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Diphyes colligerum
Diphyes colligerum (Deflandre and Cookson, 1955, p.278-279, pl.7, fig.3) Cookson, 1965a, p.86-87. Emendations: Cookson, 1965a, p.86 and Goodman and Witmer, 1985, p.77-78, both as Diphyes colligerum.
Originally Hystrichosphaeridium, subsequently Baltisphaeridium (acritarch), thirdly (and now) Diphyes.
Taxonomic junior synonym: Diphyes pseudoficusoides according to Fensome et al. (2009, p.30).
Holotype: Deflandre and Cookson, 1955, pl.7, fig.3.
Hypotypes: Cookson, 1965, pl.9, fig.1-2
Locus typicus: Dilwyn Clay, SE of mouth of Gellibrand River, Victoria, Australia
Age: Early Eocene.
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G.L. Williams short notes on species, Mesozoic-Cenozoic dinocyst course, Urbino, Italy, May 17-22, 1999 - LPP VIEWER CD-ROM 99.5.
Diphyes colligerum (Deflandre and Cookson, 1955) Cookson, 1965a, emend. Cookson, 1965a, emend. Goodman and Witmer, 1985. Processes numerous, tubular, straight or slightly flexuous, with a conical base and a closed capitate or slightly furcate apex. At one pole is a large, hollow cylindro-conical outgrowth closed by a rounded conical apex. This large process may have short conical knobs. Goodman and Witmer (1985, p.77-78) in emending the species, noted that there are four types of archeopyle. These are type P3", type 2P3-4", apical type (4A), or combination type (4A)P. The most common are the apical and 3" types. Operculum is always free. Size: overall 56-59 µm, shell 30-33 µm, processes 13 µm.
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Original description: Deflandre and Cookson 1955, p. 278-279: Hystrichosphaeridium colligerum
Diagnosis: Shell globular, almost spherical; processes numerous, tubular, straight or slightly flexuous, with a conical base and a closed capitate or slightly furcate apex. At one pole of the shell there is a large, hollow, cylindro-conical outgrowth closed by a rounded conical apex. This outgrowth, like the small processes, does not communicate with the interior of the shell. It is frequently ornamented distally with a circle of short, blunt, hollow, conical knobs. Membrane of shell finely granular.
Dimensions: holotype: diameter of shell 33 Ám, overall diameter 56 Ám, large process 20 by 13 Ám, small processes c. 13 Ám long. Range: shell 30-33 Ám, overall diameter: 56-59 Ám, appendages c. 13 Ám.
Emended description: Cookson, 1965, p. 86
Anterior portion of shell almost spherical, dorso-ventral, with numerous broadly- based, stiff or flexuous, usually unbranched tubular appendages of varying lengths and widths that narrow towards small bluntly- pointed, capitate or somewhat funnel-shaped apices. On one surface, taken as ventral, some of the appendages are arranged in two short longitudinal rows one on either side of the mid-line (Pl.9, fig.1,3); on the dorsal surface they are evenly distributed (Pl.9,fig.2,4); an archeopyle occupies a large part of the apex (Pl.9,fig.5). The appendages are hollow at and near their bases and, contrary to the statement of Deflandre and Cookson (1955), their cavities appear to communicate with that of the shell (Pl.9, fig.10). However, it is practically impossible to prove this point, as well as to trace the cavity along the whole length of the longer appendages owing to the narrowness of their distal portions and the relative thickness of their walls. The wall of the anterior portion of the shell, c. 1.5 Ám thick, is two-layered. The outer layer is densely granular in surface view the granules appearing as minute rods in optical section under oil immersion; similar rods are frequently evident on the walls of the appendages.
Posterior portion of the shell usually smaller and thinner-walled than the anterior portion, roughly bell-shaped with straight or convex sides and, in well preserved specimens, a centrally-placed, cone-like projection with a small circular opening (Pl.9,fig.12). The surface is devoid of appendages, apart from one or two circles of small, knob or spine-like thickenings in the vicinity of the antapical opening (Pl.9,fig.7,8,11). The wall, c. 1 Ám thick, is smooth in optical section, finely granular in surface view, the granules being arranged in more or less clearly defined longitudinal rows (Pl.9,fig.1).
Dimensions: Hypotypes: 1-overall length of shell 55 Ám, width of anterior part 31 Ám, width of posterior part c. 26 Ám; 2--overall length of shell 57 Ám, width of anterior part c. 33 Ám, width of posterior part 26 Ám.
