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Reproduced from the
Vol. 20, No. 6, November, 1946

With the kind permission of SEPM Society for Sedimentary Geology, February 2003


International Ecuadorean Petroleum Co., Guayaquil, Ecuador

Abstract—Three new species of arenaceous Foraminifera are described from Tertiary formations of Ecuador: Technitella archaeonitida, Planulina wheeleri, and Palmerinella thalmanni.

The wealth and variety of the foraminiferal faunas of the Ecuador Tertiaries are not adequately indicated in the published literature. Pending the opportunity to compile more comprehensive papers, we present here descriptions of three outstandingly interesting forms encountered in the course of zonational studies. This paper is published with the kind permission of the directorate of International Ecuadorean Petroleum Company. We gratefully express acknowledgment to Dr. J. A. Cushman for technical advice and assistance with the illustrations, to Dr. H. E. Thalmann for comments on the specimens, to Dr. Brooks F. Ellis for supplying photostat copies of obscure references, and to Mr. K. W. Barr for transcribing some early references to Technitella. Reliance has been placed on the “Catalogue of Foraminifera” by Ellis and Messina (1940) for all references to original type descriptions.

Family Saccamminidae

Subfamily Pelosininae

Genus Technitella Norman 1878

Type‑species: T. legumen Norman 1878, earliest species, first citation by Cushman 1910.

The arenaceous foraminiferal genus Technitella is rarely mentioned in current literature as almost without exception the known species have been described from Recent material. A few decades ago, when microscopy was the hobby of men of leisure and the results of oceanographic cruises such as those of the “Challenger” were widely publicized, the genus received considerable notice because of the outstanding elegance of the tests made by its different species. Later Heron-Allen emphasized the remarkable selective powers and constructive skill shown by the genus as evidence of some form of intelligence in the Protozoa. Typical of his attitude is the following extract from a paper (1915) which excited lively discussion:

The intelligence which, I am bold enough to claim, is displayed by the arenaceous Foraminifera in their house‑building takes two forms: first the exclusive selection of certain materials, and second, the manner in which they were used. The exclusive selection of Echinoderm plates by Technitella thompsoni Heron‑Allen and Earland, for the construction of its test, is a most remarkable illustration of this selective power, for in neither of the dredgings in which it has been found do echinoderm plates such as are used in its construction abound… The genus Technitella, as the name “Little Workman” implies. supplies us with one of the most remarkable illustrations of purpose and intelligence hitherto encountered in the Foraminifera. The most familiar species of the genus, Technitella legumen Norman, constructing its test only of spongespicules selected from the mass of mud, sand, molluscan and foraminiferal debris in which it lives, builds its shell wall in two layers, the outer one set parallel to the long axis of the shell, the inner one at right‑angles to it. As we observed when we first discovered this feature, “we thus get as close an approximation to woof and warp as is possible in a rigid non‑flexible material, and it is obvious that the strength of the test must be enormously increased by the crossing of the two layers…” In the papers to which I have referred … it is shown that there is as wide a range of skill displayed by the Foraminifera both in choice of material and in actual construction as by builders in the higher scales of life, not even excepting man.

There seem to be only two references to Technitella in the fossil state, both from Mesozoic rocks in Europe. In the earlier of these Rychlicki (1912) records, without comment or figure, “Ein Stück von Technitella raphanus Brad.” from the Upper Senonian of Leszcyzny in the Carpathians. The second reference is by Pfender (1938) to an organism studied, in sections of Valanginian limestone from Provence. This is a dubious record, not confirmed by the single figure published. Pfender’s own opinion is expressed thus:

“(Cette éspèce) … est encore plus malaise a definir car il ne semble presenter que sous forme de debris: des filaments, de 15µ d’epaisseur généralement étroitment juxtaposées en petits paquets épars. Cependant il ne s’agit certainement pas d’une Algue; je comparerais de preférence ces restes à des debris du test d’un Foraminifère tel que Technitella legumen Norman 1878…”

The type description of the genus is:

Test elliptical, cylindrical or subfusiform, composed of broken fragments of sponge spicula arranged parallel to the axis and enclosed entirely, or rarely only partially, in the body wall. Unattached below and closed. A tubular mouth-opening formed by contraction for a short distance of the body walls so as to form a short tube.

Later authors have treated some of these features as of merely specific rank. As generic characters, emphasis has been laid on a high degree of selectivity in the material of the test and an unusual precision of construction, Most, but not all, of the described species are constructed of sponge spicules.

