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Phacus

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494:, although some species have two, which emerges anteriorly. The flagellum is responsible for cell movement by gyrating in the direction of travel allowing the cell to glide and swim in the water. Furthermore, some flagella vary in length from short all the way to the length of the cell in other species. Although many members of the genus have two flagella, only one is ever used for movement. The other one is usually too short and does not exit the invagination of the posterior area known as the flagellar pocket. They are located within a posterior structure called the flagellar apparatus, also known as the 502:
body is referred to as paramylon and functions as a storage substance. The paramylon is a carbohydrate energy store reserve and it is quite different from other algal carbohydrate stores due to the high amounts of crystallization present. The nucleus of the cell is generally positioned towards the middle of the cell and is adjacent to the paramylon reserve. Within the nucleus are permanently condensed chromosomes, which can easily be viewed under a light microscope. Just like other euglenids, several species of
54: 463:, the MTR is a microtubule organizing center and is connected to a reservoir membrane by a striated fiber. Many species also possess an elongated caudal process with extended pellicle strips. Throughout the cytoplasm of the cell, chloroplasts are spread out fairly evenly and in high numbers. They are typically discoid and are regularly disposed of and replaced. Chloroplasts that are present in large numbers are typically smaller, are without 33: 384:), and the overall structure of the cells. Regardless of a large or small change in organic enrichment, studies show a consistency to these morphological changes. However, the amount of change that occurs varies between species and is dependent on the specific organic nutrients present. If the amount of organic nutrients in the genus’ habitat is insufficient, occasionally they form resting 448:
its rigid cytoskeleton (although some species have semi-rigid or plastic cytoskeleton) made up of pellicular strips and its predominantly flat, leaf-shaped structure. Many different species express secondary fusion of these pellicular strips and many of the strips have a variety of shapes including S-shaped, A-shaped, M-shaped or plateau.
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arrangement (either helical or longitudinal) has also changed throughout evolution. Furthermore, it has been argued that certain behavioral and locomotor traits which previously existed for predatory feeding have no longer been selected. This seemed to have had an effect on the number of strips species of
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have physiological traits that closely resemble other related genera. Even the most minor difference in certain physiological properties caused great confusion in determining the proper classification for many species. This occurred more prominently during the time of Dujardin because the methodology
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unicellular organisms, meaning that they are capable of producing their own food. Although the genus primarily receives their nutrients through photosynthesis, they are also capable of feeding on certain kinds of alga and bacteria using a feeding apparatus located on their underside. Many species of
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Being in an organically enriched freshwater environment is essential for the development of these species. Different studies have shown that the addition or removal of certain organic elements can have profound effects on cell development. In studies using beef extract to increase organic content of
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as being monophyletic depending on the clade studied, but the general consensus through multiple repeated analyses is that the genus is polyphyletic. This possible discrepancy has caused certain scientists to suggest possibly redefining this genus as two separate genera. In accordance with that, it
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Aside from the flagella, the flagellar apparatus also contains two basal bodies connected by a striated fiber, three asymmetric microtubular roots, and other connective fibers. The genus only has one large anisotropic body unlike many other euglenid genera, which commonly have two. This anisotropic
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within the phylum Euglenozoa, though the majority of genera within the phylum actually do not contain chloroplasts and are colourless. Generally these species are small, free swimming and exhibit a vibrant green colour. What sets the genus apart from other photosynthetic species is the presence of
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of ancestral species with the rigid pellicles of their descendants. Another feature believed to have evolved, are the longitudinal strips that appear on most species. It appears that the number of those strips has either increased or decreased over time depending on certain species and that their
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Other differences among species include: the presence or lack of haplopyrenoids within the chloroplasts, position of the nucleus, a large or small endosome, shape of the cytoskeleton, few to several paramylon discoid grains, the presence of lateral caps and presence of oblique truncated poles. In
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generally have. The changing numbers of strips and the clustered patterns associated are not actually adaptive themselves, but may have evolved due to the cell becoming more flat and more rigid over time. Those traits are believed to have evolved in order to adapt to a more planktonic lifestyle.
