部分もある。門の範囲等については大きく変わっているところは少ない。
ウィキメディア・コモンズには、動物に関連するカテゴリがあります。 |
ウィキスピーシーズに動物界に関する情報があります。 |
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Animals Temporal range: Ediacaran – Recent PreЄ
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Scientific classification | |
Domain: | Eukaryota |
(Unranked) | Opisthokonta |
(Unranked) | Holozoa |
(Unranked) | Filozoa |
Kingdom: | Animalia Linnaeus, 1758 |
Phyla | |
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Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and independently. All animals are also heterotrophs, meaning they must ingest other organisms or their products for sustenance.
Most known animal phyla appeared in the fossil record as marine species during the Cambrian explosion, about 542 million years ago.
Contents
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The word "animal" comes from the Latin word animalis, meaning "having breath".[1] In everyday colloquial usage, the word often refers to non-human members of kingdom Animalia. Sometimes, only closer relatives of humans such as mammals and other vertebrates are meant in colloquial use.[2] The biological definition of the word refers to all members of the kingdom Animalia, encompassing creatures as diverse as sponges, jellyfish, insects and humans.[3]
Animals have several characteristics that set them apart from other living things. Animals are eukaryotic and mostly multicellular,[4] which separates them from bacteria and most protists. They are heterotrophic,[5] generally digesting food in an internal chamber, which separates them from plants and algae.[6] They are also distinguished from plants, algae, and fungi by lacking rigid cell walls.[7] All animals are motile,[8] if only at certain life stages. In most animals, embryos pass through a blastula stage,[9] which is a characteristic exclusive to animals.
With a few exceptions, most notably the sponges (Phylum Porifera) and Placozoa, animals have bodies differentiated into separate tissues. These include muscles, which are able to contract and control locomotion, and nerve tissues, which send and process signals. Typically, there is also an internal digestive chamber, with one or two openings.[10] Animals with this sort of organization are called metazoans, or eumetazoans when the former is used for animals in general.[11]
All animals have eukaryotic cells, surrounded by a characteristic extracellular matrix composed of collagen and elastic glycoproteins.[12] This may be calcified to form structures like shells, bones, and spicules.[13] During development, it forms a relatively flexible framework[14] upon which cells can move about and be reorganized, making complex structures possible. In contrast, other multicellular organisms, like plants and fungi, have cells held in place by cell walls, and so develop by progressive growth.[10] Also, unique to animal cells are the following intercellular junctions: tight junctions, gap junctions, and desmosomes.[15]
Nearly all animals undergo some form of sexual reproduction.[16] They have a few specialized reproductive cells, which undergo meiosis to produce smaller, motile spermatozoa or larger, non-motile ova.[17] These fuse to form zygotes, which develop into new individuals.[18]
Many animals are also capable of asexual reproduction.[19] This may take place through parthenogenesis, where fertile eggs are produced without mating, budding, or fragmentation.[20]
A zygote initially develops into a hollow sphere, called a blastula,[21] which undergoes rearrangement and differentiation. In sponges, blastula larvae swim to a new location and develop into a new sponge.[22] In most other groups, the blastula undergoes more complicated rearrangement.[23] It first invaginates to form a gastrula with a digestive chamber, and two separate germ layers — an external ectoderm and an internal endoderm.[24] In most cases, a mesoderm also develops between them.[25] These germ layers then differentiate to form tissues and organs.[26]
All animals are heterotrophs, meaning that they feed directly or indirectly on other living things.[27] They are often further subdivided into groups such as carnivores, herbivores, omnivores, and parasites.[28]
Predation is a biological interaction where a predator (a heterotroph that is hunting) feeds on its prey (the organism that is attacked).[29] Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of the prey.[30] The other main category of consumption is detritivory, the consumption of dead organic matter.[31] It can at times be difficult to separate the two feeding behaviours, for example, where parasitic species prey on a host organism and then lay their eggs on it for their offspring to feed on its decaying corpse. Selective pressures imposed on one another has led to an evolutionary arms race between prey and predator, resulting in various antipredator adaptations.[32]
Most animals indirectly use the energy of sunlight by eating plants or plant-eating animals. Most plants use light to convert inorganic molecules in their environment into carbohydrates, fats, proteins and other biomolecules, characteristically containing reduced carbon in the form of carbon-hydrogen bonds. Starting with carbon dioxide (CO2) and water (H2O), photosynthesis converts the energy of sunlight into chemical energy in the form of simple sugars (e.g., glucose), with the release of molecular oxygen. These sugars are then used as the building blocks for plant growth, including the production of other biomolecules.[10] When an animal eats plants (or eats other animals which have eaten plants), the reduced carbon compounds in the food become a source of energy and building materials for the animal.[33] They are either used directly to help the animal grow, or broken down, releasing stored solar energy, and giving the animal the energy required for motion.[34][35]
Animals living close to hydrothermal vents and cold seeps on the ocean floor are not dependent on the energy of sunlight.[36] Instead chemosynthetic archaea and bacteria form the base of the food chain.[37]
Animals are generally considered to have evolved from a flagellated eukaryote.[39] Their closest known living relatives are the choanoflagellates, collared flagellates that have a morphology similar to the choanocytes of certain sponges.[40] Molecular studies place animals in a supergroup called the opisthokonts, which also include the choanoflagellates, fungi and a few small parasitic protists.[41] The name comes from the posterior location of the flagellum in motile cells, such as most animal spermatozoa, whereas other eukaryotes tend to have anterior flagella.[42]
The first fossils that might represent animals appear in the Trezona Formation at Trezona Bore, West Central Flinders, South Australia.[43] These fossils are interpreted as being early sponges. They were found in 665-million-year-old rock.[43]
The next oldest possible animal fossils are found towards the end of the Precambrian, around 610 million years ago, and are known as the Ediacaran or Vendian biota.[44] These are difficult to relate to later fossils, however. Some may represent precursors of modern phyla, but they may be separate groups, and it is possible they are not really animals at all.[45]
Aside from them, most known animal phyla make a more or less simultaneous appearance during the Cambrian period, about 542 million years ago.[46] It is still disputed whether this event, called the Cambrian explosion, represents a rapid divergence between different groups or a change in conditions that made fossilization possible.