Technitella Archaeonitida

Stainforth and Stevenson, n. sp. Plate 86, figures 1-3

This species is ellipsoidal and in its usual flattened state of preservation appears from 1 to 3 times as long as broad. The holotype measures 1.25 x 0.69 mm., associated specimens range in length from 0.56 to 1.50 mm. and in breadth from 0.22 to 0.77 mm. The test is made of closely spaced sponge spicules set parallel to the long axis in a minimum of non‑calcareous cement. The diameter of the spicules chosen varies from one specimen to another, but it is usually uniform in any one specimen. Tests made of very fine spicules predominate. The small aperture is terminal, a simple circular opening formed by the ends of spicules neatly arranged to lie flush with one another. The aboral end is equally neatly made, either with long spicules forming a point or with a few fragmentary ones fitted into the smooth contour of the test. Examination of thin sections and broken specimens failed to reveal any layer of transverse spicules such as was shown by Heron‑Allen and Earland to be typical of T. legumen Norman.

 Affinities and differences.—The species here described differs from T. asciformis Pearcey (1914) and T. legumen Norman (19—) in its lack of an apertural neck; from T. hystrix Chapman and Parr (1937) and T. melo Norman (19—) in its smooth contour compared with their echinate forms; from T. mestayeri Cushman (1919) and T. raphanus Brady (1884) in being ellipsoidal in contrast to their elongate, almost tubiform shape; from T. bradyi Earland (1934) in which the sponge spicules appear to be used almost adventitiously, in its higher degree of selectivity and neatness of construction; from T. thompsoni Heron-Allen and Earland (19—) in its construction of sponge spicules, not echinoderm ossicles, and likewise from the three species made of fine sand viz. T. candida, T. flexibilis and T. globulus Wiesner (23).

Of the described species this leaves only T. nitida Heron-Allen and Earland (1932), which shows little significant morphological difference from the Ecuador species. The holotype of T. nitida measured 1.5 x 0.8 mm., which compares closely with the larger specimens of T. archaeonitida. The type description of T. nitida specifies the presence only of longitudinally placed spicules and this is also true of the new species. T. nitida, however, is described as having “a large simple aperture surrounded by a slightly thickened and everted neck,” and it definitely differs in this respect from T. archaeonitida, which has no neck. Another reason for considering the two species to be distinct lies in their relative ages. Though the Ecuadoran species is quite common in beds of Lower Oligocene age, no signs of it have been found in ecologically similar beds younger than Middle Oligocene, whereas T. nitida is only known as a recent form.

Occurrence.—In the Playa Rica formation as described by Olsson (1942), which includes Upper Eocene to early Middle Oligocene beds. It is most frequent in the younger part of the formation. The holotype was collected from sample I.E.P.C. No. 17117 at the confluence of the Estero Tumbavero tributary with the main stream of the Rio Zapallo Grande, some 3 km. E.S.E. from the village of Telembi on Rio Cayapas in northwest Ecuador. The geographic coordinates of the type locality are 00° 47¢ 43˝ North: 78° 54¢ 21˝ West.

Type material.—The holotype is deposited at the Cushman Laboratory for Foraminiferal Research, in Slide No. 45587. In the same collection Slide No. 45586 contains the other figured specimens and Slide No. 45588 contains topotype material. Other topotypes have been forwarded to the following institutions: American Museum of Natural History; Colorado School of Mines; Walker Museum, University of Chicago; Stanford University; British Museum of Natural History.

Family Anomalinidae

Subfamily Anomalininae

Genus Planulina d’Orbigny 1826

Genotype P. ariminensis d’Orbigny, 1826.

This is the usually accepted classification, taken from Cushman’s textbook (1940). Galloway (1933) refers the genus to the Family Rotaliidae Reuss 1860; Subfamily Cibicidinae Galloway 1933; genus and genotype as above. More recently Brötzen (1942) has offered an interesting re-classification of the rotaliiform Foraminifera, in which Planulina is referred to the Family-group Rotaliiformes; Family Valvulineriidae; Subfamily Cibicidinae; genus and genotype as before.

