46-474: Incertae sedis Holozoa (from Ancient Greek ὅλος (holos) 'whole' and ζῷον (zoion) 'animal') is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa (or animals) and
92-468: A paper published in the journal Current Biology . The study used complete mitochondrial genomes of a choanoflagellate ( Monosiga brevicollis ) and an ichthyosporean ( Amoebidium parasiticum ) to firmly resolve the position of Ichthyosporea as the sister group to Choanoflagellata+Metazoa. This clade was named Holozoa (from Ancient Greek ὅλος (holos) 'whole' and ζῷον (zoion) 'animal'), meaning 'whole animal', referencing
138-440: A paraphyletic group rather than a true clade. Modern cladistic approaches to eukaryotic classification prioritise monophyletic groupings over traditional ranks, which are increasingly perceived as redundant and superfluous. Because Holozoa is a clade, its use is preferred over the paraphyletic taxon Choanofila. Ancient Greek language Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes
184-537: A pitch accent . In Modern Greek, all vowels and consonants are short. Many vowels and diphthongs once pronounced distinctly are pronounced as /i/ ( iotacism ). Some of the stops and glides in diphthongs have become fricatives , and the pitch accent has changed to a stress accent . Many of the changes took place in the Koine Greek period. The writing system of Modern Greek, however, does not reflect all pronunciation changes. The examples below represent Attic Greek in
230-519: A clade that contains fungi and their protist relatives ( Holomycota ), are part of the larger supergroup of eukaryotes known as Opisthokonta . Holozoa diverged from their opisthokont ancestor around 1070 million years ago (Mya). The choanoflagellates, animals and filastereans group together as the clade Filozoa . Within Filozoa, the choanoflagellates and animals group together as the clade Choanozoa . Based on phylogenetic and phylogenomic analyses,
276-404: A crucial role in understanding the evolutionary steps leading to the emergence of multicellular animals from single-celled ancestors. Recent genomic studies have shed light on the evolutionary relationships between the various holozoan lineages , revealing insights into the origins of multicellularity . Some fossils of possible metazoans have been reinterpreted as holozoan protists. Holozoa
322-472: A lack of contemporaneous evidence. Several theories exist about what Hellenic dialect groups may have existed between the divergence of early Greek-like speech from the common Proto-Indo-European language and the Classical period. They have the same general outline but differ in some of the detail. The only attested dialect from this period is Mycenaean Greek , but its relationship to the historical dialects and
368-419: A lesser degree. Pamphylian Greek , spoken in a small area on the southwestern coast of Anatolia and little preserved in inscriptions, may be either a fifth major dialect group, or it is Mycenaean Greek overlaid by Doric, with a non-Greek native influence. Regarding the speech of the ancient Macedonians diverse theories have been put forward, but the epigraphic activity and the archaeological discoveries in
414-543: A prefix /e-/, called the augment . This was probably originally a separate word, meaning something like "then", added because tenses in PIE had primarily aspectual meaning. The augment is added to the indicative of the aorist, imperfect, and pluperfect, but not to any of the other forms of the aorist (no other forms of the imperfect and pluperfect exist). The two kinds of augment in Greek are syllabic and quantitative. The syllabic augment
460-671: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek , and Koine may be classified as Ancient Greek in a wider sense. There were several regional dialects of Ancient Greek; Attic Greek developed into Koine. Ancient Greek was a pluricentric language , divided into many dialects. The main dialect groups are Attic and Ionic , Aeolic , Arcadocypriot , and Doric , many of them with several subdivisions. Some dialects are found in standardized literary forms in literature , while others are attested only in inscriptions. There are also several historical forms. Homeric Greek
506-548: A significant discovery: many genetic characteristics previously thought as unique to animals can also be found in these unicellular relatives. This suggests that the origin of multicellular animals did not happen solely because of the appearance of new genes (i.e. innovation), but because of pre-existing genes that were adapted or utilized in new ways (i.e. co-option). For example: Additionally, many biological processes seen in animals are already present in their unicellular relatives, such as sexual reproduction and gametogenesis in
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#1732884365731552-609: A standard subject of study in educational institutions of the Western world since the Renaissance . This article primarily contains information about the Epic and Classical periods of the language, which are the best-attested periods and considered most typical of Ancient Greek. From the Hellenistic period ( c. 300 BC ), Ancient Greek was followed by Koine Greek , which is regarded as