Range: Overall length: 36-57 Ám; width, anterior part: 29-36 Ám; posterior part: 13-26 Ám; appendages: 5-21 Ám long.
Emended diagnosis: Davey and Williams, 1966, p. 96
Ovoidal to spherical central body with wall composed of thin endophragm and outer finely reticulate periphragm. Processes composed of periphragm, numerous, simple, hollow, open or closed distally. One single broad antapical process. Total number of processes exceeding 50.
Dimensions: Holotype: diameter of central body 33 Ám, overall diameter 56 Ám, length of antapical process 20 Ám, width of antapical process 13 Ám, length of small processes approx. 13 Ám. Range: diameter of central body 30-33 Ám, overall diameter 56-59 Ám, length of small processes approx. 13 Ám. Figured specimens diameter of central body 33 by 37.5 Ám, length of antapical process 16 Ám, width of antapical process 15 Ám, length of small processes 11-15 Ám. Range of English specimens: diameter of central body 29-41 Ám, length of antapical process 16-21 Ám, width of antapical process 8-15 Ám, length of small processes 8-15 Ám.
Description: An apical archaeopyle and the large antapical process enable easy orientation of specimens of this species. The antapical process is hollow, cylindro-conical, occasionally closed, and bears small tubules, 2-3 Ám long, towards the distal extremity. The tubules can take the form of simple conical protuberances or can have slightly bifurcate extremities. The smaller processes are commonly simple, occasionally united proximally, and may be open or closed distally. They are slender but do vary in width, and in specimens possessing open processes they taper to a distal neck before terminating with a sightly expanded opening. The distal margin may be finely serrate or entire. Both types of processes are slightly fibrous and do not communicate with the interior of the central body. There are commonly 4, regularly distributed, processes for each precingular plate area; 2-4 for the postcingular plate areas and a constant 2 in the cingular plate areas.
Revised description: Goodman and Witmer, 1985, p. 77-78
Subspheroidal bi-layered cyst with large, rectangular antapical process, and numerous slender nontabular to intratabular processes over remainder of body; endophragm and periphragm appressed except beneath processes. Antapical process open distally with an aperture of diameter less than that of the process and may or may not bear several small, typically open-tipped spinules near its distal margin; basal margin typically bears longitudinal striations. Other processes are considerably smaller and either taper continuously to a closed, capitate tip, or they may taper for the greater length of the process and terminate in a distally open and flared, or buccinate, tip. Both open and closed processes generally do not occur on single specimens. Smaller processes may be smooth or finely granular.
Archeopyle variable; four types involving different combinations and numbers of apical and/or precingular paraplates are known to date. Archeopyles may be either precingular, Type P (3''), or Type 2P (3''- 4''); apical, Type 4A (1'-4'); or combination: Type 4AP (1'-4', 3''). Types P and 4A are most common, while Types 4AP and 2P are relatively rare. Operculum is free for all four variations, and simple for 4A apical variation; it could not be determined whether the opercula for Type 2P and 4AP archeopyles were simple or compound. Archeopyle margin on specimens with Type 4A and 4AP archeopyles show sixth precingular notch rather than the usual parasulcal notch (Plate 6, figs. 1-2), and corresponding extension on operculum is formed by fourth apical rather than first apical paraplate. Paratabulation typically expressed only by principal and accessory archeopyle sutures; outer wall bears no other archeopyle features. A gonyaulacacean paratabulation can be inferred for rare specimens which show fusion of process bases and clustering of adjacent, apparently intratabular processes, as well as a polygonal outline to the base of the antapical process. These features, in conjunction with details of archeopyle (and operculum) morphology, indicate a formula of 1-2pr, 4', 6'', Xc, 6''', 1p, 1''''. Sulcal processes are present but difficult to identify on most specimens.
Endophragm up to 1 Ám thick, and appears externally smooth. Periphragm may be up to 0.5 Ám thick, and is externally smooth to finely granulose. Paracingulum typically expressed, or may be represented by equatorial alignment of processes. Parasulcus, except for paraplates as, typically not expressed as a distinct feature; rare specimens bear smaller-than-normal processes in mid-ventral region. On very rare specimens the position of these processes, in conjunction with other features (e.g., shape and antapical process), may be interpreted to suggest five or six sulcal processes in a standard gonyaulacacean arrangement.
Affinities:
Davey and Williams, 1966, p. 97
In the original diagnosis, the smaller processes are said to be closed. The figure by Deflandre & Cookson (1955, pl. 7, fig. 3) seems to indicate, however, that the processes are open, as in the examples from the London Clay.
