EUGLENOPHYTA 259

6.15, 6.16). During the warm summer months the damselfly nymphs and C. libellee live separately. With the onset of winter, the Colacium cells attach to the cuticle of the rectum of the damselfly larvae, forming a conspicuous green plug that colors the terminal four segments of the abdomen dark green. In the rectum, the Colacium is in a palmelloid state, lacking a flagellar swelling and eyespot. As the peripheral waters of the lake freeze, the damselfly nymphs move to deeper water. Here the damselfly nymphs form a protected, motile, translucent microhabitat for the Colacium; the damselfly nymphs probably also provide a source of nutrients for the Colacium cells, which helps the alga to survive the unfavorable winter conditions. In spring, the damselfly nymphs swim to warmer water, at which time the Colacium cells swim out of the rectum to establish a free-living existence in the summer. If algal-free damselfly nymphs are placed in water with C. libellae, the alga will establish itself within 36 hours in the damselfly nymph rectum. Other species are not able to infect the nymphs (Willey et al., 1970, 1973; Willey, 1972; Rosowski and Willey, 1975).

REFERENCES

Al-Dhaheri, R. S., and Willey, P. L. (1996). Colonization and reproduction of the epibiotic flagellate Colacium vesiculosum (Euglenophyceae) in Daphnia pulex. J. Phycol. 32:770–4.

Barsanti, L., Vismara, R., Passarelli, V., and Gualtieri, P. (2001). Paramylon ( -1,3- glucan) content in wild type and WZSL mutant of Euglena gracilis.

Effects of growth conditions. J. Appl. Phycol. 13:59–65. Bastin, P., and Gull, K. (1999). Assembly and function

of complex flagellar structures illustrated by the paraflagellar rod of trypanosomes. Protist 150:113–23.

Batra, P. P., and Tollin, G. (1964). Phototaxis in Euglena. I. Isolation of the eyespot granules and identification of the eye-spot pigments. Biochem. Biophys. Acta 79:371–8.

Bäumer, D., Preisfeld, A., and Ruppel, H.G. (2001). Isolation and characterization of paramylon synthase from Euglena gracilis (Euglenophyceae). J. Phycol. 37:38–46.

Bertaux, O., and Valencia, R. (1971). Effects de la carence B12 sur les cellules synchrones de Euglena

gracilis: Blocage de la division cellulaire, polyploidie et gigantisme. J. Physiol. (London) 63:167A.

Bertaux, O., and Valencia, R. (1973). Blocage de la division cellulaire et malformations induites par carence B12 chez les cellules synchrones de Euglena gracilis Z. C. R. Séances Acad. Sci. Paris 276:753–6.

Bouck, G. B., Rogalski, A., and Valaitis, A. (1978). Surface organization and composition of Euglena. II. Flagellar mastigonemes. J. Cell Biol. 77:805–26.

Bracher, R. (1937). The light relations of Euglena limosa Gard. Part I. The influence of intensity and quality of light on phototaxy. J. Linn. Soc. Bot. 51:23–42.

Bré, M. H., and Lefort-Tran, M. (1974). Influence de l’avitaminose B12 sur les chloroplastes de l’Euglena gracilis Z. en milieu lactate. C. R. Séances Acad. Sci. Paris 278:1349–52.

Bré, M. H., Diamond, J., and Jacques, R. (1975). Factors mediating the vitamin B12 requirement of Euglena. J. Protozool. 22:432–4.

Brown, P. J. P., Leander, B. S., and Farmer, M. A. (2002). Redescription of Euglena rustica (Euglenophyceae), a rare euglenophyte from the intertidal zone.

Phycologia 41:445–52.

Carell, E. F. (1969). Studies on chloroplast development and replication in Euglena. I. Vitamin B12 and chloroplast replication. J. Cell Biol. 41:431–40.

Carell, E. F., Johnston, P. L., and Christopher, A. R. (1970). Vitamin B12 and the macromolecular composition of Euglena. J. Cell Biol. 47:525–30.

Chaly, N., Lord, A., and Lafontaine, J. G. (1977). A lightand electron-microscope study of nuclear structure throughout the cell cycle in the euglenoid Astasia longa (Jahn). J. Cell Sci. 27:23–45.

Cogburn, J. N., and Schiff, J. A. (1984). Purification and properties of the mucus of Euglena gracilis (Euglenophyceae). J. Phycol. 20:533–44.

Davis, B., and Merrett, M. J. (1974). The effect of light on the synthesis of mitochondrial enzymes in divi- sion-synchronized Euglena cultures. Plant Physiol. 53:575–80.

Dodge, J. D. (1975). The fine structure of Trachelmonas (Euglenophyceae). Arch. Protistenk. 117:65–77.