Planulina Wheeleri

Stainforth and Stevenson, n. sp. Plate 86, figures 4-6

Description.—Shell vitreous, very finely perforate. General shape compressed, almost discoidal. Initial portion trochoid, later becoming evolute and almost planispiral. Chambers 13 to 20 in the final whorl, sutures recurved, chambers slightly inflated along the sutures. On the dorsal side the small circular nucleoconch is faintly discernible, tightly coiled, though uncoiling is so rapid that in most adult specimens the inner edge of the final whorl barely overlaps the previous coil. On the ventral side only the chambers of the final whorl are visible, surrounding a small circular umbilical pit which tends to be accentuated by a low rim of clear shell material. The outline is undulose and not a precise spiral, its irregularities coinciding with variations in chamber‑breadth. The diameter of the holotype is 1.15 mm. and its median thickness 0.28 mm.

The aperture appears to be a very fine crescent‑shaped slit at the base of the apertural face, possibly running back along the spiral suture under the last three or four chambers. Despite having examined about two hundred specimens we have not found one in which the aperture is clearly visible. The above description is based on the composite impression gained by the examination of many specimens, and is supported by thin‑section studies. Some of the broken specimens show what appears to be an apertural pore at the outer tip of the apertural face, but unbroken specimens do not show this feature, nor is it apparent in any of the thin‑sections.

Comments.—On first inspection we were inclined to believe that this species belonged to a new genus of the Anomalinidae, but on detailed examination it could not be separated with certainty from Planulina. It is a peculiar species, possibly modified for life in a littoral facies.

This species seems most properly referred to Planulina on the principal morphological features of the test especially the form of coiling and the aperture. We have failed, however, to find any described species of Planulina with so finely perforate a test, so many chambers, or such an obscure aperture.

The species is named for O. C. Wheeler, director of International Petroleum Company, Toronto.

Occurrence.—The holotype is from calcareous sandstones outcropping in bluffs immediately south of the village of San Pedro, about 40 Km. (not miles as stated by Barker) northeast of Santa Elena point in southwest Ecuador. Geographic coordinates of the type locality are 01° 57¢ 03˝ South: 79° 53¢ 37˝ West This locality has been mentioned in the literature by Barker (1932) as the provenance of Miogypsina aff. panamensis Cushman, later re‑described as Miogypsina (Miolepidocyclina) ecuadoriensis Tan (1936). Barker gave the age of the deposits as Aquitanian‑Burdigalian. (Above references fide Thalmann (1945). Currently the age of the San Pedro sandstones is considered to be Upper Oligocene.

The species is common in fossiliferous parts of the sandstones at the type locality. It was also found commonly in a lenticle of foraminiferal sand‑cored in a well near Bajada in southwestern Ecuador—age of this occurrence low in the Upper Oligocene. It is noteworthy that the species was completely absent from the shales in which this lenticle of sand was included, and the inference may be drawn that the species preferred a littoral environment.

Type material.—The holotype of the species has been deposited in the Cushman Laboratory for Foraminiferal Research in Slide No. 45583. Topotypes have been deposited in the same collection in Slide No. 45582 and also in the following institutions: American Museum of Natural History; Colorado School of Mines; Walker Museum, University of Chicago; Stanford University; British Museum of Natural History. Also specimens from the Bajada well have been deposited in the Cushman collection in Slides Nos. 45593, 45594.

Family Anomalinidae

Subfamily Anomalininae

Genus Palmerinella Bermúdez 1934

Genotype P. palmerae Bermúdez 1934

The genus Palmerinella rests on a single species described from Recent beach sand in Cuba. On the evidence of the Ecuadorean species described below we are able to extend the geological range of the genus back into the basal Miocene, and to amend the original description.

Description of genotype.—(translation):

Shell in the form of a very low spiral, initial chambers visible from the dorsal side and also from the ventral side through a thin umbonal deposit. Wall vitreous, calcareous, somewhat coarsely perforate. Elongate aperture in the face of the last chamber, outlined, except at the base, by a distinct collar.

This new genus is a modification of the evolute genus Anomalina, approaching in its manner of growth to the genus Planulina, but slightly trochoid. The characteristic which distinguishes it is the oval elongate aperture in the septal face, surrounded, except at the base, by a distinct collar.

On the evidence of the species described below two modifications should be made in the definition of the genus. First, the wall may be finely perforate; Bermúdez’ descriptions were slightly contradictory since he described the genotype as “algo toscamente perforada” but the type species as “finamente pero conspicuamente perforada.” Second, the initial chambers are not always visible from the ventral side.

