46:
33:
511:, enabling them to function more efficiently and making them more tolerant against toxic compounds associated with photosynthesis. This changed the metabolism of the amoeba so much that it could no longer feed on microbes like its ancestors, and it became completely dependent on its endosymbiont, which in turn has lost so many genes it can no longer survive outside its host cell.
498:
The chromatophore genome has gone through a reduction, and is now just one third the size of the genome of its closest free living relatives, but still 10-fold larger than most plastid genomes. Some of the genes have been lost, others have migrated to the amoeba's nucleus through endosymbiotic gene
544:
also have at least two cyanobacterial-like genes, which were probably integrated into their genome through horizontal gene transfer from its cyanobacterial prey. Similar genes could have made the photosynthetic species pre-equipped to accept the chromatophore. The presence of extant heterotrophic
620:
904:"Marine Heterotrophic Amoebae, Flagellates and Heliozoa From Belize (Central America) and Tenerife (Canary Islands), With Descriptions of New Species, Luffisphaera Bulbochaete N. Sp., L. Longihastis N. Sp., L. Turriformis N. Sp. and Paulinella Intermedia N. Sp"
517:
show both a very slow growth rate and sensitivity to light, divide every 6–7 days, and prefer low light conditions, probably as a protection against oxidative stress and other light related stress as it doesn't have the same degree of
499:
transfer. It is estimated that 0.3-0.8% of
Paulinellas genes were derived from its endosymbiont, in addition to a small amount of genes from other organisms. Other genes have degenerated due to
573:
Lauterborn, R. (1895). "Protozoenstudien. II. Paulinella chromatophora nov. gen. nov. spec., ein beschalter
Rhizopode des Süßwassers mit blaugrünen chromatophorenartigen Einschlüssen".
424:
that is often referred to as a 'cyanelle' or chromatophore, and it represents the only known primary endosymbiosis event of photosynthetic cyanobacteria (other than the origin of
2335:
2504:"Ultrastructure and Ecology of Calycomonas ovalis Wulff, 1919, (Chrysophyceae) and Its Redescription as a Testate Rhizopod, Paulinella ovalis N. Comb. (Filosea: Euglyphina) 1"
1111:"Ultrastructure and Ecology of Calycomonas ovalis Wulff, 1919, (Chrysophyceae) and Its Redescription as a Testate Rhizopod, Paulinella ovalis N. Comb. (Filosea: Euglyphina) 1"
2321:
3839:
2130:"Impact of light intensity and quality on chromatophore and nuclear gene expression in Paulinella chromatophora, an amoeba with nascent photosynthetic organelles"
1045:"Diversity of the Photosynthetic Paulinella Species, with the Description of Paulinella micropora sp. nov. and the Chromatophore Genome Sequence for strain KR01"
1014:
451:
originates from), who diverged about 500 million years ago from the ancestors of its sister clade that consist of the living members of the cyanobacteria genera
3865:
549:
a valuable model for unravelling early stages of primary endosymbiosis event and studying the post symbiotic genome evolution of both the plastid and the host.
463:, was permanently established within the amoeba. It is estimated the last common ancestor of extant photosynthetic species lived about 60 million years ago.
723:"The ancestor of the Paulinella chromatophore obtained a carboxysomal operon by horizontal gene transfer from a Nitrococcus-like gamma-proteobacterium"
787:
Komm. Z. Wissensch. Untersuch. D. Deutschen Meere in Kiel und D. Biologischen
Anstalt Auf Helgoland. Wissensch. Meeresuntersuch., N.F., Abt. Kiel, Bd
3813:
3852:
694:
1615:
1605:
591:
M.D. Guiry in Guiry, M.D. & Guiry, G.M. 2013. AlgaeBase. World-wide electronic publication, National
University of Ireland, Galway.
507:. Some of the genes the nucleus received from the chromatophore were multiplied many times over through a "copy-paste" mechanism called
1043:
Lhee, Duckhyun; Yang, Eun Chan; Kim, Jong Im; Nakayama, Takuro; Zuccarello, Giuseppe; Andersen, Robert A.; Yoon, Hwan Su (April 2017).
491:, which in addition to a photosynthetic symbiont that is a captured green algae, also has a photosynthetic prokaryote as a symbiont; a
2650:
2339:
503:– accumulations of harmful mutations due to genetic isolation, and have probably been replaced with genes from other microbes through
2218:
Nowack, Eva C. M.; Price, Dana C.; Bhattacharya, Debashish; Singer, Anna; Melkonian, Michael; Grossman, Arthur R. (25 October 2016).
522:
mechanisms found in organisms with a photosynthetic apparatus of
Archaeplastid origin, which has a much longer evolutionary history.
3904:
2684:
2220:"Gene transfers from diverse bacteria compensate for reductive genome evolution in the chromatophore of Paulinella chromatophora"
2679:
474:
derive ultimately from a single cyanobacterium endosymbiont, which was taken in about 1.6 billion years ago by an ancestral
3857:
389:, the first two being freshwater forms and the third a marine form, which have recently (in evolutionary terms) taken on a
2763:
2753:
2736:
1748:
Nowack, Eva C. M.; Price, Dana C.; Bhattacharya, Debashish; Singer, Anna; Melkonian, Michael; Grossman, Arthur R. (2016).
2278:
689:
409:, who named it Paulinella after his stepmother Pauline. The event to permanent endosymbiosis probably occurred with a
1031:
Lineage(full) cellular organisms; Eukaryota; Rhizaria; Cercozoa; Imbricatea; Silicofilosea; Euglyphida; Paulinellidae
1289:
Lhee, Duckhyun; Ha, Ji-San; Kim, Sunju; Park, Myung Gil; Bhattacharya, Debashish; Yoon, Hwan Su (22 February 2019).
