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Diphyes
From Williams et al., 2017:
[Diphyes, Cookson, 1965a, p. 85, nom. cons.; Emendations: Davey and Williams, 1966b, p. 95-96; Goodman and Witmer, 1985, p. 76-77
Type species: as Hystrichosphaeridium colligerum, Deflandre and Cookson, 1955 (pl.7, fig.3)] ; Diphyes colligerum, emend. Cookson, 1965; emend. Davey and Williams, 1966; emend. Goodman and Witmer, 1985
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Original description: [Cookson, 1965]:
Description:
Shell composed of two parts, more or less unequal in size. The anterior part circular in outline with or without appendages. Archeopyle apical. The posterior part roughly bell-shaped to spherical with a small circular antapical opening and without typical appendages.
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Emended description:
Davey and Williams, 1966:
Diagnosis:
Chorate cysts with ovoidal to spherical central body composed of two layers. Processes numerous, 1 to 4 per plate area, hollow and either open or closed distally. Large antapical process occurring opposite apical archaeopyle.
Affinities:
The genus is here restricted to include forms possessing two types of processes - numerous fine ones and a single large antapical. D. nudum Cookson does not possess any processes are all and certainly does not belong in this genus.
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Goodman and Witmer, 1985:
Diagnosis:
Skolochorate cysts with subspheroidal body bearing numerous nontabular or intratabular processes of nearly uniform size and a single large antapical process. Paratabulation gonyaulacacean; indicated by archeopyle, shape of the antapical process base, and rarely by the tendency of adjacent process bases to fuse into clusters apparently within the limits of individual paraplates. Archeopyle variable: Type (4A), (4A)P 2P or P.
Description:
Shape: Body subspheroidal: antapical margin modified by large process with rectangular to broadly ovate outline.
Wall relationships: Endophragm and periphragm appressed except beneath processes.
Wall features: No parasutural features. Nontabular or intratabular processes faintly striate or fibrous, hollow, may be open or closed distally. Antapical process narrowly or broadly cylindrical, typically with a distal pore whose diameter is usually much less than the diameter of the process. Antapical process may bear several small, open-tipped spinules near its distal end.
Paratabulation: Typically expressed only by principal and accessory archeopyle sutures. On rare specimens, a gonyaulacacean paratabulation can be inferred from arrangement of clustered processes (whose bases may be fused) in conjunction with principal archeopyle suture and shape of the antapical process base. Formula derived from these features is 1-2pr, 4`, 6``, Xc, 6```, 1p, 1````. Some sulcal paraplates can be recognized, typically the anterior sulcal and more rarely others.
Archeopyle: Variable, may be Type P(3``), 2P(3-4``), (4A(1-4`)P(3``)), or (4A(1-4`)); archeopyle suture on apical archeopyles faintly to moderately zigzag; operculum is free.
Paracingulum: Not expressed, or may be suggested by the equatorial alignment of processes.
Parasulcus: Typically not expressed, or rarely indicated by smaller than normal processes in midventral region.
Size: Small to intermediate (main body transdiameter generally between 30 and 45 µm).
Affinities:
Diphyes is characterized principally by its large antapical process, smaller nontabular or intratabular processes over the remainder of the cyst body, and variable archeopyle types. In effect, the generic description used herein is essentially that of Stover and Evitt (1978), with the additional characteristics of variable archeopyle types and (in rare specimens) a demonstrable gonyaulacacean paratabulation in D. colligerum, the type species of the genus, contributing to the revision.
Although paratabulation has not been demonstrated in detail for species of Diphyes prior to this study, the apparent alignment of processes into five latitudinal rows for Maastrichtian specimens of D. colligerum and D. recurvatum had previously been noted (May, 1980) as a possible expression of paratabulation.
Diphyes morphotypes with apical, Type (4A), archeopyles are similar to Dapsilidinium Bujak et al., 1980 which bears processes similar to those on
Diphyes, but lacks a large antapical process.
