Phaeodaria

Marine planktonic sarcodines with arguably the most complex mineral skeleton in protists, with the largest nucleus known in eukaryotes, and virtually unstudied.

Reference: Korsun S, 2011. Phaeodaria Haeckel 1879. In: Protista. III. Handbook of Zoology (Ed. S.A. Karpov), Moscow, KMK Scientific Press Ltd., p. 121-159 + 3 color plates [in Russian]

Fig. 69. Cell setup in Phaeodaria. ап, axopodia, рп, rhizopodia; км, cell membrane; ск, tangential and radial skeletal spicules; ка, calymmal cysternae; мт, bundles of microtubules; пар, parapyle; ЦОМТ, microtubule organization center; я, nucleus with chromatin; эр, the cistern of er enclasping the nucleus; цк, capsular wall; аст, astropyle; фе, phaeodium; пс, peristome; пи, pylome. Upper right inset, schematic arrangement of microtubules in the axoneme. (After several authors).

Fig. 72. Skeletal spicule genesis in Phaeodaria (schematic representation of the events). I, Spiculogenic vacuole with a diatom frustule. II, Budding of offshoots of the vacuole. III, Parietal precipitation of silicate-containing matrix; an axial lumen is preserved in the thicker segments of the vacuole. IV, Congelation of the silicate-containing matrix and resorption of the vacuolar membrane.

Fig. 73.The symmetry of the pheodarioan cell. Schematic drawing of major constructive elements (aboral view): цк, capsule; аст, astropyle; пар, parapyle; фе, phaeodium; ск, skeleton; ан, axoneme. I. The set of organelles is radially symmetrical in more primitive phaeodarians. The presence of two parapylae makes this symmetry biradial. II. The sophisticated arrangement of cell compartments in advanced phaeodarians disrupts radial symmetry, so that the symmetry becomes bilateral. See the text for further explanation.

Fig. 74. Radiolarian calymma hypothetical arrangement. The vacuole is pumped with light ions, absorbs osmotic water, and thus fills with low-density aqueous solution (I). The large specific surface area serves this transport (II). 1, calymmar rhizopodium; 2, mitochondrion; 3, seawater; 4, lacunae; 5, calymmar vacuole.

Fig. 75. Phaeodarian life history. I, Adult trophont (я, nucleus; цк, capsular wall; фе, phaeodium; ск, radial and tangential skeletal spicules). II-IV, Polyploid nucleus fission and corresponding constriction of capsular wall. V, Skeletal elements of the parent cell are distributed between the offspring. VI, Phaeodium resorption; multipolar segregation of genomes in the polyploid nucleus. VII, Multinucleate stage. VIII, Capsular wall resorption; calymmar vacuoles collapse (not shown); multiple fission of the cell into spherical plasmodia (пм, plasma membrane). IX, Endomitoses, multipolar segregation of genomes. X, A spherical plasmodium splits into ameboid plasmodia (in Coelodendrum); ordinary karyokineses with a bipolar spindle. XI, An ameboid plasmodium splits into mononuclear “prespores”, which undergo one binary fission and produce biflagellated swarmers with a crystalloid (кр) in the cytoplasm. (After several authors).

Fig. 76. Nuclear fission in Aulacantha scolymantha. 1, capsular wall; 2, astropyle (after Cachon-Enjumet, 1961a).

Fig. 77. Schematic drawing of nuclear transformation in Phaeodaria in the course of binary fission (after Cachon-Enjumet, 1961a).

Fig. 78. Phases of the hypothetic nuclear cycle of the Phaeodaria.

Fig. 79. Concepts of phylogenetic relationships within the radiolaria (Phaeodaria, Polycystina and Acantharia) and Taxopodida, including early classifications (I-IV) and modern molecular phylogeny (V). Taxonomic ranges omitted. The original spelling of the taxa applied, ending modifications discarded.