Dubreuil, R. R., and Bouck, G. B. (1985). The membrane skeleton of a unicellular organism consists of bridged, articulating strips. J. Cell Biol. 101:1884–96.

Dunlap, J. R., Walne, P. L., and Bentley, J. (1983). Microarchitecture and elemental spatial segregation of envelopes of Trachelomonas lefevrei

(Euglenophyceae). Protoplasma 117:97–106. Gomez, M. P., Harris, J. B., and Walne, P. L. (1974).

Studies of Euglena gracilis in aging cultures. I. Light microscopy and cytochemistry. Br. Phycol. J. 9:163–74.

260 CHLOROPLAST E.R.: EVOLUTION OF ONE MEMBRANE

Gottleib, J. (1850). Ueber eine neue, mit Starkmehl isomere Substanz. Ann. Chem. Pharm. 75:51–61.

Gravilaˇ, L. (1996). Light and electron microscope studies of euglenoid nuclei. In Cytology, Genetics and Molecular Biology of Algae, ed. B. R. Chaudhary, and S. B. Agrawal, pp. 193–213. Amsterdam, The Netherlands: SPB Academic Pub.

Häder, D-P. (1987). Polarotaxis, gravitaxis and vertical phototaxis in the green flagellate, Euglena gracilis. Arch. Microbiol. 147:179–83.

Hilenski, L. L., and Walne, P. L. (1983). Ultrastructure of mucocysts in Peranema trichophorum

(Euglenophyceae). J. Protozool. 30:491–6.

Hilenski, L. L., and Walne, P. L. (1985). Ultrastructure of the flagella of the colorless phagotroph Peranema trichophorum (Euglenophyceae). I. Flagellar mastigonemes. J. Phycol. 21:114–25.

Huber-Pestalozzi, G. (1955). Euglenophycean. Das Phytoplankton des Susswassers, Vol. 16, Part 4. Stuttgart: E. Schweizerbart’sche Verlagsbuchhandlung.

Hutner, S. H., and Provasoli, L. (1955). Comparative biochemistry of flagellates. In Biochemistry and Physiology of Protozoa, ed. S. H. Hutner, and A. Lwoff, Vol. 2, pp. 1–40. New York and London: Academic Press.

Jahn, T. L., and Bovee, E. C. (1968). Locomotive and motile response in Euglena. In The Biology of Euglena, ed. D. E. Búetow, I:45–108. New York and London: Academic Press.

Kingston, M. B. (2002). Effect of subsurface nutrient supplies on the vertical migration of Euglena proxima (Euglenophyta). J. Phycol. 38:872–80.

Kiss, J. Z., Vasconcelos, A. C., and Triemer, R. E. (1987). Structure of the euglenoid storage carbohydrate, paramylon. Am. J. Bot. 74:877–82.

Krinsky, N. I., and Goldsmith, T. H. (1960). The carotenoids of the flagellated alga, Euglena gracilis. Arch. Biochem. Biophys. 91:271–9.

Leander, B. S., and Farmer, M. A. (2000). Comparative morphology of the euglenoid pellicle. I. Patterns of strips and pores. J. Eukary. Microbiol. 47:469–79.

Leander, B. S., and Farmer, M. A. (2001a). Comparative morphology of the euglenoid pellicle. II.

Diversity of strip substructure. J. Eukary. Microbiol. 48:202–17.

Leander, B. S., and Farmer, M. A. (2001b). Evolution of Phacus (Euglenophyceae) as inferred from pellicle morphology and SSU rDNA. J. Phycol. 37:143–59.

Leedale, G. F. (1959). Periodicity of mitosis and cell division in the Euglenineae. Biol. Bull. 116:162–74.

Leedale, G. F. (1967). Euglenoid Flagellates. Englewood Cliffs, NJ: Prentice Hall.

Leedale, G. F. (1970). Phylogenetic aspects of nuclear cytology in the algae. Ann. NY Acad. Sci. 175(2):429–53.

Leedale, G. F. (1975). Envelope formation and structure in the euglenoid genus Trachelomonas. Br. Phycol. J.

10:17–41.

Lonergan, T. A. (1983). Regulation of cell shape in Euglena gracilis. I. Involvement of the biological clock, respiration, photosynthesis, and cytoskeleton.

Plant Physiol. 71:719–30.

Marin, B. (2004). Origin and fate of chloroplasts in the Euglenoida. Protist 155:13–14.

Mignot, J-P. (1966). Structure et ultrastructure de quelques Euglénomonadines. Protistologica 2:51–117.

Murata, K., Okamoto, M., and Suzaki, T. (2000). Morphological change of cell-membrane-integrated crystal structure induced by cell shape change in

Euglena gracilis. Protoplasma 214:73–9.