Cookson & Eisenack (1961, pl. 2, fig. 9) figured H. colligerum with the processes definitely open. The species may well be confined to examples with open processes after re-examination of the holotype.
Cookson (1965, pl. 9, figs. 1-3) illustrates forms with distinctively larger antapical processes not unlike those of Litosphaeridium siphoniphorum (Cookson & Eisenack). Because of the larger size of this process she has incorrectly considered it to represent a second portion of the central body and has erected a new genus diagnosed as possessing bipartite cysts. The forms from the London Clay, the holotype of D. colligerum and the specimen illustrated by Cookson & Eisenack (1961, pl. 2, fig. 9) are probably more typical of the species and show without doubt that the " posterior portion of the shell " is in fact an enlarged antapical process.
This species has been recorded from the Upper Cretaceous (probably Senonian) and Eocene of Australia, and in England only from the Eocene.
This species is easily distinguishable from all other types of dinoflagellate cysts.
Goodman and Witmer 1985, p. 78-79
The distinctive morphology of Diphyes colligerum - which includes the large antapical process and the numerous other small processes, variable archaeopyle types, and two wall layers - readily distinguishes it from most other fossil cysts. Despite resemblances to species of Coronifera, Duosphaeridium, Florentinia, Silicisphaera, Dapsilidinium and Polysphaeridium in regard to gross morphological features, consistent separation should be relatively easy upon critical analysis off all morphologic characters. Convergence of certain Diphyes colligerum morphotypes (i.e. those with extreme fusion of processes) and specimens of some species of Hystrichokolpoma may make differentiation of these genera somewhat difficult; perhaps the customary generic boundaries between the two need to be reevaluated.
Diphyes colligerum is distinguished from D. appendiculare on the basis of size and process type. The latter is approximately two or three times as large as D. colligerum, and bears processes with relatively wider bases and which taper very rapidly to hairlike ends, corresponding to Cookson and Eisenack (1970, p. 149).
D. colligerum is distinguished from D. ficusoides by the shape of the antapical process which is bell-shaped to cylindrical on the former and fig-shaped with a biconvex margin on the latter.
Diphyes spinulum (Drugg) Stover and Evitt, D. latiusculum Matsuoka, and D. recurvatum May could prove to be synonymous with D. colligerum but formal recognition of their synonomy must await reexamination of the holotypes.
Originally Hystrichosphaeridium, subsequently Baltisphaeridium (acritarch), thirdly (and now) Diphyes.
Taxonomic junior synonym: Diphyes pseudoficusoides according to Fensome et al. (2009, p.30).
Holotype: Deflandre and Cookson, 1955, pl.7, fig.3.
Hypotypes: Cookson, 1965, pl.9, fig.1-2
Locus typicus: Dilwyn Clay, SE of mouth of Gellibrand River, Victoria, Australia
Age: Early Eocene.
--------------------------------------------------
G.L. Williams short notes on species, Mesozoic-Cenozoic dinocyst course, Urbino, Italy, May 17-22, 1999 - LPP VIEWER CD-ROM 99.5.
Diphyes colligerum (Deflandre and Cookson, 1955) Cookson, 1965a, emend. Cookson, 1965a, emend. Goodman and Witmer, 1985. Processes numerous, tubular, straight or slightly flexuous, with a conical base and a closed capitate or slightly furcate apex. At one pole is a large, hollow cylindro-conical outgrowth closed by a rounded conical apex. This large process may have short conical knobs. Goodman and Witmer (1985, p.77-78) in emending the species, noted that there are four types of archeopyle. These are type P3", type 2P3-4", apical type (4A), or combination type (4A)P. The most common are the apical and 3" types. Operculum is always free. Size: overall 56-59 µm, shell 30-33 µm, processes 13 µm.
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Original description: Deflandre and Cookson 1955, p. 278-279: Hystrichosphaeridium colligerum
Diagnosis: Shell globular, almost spherical; processes numerous, tubular, straight or slightly flexuous, with a conical base and a closed capitate or slightly furcate apex. At one pole of the shell there is a large, hollow, cylindro-conical outgrowth closed by a rounded conical apex. This outgrowth, like the small processes, does not communicate with the interior of the shell. It is frequently ornamented distally with a circle of short, blunt, hollow, conical knobs. Membrane of shell finely granular.
Dimensions: holotype: diameter of shell 33 Ám, overall diameter 56 Ám, large process 20 by 13 Ám, small processes c. 13 Ám long. Range: shell 30-33 Ám, overall diameter: 56-59 Ám, appendages c. 13 Ám.