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are commonly found in freshwater habitats all over the world. Many species of this genus have been discovered in several countries, including Japan, the United States, Portugal, Brazil, Korea and the Philippines. Different members of the genus have been found in temperatures ranging from 11.4 to
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in the database, but only 171 have been accepted taxonomically. The genus was established in 1841 and since then major discoveries have led it to become an extremely large group containing hundreds of species with varied physiological characteristics. Contemporary studies agree that
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Su, H M; Won, L J; Ho, G Y (27 September 2013). "Bacterial mixture useful for controlling proliferation of harmful algae including Microcystis, Anabaena, Aphanizomenon, Phacus, Euglena, Peridinium, Cyclotella, Chlamydomonas and Rhodomonas, comprises Lactobacillus and Lactococcus".
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actually lack a caudal process entirely. Those species are described as having rounded posteriors in place of the caudal process. Studies show that morphological changes observed in the genus are possibly due to the level of organic enrichment in their freshwater
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Kosmala, Sylwia; Bereza, Magdalena; Milanowski, Rafal; Kwiatowski, Jan; Zakrys, Bozena (October 2007). "Morphological and molecular examination of relationships and epitype establishment of Phacus pleuronectes, Phacus orbicularis, and Phacus hamelii".
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Kosmala, Sylwia; Bereza, Magdalena; Milanowski, Rafał; Kwiatowski, Jan; Zakryś, Bożena (2007-10-01). "Morphological and molecular examination of relationships and epitype establishment of Phacus pleuronectes, Phacus orbicularis, and Phacus hamelii1".
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known as a pellicle. These eukaryotes are mostly green in colour, and have a single flagellum that extends the length of their body. They are morphologically very flat, rigid, leaf-shaped, and contain many small discoid chloroplasts.
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is fully complete, the cells remain attached to one another, forming what looks like a “two-headed” organism. Prior to cytokinesis, the amount of pellicle strips each cell has is doubled in order to have an equal number between each
279:, but they all had different diagnostic criteria when classifying these organisms. Originally, Ehrenberg tried to classify three species he discovered as Euglena because of their color, but ultimately were moved to the genus 275:(Ehrenberg) was one of the first species of the genus to be discovered (1830) and was used as the holotype species when describing the genus. Ehrenberg, Hubner, Dujardin, and Klebs were some of the earliest researchers of 295:
and technology used at the time in studying these organisms were far more simplistic compared to today, leading to greater difficulty in determining critical differences between organisms. Since its inception,
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organisms are found in a range of freshwater environments (some more acidic or alkaline than others), prefer cooler temperatures, and on average exist in more neutral pH aquatic habitats. Many species of
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Milanowski, Rafał; Kosmala, Sylwia; Zakryś, Bozżena; Kwiatowski, Jan (2006-06-01). "Phylogeny of Photosynthetic Euglenophytes Based on Combined Chloroplast and Cytoplasmic SSU rDNA Sequence Analysis".
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Nannavecchia, Paula; Tolivia, Analia; Conforti, Visitacion (March 2014). "Ultrastructural alterations in Phacus brachykentron (Euglenophyta) due to excess of organic matter in the culture medium".
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Nudelman, MarĂ­a Alejandra; Rossi, MarĂ­a Susana; Conforti, VisitaciĂłn; Triemer, Richard E. (2003-02-01). "Phylogeny of Euglenophyceae Based on Small Subunit rDNA Sequences: Taxonomic Implications".
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or ingestion organelle, allowing the organism to feed when bacteria enter inside. The microtubules are arranged in a peculiar doublet and triplet pattern in the upper canal. In certain species of
261:. The reason behind the separation was in order to create a group that correctly organizes their established morphological characteristics such as their rigid, flat, leaf-shape and small discoid 1117:
Shin, Woongghi; Boo, Sung Min; Triemer, Richard E. (2001-10-01). "Ultrastructure of the Basal Body Complex and Putative Vestigial Feeding Apparatus in Phacus Pleuronectes (euglenophyceae)".
