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Freboldinium regulum
From Fensome et al., 2019:
Freboldinium regulum Below, 1990, p.16–20, pl.1, figs.1–12; text-figs.4a–k. Holotype: Below, 1990, pl.1, figs.1–6. Contrary to the opinion of Lentin and Williams (1993, p.227), this name is validly published. Below (1987a, p.6) stated that, unless otherwise indicated, all the material is in his personal collection. Since Below (1987a, 1987b, 1990) are parts I to III of a single study, we agree with J. Jansonius (personal communication) that Below's (1987a) repository statement refers to all three publications, fulfilling the requirement of ICN Article 40.7 (see also the discussion on Article 40.7 in the Introduction). Age: Toarcian.
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Original description Below, 1990:
Description:
Vesicle/plate arrangement of the dinoflagellate Amphiesma cop, pop, cap, 4'-(5'), 3a, (5")-6"-(7"), 7c, 6'", l'"'一(2""),as, FM, Is, rs, ps; tegulation 4*-5*, 3**, 6***, 0****-2****, 7c, 3+, 4++-5++, 3+++; growth of thecal plates according to a peridinoidal pattern;
Zygote cyst habitus proximate, acavate, spheroidal to rounded polyhedral, coniform to tholiform epicyst distinctly smaller than the semiovaloidal to conical hypocyst with its rounded or flattened antapex, maximum diameter at the posterior cingulate margin; wall composed of a thin pedium and thin luxuria forming the finate septa; surface smooth, scabrate, shagrinate, and rarely with rings reflecting trichocyst pores, low, smooth, distally straight or irregularly finely undulate, serrate finate septa. Arrangement scheme XPR/cop, pop, cap, 4'-5', 3a, 5"-7", 7c, 6'", 1""-2'"', as, FM, ls, rs, ps; PR undifferentiated or consisting of a very small, circular to oval cop, which is congruently enclosed by a small pop, tiny cap ventrally adjoining, usually non-finate; variants of epicystal arrangement; 5 apical/3 anterior intercalaries/7 preongular: steno VII l' (VVL) VII 2' (L), camerate 3' (D), VI4' (R), VII5' (VR); tiny caroidal la (DL), large VII2a (D) and very small caroidal 3a (RDR), la and 3a often against 2a non-finate; anterior geniculate V-nE 1"(VVL), anterior bigcniculate VI2" (L), anterior linear IV-nE3" (DL), anterior linear 4" (DR), Finis 3" /4" dorsal anterior trigeniculate VH-nE 5" (R VR), anterior line are IV-nE 6" (VVR), very small anterior geniculate V-nE 7'' (V); or 4 apical / 3 antapical intercalary / 6 precingulate: steno VIII l' (V), VIII2' (L), camerate 3' (D), VI4' (RDR); tiny caroidal LA (DL), large vn2a (D) and very small caroidal 3a (RDR), la and 3a often against 2a nonfinate; anterior geniculate VnE 1', (VVL), anterior bigeniculate VI-nE 2" (L), anterior linear JV-nE 3" (DDL), anterior linear IV-nE 4" (DR), finis 3" /4" dorsal, anterior trigeniculate 5" (RVR), anterior linear IV-nE 6"; or 4 apical/3 anterior intercalaries/5 precingular: steno VI-nE 1' (V ) metasert or insert, VIl-nE 2' (VL), small camerate 3' (DDL), VII-nE 4' (R); tiny caroidal la (L), large VII2a (DOR) and very small caroidal 3a (R), Ia and 3a often against 2a nonfinate; anterior geniculate V-nE 1", very large, anterior bigeniculate VI-nE 2" (L), anterior linear IV-nE 3" (DDR), anterior trigeniculate 4" (RVR), anterior linear IV-nES" (VVR); Cingulum left-handed spiral, heptapartite, lati lc to 6c and iso 7c; 6 postcingulates: V-nE 1'" (V), posterior linear; V-nE 2'" (VL) with contact to the sulcus; posterior linear IV-nE3"' (LDL), posterior linear IV-nE4'" (D), posterior linear IV-nE5"' (RDR), posterior linear V-nE6'" (VR); 1 pole-side VI 1'"' or rarely with 2 antapics: VI 1"" (VL) and pole-side VI 2"", but then 3'" posteriorly geniculate; sulcus partit, small vias, vFM; very small vis and very small IVrs as well as steno omega-shaped VIps; archaeopyle precingular/apical, 3" + 4" + 1a + 2a + 3a + 3';
Operculum solvate (pars) and foederat (pars), opercular plates secat, general opercular formula 3"s + 4''s + (1a + 2a + 3a + 3')s.