Palmerinella Thalmanni

Stainforth and Stevenson, n. sp. Plate 86, figures 7‑10

Description.—Small, vitreous to transparent, finely but plainly perforate. Composed of few chambers, usually 8 fully developed in the final whorl. All chambers visible on the dorsal side, only those of the final whorl on the ventral side. Two to 2 1/2 whorls on the dorsal side. Sutures limbate and transparent, usually strongly raised between the early chambers, tending to become depressed between the final chambers. Strongly raised and slightly curved early sutures coalesce into the umbo and peripheral edge. Later sutures almost straight. Sutures of successive whorls tend to be in alignment. Periphery ranges from a square to a subacute edge. Aperture a narrow, parallel‑sided to slightly tapering slit extending from inner to outer margin in center of apertural face. In younger specimens aperture is surrounded, except at the base, by a distinct collar, but in more mature specimens this feature tends to disappear. Entire apertural face rimmed by a raised extension of the peripheral edge. Umbonal plug usually strongly developed on dorsal side but absent or weakly developed on ventral side. Coiling is normally planispiral, though larger specimens tend to be trochoid. Coiling is normally sinistral on the dorsal side and dextral coiling is unusual. Diameter 0.20 to 0.30 mm., thickness 0.07 to 0.15 mm.

Variation.—Two hundred specimens chosen for this study exhibited the following variations: the umbonal plug may be strongly or weakly developed on both dorsal and ventral sides, or on the dorsal side alone, and it may be either opaque or translucent; raised limbate sutures may be strongly or weakly developed on both dorsal and ventral sides, on the dorsal side only, or the sutures may all be depressed; in some large specimens the last three or four chambers may be lobate at the periphery. Probably an environmental effect is the occasional addition of adventitious shell material in the. form of secondary peripheral flanges, umbilical nodes and enlarged sutures.

Specimens of P. palmerae Bermúdez from the Recent and Upper Miocene of Cuba and the Pliocene and Upper Miocene of the Dominican Republic, kindly sent to us by P. J. Bermúdez, show a tendency to vary in similar respects.

Affinities and differences.—This new species is undoubtedly closely related to the only other described species, P. palmerae Bermúdez, as shown by their similarity of general form and in particular their peculiar aperture. In specific features P. thalmanni n. sp. differs from P. palmerae in that the chambers of the former are fewer, shorter, straighter and broader; the dorsal side is less evolute; the sutures are usually more raised; and a more prominent umbonal plug and peripheral flange are present.

This new species is named for Dr. Hans E. Thalmann, with whom the writers had the pleasure of working when he was chief paleontologist of the International Ecuadorean Petroleum Company.

Occurrence.—Basal Miocene, Province of Manabí, Ecuador, from shale outcrops in gullied hillsides bordering the town of Charapotó on its north side. This town is located near the Pacific sea‑coast in central Ecuador at 00° 50¢ 14.7˝ South latitude and 80° 29¢ 31.2˝ West longitude.

P. thalmanni is abundant throughout a shale formation overlying the “Manta shale” (late Middle Oligocene) the fauna of which was described by Galloway and Morrey (1929).

The genus Palmerinella was first recorded from Recent beach sand in Cuba. In a personal communication P. J. Bermúdez informed us that the genotype also occurs in the Upper Miocene of Cuba and in the Upper Miocene and Pliocene of the Dominican Republic, in shallow water sediments. The new species here recorded from Ecuador extends the range of the genus downward into the base of the Miocene.

Type material.—Co‑types have been deposited in the Cushman Laboratory for Foraminiferal Research, in Slide No. 45584 and in the same collection Slide No. 45585 contains the figured specimens. Topotypes have been sent to the following institutions: American Museum of Natural History; Colorado School of Mines; Walker Museum, University of Chicago; Stanford University, British Museum of Natural History.


Barker, R. W., 1932, Three species of larger foraminifera from S. W. Ecuador. Geol. Mag., vol. 59, no. 816, pp. 277‑281. London.

Bermúdez, P. J., 1934, Un género y especie nueva de foraminiferos vivientes de Cuba. Mem. Soc. Cubana Hist. Nat., vol. 8, pp. 83-86, Havana.

Brady, H. B., 1884, Report on the foraminifera dredged by H. M. S. Challenger during the years 1873‑1876, Rept. Challenger Expdn., Zool. pt. 22, vol. 9, p. 247, London.

Brötzen, F., 1942, Die Foraminiferen‑gattung Gavelinella nov. gen. und die Systematik der Rotaliiformes, Stockholm.

Chapman, F. and Pake, W. J., 1937, Foraminifera, Australasian Antarctic Expdn. 1911‑1914, Sci. Repts., Sydney.

Cushman, J. A., 1910, A monograph of the foraminifera of the North Pacific Ocean: Part I U. S. Nat. Mus. Bull, no. 71, pt. 1, P. 48, Washington.

