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.
217:
703:
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
294:
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
407:
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
371:
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
661:
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
501:
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
447:
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
702:
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
572:
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
707:
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.
314:
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
191:
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
1638:
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".
593:
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
1548:
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".
927:
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".
172:
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.
621:
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,
388:. If this occurs, the cells would expand (swell) and become more rounded, and also lose their flagella. This increase in size forces the cell to increase the number of paramylon storage granules and develop a
323:
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
1386:
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".
1599:
Nannavecchia, Paula; Tolivia, Analia; Conforti, Visitacion (March 2014). "Ultrastructural alterations in Phacus brachykentron (Euglenophyta) due to excess of organic matter in the culture medium".
884:
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".
459:
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)".
518:
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
1313:
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.
352:
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
1817:
626:. In addition, each daughter cell will contain half the number of the newly formed strips and half the number of the old strips present prior to cell division.
392:
mucilaginous wall for protection until it enters a more habitable environment. In addition, cell division continues to take place even as a reproductive cyst.
1020:
Pereira, Mário Jorge; Azeiteiro, Ulisses M.M (2003). "Ecological notes on the species of Phacus
Dujardin (Euglenophyta) from the central region of Portugal".
694:
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
645:
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)".
376:
were observed to have clear morphological changes different from the controls. These changes include: increasing thickness of the cell, increase in
1434:
Esson, Heather J.; Leander, Brian S. (2010-01-01). "Evolution of Distorted Pellicle Patterns in Rigid Photosynthetic Euglenids (Phacus Dujardin)".
1791:
1578:
Pereira, MJ; Azeiteiro, UMM (May 2003). "Ecological notes on the species of Phacus Dujardin (Euglenophyta) from the central region of Portugal".
328:
are considered to be euplanktonic (free-floating organisms or open water plankton) because they are commonly found together with other genera of
364:
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
183:
habitats around the globe and include several hundred species that continue to be discovered to this day. Currently, there are 564
1850:
1521:
553:
Dujardin, have also observed the presence of ellipsoid, tiny disk-shaped or flat-shaped paramylon grains. In some species of
1804:
1796:
268:
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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
348:, and many different kinds of algae, which are typically found in similar aquatic habitats. The planktonic environments
641:, the order Euglenales, class Euglenoidea, and finally the phylum Euglenozoa. Certain ancestral information regarding
368:
are not common inhabitants of stagnant environments because those areas often do not have proper organic composition.
674:
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
53:
1073:
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
598:. These morphological differences, given the massive size of the genus, have led to certain confusion in
1855:
1832:
1739:
1654:
614:
534:
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
775:
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
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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
32:
1655:
A history of Infusoria, including the Desmidiaceae and Diatomaceae, British and foreign
1485:
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
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565:), and in others there are multiple plates with different morphologies that exist (
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476:
471:, like all photosynthetic euglenoids, obtained their plastids through secondary
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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
158:
135:
85:
779:(Phacaceae, Euglenophyceae) and the Descriptions of Seven New Species".
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in certain bodies of water can actually indicate the level of organic
1326:
1164:(Euglenophyceae) as Inferred From Pellicle Morphology and SSU rDNA".
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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.
1664:
1160:
Leander, Brian S.; Farmer, Mark A. (2001-02-06). "Evolution of
1744:
1522:"Algae can function as indicators of water pollution | WALPA"
1211:(Euglenophyceae): an initial datum point for the phacaceae".
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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.
649:. What those scientists were trying to determine is what the
833:
316:
418:
Diaptomus, Tropocyclops, Epischura, Daphnia, Diaphanosoma,
486:
Most of these organisms also possess a semi-rectangular
249:
The genus was first established by the French biologist
455:
reinforced-pocket (MTR). This pocket acts as a sort of
229:
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
287:
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
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7:
1833:7a09319b-6cbc-4990-aa0d-7fe30bb30601
1178:10.1046/j.1529-8817.2001.037001143.x
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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 °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
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1557:(5): 1071–1082.
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865:fmp.conncoll.edu
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1596:
1575:
1543:
1540:
1538:
1537:
1512:
1493:(8): 421–428.
1477:
1421:
1394:(3): 721–730.
1375:
1340:
1321:(2): 156–169.
1305:
1270:(5): 363–368.
1254:
1219:(3): 404–411.
1199:
1172:(1): 143–159.
1152:
1125:(5): 913–921.
1104:
1058:
1039:
1009:
990:(1–2): 37–68.
963:
919:
892:(1): 226–235.
876:
849:
822:
787:(5): 948–959.
763:
761:
758:
741:
738:
713:
712:Fossil history
710:
691:
688:
671:
668:
631:
628:
607:
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515:
512:
433:
430:
428:
425:
405:photosynthetic
397:
394:
390:polysaccharide
308:
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251:FĂ©lix Dujardin
246:
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220:
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99:Euglenophyceae
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59:
58:
45:
44:
37:
36:
28:
27:
16:Genus of algae
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1075:Hydrobiologia
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984:Nova Hedwigia
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718:fossil record
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473:endosymbiosis
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354:phytoplankton
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341:Trachelomonas
338:
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285:morphological
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1640:
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1600:
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1550:
1529:. Retrieved
1525:
1515:
1490:
1486:
1480:
1442:(1): 19–32.