Species of Polysphaeridium Davey & Williams, 1966, also lack a large antapical process, and they exhibit panepicystal archeopyle (one that includes all the epicystal paraplates).
Hystrichokolpoma Klumpp, 1953 has large intratabular precingular and postcingular processes.
Coronifera Cookson & Eisenack, 1958 resembles in general form those morphotypes of Diphyes having a Type P archeopyle, but can be differentiated by its more numerous and more densely distributed nontabular processes, by its single wall layer (autophragm), and by the shape of its antapical process. In Diphyes, the antapical process tends to be more broadly cylindrical and is typically boxlike in longitudinal section whereas in Coronifera it is more trumpet-shaped with a rapidly flaring, often serrated distal margin.
Both Silicisphaera Davey & Verdier, 1976 and Florentinia Davey & Verdier, 1973 superficially resemble some morphotypes of Diphyes. Both genera
may have either a precingular, Type P, or combination, Type (4A)P, archeopyle. However, Silicisphaera lacks a large antapical process, and Florentinia bears large precingular and postcingular processes. Stover and Evitt (1978) state that consistent separation of Florentinia and Silicisphaera may be difficult, although there appears to be little chance of confusing them with Diphyes. Those specimens of Diphyes with an apical, Type (4A), archeopyle tend to have fused process bases, and neither Florentinia nor Silicisphaera have an apical archeopyle.
Diphyes is very similar to Duospheridium Loeblich & Loeblich, 1968 in many aspects of gross morphology. Both have a spheroidal main body, a large antapical process, and an apical archeopyle. However, Duosphaeridium lacks processes on the anterior body, generally has a considerably more inflated antapical process, and does not develop other archeopyle types as does Diphyes.
Cysts of Diphyes are not known to occur in Recent marine sediments, and cyst-theca relationships to living motile stage dinoflagellates therefore
cannot be established.
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Modified description:
Stover and Evitt 1978, p. 38-39:
Synopsis:
Cysts skolochorate, body subspherical with numerous nontabular processes of essentially uniform size and a single large antapical process; paratabulation indicated by archeopyle only; latter apical, Type tA.
Description:
Shape: Body subspherical; antapical margin modified by large process with a somewhat rectangular to broadly ovate outline.
Wall relationships: Endophragm and periphragm appressed except beneath processes.
Wall features: No parasutural features. Nontabular processes faintly striate or fibrous, hollow, normally open distally. Antapical process narrowly to broadly cylindrical with a distal pore whose diameter is usually much less than the diameter of the process.
Paratabulation: Expressed solely by principal and accessory archeopyle sutures.
Archeopyle: Apical, Type tA; principal archeopyle suture faintly to moderate zigzag; operculum free.
Paracingulum: Not expressed or suggested by the equatorial alignment of processes.
Parasulcus: Not expressed.
Size: Small to intermediate.
Affinities:
The most diagnostic feature of Diphyes is the distinctive, large antapical process. Another genus possessing this feature and nontabular processes is Coronifera, which differs from Diphyes in having a precingular archeopyle rather than an apical archeopyle, and normally closed processes (except the antapical one). In Duosphaeridium, the antapical structure may be larger and more inflated than in Diphyes, and no processes are present on the anterior part. Polysphaeridium, which has processes like those on Diphyes, lacks the large antapical process.
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Notes:
G.L. Williams short notes on species, Mesozoic-Cenozoic dinocyst course, Urbino, Italy, May 17-22, 1999 - LPP VIEWER CD-ROM 99.5.
Diphyes Cookson, 1965, emend. Davey and Williams, 1966b, emend. Goodman and Witmer, 1985. Emended diagnosis, from Goodman and Witmer (1985, p.76), Skolochorate cysts with subspheroidal body bearing numerous nontabular or intratabular processes of nearly uniform size and a single large antapical process. Paratabulation gonyaulacacean; indicated by archeopyle, shape of the antapical process base, and rarely by the tendency of adjacent process bases to fuse into clusters apparently with the limits of individual paraplates. Archeopyle variable: type (4A), (4A)P, 2P, or P.