45:
244:
3870:
2016:"Endosymbiotic ratchet accelerates divergence after organelle origin: The Paulinella model for plastid evolution"
1485:"Cymbomonas tetramitiformis - a peculiar prasinophyte with a taste for bacteria sheds light on plastid evolution"
1358:
435:
3932:
959:"Paulinella longichromatophora sp. nov., a New Marine Photosynthetic Testate Amoeba Containing a Chromatophore"
504:
326:
197:
2179:"Endosymbiotic gene transfer and transcriptional regulation of transferred genes in Paulinella chromatophora"
854:
311:
2768:
2361:
Gabr, Arwa; Zournas, Apostolos; Stephens, Timothy G.; Dismukes, G. Charles; Bhattacharya, Debashish (2022).
855:"A Description of Paulinella indentata N. Sp. (Filosea: Euglyphina) from Subtidal Coastal Benthic Sediments"
304:
297:
283:
236:
2128:
Zhang, Ru; Nowack, Eva C. M.; Price, Dana C.; Bhattacharya, Debashish; Grossman, Arthur R. (1 April 2017).
1632:
228:
3740:
2643:
1898:
958:
268:
260:
252:
220:
205:
182:
2503:
903:
785:
Lohmann, H. (1908). "Untersuchungen zur
Feststellung des vollständigen Gehaltes des Meeres an Plankton".
428:), although primary endosymbiosis with a non-photosynthetic cyanobacterial symbiont have occurred in the
213:
174:
3891:
2731:
1750:"Gene transfers from diverse bacteria compensate for reductive genome evolution in the chromatophore of
1346:
807:"Heterotrophic Amoebae, Flagellates and Heliozoa from the Tvärminne Area, Gulf of Finland, in 1988–1990"
318:
290:
189:
3670:
3589:
540:
is a marine heterotrophic species of
Paulinella that has been shown to eat cyanobacteria and bacteria.
275:
3909:
3572:
3395:
2580:
2427:
2231:
2076:
2065:"A microbial eukaryote with a unique combination of purple bacteria and green algae as endosymbionts"
1960:
1824:
1765:
1696:
1685:"Genomic divergence within non-photosynthetic cyanobacterial endosymbionts in rhopalodiacean diatoms"
1560:
1496:
1302:
1183:
806:
734:
633:
3675:
3584:
3578:
3385:
3156:
3031:
2873:
3937:
3659:
3124:
2925:
1811:
Sánchez-Baracaldo, Patricia; Raven, John A.; Pisani, Davide; Knoll, Andrew H. (12 September 2017).
500:
3618:
3528:
3481:
3390:
3041:
3036:
2817:
2748:
2726:
2714:
2689:
2542:
2415:
2396:
2363:
2296:
2159:
1586:
1465:
1350:
932:
883:
649:
336:
40:
3792:
2878:
3896:
2920:
2822:
3878:
3844:
3800:
3428:
3144:
3139:
3099:
2995:
2758:
2721:
2709:
2699:
2636:
2614:
2596:
2523:
2484:
2443:
2388:
2259:
2200:
2151:
2110:
2092:
2045:
2037:
1996:
1978:
1921:
1903:
1860:
1842:
1793:
1730:
1712:
1665:
1657:
1611:
1578:
1522:
1457:
1449:
1400:
1382:
1328:
1291:"Evolutionary dynamics of the chromatophore genome in three photosynthetic Paulinella species"
1268:
1217:
1199:
1072:
1064:
986:
978:
924:
875:
827:
762:
487:
406:
140:
2015:
3883:
3706:
3523:
3302:
3059:
2673:
2604:
2588:
2515:
2474:
2435:
2380:
2286:
2249:
2239:
2190:
2141:
2100:
2084:
2027:
2014:
Bhattacharya, Debashish; Etten, Julia Van; Benites, L. Felipe; Stephens, Timothy G. (2023).
1986:
1968:
1929:
1911:
1893:
1850:
1832:
1783:
1773:
1720:
1704:
1647:
1568:
1512:
1504:
1439:
1408:
1390:
1372:
1318:
1310:
1258:
1248:
1207:
1191:
1140:
1122:
1056:
970:
916:
867:
819:
752:
742:
641:
621:"Six new marine species of the genus Paulinella (Rhizopoda: Filosea, or Rhizaria: Cercozoa)"
529:(those regions not affected by endosymbiotic gene transfer) are most closely related to the
3778:
2567:
Gabr, Arwa; Grossman, Arthur R.; Bhattacharya, Debashish (August 2020). Palenik, B. (ed.).
3710:
3247:
2177:
Nowack, E. C.; Vogel, H.; Groth, M.; Grossman, A. R.; Melkonian, M.; Glöckner, G. (2011).
1444:
1427:
1170:
Gabr, Arwa; Grossman, Arthur R.; Bhattacharya, Debashish (5 May 2020). Palenik, B. (ed.).