Emended description: Cookson, 1965, p. 86
Anterior portion of shell almost spherical, dorso-ventral, with numerous broadly- based, stiff or flexuous, usually unbranched tubular appendages of varying lengths and widths that narrow towards small bluntly- pointed, capitate or somewhat funnel-shaped apices. On one surface, taken as ventral, some of the appendages are arranged in two short longitudinal rows one on either side of the mid-line (Pl.9, fig.1,3); on the dorsal surface they are evenly distributed (Pl.9,fig.2,4); an archeopyle occupies a large part of the apex (Pl.9,fig.5). The appendages are hollow at and near their bases and, contrary to the statement of Deflandre and Cookson (1955), their cavities appear to communicate with that of the shell (Pl.9, fig.10). However, it is practically impossible to prove this point, as well as to trace the cavity along the whole length of the longer appendages owing to the narrowness of their distal portions and the relative thickness of their walls. The wall of the anterior portion of the shell, c. 1.5 Ám thick, is two-layered. The outer layer is densely granular in surface view the granules appearing as minute rods in optical section under oil immersion; similar rods are frequently evident on the walls of the appendages.
Posterior portion of the shell usually smaller and thinner-walled than the anterior portion, roughly bell-shaped with straight or convex sides and, in well preserved specimens, a centrally-placed, cone-like projection with a small circular opening (Pl.9,fig.12). The surface is devoid of appendages, apart from one or two circles of small, knob or spine-like thickenings in the vicinity of the antapical opening (Pl.9,fig.7,8,11). The wall, c. 1 Ám thick, is smooth in optical section, finely granular in surface view, the granules being arranged in more or less clearly defined longitudinal rows (Pl.9,fig.1).
Dimensions: Hypotypes: 1-overall length of shell 55 Ám, width of anterior part 31 Ám, width of posterior part c. 26 Ám; 2--overall length of shell 57 Ám, width of anterior part c. 33 Ám, width of posterior part 26 Ám.
Range: Overall length: 36-57 Ám; width, anterior part: 29-36 Ám; posterior part: 13-26 Ám; appendages: 5-21 Ám long.
Emended diagnosis: Davey and Williams, 1966, p. 96
Ovoidal to spherical central body with wall composed of thin endophragm and outer finely reticulate periphragm. Processes composed of periphragm, numerous, simple, hollow, open or closed distally. One single broad antapical process. Total number of processes exceeding 50.
Dimensions: Holotype: diameter of central body 33 Ám, overall diameter 56 Ám, length of antapical process 20 Ám, width of antapical process 13 Ám, length of small processes approx. 13 Ám. Range: diameter of central body 30-33 Ám, overall diameter 56-59 Ám, length of small processes approx. 13 Ám. Figured specimens diameter of central body 33 by 37.5 Ám, length of antapical process 16 Ám, width of antapical process 15 Ám, length of small processes 11-15 Ám. Range of English specimens: diameter of central body 29-41 Ám, length of antapical process 16-21 Ám, width of antapical process 8-15 Ám, length of small processes 8-15 Ám.
Description: An apical archaeopyle and the large antapical process enable easy orientation of specimens of this species. The antapical process is hollow, cylindro-conical, occasionally closed, and bears small tubules, 2-3 Ám long, towards the distal extremity. The tubules can take the form of simple conical protuberances or can have slightly bifurcate extremities. The smaller processes are commonly simple, occasionally united proximally, and may be open or closed distally. They are slender but do vary in width, and in specimens possessing open processes they taper to a distal neck before terminating with a sightly expanded opening. The distal margin may be finely serrate or entire. Both types of processes are slightly fibrous and do not communicate with the interior of the central body. There are commonly 4, regularly distributed, processes for each precingular plate area; 2-4 for the postcingular plate areas and a constant 2 in the cingular plate areas.
Revised description: Goodman and Witmer, 1985, p. 77-78
Subspheroidal bi-layered cyst with large, rectangular antapical process, and numerous slender nontabular to intratabular processes over remainder of body; endophragm and periphragm appressed except beneath processes. Antapical process open distally with an aperture of diameter less than that of the process and may or may not bear several small, typically open-tipped spinules near its distal margin; basal margin typically bears longitudinal striations. Other processes are considerably smaller and either taper continuously to a closed, capitate tip, or they may taper for the greater length of the process and terminate in a distally open and flared, or buccinate, tip. Both open and closed processes generally do not occur on single specimens. Smaller processes may be smooth or finely granular.