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Although the general morphology of the genus is considered to be well established, given the large number of species there are critical morphological differences observed worth noting. The
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Monfils, Anna K; Triemer, Richard E; Bellairs, Emily F (2011-02-28). "Characterization of paramylon morphological diversity in photosynthetic euglenoids (Euglenales, Euglenophyta)".
237:, meaning lentil or lens. This may be due to the general round or oval shape of the many species that are part of this genus. Its origins date back to the nineteenth century, in 756:
organisms are present, that is indicative of high organic pollution. This particular trait allows scientists to determine the health of different bodies of water.
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are generally found include swamps, ditches, trenches, ponds and even in many rice fields throughout North America and around the world. They are a small part of
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mucilaginous wall for protection until it enters a more habitable environment. In addition, cell division continues to take place even as a reproductive cyst.
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Pereira, Mário Jorge; Azeiteiro, Ulisses M.M (2003). "Ecological notes on the species of Phacus Dujardin (Euglenophyta) from the central region of Portugal".
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The literature mentioned above have also looked into the evolutionary history of the genus’ morphology. It is believed that the rigidity of the cells has
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has been debated. Many studies looked at the genes of its many species by examining small subunit rRNA (SSU) sequences and arranging certain species into
724:, like most euglenoids, is very scarce, and little information is actually known of their geographical origins. However, there have been reports that 530:). As well, the shape of the cell in some species are completely flat, while many have also been described as helically twisted, straight or curved. 1765: 1348:
Kim, Jong Im; Shin, Woongghi; Triemer, Richard E. (2010). "Multigene Analyses of Photosynthetic Euglenoids and New Family, Phacaceae (Euglenales)".
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were observed to have clear morphological changes different from the controls. These changes include: increasing thickness of the cell, increase in
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Esson, Heather J.; Leander, Brian S. (2010-01-01). "Evolution of Distorted Pellicle Patterns in Rigid Photosynthetic Euglenids (Phacus Dujardin)".
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Pereira, MJ; Azeiteiro, UMM (May 2003). "Ecological notes on the species of Phacus Dujardin (Euglenophyta) from the central region of Portugal".
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are considered to be euplanktonic (free-floating organisms or open water plankton) because they are commonly found together with other genera of
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stability. Their keen ability to colonize in huge numbers makes them able to survive and thrive in areas where certain algae cannot. However,
860: 241:, where it first received publication and establishment by Dujardin. The genus name is currently treated in literature as masculine. 290:
Since the establishment of the genus, there has been difficulty in classifying organisms due to morphology because many species of
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habitats around the globe and include several hundred species that continue to be discovered to this day. Currently, there are 564
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Dujardin, have also observed the presence of ellipsoid, tiny disk-shaped or flat-shaped paramylon grains. In some species of
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are known to be prey for a variety of marine and freshwater genera. The best known predators of the genus are planktonic
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are not common inhabitants of stagnant environments because those areas often do not have proper organic composition.
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A major genetic change in the genus occurred in its chloroplast genome throughout its evolution. This resulted in a
1878: 1822: 1888: 1262:Žižka, Zdeněk (2014-09-01). "Anisotropic structures of some microorganisms studied by polarization microscopy". 545:
due to the fusion of the genus’ pellicular strips. These varied morphological shapes make defining the genus as
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Conforti, VisitaciĂłn (1998-05-01). "Morphological changes of Euglenophyta in response to organic enrichment".