519:
508:
492:
453:
405:
was discovered in sediments of the river Rhine on
Christmas Eve 1894 by German biologist
32:
482:
events, and later tertiary and quaternary endosymbiosis, etc). The only exception is the
2584:
2431:
2235:
2080:
1964:
1828:
1769:
1700:
1564:
1500:
1306:
1187:
738:
637:
3665:
3476:
3380:
3109:
2985:
2845:
2801:
2704:
2609:
2568:
2519:
2254:
2219:
2105:
2064:
1991:
1948:
1916:
1879:
1855:
1812:
1788:
1749:
1725:
1684:
1517:
1395:
1323:
1290:
1263:
1236:
1212:
1171:
1126:
920:
871:
757:
722:
475:
448:
418:
390:
83:
70:
3926:
3566:
3538:
3354:
3151:
3051:
2865:
2850:
2400:
2300:
1884:
1469:
1363:
1354:
1110:
936:
530:
479:
459:
123:
2163:
1590:
887:
653:
3689:
3607:
3546:
3518:
3499:
3488:
3470:
3461:
3451:
3287:
3252:
3239:
3229:
3134:
3119:
3114:
3104:
3086:
2972:
2962:
2897:
2840:
2832:
823:
398:
394:
371:
148:
3805:
2336:"Uncovering An Evolutionary Process That Enabled Diversity of Plant Life on Earth"
1377:
1060:
974:
3826:
3772:
3561:
3556:
3423:
3418:
3405:
3297:
3257:
2855:
2322:"Microbe study sheds light on a critical step in the evolution of life on Earth"
467:
444:
440:
425:
367:
3763:
1708:
1314:
3698:
3624:
3508:
3413:
3372:
3362:
3344:
3292:
3267:
3262:
3094:
3064:
3018:
3000:
2990:
2439:
2291:
1573:
1548:
1508:
645:
410:
363:
113:
103:
2600:
2527:
2096:
2041:
1982:
1907:
1846:
1716:
1661:
1453:
1386:
1203:
1068:
982:
928:
879:
831:
3787:
3551:
3513:
3216:
3026:
2943:
2244:
2195:
2178:
1973:
1878:
Luis Delaye; Cecilio
Valadez-Cano; Bernardo Pérez-Zamorano (15 March 2016).
1837:
1778:
1253:
1044:
471:
414:
57:
2618:
2502:
Johnson, Paul W.; Hargraves, Paul E.; Sieburth, JOHN McN. (November 1988).
2488:
2479:
2462:
2447:
2392:
2263:
2204:
2155:
2114:
2088:
2049:
2032:
2000:
1925:
1864:
1797:
1734:
1669:
1582:
1526:
1461:
1428:"The Genomics and Cell Biology of Host-Beneficial Intracellular Infections"
1404:
1359:"Evaluating support for the current classification of eukaryotic diversity"
1332:
1272:
1221:
1109:
Johnson, Paul W.; Hargraves, Paul E.; Sieburth, JOHN McN. (November 1988).
1076:
990:
766:
747:
358:
is a genus of at least eleven species including both freshwater and marine
2628:
1933:
1412:
439:. The endosymbiotic event happened about 90–140 million years ago when an
3757:
3718:
3714:
3612:
3224:
3199:
2935:
2790:
2743:
2659:
359:
93:
2063:
Muñoz-Gómez, Sergio A.; Kreutz, Martin; Hess, Sebastian (11 June 2021).
1349:; Erika Barbero; Elyse Lasser; Micah Dunthorn; Debashish Bhattacharya;
3831:
3818:
3702:
3328:
1652:
483:
421:
2592:
2384:
2146:
2129:
1195:
3722:
2569:"Paulinella, a model for understanding plastid primary endosymbiosis"
1172:"Paulinella, a model for understanding plastid primary endosymbiosis"
1092:
429:
3734:
1607:
One Plus One Equals One: Symbiosis and the evolution of complex life
852:
Hannah, Fiona; Rogerson, Andrew; Anderson, O. Roger (January 1996).
2364:"Evidence for a robust photosystem II in the photosynthetic amoeba
1484:
626:
Journal of the Marine Biological Association of the United Kingdom
1813:"Early photosynthetic eukaryotes inhabited low-salinity habitats"
1549:"Evolution: Protein Import in a Nascent Photosynthetic Organelle"
2463:"Diversity and evolutionary history of plastids and their hosts"
3738:
2632:
721:
Marin, B; Nowack, EC; Glöckner, G; Melkonian, M (5 June 2007).
478:(and subsequently adopted into other eukaryote groups through
1021:. Bethesda, MD: National Center for Biotechnology Information
377:
Its most famous members are the three photosynthetic species
1090:
Wulff, A. (1919). "Ueber das Kleinplankton der Barentssee".
1633:"The monoplastidic bottleneck in algae and plant evolution"
592:
366:
it is covered by rows of siliceous scales, and use filose
1949:"Hold the salt: Freshwater origin of primary plastids"
1899:
10.1371/CURRENTS.TOL.E68A099364BB1A1E129A17B4E06B0C6B
1483:
Gagat, Przemysław; Mackiewicz, Paweł (January 2017).
495:
with a reduced genome, instead of a cyanobacterium.
3747:
3649:
3598:
3537:
3498:
3460:
3449:
3404:
3371:
3353:
3342:
3325:
3280:
3238:
3215:
3208:
3197:
3085:
3050:
3017:
2971:
2960:
2934:
2913:
2906:
2895:
2864:
2831:
2810:
2800:
2787:
1683:Nakayama, Takuro; Inagaki, Yuji (12 October 2017).
2362:
902:
853:
805:
619:
2134:The Plant Journal: For Cell and Molecular Biology
1880:"How Really Ancient Is Paulinella Chromatophora?"
1631:Vries, Jan de; Gould, Sven B. (15 January 2018).
397:. As a result they are no longer able to perform
2224:Proceedings of the National Academy of Sciences
1953:Proceedings of the National Academy of Sciences
1817:Proceedings of the National Academy of Sciences
1758:Proceedings of the National Academy of Sciences
1432:Annual Review of Cell and Developmental Biology
957:Kim, Sunju; Park, Myung Gil (1 February 2016).
1284:
1282:
2644:
1235:Nakayama, Takuro; Archibald, John M. (2012).
8:
568:
566:
564:
562:
2279:"Mitochondria and the origin of eukaryotes"
1165:
1163:
1161:
3735:
3457:
3446:
3350:
3339:
3335:
3322:
3212:
3205:
3194:
3082:
3047:
3014:
2978:
2968:
2957:
2910:
2903:
2892:
2807:
2797:
2784:
2651:
2637:
2629:
613:
611:
609:
607:
605:
603:
601:
575:Zeitschrift für Wissenschaftliche Zoologie
31:
20:
2608:
2478:
2414:Macorano, Luis; Nowack, Eva C.M. (2021).