Some paleontologists suggest that animals appeared much earlier than the Cambrian explosion, possibly as early as 1 billion years ago.[47] Trace fossils such as tracks and burrows found in the Tonian era indicate the presence of triploblastic worms, like metazoans, roughly as large (about 5 mm wide) and complex as earthworms.[48] During the beginning of the Tonian period around 1 billion years ago, there was a decrease in Stromatolite diversity, which may indicate the appearance of grazing animals, since stromatolite diversity increased when grazing animals went extinct at the End Permian and End Ordovician extinction events, and decreased shortly after the grazer populations recovered. However the discovery that tracks very similar to these early trace fossils are produced today by the giant single-celled protist Gromia sphaerica casts doubt on their interpretation as evidence of early animal evolution.[49][50]
It has been estimated that 99.9% of animals that have ever existed are extinct.[51]
Phylogenetic analysis suggests that the Porifera and Ctenophora diverged before a clade that gave rise to the Bilateria, Cnidaria and Placozoa.[52]
The sponges (Porifera) were long thought to have diverged from other animals early.[53] They lack the complex organization found in most other phyla.[54] Their cells are differentiated, but in most cases not organized into distinct tissues.[55] Sponges typically feed by drawing in water through pores.[56] Archaeocyatha, which have fused skeletons, may represent sponges or a separate phylum.[57] However, a phylogenomic study in 2008 of 150 genes in 29 animals across 21 phyla revealed that it is the Ctenophora or comb jellies which are the basal lineage of animals, at least among those 21 phyla. The authors speculate that sponges—or at least those lines of sponges they investigated—are not so primitive, but may instead be secondarily simplified.[58]
Among the other phyla, the Ctenophora and the Cnidaria, which includes sea anemones, corals, and jellyfish, are radially symmetric and have digestive chambers with a single opening, which serves as both the mouth and the anus.[59] Both have distinct tissues, but they are not organized into organs.[60] There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, these animals are sometimes called diploblastic.[61] The tiny placozoans are similar, but they do not have a permanent digestive chamber.
The remaining animals form a monophyletic group called the Bilateria. For the most part, they are bilaterally symmetric, and often have a specialized head with feeding and sensory organs. The body is triploblastic, i.e. all three germ layers are well-developed, and tissues form distinct organs. The digestive chamber has two openings, a mouth and an anus, and there is also an internal body cavity called a coelom or pseudocoelom. There are exceptions to each of these characteristics, however — for instance adult echinoderms are radially symmetric, and certain parasitic worms have extremely simplified body structures.
Genetic studies have considerably changed our understanding of the relationships within the Bilateria. Most appear to belong to two major lineages: the deuterostomes and the protostomes, the latter of which includes the Ecdysozoa, Platyzoa, and Lophotrochozoa. In addition, there are a few small groups of bilaterians with relatively similar structure that appear to have diverged before these major groups. These include the Acoelomorpha, Rhombozoa, and Orthonectida. The Myxozoa, single-celled parasites that were originally considered Protozoa, are now believed to have developed from the Medusozoa as well.
Deuterostomes differ from the other Bilateria, called protostomes, in several ways. In both cases there is a complete digestive tract. However, in protostomes, the first opening of the gut to appear in embryological development (the archenteron) develops into the mouth, with the anus forming secondarily. In deuterostomes the anus forms first, with the mouth developing secondarily.[62] In most protostomes, cells simply fill in the interior of the gastrula to form the mesoderm, called schizocoelous development, but in deuterostomes, it forms through invagination of the endoderm, called enterocoelic pouching.[63] Deuterostome embryos undergo radial cleavage during cell division, while protostomes undergo spiral cleavage.[64]
All this suggests the deuterostomes and protostomes are separate, monophyletic lineages. The main phyla of deuterostomes are the Echinodermata and Chordata.[65] The former are radially symmetric and exclusively marine, such as starfish, sea urchins, and sea cucumbers.[66] The latter are dominated by the vertebrates, animals with backbones.[67] These include fish, amphibians, reptiles, birds, and mammals.[68]
In addition to these, the deuterostomes also include the Hemichordata, or acorn worms.[69] Although they are not especially prominent today, the important fossil graptolites may belong to this group.[70]
The Chaetognatha or arrow worms may also be deuterostomes, but more recent studies suggest protostome affinities.