736:. These fossils, although not certain to belong to the genus, are estimated to be over 60 million years old. 1666: 451:
The pellicles forms a shell around the cytoskeleton covering the whole cell and fuses together around the
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is actually helical in shape throughout the entire cell as opposed to flat and leaf-shaped like most
284: 728:-like microfossils have been discovered from pyriform cells, which seem to closely resemble that of 1883: 1207:
Kasiborski, Beth A.; Bennett, Matthew S.; Linton, Eric W. (2016-06-01). "The chloroplast genome of
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Kim, Jong Im; Shin, Woongghi (2014-10-01). "Molecular Phylogeny and Cryptic Diversity of the Genus
519: 507: 617:. They do so by dividing their cells longitudinally, from the apex of the cell to the base. Until 1566: 1502: 1467: 1411: 1365: 1330: 1295: 1244: 1189: 1142: 1094: 999: 953: 909: 812: 699: 48: 1718: 271:
was one of the first scientists to discover and classify members of the genus; his discovery of
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looks like based on molecular factors. Certain molecular phylogenetic analyses have described
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in 1841. Dujardin first published collective findings of the genus in the scientific journal
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Metaboly, which is the ability of some organisms to alter their shape, is not possible in
168:(also known as Euglenophyta), characterized by its flat, leaf-shaped structure, and rigid 1177: 978:
Kim, Jun Tae; Boo, S. M.; Zakrys, B. (2000). "Contribution to the knowledge of the genus
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A history of Infusoria, including the Desmidiaceae and Diatomaceae, British and foreign
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Bradley, W. H. (1929). "Fresh water algae from the Green River Formation of Colorado".
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photosynthetic Euglenoids, which are a group of single celled, chloroplast-containing
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has undergone constant reclassification and considerable re-evaluation of different
1704: 1471: 1098: 679: 565:), and in others there are multiple plates with different morphologies that exist ( 169: 1731: 995: 1612: 1809: 1778: 1698: 618: 476: 471:, like all photosynthetic euglenoids, obtained their plastids through secondary 452: 336: 262: 180: 155: 1275: 1082: 495: 413: 165: 108: 75: 1689: 1455: 1407: 1283: 1232: 1185: 1138: 1090: 949: 905: 800: 1752: 1713: 749: 695: 491: 456: 444: 377: 205: 201: 197: 65: 1630: 1463: 1291: 1240: 808: 1683: 1053:
Dujardin in Illinois, Dissertation for the Degree of Doctor of Philosophy
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and species containing fewer chloroplasts tend to have much larger ones.
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by Dujardin due to the similarity they exhibited in other more prominent
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in certain bodies of water can actually indicate the level of organic
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numerous times, and that was determined by comparing the semi-rigid
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addition, the morphology of the caudal process in many species of
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or asymmetrical rather difficult. Scientists who study the genus
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in accordance with new morphological and molecular information.
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relationships within the genus are currently poorly understood.
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Leander, Brian S.; Farmer, Mark A. (2001-02-06). "Evolution of
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of the water. It has been observed that if a large number of
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divided early on in the history of photosynthetic euglenids.
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Diaptomus, Tropocyclops, Epischura, Daphnia, Diaphanosoma,
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Most of these organisms also possess a semi-rectangular
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The genus was first established by the French biologist
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reinforced-pocket (MTR). This pocket acts as a sort of
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The genus name is believed to have originated from the
1055:. Michigan: Southern Illinois University. p. 237. 475:, where the ancestral phagocytic euglenoid engulfed a 356:
communities, but do serve important purposes such as
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characteristics such as shape and strip arrangement.