2290:
2253:
2243:
2194:
2145:
2104:
2031:
1990:
1972:
1915:
1897:
1854:
1836:
1787:
1777:
1724:
1651:
1572:
1516:
1443:
1394:
1376:
1322:
1262:
1252:
1211:
780:
778:
776:
756:
746:
401:like their non-photosynthetic relatives.
1009:
1007:
716:
714:
712:
695:Integrated Taxonomic Information System
558:
1947:Lewis, Louise A. (12 September 2017).
1426:McCutcheon, John P. (6 October 2021).
909:The Journal of Eukaryotic Microbiology
860:The Journal of Eukaryotic Microbiology
684:
682:
680:
678:
676:
674:
672:
670:
618:Nicholls, Kenneth H. (November 2009).
1445:10.1146/annurev-cellbio-120219-024122
1237:"Evolving a photosynthetic organelle"
7:
370:to crawl over the substrate of the
2520:10.1111/j.1550-7408.1988.tb04160.x
1141:"Species Search :: AlgaeBase"
1127:10.1111/j.1550-7408.1988.tb04160.x
921:10.1111/j.1550-7408.1993.tb04917.x
872:10.1111/j.1550-7408.1996.tb02464.x
470:of all other known photosynthetic
14:
2277:Callier, Viviane (8 June 2022).
44:
16:Genus of single-celled organisms
2541:Smith, David (1 January 2013).
2183:Molecular Biology and Evolution
804:Vørs, Naja (1 September 1992).
824:10.1080/00785326.1992.10429930
1:
466:This is striking because the
434:Rhopalodiaceae and the algae
1378:10.1371/JOURNAL.PGEN.0020220
1061:10.1016/j.protis.2017.01.003
975:10.1016/j.protis.2015.11.003
2508:The Journal of Protozoology
2461:Patrick J. Keeling (2004).
1547:Archibald, John M. (2017).
1115:The Journal of Protozoology
595:; searched on 4 March 2013.
3954:
2467:American Journal of Botany
2416:"Paulinella chromatophora"
1709:10.1038/s41598-017-13578-8
1315:10.1038/s41598-019-38621-8
3684:
3445:
3338:
3321:
3193:
3081:
3013:
2981:
2956:
2891:
2783:
2667:
2440:10.1016/j.cub.2021.07.028
2292:10.1146/knowable-060822-2
1574:10.1016/j.cub.2017.08.013
1509:10.1007/s13199-016-0464-1
646:10.1017/S0025315409000514
436:Braarudosphaera bigelowii
342:
335:
170:
165:
154:
147:
41:Scientific classification
39:
30:
23:
1752:Paulinella chromatophora
1604:Archibald, John (2014).
727:BMC Evolutionary Biology
593:http://www.algaebase.org
505:horizontal gene transfer
156:Paulinella chromatophora
2547:The Scientist Magazine®
2245:10.1073/pnas.1608016113
1974:10.1073/pnas.1712956114
1838:10.1073/pnas.1620089114
1779:10.1073/pnas.1608016113
1640:Journal of Cell Science
1254:10.1186/1741-7007-10-35
901:Vørs, Naja (May 1993).
488:Pseudoblepharisma tenue
480:secondary endosymbiosis
362:. Like many members of
2480:10.3732/ajb.91.10.1481
2089:10.1126/sciadv.abg4102
2033:10.1002/bies.202200165
748:10.1186/1471-2148-7-85
447:which the plastids in
3892:Paleobiology Database
2196:10.1093/molbev/msq209
1347:Laura Wegener Parfrey
1182:(4). Wiley: 837–843.
525:The nuclear genes of
387:P. longichromatophora
245:P. longichromatophora
2573:Journal of Phycology
2342:on 15 September 2022
1695:(1). Nature: 13075.
1176:Journal of Phycology
3697:organization type:
2585:2020JPcgy..56..837G
2543:"Steal My Sunshine"
2432:2021CBio...31R1024M
2426:(17): R1024–R1026.
2236:2016PNAS..11312214N
2230:(43): 12214–12219.
2081:2021SciA....7.4102M
1965:2017PNAS..114.9759L
1829:2017PNAS..114E7737S
1823:(37): E7737–E7745.
1770:2016PNAS..11312214N
1764:(43): 12214–12219.
1701:2017NatSR...713075N
1565:2017CBio...27R1004A
1559:(18): R1004–R1006.
1501:2017Symbi..71....1G
1307:2019NatSR...9.2560L
1301:(1). Nature: 2560.
1188:2020JPcgy..56..837G
739:2007BMCEE...7...85M
638:2009JMBUK..89.1415N
3331:
3327:
1689:Scientific Reports
1653:10.1242/jcs.203414
1295:Scientific Reports
509:retrotransposition
3920:
3919:
3879:Open Tree of Life
3741:Taxon identifiers
3732:
3731:
3715:heliozoan amoebae
3707:amoeboflagellates
3671:Labyrinthomyxidae
3645:
3644:
3641:
3640:
3637:
3636:
3633:
3632:
3590:Globigerinitoidea
3441:
3440:
3437:
3436:
3317:
3316:
3313:
3312:
3276:
3275:
3189:
3188:
3185:
3184:
3181:
3180:
3177:
3176:
3173:
3172:
3169:
3168:
3165:
3164:
3100:Phaeogymnocellida
3077:
3076:
3073:
3072:
3009:
3008:
2996:Paracercomonadida
2952:
2951:
2887:
2886:
2593:10.1111/jpy.13003
2473:(10): 1481–1493.
2385:10.1111/nph.18052
2283:Knowable Magazine
2147:10.1111/tpj.13488
1959:(37): 9759–9760.