598-510: A vowel or /n s r/ ; final stops were lost, as in γάλα "milk", compared with γάλακτος "of milk" (genitive). Ancient Greek of the classical period also differed in both the inventory and distribution of original PIE phonemes due to numerous sound changes, notably the following: The pronunciation of Ancient Greek was very different from that of Modern Greek . Ancient Greek had long and short vowels ; many diphthongs ; double and single consonants; voiced, voiceless, and aspirated stops ; and
644-453: Is a clade that includes animals and their closest relatives, as well as their common ancestor , but excludes fungi . It is defined on a branch-based approach as the clade encompassing all relatives of Homo sapiens (an animal), but not Neurospora crassa (a fungus ). Holozoa, besides animals, primarily comprises unicellular protist lineages of varied morphologies such as choanoflagellates , filastereans , ichthyosporeans , and
690-556: Is a literary form of Archaic Greek (derived primarily from Ionic and Aeolic) used in the epic poems , the Iliad and the Odyssey , and in later poems by other authors. Homeric Greek had significant differences in grammar and pronunciation from Classical Attic and other Classical-era dialects. The origins, early form and development of the Hellenic language family are not well understood because of
736-577: Is a potential new holozoan lineage. It has been tentatively named MASHOL (for 'marine small Holozoa'). The quest to elucidate the evolutionary origins of animals from a unicellular ancestor requires an examination of the transition to multicellularity . In the absence of a fossil record documenting this evolution, insights into the unicellular ancestor of animals are obtained from the analysis of shared genes and genetic pathways between animals and their closest living unicellular relatives. The genetic content of these single-celled holozoans has revealed
782-418: Is added to stems beginning with consonants, and simply prefixes e (stems beginning with r , however, add er ). The quantitative augment is added to stems beginning with vowels, and involves lengthening the vowel: Some verbs augment irregularly; the most common variation is e → ei . The irregularity can be explained diachronically by the loss of s between vowels, or that of the letter w , which affected
828-644: Is called 'East Greek'. Arcadocypriot apparently descended more closely from the Mycenaean Greek of the Bronze Age. Boeotian Greek had come under a strong Northwest Greek influence, and can in some respects be considered a transitional dialect, as exemplified in the poems of the Boeotian poet Pindar who wrote in Doric with a small Aeolic admixture. Thessalian likewise had come under Northwest Greek influence, though to
874-422: Is comparable to the germination stage of non-animal holozoans. They possibly represent an evolutionary grade in which palintomic cleavage (i.e. rapid cell divisions without cytoplasmic growth in between, a characteristic of animal embryonic cleavage ) was the method of dispersal and propagation . Prior to 2002, a relationship between Choanoflagellata , Ichthyosporea and the animal - fungi divergence
920-448: Is considered by some linguists to have been closely related to Greek . Among Indo-European branches with living descendants, Greek is often argued to have the closest genetic ties with Armenian (see also Graeco-Armenian ) and Indo-Iranian languages (see Graeco-Aryan ). Ancient Greek differs from Proto-Indo-European (PIE) and other Indo-European languages in certain ways. In phonotactics , ancient Greek words could end only in
966-745: The Greek region of Macedonia during the last decades has brought to light documents, among which the first texts written in Macedonian , such as the Pella curse tablet , as Hatzopoulos and other scholars note. Based on the conclusions drawn by several studies and findings such as Pella curse tablet , Emilio Crespo and other scholars suggest that ancient Macedonian was a Northwest Doric dialect , which shares isoglosses with its neighboring Thessalian dialects spoken in northeastern Thessaly . Some have also suggested an Aeolic Greek classification. The Lesbian dialect
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#17328843657311012-532: The choanoflagellate Salpingoeca rosetta and several types of multi cellular differentiation . A billion -year-old freshwater microscopic fossil named Bicellum brasieri is possibly the earliest known holozoan. It shows two differentiated cell types or life cycle stages . It consists of a spherical ball of tightly packed cells (stereoblasts) enclosed in a single layer of elongated cells . There are also two populations of stereoblasts with mixed shapes, which have been interpreted as cellular migration to
1058-417: The cladogram of Holozoa is shown below: Cristidiscoidea [REDACTED] Fungi [REDACTED] Dermocystida [REDACTED] Ichthyophonida [REDACTED] Corallochytrium [REDACTED] Syssomonas [REDACTED] Tunicaraptor Filasterea [REDACTED] Choanoflagellata [REDACTED] Metazoa [REDACTED] Uncertainty remains around the relationship of
1104-501: The present , future , and imperfect are imperfective in aspect; the aorist , present perfect , pluperfect and future perfect are perfective in aspect. Most tenses display all four moods and three voices, although there is no future subjunctive or imperative. Also, there is no imperfect subjunctive, optative or imperative. The infinitives and participles correspond to the finite combinations of tense, aspect, and voice. The indicative of past tenses adds (conceptually, at least)