Archeopyle variable; four types involving different combinations and numbers of apical and/or precingular paraplates are known to date. Archeopyles may be either precingular, Type P (3''), or Type 2P (3''- 4''); apical, Type 4A (1'-4'); or combination: Type 4AP (1'-4', 3''). Types P and 4A are most common, while Types 4AP and 2P are relatively rare. Operculum is free for all four variations, and simple for 4A apical variation; it could not be determined whether the opercula for Type 2P and 4AP archeopyles were simple or compound. Archeopyle margin on specimens with Type 4A and 4AP archeopyles show sixth precingular notch rather than the usual parasulcal notch (Plate 6, figs. 1-2), and corresponding extension on operculum is formed by fourth apical rather than first apical paraplate. Paratabulation typically expressed only by principal and accessory archeopyle sutures; outer wall bears no other archeopyle features. A gonyaulacacean paratabulation can be inferred for rare specimens which show fusion of process bases and clustering of adjacent, apparently intratabular processes, as well as a polygonal outline to the base of the antapical process. These features, in conjunction with details of archeopyle (and operculum) morphology, indicate a formula of 1-2pr, 4', 6'', Xc, 6''', 1p, 1''''. Sulcal processes are present but difficult to identify on most specimens.
Endophragm up to 1 Ám thick, and appears externally smooth. Periphragm may be up to 0.5 Ám thick, and is externally smooth to finely granulose. Paracingulum typically expressed, or may be represented by equatorial alignment of processes. Parasulcus, except for paraplates as, typically not expressed as a distinct feature; rare specimens bear smaller-than-normal processes in mid-ventral region. On very rare specimens the position of these processes, in conjunction with other features (e.g., shape and antapical process), may be interpreted to suggest five or six sulcal processes in a standard gonyaulacacean arrangement.
Affinities:
Davey and Williams, 1966, p. 97
In the original diagnosis, the smaller processes are said to be closed. The figure by Deflandre & Cookson (1955, pl. 7, fig. 3) seems to indicate, however, that the processes are open, as in the examples from the London Clay.
Cookson & Eisenack (1961, pl. 2, fig. 9) figured H. colligerum with the processes definitely open. The species may well be confined to examples with open processes after re-examination of the holotype.
Cookson (1965, pl. 9, figs. 1-3) illustrates forms with distinctively larger antapical processes not unlike those of Litosphaeridium siphoniphorum (Cookson & Eisenack). Because of the larger size of this process she has incorrectly considered it to represent a second portion of the central body and has erected a new genus diagnosed as possessing bipartite cysts. The forms from the London Clay, the holotype of D. colligerum and the specimen illustrated by Cookson & Eisenack (1961, pl. 2, fig. 9) are probably more typical of the species and show without doubt that the " posterior portion of the shell " is in fact an enlarged antapical process.
This species has been recorded from the Upper Cretaceous (probably Senonian) and Eocene of Australia, and in England only from the Eocene.
This species is easily distinguishable from all other types of dinoflagellate cysts.
Goodman and Witmer 1985, p. 78-79
The distinctive morphology of Diphyes colligerum - which includes the large antapical process and the numerous other small processes, variable archaeopyle types, and two wall layers - readily distinguishes it from most other fossil cysts. Despite resemblances to species of Coronifera, Duosphaeridium, Florentinia, Silicisphaera, Dapsilidinium and Polysphaeridium in regard to gross morphological features, consistent separation should be relatively easy upon critical analysis off all morphologic characters. Convergence of certain Diphyes colligerum morphotypes (i.e. those with extreme fusion of processes) and specimens of some species of Hystrichokolpoma may make differentiation of these genera somewhat difficult; perhaps the customary generic boundaries between the two need to be reevaluated.
Diphyes colligerum is distinguished from D. appendiculare on the basis of size and process type. The latter is approximately two or three times as large as D. colligerum, and bears processes with relatively wider bases and which taper very rapidly to hairlike ends, corresponding to Cookson and Eisenack (1970, p. 149).
D. colligerum is distinguished from D. ficusoides by the shape of the antapical process which is bell-shaped to cylindrical on the former and fig-shaped with a biconvex margin on the latter.
Diphyes spinulum (Drugg) Stover and Evitt, D. latiusculum Matsuoka, and D. recurvatum May could prove to be synonymous with D. colligerum but formal recognition of their synonomy must await reexamination of the holotypes.