1617:978-0-19-163628-8
1357:(December 2006).
1351:David J Patterson
1196:10.1111/jpy.13003
407:Robert Lauterborn
351:
350:
143:
3945:
3913:
3912:
3900:
3899:
3887:
3886:
3874:
3873:
3861:
3860:
3848:
3847:
3835:
3834:
3822:
3821:
3809:
3808:
3796:
3795:
3783:
3782:
3781:
3768:
3767:
3766:
3736:
3524:Silicoloculinida
3458:
3454:
3447:
3351:
3347:
3340:
3336:
3332:
3323:
3303:Plasmodiophorida
3213:
3206:
3202:
3195:
3083:
3060:Thaumatomonadida
3048:
3015:
2979:
2969:
2965:
2958:
2911:
2904:
2900:
2893:
2808:
2798:
2794:
2785:
2696:
2676:
2653:
2646:
2639:
2630:
2623:
2622:
2612:
2564:
2558:
2557:
2555:
2553:
2538:
2532:
2531:
2499:
2493:
2492:
2482:
2458:
2452:
2451:
2411:
2405:
2404:
2370:
2358:
2352:
2351:
2349:
2347:
2338:. Archived from
2332:
2326:
2325:
2318:
2312:
2311:
2309:
2307:
2294:
2274:
2268:
2267:
2257:
2247:
2215:
2209:
2208:
2198:
2174:
2168:
2167:
2149:
2125:
2119:
2118:
2108:
2075:(24): eabg4102.
2069:Science Advances
2060:
2054:
2053:
2035:
2011:
2005:
2004:
1994:
1976:
1944:
1938:
1937:
1919:
1901:
1875:
1869:
1868:
1858:
1840:
1808:
1802:
1801:
1791:
1781:
1745:
1739:
1738:
1728:
1680:
1674:
1673:
1655:
1646:(2): jcs203414.
1637:
1628:
1622:
1621:
1601:
1595:
1594:
1576:
1544:
1538:
1537:
1535:
1533:
1520:
1480:
1474:
1473:
1447:
1423:
1417:
1416:
1398:
1380:
1343:
1337:
1336:
1326:
1286:
1277:
1276:
1266:
1256:
1232:
1226:
1225:
1215:
1167:
1156:
1155:
1153:
1151:
1137:
1131:
1130:
1106:
1100:
1099:
1087:
1081:
1080:
1040:
1034:
1033:
1028:
1026:
1011:
1002:
1001:
999:
997:
954:
948:
947:
945:
943:
906:
898:
892:
891:
857:
849:
843:
842:
840:
838:
809:
801:
795:
794:
782:
771:
770:
760:
750:
718:
707:
706:
704:
702:
686:
665:
664:
662:
660:
632:(7): 1415–1425.
623:
615:
596:
589:
583:
582:
570:
527:P. chromatophora
501:Muller's ratchet
493:purple bacterium
445:β-cyanobacterium
441:α-cyanobacterium
413:. The resulting
403:P. chromatophora
379:P. chromatophora
327:P. trinitatensis
198:P. chromatophora
139:
49:
48:
35:
21:
3953:
3952:
3948:
3947:
3946:
3944:
3943:
3942:
3933:Rhizaria genera
3923:
3922:
3921:
3916:
3908:
3903:
3895:
3890:
3882:
3877:
3869:
3864:
3856:
3851:
3843:
3838:
3830:
3825:
3817:
3812:
3804:
3799:
3791:
3786:
3777:
3776:
3771:
3762:
3761:
3756:
3743:
3733:
3728:
3711:testate amoebae
3680:
3629:
3594:
3573:Globotruncanida
3533:
3494:
3450:
3433:
3400:
3396:Symphyacanthida
3367:
3343:
3326:
3309:
3272:
3248:Claustrosporida
3234:
3198:
3161:
3069:
3046:
3005:
2961:
2948:
2930:
2896:
2883:
2860:
2827:
2788:
2779:
2778:
2694:
2672:
2663:
2657:
2627:
2626:
2566:
2565:
2561:
2551:
2549:
2540:
2539:
2535:
2501:
2500:
2496:
2460:
2459:
2455:
2420:Current Biology
2413:
2412:
2408:
2373:New Phytologist
2360:
2359:
2355:
2345:
2343:
2334:
2333:
2329:
2320:
2319:
2315:
2305:
2303:
2276:
2275:
2271:
2217:
2216:
2212:
2176:
2175:
2171:
2127:
2126:
2122:
2062:
2061:
2057:
2013:
2012:
2008:
1946:
1945:
1941:
1877:
1876:
1872:
1810:
1809:
1805:
1747:
1746:
1742:
1682:
1681:
1677:
1635:
1630:
1629:
1625:
1618:
1603:
1602:
1598:
1553:Current Biology
1546:
1545:
1541:
1531:
1529:
1482:
1481:
1477:
1425:
1424:
1420:
1345:
1344:
1340:
1288:
1287:
1280:
1234:
1233:
1229:
1169:
1168:
1159:
1149:
1147:
1139:
1138:
1134:
1108:
1107:
1103:
1089:
1088:
1084:
1042:
1041:
1037:
1024:
1022:
1013:
1012:
1005:
995:
993:
956:
955:
951:
941:
939:
900:
899:
895:
851:
850:
846:
836:
834:
803:
802:
798:
784:
783:
774:
720:
719:
710:
700:
698:
688:
687:
668:
658:
656:
617:
616:
599:
590:
586:
572:
571:
560:
555:
545:lineages makes
520:photoprotection
476:archaeplastidan
454:Prochlorococcus
443:(rather than a
312:P. subsphaerica
161:
160:Lauterborn 1895
158:
138:
43:
17:
12:
11:
5:
3951:
3949:
3941:
3940:
3935:
3925:
3924:
3918:
3917:
3915:
3914:
3901:
3888:
3875:
3862:
3849:
3836:
3823:
3810:
3797:
3784:
3769:
3753:
3751:
3745:
3744:
3739:
3730:
3729:
3727:
3726:
3719:sporozoan-like
3695:
3692:
3685:
3682:
3681:
3679:
3678:
3676:Rhizoplasmidae
3673:
3668:
3666:Gymnosphaerida
3663:
3655:
3653:
3651:Incertae sedis
3647:
3646:
3643:
3642:
3639:
3638:
3635:
3634:
3631:
3630:
3628:
3627:
3622:
3615:
3610:
3604:
3602:
3600:Incertae sedis
3596:
3595:
3593:
3592:
3587:
3585:Guembelitriida
3581:
3579:Heterohelicida
3575:
3569:
3564:
3559:
3554:
3549:
3543:
3541:
3535:
3534:
3532:
3531:
3526:
3521:
3516:
3511:
3504:
3502:
3496:
3495:
3493:
3492:
3485:
3479:
3477:Xenophyophorea
3474:
3467:
3465:
3455:
3443:
3442:
3439:
3438:
3435:
3434:
3432:
3431:
3426:
3421:
3416:
3410:
3408:
3402:
3401:
3399:
3398:
3393:
3388:
3386:Chaunocanthida
3383:
3381:Arthracanthida
3377:
3375:
3369:
3368:
3366:
3365:
3359:
3357:
3348:
3333:
3319:
3318:
3315:
3314:
3311:
3310:
3308:
3307:
3306:
3305:
3300:
3290:
3284:
3282:
3278:
3277:
3274:
3273:
3271:
3270:
3265:
3260:
3255:
3250:
3244:
3242:
3236:
3235:
3233:
3232:
3227:
3221:
3219:
3210:
3203:
3191:
3190:
3187:
3186:
3183:
3182:
3179:
3178:
3175:
3174:
3171:
3170:
3167:
3166:
3163:
3162:
3160:
3159:
3157:Ventricleftida
3154:
3149:
3148:
3147:
3142:
3137:
3129:
3128:
3127:
3122:
3117:
3112:
3110:Phaeosphaerida
3107:
3102:
3091:
3089:
3079:
3078:
3075:
3074:
3071:
3070:
3068:
3067:
3062:
3056:
3054:
3045:
3044:
3039:
3034:
3032:Spongomonadida
3029:
3023:
3021:
3011:
3010:
3007:
3006:
3004:
3003:
2998:
2993:
2988:
2986:Glissomonadida
2982:
2976:
2966:
2954:
2953:
2950:
2949:
2947:
2946:
2940:
2938:
2932:
2931:
2929:
2928:
2923:
2917:
2915:
2908:
2901:
2889:
2888:
2885:
2884:
2882:
2881:
2876:
2874:Chlorarachnida
2870:
2868:
2862:
2861:
2859:
2858:
2853:
2848:
2846:Desmothoracida
2843:
2837:
2835:
2829:
2828:
2826:
2825:
2820:
2814:
2812:
2805:
2802:Reticulofilosa
2795:
2781:
2780:
2777:
2776:
2775:
2774:
2773:
2772:
2769:Mesomycetozoea
2766:
2761:
2751:
2741:
2740:
2739:
2734:
2729:
2724:
2719:
2718:
2717:
2705:Diaphoretickes
2702:
2697:
2692:
2687:
2682:
2677:
2669:
2668:
2665:
2664:
2662:classification
2658:
2656:
2655:
2648:
2641:
2633:
2625:
2624:
2579:(4): 837–843.
2559:
2533:
2514:(4): 618–626.
2494:
2453:
2406:
2379:(3): 934–945.
2353:
2327:
2313:
2269:
2210:
2189:(1): 407–422.
2169:
2140:(2): 221–234.
2120:
2055:
2026:(1): 2200165.
2006:
1939:
1870:
1803:
1740:
1675:
1623:
1616:
1610:. OUP Oxford.
1596:
1539:
1475:
1438:(1): 115–142.
1418:
1338:
1278:
1227:
1157:
1132:
1121:(4): 618–626.
1101:
1082:
1055:(2): 155–170.
1035:
1003:
949:
915:(3): 272–287.