The Ecdysozoa are protostomes, named after the common trait of growth by moulting or ecdysis.[71] The largest animal phylum belongs here, the Arthropoda, including insects, spiders, crabs, and their kin. All these organisms have a body divided into repeating segments, typically with paired appendages. Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits.
The ecdysozoans also include the Nematoda or roundworms, perhaps the second largest animal phylum. Roundworms are typically microscopic, and occur in nearly every environment where there is water.[72] A number are important parasites.[73] Smaller phyla related to them are the Nematomorpha or horsehair worms, and the Kinorhyncha, Priapulida, and Loricifera. These groups have a reduced coelom, called a pseudocoelom.
The remaining two groups of protostomes are sometimes grouped together as the Spiralia, since in both embryos develop with spiral cleavage.
The Platyzoa include the phylum Platyhelminthes, the flatworms.[74] These were originally considered some of the most primitive Bilateria, but it now appears they developed from more complex ancestors.[75] A number of parasites are included in this group, such as the flukes and tapeworms.[74] Flatworms are acoelomates, lacking a body cavity, as are their closest relatives, the microscopic Gastrotricha.[76]
The other platyzoan phyla are mostly microscopic and pseudocoelomate. The most prominent are the Rotifera or rotifers, which are common in aqueous environments. They also include the Acanthocephala or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and possibly the Cycliophora.[77] These groups share the presence of complex jaws, from which they are called the Gnathifera.
The Lophotrochozoa include two of the most successful animal phyla, the Mollusca and Annelida.[78][79] The former, which is the second-largest animal phylum by number of described species, includes animals such as snails, clams, and squids, and the latter comprises the segmented worms, such as earthworms and leeches. These two groups have long been considered close relatives because of the common presence of trochophore larvae, but the annelids were considered closer to the arthropods because they are both segmented.[80] Now, this is generally considered convergent evolution, owing to many morphological and genetic differences between the two phyla.[81]
The Lophotrochozoa also include the Nemertea or ribbon worms, the Sipuncula, and several phyla that have a ring of ciliated tentacles around the mouth, called a lophophore.[82] These were traditionally grouped together as the lophophorates.[83] but it now appears that the lophophorate group may be paraphyletic,[84] with some closer to the nemerteans and some to the molluscs and annelids.[85][86] They include the Brachiopoda or lamp shells, which are prominent in the fossil record, the Entoprocta, the Phoronida, and possibly the Bryozoa or moss animals.[87]
Because of the great diversity found in animals, it is more economical for scientists to study a small number of chosen species so that connections can be drawn from their work and conclusions extrapolated about how animals function in general. Because they are easy to keep and breed, the fruit fly Drosophila melanogaster and the nematode Caenorhabditis elegans have long been the most intensively studied metazoan model organisms, and were among the first life-forms to be genetically sequenced. This was facilitated by the severely reduced state of their genomes, but as many genes, introns, and linkages lost, these ecdysozoans can teach us little about the origins of animals in general. The extent of this type of evolution within the superphylum will be revealed by the crustacean, annelid, and molluscan genome projects currently in progress. Analysis of the starlet sea anemone genome has emphasised the importance of sponges, placozoans, and choanoflagellates, also being sequenced, in explaining the arrival of 1500 ancestral genes unique to the Eumetazoa.[88]
An analysis of the homoscleromorph sponge Oscarella carmela also suggests that the last common ancestor of sponges and the eumetazoan animals was more complex than previously assumed.[89]
Other model organisms belonging to the animal kingdom include the house mouse (Mus musculus) and zebrafish (Danio rerio).
Aristotle divided the living world between animals and plants, and this was followed by Carolus Linnaeus (Carl von Linné), in the first hierarchical classification.[90] Since then biologists have begun emphasizing evolutionary relationships, and so these groups have been restricted somewhat. For instance, microscopic protozoa were originally considered animals because they move, but are now treated separately.
In Linnaeus's original scheme, the animals were one of three kingdoms, divided into the classes of Vermes, Insecta, Pisces, Amphibia, Aves, and Mammalia. Since then the last four have all been subsumed into a single phylum, the Chordata, whereas the various other forms have been separated out. The above lists represent our current understanding of the group, though there is some variation from source to source.
This section requires expansion. (October 2012) |
Animals are regarded by philosophers like Peter Singer as ethical subjects. Animal rights movements have worked to promote animal welfare.
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リンク元 | 「animal」 |
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