1150-463: The protists Choanoflagellata , Filasterea , Pluriformea and Ichthyosporea . Along with fungi and some other groups, Holozoa is part of the Opisthokonta , a supergroup of eukaryotes . Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic . The holozoan protists play
1196-1031: The 5th century BC. Ancient pronunciation cannot be reconstructed with certainty, but Greek from the period is well documented, and there is little disagreement among linguists as to the general nature of the sounds that the letters represent. /oː/ raised to [uː] , probably by the 4th century BC. Greek, like all of the older Indo-European languages , is highly inflected. It is highly archaic in its preservation of Proto-Indo-European forms. In ancient Greek, nouns (including proper nouns) have five cases ( nominative , genitive , dative , accusative , and vocative ), three genders ( masculine , feminine , and neuter ), and three numbers (singular, dual , and plural ). Verbs have four moods ( indicative , imperative , subjunctive , and optative ) and three voices (active, middle, and passive ), as well as three persons (first, second, and third) and various other forms. Verbs are conjugated through seven combinations of tenses and aspect (generally simply called "tenses"):
1242-490: The Archaic period of ancient Greek (see Homeric Greek for more details): Μῆνιν ἄειδε, θεά, Πηληϊάδεω Ἀχιλῆος οὐλομένην, ἣ μυρί' Ἀχαιοῖς ἄλγε' ἔθηκε, πολλὰς δ' ἰφθίμους ψυχὰς Ἄϊδι προΐαψεν ἡρώων, αὐτοὺς δὲ ἑλώρια τεῦχε κύνεσσιν οἰωνοῖσί τε πᾶσι· Διὸς δ' ἐτελείετο βουλή· ἐξ οὗ δὴ τὰ πρῶτα διαστήτην ἐρίσαντε Ἀτρεΐδης τε ἄναξ ἀνδρῶν καὶ δῖος Ἀχιλλεύς. The beginning of Apology by Plato exemplifies Attic Greek from
1288-656: The Classical period of ancient Greek. (The second line is the IPA , the third is transliterated into the Latin alphabet using a modern version of the Erasmian scheme .) Ὅτι [hóti Hóti μὲν men mèn ὑμεῖς, hyːmêːs hūmeîs, Dermocystida Dermocystida is an order of parasitic eukaryotes . Sphaerothecum Dermocystidium Valentines Rhinosporidium Amphibiocystidium Dermotheca Dermosporidium This Holozoa -related article
1334-550: The aorist. Following Homer 's practice, the augment is sometimes not made in poetry , especially epic poetry. The augment sometimes substitutes for reduplication; see below. Almost all forms of the perfect, pluperfect, and future perfect reduplicate the initial syllable of the verb stem. (A few irregular forms of perfect do not reduplicate, whereas a handful of irregular aorists reduplicate.) The three types of reduplication are: Irregular duplication can be understood diachronically. For example, lambanō (root lab ) has
1380-419: The augment when it was word-initial. In verbs with a preposition as a prefix, the augment is placed not at the start of the word, but between the preposition and the original verb. For example, προσ(-)βάλλω (I attack) goes to προσ έ βαλoν in the aorist. However compound verbs consisting of a prefix that is not a preposition retain the augment at the start of the word: αὐτο(-)μολῶ goes to ηὐ τομόλησα in
1426-438: The center of Greek scholarship, this division of people and language is quite similar to the results of modern archaeological-linguistic investigation. One standard formulation for the dialects is: West vs. non-West Greek is the strongest-marked and earliest division, with non-West in subsets of Ionic-Attic (or Attic-Ionic) and Aeolic vs. Arcadocypriot, or Aeolic and Arcado-Cypriot vs. Ionic-Attic. Often non-West
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1472-434: The classification of eukaryotes. In classifications that use traditional taxonomic ranks (e.g. kingdom, phylum, class), all holozoan protists are classified as subphylum Choanofila (phylum Choanozoa , kingdom Protozoa ) while the animals are classified as a separate kingdom Metazoa or Animalia. This classification excludes animals, even though they descend from the same common ancestor as choanofilan protists, making it
1518-611: The dialect of Sparta ), and Northern Peloponnesus Doric (including Corinthian ). All the groups were represented by colonies beyond Greece proper as well, and these colonies generally developed local characteristics, often under the influence of settlers or neighbors speaking different Greek dialects. After the conquests of Alexander the Great in the late 4th century BC, a new international dialect known as Koine or Common Greek developed, largely based on Attic Greek , but with influence from other dialects. This dialect slowly replaced most of
1564-481: The distinct genera Corallochytrium , Syssomonas , and Tunicaraptor . The first sequenced unicellular holozoan genome was that of Monosiga brevicollis , a choanoflagellate . It measures around 41.6 mega–base-pairs (Mbp) and contains around 9200 coding genes , making it comparable in size to the genome of filamentous fungi . Animal genomes are usually larger (e.g. human genome , 2900 Mbp; fruit fly , 180 Mbp), with some exceptions. Holozoa, along with
1610-663: The forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek ( c. 1400–1200 BC ), Dark Ages ( c. 1200–800 BC ), the Archaic or Epic period ( c. 800–500 BC ), and the Classical period ( c. 500–300 BC ). Ancient Greek was the language of Homer and of fifth-century Athenian historians, playwrights, and philosophers . It has contributed many words to English vocabulary and has been