893:
844:
796:
772:
708:
666:
597:
584:
557:
556:
554:
551:
449:Archaeplastida
419:photosynthetic
391:cyanobacterium
349:
348:
340:
339:
333:
332:
331:
330:
323:
315:
308:
305:P. subcarinata
301:
298:P. sphaeroides
294:
287:
284:P. quinqueloba
280:
272:
265:
257:
249:
241:
237:P. lauterborni
233:
225:
217:
210:
202:
194:
186:
179:
168:
167:
163:
162:
159:
152:
151:
145:
144:
131:
127:
126:
121:
117:
116:
111:
107:
106:
101:
97:
96:
91:
87:
86:
81:
74:
73:
71:Diaphoretickes
68:
61:
60:
55:
51:
50:
37:
36:
28:
27:
15:
13:
10:
9:
6:
4:
3:
2:
3950:
3939:
3936:
3934:
3931:
3930:
3928:
3911:
3906:
3902:
3898:
3893:
3889:
3885:
3880:
3876:
3872:
3867:
3863:
3859:
3854:
3850:
3846:
3841:
3837:
3833:
3828:
3824:
3820:
3815:
3811:
3807:
3802:
3798:
3794:
3789:
3785:
3780:
3774:
3770:
3765:
3759:
3755:
3754:
3752:
3750:
3746:
3742:
3737:
3724:
3720:
3716:
3712:
3708:
3704:
3700:
3696:
3693:
3691:
3687:
3686:
3683:
3677:
3674:
3672:
3669:
3667:
3664:
3662:
3661:
3657:
3656:
3654:
3652:
3648:
3626:
3623:
3621:
3620:
3616:
3614:
3611:
3609:
3606:
3605:
3603:
3601:
3597:
3591:
3588:
3586:
3582:
3580:
3576:
3574:
3570:
3568:
3567:Globigerinida
3565:
3563:
3560:
3558:
3555:
3553:
3550:
3548:
3545:
3544:
3542:
3540:
3539:Globothalamea
3536:
3530:
3527:
3525:
3522:
3520:
3517:
3515:
3512:
3510:
3506:
3505:
3503:
3501:
3497:
3491:
3490:
3486:
3483:
3480:
3478:
3475:
3472:
3469:
3468:
3466:
3463:
3459:
3456:
3453:
3448:
3444:
3430:
3427:
3425:
3422:
3420:
3417:
3415:
3412:
3411:
3409:
3407:
3403:
3397:
3394:
3392:
3389:
3387:
3384:
3382:
3379:
3378:
3376:
3374:
3370:
3364:
3361:
3360:
3358:
3356:
3355:Sticholonchea
3352:
3349:
3346:
3341:
3337:
3334:
3330:
3324:
3320:
3304:
3301:
3299:
3296:
3295:
3294:
3291:
3289:
3286:
3285:
3283:
3279:
3269:
3266:
3264:
3261:
3259:
3256:
3254:
3251:
3249:
3246:
3245:
3243:
3241:
3237:
3231:
3228:
3226:
3223:
3222:
3220:
3218:
3214:
3211:
3207:
3204:
3201:
3196:
3192:
3158:
3155:
3153:
3152:Tectofilosida
3150:
3146:
3143:
3141:
3138:
3136:
3133:
3132:
3130:
3126:
3125:Phaeoconchida
3123:
3121:
3118:
3116:
3113:
3111:
3108:
3106:
3103:
3101:
3098:
3097:
3096:
3093:
3092:
3090:
3088:
3084:
3080:
3066:
3063:
3061:
3058:
3057:
3055:
3053:
3052:Silicofilosea
3049:
3043:
3040:
3038:
3035:
3033:
3030:
3028:
3025:
3024:
3022:
3020:
3016:
3012:
3002:
3001:Cercomonadida
2999:
2997:
2994:
2992:
2989:
2987:
2984:
2983:
2980:
2977:
2974:
2970:
2967:
2964:
2959:
2955:
2945:
2942:
2941:
2939:
2937:
2933:
2927:
2926:Metromonadida
2924:
2922:
2919:
2918:
2916:
2912:
2909:
2905:
2902:
2899:
2894:
2890:
2880:
2877:
2875:
2872:
2871:
2869:
2867:
2866:Chlorarachnea
2863:
2857:
2854:
2852:
2851:Leucodictyida
2849:
2847:
2844:
2842:
2839:
2838:
2836:
2834:
2830:
2824:
2821:
2819:
2816:
2815:
2813:
2809:
2806:
2803:
2799:
2796:
2792:
2786:
2782:
2770:
2767:
2765:
2762:
2760:
2757:
2756:
2755:
2752:
2750:
2747:
2746:
2745:
2742:
2738:
2735:
2733:
2732:Stramenopiles
2730:
2728:
2725:
2723:
2720:
2716:
2713:
2712:
2711:
2708:
2707:
2706:
2703:
2701:
2698:
2695:(major groups
2693:
2691:
2688:
2686:
2683:
2681:
2678:
2675:
2671:
2670:
2666:
2661:
2654:
2649:
2647:
2642:
2640:
2635:
2634:
2631:
2620:
2616:
2611:
2606:
2602:
2598:
2594:
2590:
2586:
2582:
2578:
2574:
2570:
2563:
2560:
2548:
2544:
2537:
2534:
2529:
2525:
2521:
2517:
2513:
2509:
2505:
2498:
2495:
2490:
2486:
2481:
2476:
2472:
2468:
2464:
2457:
2454:
2449:
2445:
2441:
2437:
2433:
2429:
2425:
2421:
2417:
2410:
2407:
2402:
2398:
2394:
2390:
2386:
2382:
2378:
2374:
2369:
2367:
2357:
2354:
2341:
2337:
2331:
2328:
2323:
2317:
2314:
2302:
2298:
2293:
2288:
2284:
2280:
2273:
2270:
2265:
2261:
2256:
2251:
2246:
2241:
2237:
2233:
2229:
2225:
2221:
2214:
2211:
2206:
2202:
2197:
2192:
2188:
2184:
2180:
2173:
2170:
2165:
2161:
2157:
2153:
2148:
2143:
2139:
2135:
2131:
2124:
2121:
2116:
2112:
2107:
2102:
2098:
2094:
2090:
2086:
2082:
2078:
2074:
2070:
2066:
2059:
2056:
2051:
2047:
2043:
2039:
2034:
2029:
2025:
2021:
2017:
2010:
2007:
2002:
1998:
1993:
1988:
1984:
1980:
1975:
1970:
1966:
1962:
1958:
1954:
1950:
1943:
1940:
1935:
1931:
1927:
1923:
1918:
1913:
1909:
1905:
1900:
1895:
1891:
1887:
1886:
1885:PLOS Currents
1881:
1874:
1871:
1866:
1862:
1857:
1852:
1848:
1844:
1839:
1834:
1830:
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3690:paraphyletic
3660:Lapot gusevi
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3619:Heterogromia
3617:
3608:Involutinida
3599:
3547:Textulariida
3529:Ammodiscidae
3519:Spirillinida
3500:Tubothalamea
3489:Reticulomyxa
3487:
3482:Astrorhizida
3471:Allogromiida
3462:Monothalamea
3452:Foraminifera
3406:Polycystinea
3391:Holocanthida
3288:Vampyrellida
3253:Haplosporida
3240:Ascetosporea
3230:Reticulosida
3135:Cryomonadida
3120:Phaeogromida
3115:Phaeocalpida
3105:Phaeocystida
3087:Thecofilosea
3042:Variglissida
3037:Marimonadida
2973:Sarcomonadea
2963:Ventrifilosa
2914:Metromonadea
2898:Monadofilosa
2841:Cryptofilida
2833:Granofilosea
2818:Aquavolonida
2754:Opisthokonta
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1371:(12): e220.