Holotype margin:
Length 39 μm, width 30 μm, thickness 29 μm.
Note:
The few specimens of the species found so far are notable for their varied arrangement scheme. Specimens with 4 apicals and 5 or 6 precingulates are common; only one individual shows 5 apicals and 7 precingulates. The number of anterior intercalaries is consistently 3, but the tiny areas 1a and 3; are often not defined by 2a and therefore easily overlooked. The pattern 5', 3a, 7'' can be converted into the pattern 4', 3a, 6' by fusion of the theca plates 1' + 5' and 7'' + 6'; the position of 2', 3', 4': 1a, 2a, 3a, 1" to 6" remains unchanged. The reduction of the number of precingulate areas from 6 to 5 apparently occurred through fusion of 3'' and 4'' or displacement of 4'' by 3''. 2'' expands significantly counterclockwise, while the first and last two areas of the precingulate ring (individually the 4th and 5th areas, but homologous to the 5th and 6th areas of the pattern with 6 precingulates) retain their position. In contrast, the hypocystal pattern is almost stable. Usually, 6 postcingulates and only 1 (!) pole-standing antapic are visible. The latter is 1" for the individual, but corresponds to 2"" of the usual gonyaulacoid or peridinoid arrangement scheme.
2"' borders the sulcus and overlaps the pole-side antapic area. Very rarely, a further area is separated between 2'', and the pole-side area (individually 1'''' but also generally homologous to l'''' of the gonyaulacoid and peridinoid tabulation scheme), which separates 2"' from 2"", but does not extend to 1"', so that 2',' continues to border the sulcus. The contact of 2'" with the sulcus means an interruption of the secposterior tegula series, in the first case also with a reduction of this series by one plate. The following area contacts apply:
PR (1', 2', 4', 5'),
l' (as, 1", 2', PR, 5', 7"),
2' (l", 2", la, 3', 4', PR, l'),
3'(la, 2a, 3a, 4', 2'),
4' (3a, 5", 5', PR, 2', 3'), 5'(PR, 4', 5'', 6'', 7'', l'),
la (2", 2a, 3', 2'),
2a (2", 3", 4", 5", 3a, 3', la),
3a (5", 4', 3', 2a),
l" (c, 211 , 2', l', as),
2" (c, 3", 2a, la, 2', 1''),
or
PR (l', 2', 4'),
l'(as, 1", 2', PR, 4', 5", 6"), 2'(1", 2", la, 3', 4', PR, l'), 3'(la, 2a, 3a, 4', 2'),
4'(5", l', PR, 3', 3a),
la (2", 2a, 3', 2'),
2a (2", 3", 4", 5", 3a, 3', la}, 3a (5", 4', 3', 2a),
1" (c, 2", 2', l", as),
2" (c, 3", 2a, la, 2', l"),
3" (c, 4", 2a, 2"),
4" (c, 5", 2a, 3"),
5" (c, 6", l', 4', 3a, 2a, 4"),
6(c, as, l', 5"),
as (c, l", l', 6"),
or
PR (2', 3', 4'),
1' (as, 1", 2', 4', 4", 5"),
2' (l", 2", la, 3', PR, l'),
3' (la, 2a, 3a, 4', PR, 2'),
4' (4'', 5'', l', 2', 3', PR, 3a),
1a (2'', 2a, 3', 2'),
2a (2'', 3'', 4'', 3a, 3', 1a),
3a (4'', 4', 3', 2a),
1'' (c, 2'', 2', 1', as),
2'' (c, 3'', 2a, 1a, 2', 1''),
3'' (c, 4'', 2a, 2'')
4'' (c, 5", 1', 4'', 3a, 2a, 3''),
5'' (c, as, 1', 4''),
as (c, 1'', 1', 5'),
1''' (s, 2''', c),
2''' (s, 1'''', 3''', c),
3''' (2''', 1'''', 4''', c),
4''' (3''', 1'''', 5''', c),
5''' (4''', 1'''', 6''', c),
6''' (5''', 1'''', s, c),
or
l'" (s, 2'", c),
2''' (s, 1'''', 3''', c),
3''' (2''', 1'''', 2'''', 4''', c),
4''' (3''', 2'''', 5''', c),
5''' (4''', 2'''', 6''', c),
6'''(5''', 2'''', s, c).