Ngô, H. M., and Bouck, G. B. (1998). Heterogeneity and a coiled coil predication of trypanosomatid-like flagellar rod proteins in Euglena. J. Eukary. Microbiol. 45:323–33.

Nudelman, A. A., Rossa, M. S., Conforti, V., and Triemer, R. E. (2003). Phylogeny of Euglenophyceae based on small subunit rDNA sequences: taxonomic implications. J. Phycol. 39:226–35.

Olli, K. (1996). Resting cyst formation of Eutreptiella gymnastica (Euglenophyceae) in the northern coastal Baltic Sea. J. Phycol. 32:535–42.

Palmer, J. D., and Round, F. E. (1965). Persistent, verti- cal-migration rhythms in the benthic microflora. I. The effect of light and temperature on the rhythmic behaviour of Euglena obtusa. J. Mar. Biol. Assoc. UK

45:567–82.

Preisfeld, A., Berger, S., Busse, I., Liller, S., and Ruppel, H. G. (2000). Phylogenetic analyses of various euglenoid taxa (Euglenozoa) based on 18S rDNA sequence data. J. Phycol. 36:220–6.

Pringsheim, E. G. (1953). Observations on some species of Trachelomonas grown in culture. New Phytol. 52:93–113, 238–66.

Rosowski, J. R., and Willey, R. L. (1975). Colacium libellae sp. nov. (Euglenophyceae), a photosynthetic inhabitant of the larval damselfly rectum. J. Phycol. 11:310–15.

Saito, A., Suetomo, Y., Arikawa, M., et al. (2003) Gliding movement in Peranema trichophorum is powered by flagellar surface motility. Cell Motil. Cytoskel. 55:244–53.

Sekiguchi, H., Moriya, M., Nakayama, T., and Inouye, I. (2002). Vestigial chloroplasts in heterotrophic stramenopiles Pteridomonas danica and Ciliophrys infusionum (Dictyochophyceae). Protist 153:157–67.

EUGLENOPHYTA 261

Shin, W., and Triemer, R. E. (2004). Phylogenetic analysis on the genus Euglena (Euglenophyceae) with particular reference to the species Euglena viridis. J.

Phycol. 40:759–71.

Sperling Pagni, P. G., Walne, P. L., and Pagni, R. M. (1981). On the occurrence of -carotene in isolated stigmata of Euglena gracilis var. bacillaris. Phycologia

20:431–4.

Surek, B., and Melkonian, M. (1986). A cryptic cytosome is present in Euglena. Protoplasma 133:39–49.

Talke, S., and Preisfield, A. (2002). Molecular evolution of euglenozoan paraxonemal rod genes par1 and par2 coincides with phylogenetic reconstruction based on small subunit rDNA data. J. Phycol. 38:995–1003.

Throndsen, J. (1969). Flagellates of Norwegian coastal waters. Nytt Mag. Bot. 16:161–216.

Throndsen, J. (1973). Fine structure of Eutreptiella gymnastica (Euglenophyceae). Norw. J. Bot. 20:271–80.

Walne, P. L. (1980). Euglenoid flagellates. In Phytoflagellates, ed. E. Cox, pp. 165–212. North Holland: Elsevier.

West, L. K., and Walne, P. L. (1980). Trachelomonas hispada var. coronata (Euglenophyceae). II. Envelope substructure. J. Phycol. 16:498–506.

West, L. K., Walne, P. L., and Bentley, J. (1980).

Trachelomonas hispida var. coronata (Euglenophyceae). III. Envelope elemental composition and mineralization. J. Phycol. 16:582–91.

Willey, R. L. (1972). The damselfly (Odonata) hindgut as a host for the euglenoid Colacium. Trans. Am. Microsc. Soc. 91:585–93.

Willey, R. L. (1984). Fine structure of the mucocysts of

Colacium calvum (Euglenophyceae). J. Phycol. 20:426–30.

Willey, R. L., and Wibel, R. G. (1985a). The reservoir cytoskeleton and a possible cytosomal homologue in Colacium (Euglenophyceae). J. Phycol. 21:570–7.

Willey, R. L., and Wibel, R. G. (1985b). A cytosome/cytopharynx in green euglenoid flagellates (Euglenales) and its phylogenetic implications.

BioSystems 18:369–76.

Willey, R. L., Bowen, W. R., and Durban, E. M. (1970). Symbiosis between Euglena and damselfly nymphs is seasonal. Science 170:80–1.

Willey, R. L., Durban, E. M., and Bowen, W. R. (1973). Ultrastructural observations of a Colacium palmella: The reservoir, eyespot, and flagella. J. Phycol. 9:211–15.