Paragloborotalia kugleri

Classification: pf_cenozoic -> Globigerinidae -> Paragloborotalia -> Paragloborotalia kugleri
Sister taxa: P. acrostoma, P. incognita, P. pseudocontinuosa, P. semivera, P. kugleri, P. pseudokugleri, P. mayeri, P. siakensis, P. birnageae, P. continuosa, P. opima, P. nana, P. griffinoides, P. sp.,


Citation: Paragloborotalia kugleri (Bolli, 1957)
Rank: species
Basionym: Globorotalia kugleri Bolli, 1957
Taxonomic discussion:

When Paragloborotalia kugleri was described by Bolli (1957), he included within the concept forms which were later separated as pseudokugleri by Blow (1969) (see also Postuma, 1971). Most authors since the 1990s have followed Blow, as we do here, although influential works by Stainforth and others (1975), Kennett and Srinivasan (1983), and Bolli and Saunders (1985) continued to recognize only kugleri. Spezzaferri (1991) and Rögl (1996) recognized spine holes, and Spezzaferri (1991; pl. 1, fig. 6) illustrated calcite build-ups (‘rises’) at the intersections of cancellate ridges that may represent gametogenic calcification over spine holes as observed in Trilobatus sacculifer (e.g., Hemleben and Olsson, 2006). However, Pearson and Wade (2009) considered that evidence questionable and failed to find evidence of spine holes or spines embedded in the wall in well-preserved populations from Trinidad. Hence they, and Aze and others (2011), regarded the species as probably nonspinose and referred it, and the closely related pseudokugleri, only questionably to the genus Paragloborotalia. Our ongoing investigations have produced unequivocal evidence of true spines in pseudokugleri (Plate 5.9, Figs. 4, 8), supporting the original evidence of Spezzaferri (1991) and Rögl (1996) and confirming the generic assignment. Like most species of Paragloborotalia, P. kugleri was subject to heavy gametogenetic calcification (Hemleben and others, 1989) making spines often difficult to detect. [Leckie et al. 2018]

Catalog entries: Globorotalia kugleri

Type images:

Distinguishing features:

Like P. pseudokugleri but the final chamber shows a distinct pinching; the chambers are less inflated and more appressed; the outline is more ovate and less lobulate; spiral side sutures are more strongly curved and less depressed.

NB These concise distinguishing features statements are used in the tables of daughter-taxa to act as quick summaries of the differences between e.g. species of one genus.
They are being edited as the site is developed and comments on them are especially welcome.


Diagnostic characters:

Paragloborotalia kugleri would appear to have evolved from P. pseudokugleri by gradual evolutionary transition (Blow, 1969, 1979; Chaproniere, 1981; Keller, 1981; Premoli Silva and Spezzaferri, 1990; Spezzaferri, 1991; Leckie and others, 1993; Pearson, 1995; Pearson and Chaisson, 1997). No detailed morphometric study has yet been attempted, but the overall features of the evolutionary trend are generally agreed by workers. Because it is a gradual transition, the distinction of the two taxa is an arbitrary divide, and the lowermost specimens of kugleri in a given section are usually rare and found among an intergrading population that is predominantly assigned to pseudokugleri. The frequency of kugleri versus pseudokugleri typically increases up-section, with the pseudokugleri morphospecies often persisting through much of the range of kugleri.

Because kugleri is an important zone fossil whose lowermost occurrence defines Zone M1 and approximates the Oligocene/Miocene boundary (Steininger and others, 1997), it is important to establish clear criteria for distinguishing the morphospecies. The method for doing this should involve careful consideration of the morphology of two holotype specimens (which are shown here in SEM for the first time on their respective plates) and trying to draw the line half way between them, rather than by trying to develop an idealized view of what a quintessential kugleri and pseudokugleri should look like. This point is stressed because the pseudokugleri holotype is quite an ‘advanced’ form for the morphospecies, and shows various features that tend to be more clearly associated with kugleri, especially the number of chambers in the final whorl and curved spiral sutures, as discussed further below. A mosaic of characters is involved when trying to draw a subjective line in the evolutionary transition between the morphospecies. It is a multidimensional problem that ultimately may benefit from a quantitative, morphometric approach using multivariate statistics. At present, however, we rely on our subjective appreciation of shape to accomplish this task.