[Diphyes, Cookson, 1965a, p. 85, nom. cons.; Emendations: Davey and Williams, 1966b, p. 95-96; Goodman and Witmer, 1985, p. 76-77
Type species: as Hystrichosphaeridium colligerum, Deflandre and Cookson, 1955 (pl.7, fig.3)] ; Diphyes colligerum, emend. Cookson, 1965; emend. Davey and Williams, 1966; emend. Goodman and Witmer, 1985
-------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Original description: [Cookson, 1965]:
Description:
Shell composed of two parts, more or less unequal in size. The anterior part circular in outline with or without appendages. Archeopyle apical. The posterior part roughly bell-shaped to spherical with a small circular antapical opening and without typical appendages.
-------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Emended description:
Davey and Williams, 1966:
Diagnosis:
Chorate cysts with ovoidal to spherical central body composed of two layers. Processes numerous, 1 to 4 per plate area, hollow and either open or closed distally. Large antapical process occurring opposite apical archaeopyle.
Affinities:
The genus is here restricted to include forms possessing two types of processes - numerous fine ones and a single large antapical. D. nudum Cookson does not possess any processes are all and certainly does not belong in this genus.
------------------------------------------
Goodman and Witmer, 1985:
Diagnosis:
Skolochorate cysts with subspheroidal body bearing numerous nontabular or intratabular processes of nearly uniform size and a single large antapical process. Paratabulation gonyaulacacean; indicated by archeopyle, shape of the antapical process base, and rarely by the tendency of adjacent process bases to fuse into clusters apparently within the limits of individual paraplates. Archeopyle variable: Type (4A), (4A)P 2P or P.
Description:
Shape: Body subspheroidal: antapical margin modified by large process with rectangular to broadly ovate outline.
Wall relationships: Endophragm and periphragm appressed except beneath processes.
Wall features: No parasutural features. Nontabular or intratabular processes faintly striate or fibrous, hollow, may be open or closed distally. Antapical process narrowly or broadly cylindrical, typically with a distal pore whose diameter is usually much less than the diameter of the process. Antapical process may bear several small, open-tipped spinules near its distal end.
Paratabulation: Typically expressed only by principal and accessory archeopyle sutures. On rare specimens, a gonyaulacacean paratabulation can be inferred from arrangement of clustered processes (whose bases may be fused) in conjunction with principal archeopyle suture and shape of the antapical process base. Formula derived from these features is 1-2pr, 4`, 6``, Xc, 6```, 1p, 1````. Some sulcal paraplates can be recognized, typically the anterior sulcal and more rarely others.
Archeopyle: Variable, may be Type P(3``), 2P(3-4``), (4A(1-4`)P(3``)), or (4A(1-4`)); archeopyle suture on apical archeopyles faintly to moderately zigzag; operculum is free.
Paracingulum: Not expressed, or may be suggested by the equatorial alignment of processes.
Parasulcus: Typically not expressed, or rarely indicated by smaller than normal processes in midventral region.
Size: Small to intermediate (main body transdiameter generally between 30 and 45 µm).
Affinities:
Diphyes is characterized principally by its large antapical process, smaller nontabular or intratabular processes over the remainder of the cyst body, and variable archeopyle types. In effect, the generic description used herein is essentially that of Stover and Evitt (1978), with the additional characteristics of variable archeopyle types and (in rare specimens) a demonstrable gonyaulacacean paratabulation in D. colligerum, the type species of the genus, contributing to the revision.
Although paratabulation has not been demonstrated in detail for species of Diphyes prior to this study, the apparent alignment of processes into five latitudinal rows for Maastrichtian specimens of D. colligerum and D. recurvatum had previously been noted (May, 1980) as a possible expression of paratabulation.
Diphyes morphotypes with apical, Type (4A), archeopyles are similar to Dapsilidinium Bujak et al., 1980 which bears processes similar to those on
Diphyes, but lacks a large antapical process.