—, 1919, Recent foraminifera from off New Zealand, U. S. Nat. Mus. Proc. no. 2302, vol. 56 (1920), Washington.

—, 1940, Foraminifera: their classification and economic use (Third edition) Cambridge.

Earland, A., 1934, Foraminifera; Part III—The Falkland Islands sector of the Antarctic (excluding South Georgia), “Discovery” Reports, vol. 10 (1935), p. 65, Cambridge University Press, England.

Ellis, B. F., and Messina, A. R., 1940 et seq., Catalogue of Foraminifera, Amer. Mus. Nat. Hist., New York.

Galloway, J. J., 1933, A manual of Foraminifera, Bloomington, Ind.

— and Morrey, M., 1929, A Lower Tertiary foraminiferal fauna from Manta, Ecuador, Bull. Amer. Pal., vol. 15, no. 55, pp. 1‑56, Ithaca, N. Y.

Heron‑Allen, E., 1915, A short statement upon the theory and the phenomena of purpose and intelligence exhibited by the Protozoa, as illustrated by selection and behaviour in the foraminifera. Roy. Micr. Soc., Jour. Vol. (1915), pp. 547-557, London.

— and Earland, A., — On a new species of Technitella from the North Sea, with some observations upon selective power as exercised by certain species of arenaceous foraminifera, Queckett Micr. Club, Jour. ser. 2, vol. 10, p. 403.

—, 1932, Foraminifera: Part I‑The ice‑free area of the Falkland Islands and adjacent areas, “Discovery” Reports, vol. 4, p. 328, Cambridge University Press.

Norman, A. M., 1878, On the genus Haliphysema, with a description of several forms apparently allied to it, Ann. Mag. Nat. Hist., ser. 5, vol. 1, pp. 279‑281.

Olsson, A. A., 1942, Tertiary deposits of northwestern South America and Panama, 8th Amer. Sci. Congress, Proc., Geol. Sci., vol. 4, Washington.

Pearcey, F. G., 1914, Foraminifera of the Scottish National Antarctic Expedition, Roy. Soc. Edinburgh, Trans. Vol. 49, pt. 4, no. 19, p. 1002, Edinburgh.

Pfender, J., 1938, Les foraminifères du Valanginien provençal, Soc. Géol. de France, Bull. sér. 5, tome 8, fasc. 3‑4, p. 231, Paris.

Rhychlicki, J., 1912, Die Foraminiferenfauna der karpatischen untersenonen Mergel von Leszcyzny, Acad. Sci. Cracovie, Bull. Inter. ser. A, no. 7a, p. 756, Cracovie, Poland.

Tan Sin Hok, 1936, Zur Kenntnis der Miogypsiniden De Ingenieur en Nederl. Indie, vol. 3, pt. 4, pp. 45-61, Bandoeng, Java.

Thalmann, H. E., 1945, Resumen de las investigaciones micropaleontológicas en el Ecuador, Ecuador Petrolero, vol. 1, no. 1, p. 22, Quito, Ecuador.

Wiesner, H., 1931, Die Foraminiferen der deutsch Südpolar‑Expedition 1901‑1903. In: Drygalsni, E. Von,—Deutsche Südpolar‑Expedition 1901‑1903, Bd. 20 (Zool. Bd. 12),  pp. 53‑165, Berlin and Leipzig.

Explanation of Plate 86

Figs. 1-3Technitella archaeonitida Stainforth and Stevenson n. sp. 1, Side view of holotype, x 30. Cushman Collection 45587. 2, Apertural view of topotype, x 40. Cushman Collection 45586. 3, Side view of another topotype showing construction of unusually thick sponge spicules, x 30. Cushman Collection 45586. All three specimens from the Lower Oligocene Playa Rica formation, northwest Ecuador.

       4-6 Planulina wheeleri Stainforth and Stevenson n. sp. Holotype, x 30. Cushman Collection 45583. 4, Ventral aspect, 5, Peripheral aspect, 6, Dorsal aspect. From Upper Oligocene San Pedro sandstone, southwest Ecuador.

        7-10 Palmerinella thalmanni Stainforth and Stevenson n. sp. Co‑types, Cushman Collection 45585. 7, Dorsal aspect of a small specimen, x 75; 8, Dorsal aspect, x 65. 9, Peripheral aspect, x 75, and 10, Ventral aspect, x 75, of a larger specimen. From Basal Miocene shale near Charapotó, west central Ecuador.