Before proceeding, it is worth mentioning that some authors (Premoli Silva and Spezzaferri, 1990; Spezzaferri and Premoli Silva, 1991; Spezzaferri, 1994; Rögl, 1996) have recognized on their range charts a third morphotype in open nomenclature as “pseudokugleri - kugleri transition”. To operate that distinction, it would be necessary (using the same principles described above) to define arbitrary divisions between 1) pseudokugleri s.s. and “pseudokugleri - kugleri transition”; and 2) between “pseudokugleri - kugleri transition” and kugleri s.s. Moreover, the lowermost kugleri s.s. is likely to be drawn at somewhat higher level if transitional forms are entered in to a separate category. We have elected not to do this and assign all ‘transitional’ forms to one or the other morphospecies.

The principal morphological changes that occur from ‘primitive’ pseudokugleri to ‘advanced’ kugleri are as follows;

  1. An increase in the number of chambers in the final whorl from 5-6 to 7 and more rarely 8,
  2. A concomitant reduction in the rate of chamber enlargement, making successive chambers more equal in size,
  3. Reduced inflation and increasing compression and appression of the chambers,
  4. A concomitant reduction in the lobateness of the periphery, which ranges from ovate to more circular,
  5. A transition from straight to strongly curved sutures on the spiral side,
  6. An increase in the tendency for weakly curved sutures on the umbilical side,
  7. Less marked depression of the spiral side sutures,
  8. Flattening of the spiral side generally, although some ‘advanced’ forms are convex on both sides and biconvex - lensoidal in overall morphology,
  9. Increased pinching of the periphery of the final one or more chambers, which go from sub-rounded to sub-acute,
  10. A slight increase in average size, after an initially more rapid increase near the very beginning of the stratigraphic range (Spezzaferri, 1994).

As mentioned above, the holotype of pseudokugleri is a considerable way along this transition. It has 7 chambers in the final whorl, which expand fairly slowly in size; fairly inflated chambers and a distinctly lobulate periphery with, nevertheless moderate chamber appression; slightly curved sutures on the spiral side throughout ontogeny (see the SEMs of the holotype, Pl. 5.9, Figs. 1-3; these are seemingly exaggerated on Bolli’s original drawing; Blow (1969) on the other hand, originally described the sutures as straight); straight sutures on the umbilical side; moderately depressed spiral side sutures with a moderately flattened spiral side; and a fully rounded periphery. A rounded periphery is the key feature of pseudokugleri s.s. (e.g., Leckie and others, 1993). Pearson and Wade (2009) collected more specimens from the ‘cotype locality’ at Mosquito Creek, Trinidad, which show a similar set of characters. The holotype of kugleri shows 8 chambers of gradually increasing size in the final whorl, although the final chamber is unusually large; low chamber inflation and high appression making the periphery almost entire and non-lobulate; strongly curved spiral side sutures and fairly straight umbilical side sutures; a very flat spiral side and a somewhat pinched/subacute periphery to the final chamber. An asymmetrically subacute periphery is a key feature of kugleri s.s.

It is clear from this that the number of chambers in the final whorl and the presence of curved spiral sutures are of less importance in distinguishing the morphospecies than some of the other features. Accordingly, we suggest the following guidelines for distinguishing kugleri: most importantly, the final chamber should show a distinct pinching and not be broadly rounded (contra Rögl, 1996); the chambers are less inflated and more appressed and the outline is more ovate and less lobulate; and spiral side sutures are more strongly curved and less depressed than is typical in pseudokugleri; and the spiral side is generally flatter. A combination of these features allows for the most reliable means to differentiate the two taxa and recognize the base of Zone M1 near the Oligocene/Miocene boundary.

Finally, although the spiral side in kugleri is usually flat, it can also be convex, rendering the test markedly biconvex (e.g., Premoli Silva and Spezzaferri, 1990, pl. 3, fig. 5a-c; Spezzaferri, 1991, fig. 1a-d). These variants have sometimes been identified as Globorotalia (Turborotalia) mendacis Blow but the holotype of that species is regarded here as a synonym of P. birnageae. Paragloborotalia kugleri is distinguished from P. birnageae by its more asymmetrically subacute peripheral margin, more ovate equatorial outline, flatter spiral side, and higher aperture with smaller lip (P. kugleri lacks an appressed final chamber with flap-like flange or apertural plate). It is distinguished from Fohsella? peripheroronda by its generally greater number of chambers in the final whorl (typically 7 compared with 6) and slower rate of chamber expansion giving kugleri a more circular equatorial outline compared with a more ovate outline of peripheroronda. [Leckie et al. 2018]

Wall type: Normal perforate, coarsely cancellate, sparsely spinose in life, heavy gametogenetic calcification is often present.