1439:
1435:
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1257:
1216:
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885:
879:
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864:
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837:
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743:
733:
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436:
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381:
373:
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349:
335:
331:Euglenophyta
329:
325:
320:
311:
310:
296:
291:
289:
280:
276:
272:
267:
263:chloroplasts
258:
254:
248:
234:
228:
221:
206:phylogenetic
202:polyphyletic
198:monophyletic
193:
188:
176:
175:
170:cytoskeleton
146:
145:
144:
129:
128:
92:
40:
24:
18:
1779:iNaturalist
1699:Wikispecies
1622:11336/19526
1028:: S33–S48.
734:Lepocinclis
619:cytokinesis
547:symmetrical
538:organisms.
453:microtubule
441:microscopic
427:Description
422:Holopedium.
414:crustaceans
337:Lepocinclis
156:unicellular
1884:Euglenozoa
1873:Categories
1531:2017-07-25
1315:Phycologia
870:2017-07-25
843:2017-07-25
760:References
606:Life cycle
602:taxonomy.
496:basal body
477:green alga
445:autotrophs
432:Morphology
181:freshwater
166:Euglenozoa
109:Euglenales
76:Euglenozoa
1714:AlgaeBase
1607:: 36–41.
1456:1550-7408
1408:1529-8817
1284:0015-5632
1233:1529-8817
1186:1529-8817
1139:1529-8817
1091:0018-8158
950:1529-8817
906:1529-8817
801:1529-8817
750:pollution
700:pellicles
690:Evolution
639:Phacaceae
630:Phylogeny
615:asexually
498:complex.
492:flagellum
465:pyrenoids
457:cytostome
378:paramylon
212:Etymology
161:, of the
159:excavates
119:Phacaceae
86:Euglenida
66:Eukaryota
1684:Wikidata
1631:24507124
1571:85569422
1464:19878404
1416:86618135
1370:86347770
1335:85569364
1300:10416537
1292:24557733
1249:30363801
1241:27273533
1194:14334118
1147:84493597
1004:81548264
958:85569422
914:85275367
861:"Phacus"
817:23561445
809:26988648
670:Genetics
596:habitats
362:food web
136:Dujardin
115:Family:
72:Phylum:
62:Domain:
1843:5325077
1771:5424921
1690:Q144584
1507:2480453
1472:2010902
1099:6771635
696:evolved
684:introns
680:nucleus
555:Phacus,
488:eyespot
481:plastid
396:Feeding
345:Euglena
259:Euglena
196:is not
185:species
125:Genus:
105:Order:
82:Class:
1856:163339
1840:uBio:
1830:NZOR:
1784:200867
1758:1PHCUG
1705:Phacus
1675:Phacus
1629:
1569:
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1470:
1462:
1454:
1414:
1406:
1368:
1333:
1298:
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1162:Phacus
1145:
1137:
1097:
1089:
1051:Phacus
1002:
980:Phacus
956:
948:
912:
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815:
807:
799:
777:Phacus
754:Phacus
746:Phacus
730:Phacus
726:Phacus
722:Phacus
705:Phacus
664:Phacus
659:Phacus
655:Phacus
647:clades
643:Phacus
635:Phacus
611:Phacus
600:Phacus
575:Phacus
551:Phacus
543:Phacus
536:Phacus
520:sulcus
504:Phacus
469:Phacus
461:Phacus
437:Phacus
410:Phacus
401:Phacus
374:Phacus
366:Phacus
350:Phacus
326:Phacus
321:Phacus
312:Phacus
297:Phacus
292:Phacus
281:Phacus
277:Phacus
239:France
235:Phakos
222:Phacus
194:Phacus
189:Phacus
177:Phacus
163:phylum
147:Phacus
138:, 1841
130:Phacus
41:Phacus
25:Phacus
1851:WoRMS
1823:96783
1745:11710
1719:43655
1567:S2CID
1503:JSTOR
1468:S2CID
1412:S2CID
1366:S2CID
1331:S2CID
1296:S2CID
1245:S2CID
1190:S2CID
1143:S2CID
1095:S2CID
1000:S2CID
954:S2CID
910:S2CID
813:S2CID
386:cysts
233:word
231:Greek
152:genus
150:is a
93:Clade
1818:NCBI
1797:9766
1792:ITIS
1766:GBIF
1753:EPPO
1732:6MBX
1627:PMID
1460:PMID
1452:ISSN
1404:ISSN
1288:PMID
1280:ISSN
1237:PMID
1229:ISSN
1182:ISSN
1135:ISSN
1087:ISSN
946:ISSN
902:ISSN
805:PMID
797:ISSN
720:for
716:The
589:and
581:and
559:cell
420:and
403:are
360:and
301:taxa
43:sp.
1805:NBN
1740:EoL
1727:CoL
1617:hdl
1609:doi
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1559:doi
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1358:doi
1323:doi
1272:doi
1221:doi
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