Species of Polysphaeridium Davey & Williams, 1966, also lack a large antapical process, and they exhibit panepicystal archeopyle (one that includes all the epicystal paraplates).
Hystrichokolpoma Klumpp, 1953 has large intratabular precingular and postcingular processes.
Coronifera Cookson & Eisenack, 1958 resembles in general form those morphotypes of Diphyes having a Type P archeopyle, but can be differentiated by its more numerous and more densely distributed nontabular processes, by its single wall layer (autophragm), and by the shape of its antapical process. In Diphyes, the antapical process tends to be more broadly cylindrical and is typically boxlike in longitudinal section whereas in Coronifera it is more trumpet-shaped with a rapidly flaring, often serrated distal margin.
Both Silicisphaera Davey & Verdier, 1976 and Florentinia Davey & Verdier, 1973 superficially resemble some morphotypes of Diphyes. Both genera
may have either a precingular, Type P, or combination, Type (4A)P, archeopyle. However, Silicisphaera lacks a large antapical process, and Florentinia bears large precingular and postcingular processes. Stover and Evitt (1978) state that consistent separation of Florentinia and Silicisphaera may be difficult, although there appears to be little chance of confusing them with Diphyes. Those specimens of Diphyes with an apical, Type (4A), archeopyle tend to have fused process bases, and neither Florentinia nor Silicisphaera have an apical archeopyle.
Diphyes is very similar to Duospheridium Loeblich & Loeblich, 1968 in many aspects of gross morphology. Both have a spheroidal main body, a large antapical process, and an apical archeopyle. However, Duosphaeridium lacks processes on the anterior body, generally has a considerably more inflated antapical process, and does not develop other archeopyle types as does Diphyes.
Cysts of Diphyes are not known to occur in Recent marine sediments, and cyst-theca relationships to living motile stage dinoflagellates therefore
cannot be established.
-------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Modified description:
Stover and Evitt 1978, p. 38-39:
Synopsis:
Cysts skolochorate, body subspherical with numerous nontabular processes of essentially uniform size and a single large antapical process; paratabulation indicated by archeopyle only; latter apical, Type tA.
Description:
Shape: Body subspherical; antapical margin modified by large process with a somewhat rectangular to broadly ovate outline.
Wall relationships: Endophragm and periphragm appressed except beneath processes.
Wall features: No parasutural features. Nontabular processes faintly striate or fibrous, hollow, normally open distally. Antapical process narrowly to broadly cylindrical with a distal pore whose diameter is usually much less than the diameter of the process.
Paratabulation: Expressed solely by principal and accessory archeopyle sutures.
Archeopyle: Apical, Type tA; principal archeopyle suture faintly to moderate zigzag; operculum free.
Paracingulum: Not expressed or suggested by the equatorial alignment of processes.
Parasulcus: Not expressed.
Size: Small to intermediate.
Affinities:
The most diagnostic feature of Diphyes is the distinctive, large antapical process. Another genus possessing this feature and nontabular processes is Coronifera, which differs from Diphyes in having a precingular archeopyle rather than an apical archeopyle, and normally closed processes (except the antapical one). In Duosphaeridium, the antapical structure may be larger and more inflated than in Diphyes, and no processes are present on the anterior part. Polysphaeridium, which has processes like those on Diphyes, lacks the large antapical process.
-------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Notes:
G.L. Williams short notes on species, Mesozoic-Cenozoic dinocyst course, Urbino, Italy, May 17-22, 1999 - LPP VIEWER CD-ROM 99.5.
Diphyes Cookson, 1965, emend. Davey and Williams, 1966b, emend. Goodman and Witmer, 1985. Emended diagnosis, from Goodman and Witmer (1985, p.76), Skolochorate cysts with subspheroidal body bearing numerous nontabular or intratabular processes of nearly uniform size and a single large antapical process. Paratabulation gonyaulacacean; indicated by archeopyle, shape of the antapical process base, and rarely by the tendency of adjacent process bases to fuse into clusters apparently with the limits of individual paraplates. Archeopyle variable: type (4A), (4A)P, 2P, or P.