Classification

Phylum Phaeodaria
1. Order Phaeocystida: skeleton of isolated spicules.
    1.1. Fam. Aulacanthidae
    1.2. Fam. Astracanthidae
2. Order Phaeosphaerida: skeleton is a subspherical lattice with trigonal, tetragonal or polygonal mesh; tangential rods have a lumen.
    2.1. Fam. Aulosphaeridae
    2.2. Fam. Sagosphaeridae
    2.3. Fam. Cannosphaeridae
3. Order Phaeocalpida: the porous spherical shell with radial spines has a single large opening (pylome) and thus has radial symmetry of infinite order.
    3.1. Fam. Castanellidae
    3.2. Fam. Circoporidae
    3.3. Fam. Polypyramidae
    3.4. Fam. Porospathidae
    3.5. Fam. Tuscaroridae
4. Order Phaeogromida: bilaterally or radially symmetrical shell has fine pores, a single large opening (pylome), and a few radial processes originating in certain positions.
    4.1. Fam. Lirellidae
    4.2. Fam. Challengeridae
    4.3. Fam. Medusettidae
5. Order Phaeogymnocellida: skeleton either lacking or present only at the astropyle.
    5.1. Fam. Atlanticellidae
    5.2. Fam. Phaeodinidae
    5.3. Fam. Phaeosphaeridae
6. Order Phaeoconchida: a bivalve shell is present.
    6.1. Fam. Concharidae
    6.2. Fam. Coelodendridae

Fig. 80. Schematic drawing of the distal part of radial spicule in different Aulacanthidae genera.

Fig. 81. Families Aulacanthidae (I), Circoporidae (II) and Castanellidae (III–VI).

I, Family 1.1 Aulacanthidae: species with two capsules Aulographis furcula; the phaeodium and the two central capsules above it are deeply colored; note radial spicules with terminal branches (рс) and numerous tangential spicules (тс). II, Family 3.2 Circoporidae: Circospathis sexfurca. III-VI, Family 3.1 Castanellidae: III, Castanidium variabile; IV, Castanidium podacontium, note radial spines mounted on pyramidal mounds; V, Castanella sloggetti; VI, Circocastanea margarita. Photo: all specimens from Bering Sea; I, ethanol fixation, carmine dye, inverted compound microscope Nikon TS100, by V.Zlatogurski; II-VI, dry specimens, dissecting microscope Leica M205 C, optical sections stacked using Helicon Focus software, by I.Kuznetsov. Scale bar I 1 mm, others 0.1 mm.

Fig. 82. Family 1.1 Aulacanthidae. I. Auloceros arborescens. II. Aulokleptes ramosus. III. The distal part of radial spicules of Aulographonium spp. IV. Aulospathis variabilis. Note radial spicules with terminal and side branches, tangential spicules, phaeodium and capsule with nucleus. V. The distal part of radial spicules of Aulodendron spp. Scale bar 0.1 mm. Line drawings from Reschetnjak (1966).

Fig. 84. Order 2. Phaeosphaerida. I. Aulastrum spinosum. II. Aulatractus fusiformis. III. Aulosphaera dendrophora. IV. Sagenoscena irmingeriana. V. Sagoscena elegans. VI. Auloscena verticillus. VII. Sagenoarium chuni. VIII. Coelacantha dogieli. Porous inner shell with a broad pylome. Spokes radiating from the inner shell attach to the midpoints of the tangential rods that compose the concentric latticed outer shell. Scale bar 0.1 mm. Line drawings: III from Haeckel (1887) and others from Reschetnjak (1966).

Fig. 85. Main diagnostic characters of Castanellidae genera. 1, main spines; 2, supplementary spines; 3, teeth.

Fig. 86. Order 3. Phaeocalpida. I, Castanissa megastomata. II, Castanarium antarcticum. III, Castanarium favosum. IV, Castanura primitiva. V, Castanea henseni. VI, Haeckeliana irregularis. Scale bar 0.1 mm. Line drawings from Reschetnjak (1966).

Fig. 87. Order 3. Phaeocalpida. I. Porospathis holostoma. II, Polypyramis fenestrata. III, Tuscarora bisternaria, 3-4 oral processes around the peristome, 3-4 somatic processes in equatorial position (distal parts not shown), phaeodium, a capsule with the surrounding frothy calymma. IV, Tuscarantha luciae. V, Tuscarilla scutellum. VI, Tuscarilla ampula. Scale bar I-II 0.1 mm, III-VI 1 mm. Line drawings from Reschetnjak (1966).

Fig. 88. Main diagnostic characters of Tuscaroridae genera.