1673: 678:- possibly due to gene loss or transfer to the 557:a single plate dominates the interiors of the 526:), and in others it is deep and longitudinal ( 334:. These species include members of the genera 834:"Algaebase :: Listing the World's Algae" 8: 255:Histoire naturelle des Zoophytes, Infusoires 1661: 982:Dujardin 1841 (Euglenophyceae) in Korea". 31: 20: 1620: 522:for example, in many species is shallow ( 479:, and the resulting organism became the 257:in 1841, separating them from the genus 764: 585:have very a blunt caudal process while 490:, often reddish in color, and a single 1601:Ecotoxicology and Environmental Safety 1487:Bulletin of the Torrey Botanical Club 1429: 1427: 1425: 1381: 1379: 1112: 1110: 1108: 7: 1833:7a09319b-6cbc-4990-aa0d-7fe30bb30601 1178:10.1046/j.1529-8817.2001.037001143.x 1068: 1066: 1064: 1062: 1015: 1013: 973: 971: 969: 967: 855: 853: 828: 826: 770: 768: 686:, and large genomic rearrangements. 1649:http://eol.org/pages/11710/overview 380:bodies (both in size and number in 1436:Journal of Eukaryotic Microbiology 732:or another closely related genus, 510:and have a red-pigmented eyespot. 372:certain cultures, some species of 14: 200:or holophyletic, but is actually 1563:10.1111/j.1529-8817.2007.00386.x 1448:10.1111/j.1550-7408.2009.00447.x 1400:10.1111/j.1529-8817.2006.00216.x 1362:10.1111/j.1529-8817.2010.00910.x 1131:10.1046/j.1529-8817.2001.01041.x 942:10.1111/j.1529-8817.2007.00386.x 898:10.1046/j.1529-8817.2003.02075.x 52: 1647:Phacus. (n.d.). Retrieved from 682:, an increase in the number of 439:consists of organisms that are 613:and other euglenids reproduce 1: 1592:10.1016/S1146-609X(03)00006-7 1034:10.1016/s1146-609x(03)00006-7 1653:Pritchard, A., et al. 1861: 1641:Kr2013009902-A; Kr1311837-B1 1613:10.1016/j.ecoenv.2013.12.007 1905: 1657:. London: Whitaker and co. 637:is a member of the family 1586:(Supplement 1): S33–S48. 1276:10.1007/s12223-014-0307-5 514:Differences in morphology 49:Scientific classification 47: 39: 30: 23: 1526:Nostoca Algae Laboratory 1049:A revision of the genus 1083:10.1023/a:1017049910481 996:10.1127/nova/71/2000/37 265:with absent pyrenoids. 226: 179:are commonly found in 577:is extremely varied. 416:, such as species of 319:between 6.2 and 7.5. 219: 204:. Unfortunately, the 1551:Journal of Phycology 1388:Journal of Phycology 1350:Journal of Phycology 1264:Folia Microbiologica 1213:Journal of Phycology 1166:Journal of Phycology 1119:Journal of Phycology 1077:. 369–370: 277–285. 1047:Weik, K. L. (1967). 930:Journal of Phycology 886:Journal of Phycology 781:Journal of Phycology 740:Practical importance 508:contractile vacuoles 315:21.6 Â°C, and a 245:History of knowledge 528:Phacus hordeiformis 307:Habitat and ecology 269:Christian Ehrenberg 1209:Phacus orbicularis 563:Phacus orbicularis 524:Phacus viridioryza 506:also possess many 227: 1879:Euglenozoa genera 1866: 1865: 1667:Taxon identifiers 1520:Brunn, K (2012). 1225:10.1111/jpy.12403 838:www.algaebase.org 793:10.1111/jpy.12227 662:is believed that 651:phylogenetic tree 567:Phacus curvicauda 532:Phacus helikoides 382:Phacus curvicauda 273:Phacus longicauda 143: 142: 139: 1896: 1889:Freshwater algae 1859: 1858: 1846: 1845: 1836: 1835: 1826: 1825: 1813: 1812: 1810:NHMSYS0000605213 1800: 1799: 1787: 1786: 1774: 1773: 1761: 1760: 1748: 1747: 1735: 1734: 1722: 1721: 1709: 1708: 1707: 1694: 1693: 1692: 1662: 1644: 1634: 1624: 1595: 1574: 1557:(5): 1071–1082. 1536: 1535: 1533: 1532: 1517: 1511: 1510: 1482: 1476: 1475: 1431: 1420: 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Index


Scientific classification
Edit this classification
Eukaryota
Euglenozoa
Euglenida
Euglenophyceae
Euglenales
Phacaceae
Phacus
Dujardin
genus
unicellular
excavates
phylum
Euglenozoa
cytoskeleton
freshwater
species
monophyletic
polyphyletic
phylogenetic

Greek
France
FĂ©lix Dujardin
chloroplasts
Christian Ehrenberg
morphological
taxa

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