Test morphology: Test small to medium in size; low trochospiral; 6 to typically 7, rarely 8 chambers in the final whorl, increasing slowly in size; equatorial outline slightly lobulate, circular to slightly ovate depending on the size of the final chamber which is often smaller than the penultimate chamber; in spiral view chambers arranged in 2½-3 whorls, sutures slightly depressed, strongly curved; in umbilical view chambers wedge-shaped, weakly inflated, sutures slightly depressed, radial or slightly curved, umbilicus narrow and moderately deep; aperture umbilical-extraumbilical, a low arch, frequently hooked, typically bordered by a lip; in edge view spiral side nearly flat to moderately convex, umbilical side more convex, periphery of final chamber subacute. [Leckie et al. 2018]

Size: Maximum diameter of holotype 0.30 mm (original measurement); 0.27 mm (remeasured this study); thickness 0.15 mm (this study). [Leckie et al. 2018]

Character matrix

test outline:Subcircularchamber arrangement:Trochospiraledge view:Equally biconvexaperture:Umbilical-extraumbilical
sp chamber shape:Globularcoiling axis:Lowperiphery:N/Aaperture border:Thin lip
umb chbr shape:Globularumbilicus:Narrowperiph margin shape:Narrowly roundedaccessory apertures:None
spiral sutures:Weakly depressedumb depth:Shallowwall texture:Cancellateshell porosity:Macroperforate: >2.5µm
umbilical or test sutures:Moderately depressedfinal-whorl chambers:6.0-8.0 N.B. These characters are used for advanced search. N/A - not applicable

Biogeography and Palaeobiology

Geographic distribution: According to Kennett and Srinivasan (1983) and Rögl (1996), this species is limited to the tropics and subtropics, but in fact it has been found nearly as far north as the Reykjanes Ridge and east Greenland margin in the North Atlantic Ocean (Poore, 1979; Spezzaferri, 1998). [Leckie et al. 2018]

Isotope paleobiology: Paragloborotalia kugleri has low δ18O and high δ13C relative to other species suggesting it was a mixed-layer dweller (Douglas and Savin, 1973; Biolzi, 1983; van Eijden and Ganssen, 1995; Pearson and others, 1997; Spezzaferri and Pearson, 2009). [Leckie et al. 2018]

Phylogenetic relations: Evolved by gradual transition from P. pseudokugleri (Blow, 1969, and subsequent authors; see discussion above). Many have proposed that kugleri is the direct ancestor of F.? peripheroronda and the Fohsella lineage (e.g., Fleisher, 1974, Stainforth and others, 1975; Keller, 1981; Kennett and Srinivasan, 1983; Cifelli and Scott, 1986; Chaisson and Leckie, 1993). An alternate hypothesis is that kugleri became extinct with no descendants. Olsson (1972) proposed that mayeri was the ancestor of peripheroronda, while Jenkins (1960, 1971) and Blow (1969) suggested that mayeri was the descendant of peripheroronda. [Leckie et al. 2018]

Most likely ancestor: Paragloborotalia pseudokugleri - at confidence level 4 (out of 5). Data source: Leckie et al. 2018.
Likely descendants: Fohsella peripheroronda;

Biostratigraphic distribution

Geological Range:
Notes: Base of Zone M1, by definition, to top of Zone M1, also by definition (Berggren and others, 1995; Wade and others, 2011; see discussion in Chapter 2, this volume). The base of Zone M1 occurs two meters above the ‘golden spike’ for the Neogene Period, Miocene Epoch, and Aquitanian Stage, at Lemme-Carrosio, Italy (Steininger and others, 1997) hence it is used to approximate these levels in planktonic foraminiferal biostratigraphy. [Leckie et al. 2018]
Last occurrence (top): at top of M1 zone (100% up, 21.1Ma, in Aquitanian stage). Data source: Wade et al. 2011; Leckie et al. 2018
First occurrence (base): at base of M1 zone (0% up, 23Ma, in Aquitanian stage). Data source: Wade et al. 2011; Leckie et al. 2018

Plot of occurrence data:

Primary source for this page: Leckie et al. 2018 - Olig Atlas chap.5 p.141


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Paragloborotalia kugleri compiled by the pforams@mikrotax project team viewed: 17-11-2019

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