Freboldinium regulum Below, 1990, p.16–20, pl.1, figs.1–12; text-figs.4a–k. Holotype: Below, 1990, pl.1, figs.1–6. Contrary to the opinion of Lentin and Williams (1993, p.227), this name is validly published. Below (1987a, p.6) stated that, unless otherwise indicated, all the material is in his personal collection. Since Below (1987a, 1987b, 1990) are parts I to III of a single study, we agree with J. Jansonius (personal communication) that Below's (1987a) repository statement refers to all three publications, fulfilling the requirement of ICN Article 40.7 (see also the discussion on Article 40.7 in the Introduction). Age: Toarcian.
---------------------------------------------------------------------------------------------------
Original description Below, 1990:
Description:
Vesicle/plate arrangement of the dinoflagellate Amphiesma cop, pop, cap, 4'-(5'), 3a, (5")-6"-(7"), 7c, 6'", l'"'一(2""),as, FM, Is, rs, ps; tegulation 4*-5*, 3**, 6***, 0****-2****, 7c, 3+, 4++-5++, 3+++; growth of thecal plates according to a peridinoidal pattern;
Zygote cyst habitus proximate, acavate, spheroidal to rounded polyhedral, coniform to tholiform epicyst distinctly smaller than the semiovaloidal to conical hypocyst with its rounded or flattened antapex, maximum diameter at the posterior cingulate margin; wall composed of a thin pedium and thin luxuria forming the finate septa; surface smooth, scabrate, shagrinate, and rarely with rings reflecting trichocyst pores, low, smooth, distally straight or irregularly finely undulate, serrate finate septa. Arrangement scheme XPR/cop, pop, cap, 4'-5', 3a, 5"-7", 7c, 6'", 1""-2'"', as, FM, ls, rs, ps; PR undifferentiated or consisting of a very small, circular to oval cop, which is congruently enclosed by a small pop, tiny cap ventrally adjoining, usually non-finate; variants of epicystal arrangement; 5 apical/3 anterior intercalaries/7 preongular: steno VII l' (VVL) VII 2' (L), camerate 3' (D), VI4' (R), VII5' (VR); tiny caroidal la (DL), large VII2a (D) and very small caroidal 3a (RDR), la and 3a often against 2a non-finate; anterior geniculate V-nE 1"(VVL), anterior bigcniculate VI2" (L), anterior linear IV-nE3" (DL), anterior linear 4" (DR), Finis 3" /4" dorsal anterior trigeniculate VH-nE 5" (R VR), anterior line are IV-nE 6" (VVR), very small anterior geniculate V-nE 7'' (V); or 4 apical / 3 antapical intercalary / 6 precingulate: steno VIII l' (V), VIII2' (L), camerate 3' (D), VI4' (RDR); tiny caroidal LA (DL), large vn2a (D) and very small caroidal 3a (RDR), la and 3a often against 2a nonfinate; anterior geniculate VnE 1', (VVL), anterior bigeniculate VI-nE 2" (L), anterior linear JV-nE 3" (DDL), anterior linear IV-nE 4" (DR), finis 3" /4" dorsal, anterior trigeniculate 5" (RVR), anterior linear IV-nE 6"; or 4 apical/3 anterior intercalaries/5 precingular: steno VI-nE 1' (V ) metasert or insert, VIl-nE 2' (VL), small camerate 3' (DDL), VII-nE 4' (R); tiny caroidal la (L), large VII2a (DOR) and very small caroidal 3a (R), Ia and 3a often against 2a nonfinate; anterior geniculate V-nE 1", very large, anterior bigeniculate VI-nE 2" (L), anterior linear IV-nE 3" (DDR), anterior trigeniculate 4" (RVR), anterior linear IV-nES" (VVR); Cingulum left-handed spiral, heptapartite, lati lc to 6c and iso 7c; 6 postcingulates: V-nE 1'" (V), posterior linear; V-nE 2'" (VL) with contact to the sulcus; posterior linear IV-nE3"' (LDL), posterior linear IV-nE4'" (D), posterior linear IV-nE5"' (RDR), posterior linear V-nE6'" (VR); 1 pole-side VI 1'"' or rarely with 2 antapics: VI 1"" (VL) and pole-side VI 2"", but then 3'" posteriorly geniculate; sulcus partit, small vias, vFM; very small vis and very small IVrs as well as steno omega-shaped VIps; archaeopyle precingular/apical, 3" + 4" + 1a + 2a + 3a + 3';
Operculum solvate (pars) and foederat (pars), opercular plates secat, general opercular formula 3"s + 4''s + (1a + 2a + 3a + 3')s.