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1355:Laura A Katz
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1023:. Retrieved
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1015:"Paulinella"
994:. Retrieved
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962:
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940:. Retrieved
912:
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835:. Retrieved
818:(1): 1–109.
815:
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693:
690:"Paulinella"
657:. Retrieved
629:
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269:P. osloensis
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221:P. indentata
219:
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206:P. gigantica
204:
196:
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183:P. bulloides
181:
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149:Type species
134:
133:
77:
64:
24:
18:
3827:iNaturalist
3773:Wikispecies
3723:fungus-like
3699:flagellates
3562:Carterinida
3557:Robertinida
3424:Spumellaria
3419:Nassellaria
3298:Phagomyxida
3281:Proteomyxia
3258:Mikrocytida
2856:Limnofilida
2811:Skiomonadea
1241:BMC Biology
969:(1): 1–12.
426:chloroplast
345:Calycomonas
214:P. gracilis
175:P. agassizi
3938:Imbricatea
3927:Categories
3779:Paulinella
3749:Paulinella
3625:Komokiacea
3509:Fusulinida
3414:Collodaria
3373:Acantharea
3363:Taxopodida
3345:Radiolaria
3293:Phytomyxea
3268:Paramyxida
3263:Paradinida
3095:Phaeodarea
3065:Euglyphida
3019:Imbricatea
2991:Katabiidae
2879:Minorisida
2366:Paulinella
1495:(1): 1–7.
866:(1): 1–4.
793:: 131–137.
701:28 January
581:: 537–544.
553:References
547:Paulinella
515:Paulinella
472:eukaryotes
411:cyanobiont
368:pseudopods
364:euglyphids
355:Paulinella
319:P. suzukii
291:P. riveroi
190:P. carsoni
141:Lauterborn
135:Paulinella
114:Euglyphida
104:Imbricatea
25:Paulinella
3788:AlgaeBase
3552:Rotaliida
3514:Miliolida
3217:Gromiidea
3209:Marimyxia
3131:Eothecia
3027:Krakenida
2944:Helkesida
2921:Metopiida
2823:Tremulida
2749:Amoebozoa
2727:Alveolata
2715:Cryptista
2690:Eukaryota
2601:0022-3646
2552:18 August
2528:0022-3921
2401:247106539
2306:18 August
2301:249526889
2097:2375-2548
2042:0265-9247
2020:BioEssays
1983:0027-8424
1934:Q36374426
1908:2157-3999
1847:0027-8424
1717:2045-2322
1662:0021-9533
1532:18 August
1489:Symbiosis
1470:235786110
1454:1081-0706
1413:Q21090155
1387:1553-7390
1204:0022-3646
1150:19 August
1098:: 95–125.
1069:1618-0941
1025:4 January
996:19 August
983:1434-4610
942:19 August
937:221852241
929:1066-5234
880:1066-5234
837:19 August
832:0078-5326
733:(1): 85.
659:18 August
542:P. ovalis
538:P. ovalis
534:P. ovalis
415:organelle
360:amoeboids
276:P. ovalis
58:Eukaryota
3764:Q3815918
3758:Wikidata
3694:†extinct
3613:Lagenida
3429:Orodaria
3225:Gromiida
3200:Endomyxa
3145:Matazida
3140:Ebriacea
2936:Helkesea
2907:Eoglissa
2791:Cercozoa
2744:Amorphea
2722:Rhizaria
2710:Hacrobia
2700:Excavata
2685:Bacteria
2660:Rhizaria
2619:32289879
2489:21652304
2448:34520707
2393:35211975
2264:27791007
2205:20702568
2164:39165272
2156:28182317
2115:34117067
2050:36328783
2001:28860199
1930:Wikidata
1926:28515968
1865:28808007
1798:27791007
1735:29026213
1670:28893840
1591:21228029
1583:28950079
1527:28066124
1462:34242059
1409:Wikidata
1405:17194223
1333:30796245
1273:22531210
1222:32289879
1077:28262587
991:26709891
888:84358420
767:17550603
654:86359271
337:Synonyms
166:Species
120:Family:
94:Cercozoa
90:Phylum:
54:Domain:
3845:1267643
3832:1284695
3819:3212296
3703:amoebae
3329:Retaria
2759:Animals
2680:Archaea
2610:7734844
2581:Bibcode
2428:Bibcode
2255:5087059
2232:Bibcode
2106:8195481
2077:Bibcode
1992:5604047
1961:Bibcode
1917:4866557
1856:5603991
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1766:Bibcode
1726:5638926
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1396:1713255
1324:6384880
1303:Bibcode
1264:3337241
1213:7734844
1184:Bibcode
1049:Protist
963:Protist
812:Ophelia
758:1904183
735:Bibcode
634:Bibcode
484:ciliate
432:family
422:plastid
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264:, 2009
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248:, 2016
240:, 2009
232:, 1993
224:, 1996
209:, 2009
201:, 1895
193:, 2009
178:, 2009
3905:WoRMS
3897:30414
3871:39716
3858:44014
3840:IRMNG
3793:45017
2764:Fungi
2397:S2CID
2297:S2CID
2160:S2CID
1636:(PDF)
1587:S2CID
1466:S2CID
933:S2CID
884:S2CID
650:S2CID
417:is a
78:Clade
65:Clade
3866:NCBI
3853:ITIS
3814:GBIF
3806:6JP3
2615:PMID
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