1656-556: The historical Dorians . The invasion is known to have displaced population to the later Attic-Ionic regions, who regarded themselves as descendants of the population displaced by or contending with the Dorians. The Greeks of this period believed there were three major divisions of all Greek people – Dorians, Aeolians, and Ionians (including Athenians), each with their own defining and distinctive dialects. Allowing for their oversight of Arcadian, an obscure mountain dialect, and Cypriot, far from
1702-472: The historical circumstances of the times imply that the overall groups already existed in some form. Scholars assume that major Ancient Greek period dialect groups developed not later than 1120 BC, at the time of the Dorian invasions —and that their first appearances as precise alphabetic writing began in the 8th century BC. The invasion would not be "Dorian" unless the invaders had some cultural relationship to
1748-490: The newest holozoan member, is still unresolved. Three different phylogenetic positions of Tunicaraptor have been obtained from analyses: as the sister group to Filasterea , as sister to Filozoa , or as the most basal group of all Holozoa. Environmental DNA surveys of oceans have revealed new diverse lineages of Holozoa. Most of them nest within known groups, mainly Ichthyosporea and Choanoflagellata . However, one environmental clade does not nest within any known group and
1794-499: The older dialects, although the Doric dialect has survived in the Tsakonian language , which is spoken in the region of modern Sparta. Doric has also passed down its aorist terminations into most verbs of Demotic Greek . By about the 6th century AD, the Koine had slowly metamorphosed into Medieval Greek . Phrygian is an extinct Indo-European language of West and Central Anatolia , which
1840-487: The perfect stem eilēpha (not * lelēpha ) because it was originally slambanō , with perfect seslēpha , becoming eilēpha through compensatory lengthening. Reduplication is also visible in the present tense stems of certain verbs. These stems add a syllable consisting of the root's initial consonant followed by i . A nasal stop appears after the reduplication in some verbs. The earliest extant examples of ancient Greek writing ( c. 1450 BC ) are in
1886-711: The periphery, a movement that could be explained by differential cell-cell adhesion . These occurrences are consistent with extant unicellular holozoans, which are known to form multicellular stages in complex life cycles. Proposed Ediacaran fossil " embryos " of early metazoans , discovered in the Doushantuo Formation , have been reinterpreted as non-animal protists within Holozoa. According to some authors, although they present possible embryonic cleavage , they lack metazoan synapomorphies such as tissue differentiation and nearby juveniles or adults. Instead, its development
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1932-517: The syllabic script Linear B . Beginning in the 8th century BC, however, the Greek alphabet became standard, albeit with some variation among dialects. Early texts are written in boustrophedon style, but left-to-right became standard during the classic period. Modern editions of ancient Greek texts are usually written with accents and breathing marks , interword spacing , modern punctuation , and sometimes mixed case , but these were all introduced later. The beginning of Homer 's Iliad exemplifies
1978-493: The two most basal groups, Ichthyosporea and Pluriformea . They may be sister to each other, forming the putative clade Teretosporea . Alternatively, Ichthyosporea may be the earliest-branching of the two, while Pluriformea is sister to the Filozoa clade comprising filastereans, choanoflagellates and animals. This second outcome is more strongly supported after the discovery of Syssomonas . The position of Tunicaraptor ,
2024-479: The wider animal ancestry that it contains. Holozoa has since been supported as a robust clade by every posterior analysis, even after the discovery of more taxa nested within it (namely Filasterea since 2008, and the pluriformean species Corallochytrium and Syssomonas since 2014 and 2017 respectively). As of 2019, the clade is accepted by the International Society of Protistologists, which revises
2070-475: Was Aeolic. For example, fragments of the works of the poet Sappho from the island of Lesbos are in Aeolian. Most of the dialect sub-groups listed above had further subdivisions, generally equivalent to a city-state and its surrounding territory, or to an island. Doric notably had several intermediate divisions as well, into Island Doric (including Cretan Doric ), Southern Peloponnesus Doric (including Laconian ,
2116-499: Was considered on the basis of morphology and ultrastructure . Early phylogenetic analyses gave contradicting results, because the amount of available DNA sequences was insufficient to yield unambiguous results. The taxonomic uncertainty was such that, for example, some Ichthyosporea were traditionally treated as trichomycete fungi. Holozoa was first recognized as a clade in 2002 through a phylogenomic analysis by Franz Bernd Lang, Charles J. O'Kelly and other collaborators, as part of
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