Fig. 89. Family 3.5 Tuscaroridae, Tuscaretta belknapi. I, specimen with 4 oral processes and 4 somatic processes (one broken off). II, the shell contains the phaeodium (arrow) and two calymmas (arrowheads) around the two capsules. III, cytoplasmic strands stretch from the peristome into the axial lumen of oral processes. IV, oral processes and proximal parts of somatic processes are armed with spines; bases of somatic processes have large openings (arrow). Specimens from Bering Sea, ethanol fixation, no staining, in glycerol, dissecting microscope Leica M205 C, optical sections stacked using Helicon Focus software, by I.Kuznetsov. Scale bar I 1 mm, IV 0.1 mm.

Fig. 90. I, Family 4.2 Challengeridae, Protocystis sp. II-IV, family 4.2 Challengeridae, Challengeria naresi: II, transmitted and incident light, lateral view, note absence of pharynx and dry phaeodium seen through the wall; III, note the hexagonal pore pattern; IV, incident light, oblique view, note pylome. V, Family 6.1 Concharidae, Conchopsis barca: note the serrate hinge within the suture; pores slit-shaped near the keel and rounded near the hinge. Bering Sea, dry specimens, dissecting microscope Leica M205 C, optical sections stacked using Helicon Focus software, by I.Kuznetsov. Scale bar 0.1 mm.

Fig. 92. Family 4.3 Medusettidae: Gazelletta (I). Family 5.1 Atlanticellidae: Nationaletta fragilis (II) and Planktonetta parthenopaea Cachon-Enjumet (this is a Planktonetta s.l., a morph transitional between Nationaletta and Atlanticella) (III). 1 parapylae, 2 capsule, 3 nucleus, 4 peristome, 5 astropyle, 6 phaeodium, 7 secondary shell. (From Cachon-Enjumet, 1961a)

Fig. 93. Family 5.1 Atlanticellidae. I, Planktonetta atlantica. II, skeleton of Atlanticella bicornis. 1, central capsule. 2, the peristome of the primary shell. 3, proximal parts of the oral processes subdivided internally into alveoli. 4, the secondary shell that embraces the phaeodium. 5, the float. The living position of Planktonetta (I) is probably 180°, with the float up (Fowler, 1903). Scale bar 0.1 mm. Line drawings from Reschetnjak (1966)

Fig. 94. Family 5.1 Atlanticellidae. Schematic drawing of Nationaletta. 1, the central capsule functioning as the float. 2, the peristome of the lost primary shell. 3, proximal parts of the oral processes (branching not shown). 4, the secondary shell embracing the phaeodium

Fig. 95. Family 5.1 Atlanticellidae: I, Gymnocella tregouboffi; II-III, Halocella inermis; IV-V, Halocella borgerti; VI, Lobocella tenella; VII, Lobocella proteus; VIII, Lobocella tenella; IX, Miracella ovulum. 1 parapyle, 2 nucleus, 3 astropyle. (From Cachon-Enjumet, 1961a).

Fig. 96. Family 5.2 Phaeodinidae: I-II, Phaeodina valdiviae; III, Phaeodina pelagica. Family 5.3 Phaeosphaeridae: IV, Phaeosphaera nucleoelongata; V, Phaeosphaera pigmaea; VI, Phaeopyla sphaerica. (From Cachon-Enjumet, 1961a).

Fig. 97. Family 5.3 Phaeosphaeridae: I, Phaeodactylis hollandei; II, Phaeodactylis radiata; III, Phaeodactylis fagei. (From Cachon-Enjumet, 1961a).

Fig. 98. Diagrammatic drawing of Coelodendridae skeleton. 1 capsule; 2 valves; 3 rhinocanna; 4 openning of rhinocanna; 5 galea; 6 frenulum; 7 anterior primary tube; 8 lateral primary tube; 9 verticillate style; 10 anchor threads. See text for further explanations. Line drawing from Reschetnjak (1966).

Fig. 99. Family 6.2 Coelodendridae. I, Coelodendrum ramosissimum. II, C. ramosissimum: shell, galea and the bases of the four radial processes. III. Coelotetraceras xanthacanthum. IV. Coeloplegma murrayana. Scale bar 0.1 mm. Line drawings I, III, IV from Reschetnjak (1966) and II from Haeckel, 1862.

Fig. 100. Family 6.2 Coelodendridae. I, Coelographis antarcticum, lateral view. II, Coelothyrsus cypripedium (distal portions of verticillate styles not shown). III, Coelanthemum auloceroides. Scale bar 0.1 mm. Line drawings from Reschetnjak (1966)

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