Holotype margin:
Length 39 μm, width 30 μm, thickness 29 μm.
Note:
The few specimens of the species found so far are notable for their varied arrangement scheme. Specimens with 4 apicals and 5 or 6 precingulates are common; only one individual shows 5 apicals and 7 precingulates. The number of anterior intercalaries is consistently 3, but the tiny areas 1a and 3; are often not defined by 2a and therefore easily overlooked. The pattern 5', 3a, 7'' can be converted into the pattern 4', 3a, 6' by fusion of the theca plates 1' + 5' and 7'' + 6'; the position of 2', 3', 4': 1a, 2a, 3a, 1" to 6" remains unchanged. The reduction of the number of precingulate areas from 6 to 5 apparently occurred through fusion of 3'' and 4'' or displacement of 4'' by 3''. 2'' expands significantly counterclockwise, while the first and last two areas of the precingulate ring (individually the 4th and 5th areas, but homologous to the 5th and 6th areas of the pattern with 6 precingulates) retain their position. In contrast, the hypocystal pattern is almost stable. Usually, 6 postcingulates and only 1 (!) pole-standing antapic are visible. The latter is 1" for the individual, but corresponds to 2"" of the usual gonyaulacoid or peridinoid arrangement scheme.
2"' borders the sulcus and overlaps the pole-side antapic area. Very rarely, a further area is separated between 2'', and the pole-side area (individually 1'''' but also generally homologous to l'''' of the gonyaulacoid and peridinoid tabulation scheme), which separates 2"' from 2"", but does not extend to 1"', so that 2',' continues to border the sulcus. The contact of 2'" with the sulcus means an interruption of the secposterior tegula series, in the first case also with a reduction of this series by one plate. The following area contacts apply:
PR (1', 2', 4', 5'),
l' (as, 1", 2', PR, 5', 7"),
2' (l", 2", la, 3', 4', PR, l'),
3'(la, 2a, 3a, 4', 2'),
4' (3a, 5", 5', PR, 2', 3'), 5'(PR, 4', 5'', 6'', 7'', l'),
la (2", 2a, 3', 2'),
2a (2", 3", 4", 5", 3a, 3', la),
3a (5", 4', 3', 2a),
l" (c, 211 , 2', l', as),
2" (c, 3", 2a, la, 2', 1''),
or
PR (l', 2', 4'),
l'(as, 1", 2', PR, 4', 5", 6"), 2'(1", 2", la, 3', 4', PR, l'), 3'(la, 2a, 3a, 4', 2'),
4'(5", l', PR, 3', 3a),
la (2", 2a, 3', 2'),
2a (2", 3", 4", 5", 3a, 3', la}, 3a (5", 4', 3', 2a),
1" (c, 2", 2', l", as),
2" (c, 3", 2a, la, 2', l"),
3" (c, 4", 2a, 2"),
4" (c, 5", 2a, 3"),
5" (c, 6", l', 4', 3a, 2a, 4"),
6(c, as, l', 5"),
as (c, l", l', 6"),
or
PR (2', 3', 4'),
1' (as, 1", 2', 4', 4", 5"),
2' (l", 2", la, 3', PR, l'),
3' (la, 2a, 3a, 4', PR, 2'),
4' (4'', 5'', l', 2', 3', PR, 3a),
1a (2'', 2a, 3', 2'),
2a (2'', 3'', 4'', 3a, 3', 1a),
3a (4'', 4', 3', 2a),
1'' (c, 2'', 2', 1', as),
2'' (c, 3'', 2a, 1a, 2', 1''),
3'' (c, 4'', 2a, 2'')
4'' (c, 5", 1', 4'', 3a, 2a, 3''),
5'' (c, as, 1', 4''),
as (c, 1'', 1', 5'),
1''' (s, 2''', c),
2''' (s, 1'''', 3''', c),
3''' (2''', 1'''', 4''', c),
4''' (3''', 1'''', 5''', c),
5''' (4''', 1'''', 6''', c),
6''' (5''', 1'''', s, c),
or
l'" (s, 2'", c),
2''' (s, 1'''', 3''', c),
3''' (2''', 1'''', 2'''', 4''', c),
4''' (3''', 2'''', 5''', c),
5''' (4''', 2'''', 6''', c),
6'''(5''', 2'''', s, c).