PALYNOLOGICAL CONTRIBUTIONS TO THE
CHRONOZOGY AND STRATIGRAPHY OF THE
HARTFORD BASIN IN
CONNECTICUT AND MASSACHUSETTS
BRUCE CORNET
ALFRED TRAVERSE
Department of Geosciences
The Pennsylvania State University
University Park, Pennsylvania 16802
ABSTRACT
Recent discoveries of palynoflorules at numerous localities in the Newark Group basins of the eastern
United States provide new evidence for correlation of deposits in these basins. Floras from the Shuttle
Meadow and Portland Formations of the Newark Group rocks in the Hartford Basin of Connecticut and
Massachusetts indicate that the Triassic-Jurassic boundary is located within the rocks of this basin: a
shift of Corollina from more than 90% C. meyeriana to more than 90% C. torosus occurs somewhere
between the Shuttle Meadow and Portland Formations and generally indicates a Rhaeto-Liassic age. In the
Shuttle Meadow Formation, the overall shape of the palynoflora (particularly the presence of
Convolutispora klukiforma), associated fish, paleomagnetic data, and radiometric dates, as well as
megafossil evidence of Brachyphyllum scotii and Clathropteris meniscoides, support a basal Liassic age for
this formation. Reptilean evidence in older strata of the Hartford Basin suggests that the Triassic-Jurassic
boundary may lie just below the Shuttle Meadow Formation. In addition the Portland Formation contains a
palynomorph association clearly favoring early to middle Liassic age. Comparisons are made on the basis
of studies of recently discovered Newark Group palynoflorules of early Liassic age from Virginia and of
Carnian-Norian age from New Jersey. Paleobotanical and geological evidence is summarized, showing that
the predominantly Corollina (Hirmerella) palynofloras of the Hartford Basin were associated with a
warm, seasonally wet and dry climate, and casting doubt on the idea that Corollina-producing plants
composed a swamp association within the original basin. Twenty-seven genera and 42 species are
described, as are 1 spore and 1 pollen type of uncertain taxonomic position. Eight new species are
described: Camerosporites reductiverrucatus n. sp., Corollina murphyi n. sp., Cycadopites andrewsii n.
sp., Cycadopites durhamensis n. sp., Cycadopites westfieldicus n. sp., Dictyophyllidites paramuensteri n.
sp., Foveosporites agawamensis n. sp., Verrucosisporites cheneyi n. sp.
Six new combinations are proposed: Araucariacites punctatus (Nilsson) comb. nov., Callialasporites
segmentatus (Balme) comb. nov., Corollina itunensis (Pocock) comb. nov., Corollina simplex (DanzéCorsin & Laveine) comb. nov., Cycadopites reticulatus (Nilsson) comb. nov., Granulatisporites infirmus
(Balme) comb. nov. The genera Circullina and Corollina are formally emended, as are the species
Corollina torosus (Reissinger) Klaus and Circulina simplex Malyavkina.
INTRODUCTION
Recent discoveries of palynomorphs in most of the major Newark Group basins of the eastern United
States (Text-fig. 1) provide new evidence for correlating Newark Group strata with European TriassicJurassic type sections. The Hartford Basin
TEXT-FIGURE 1. "Newark" basins of eastern North America. Asterisks indicate basins in which
palynoflorules have been found.
palynofloras of Connecticut and Massachusetts (Text-fig. 2) provide at present the most comprehensive
data for upper Newark Group correlation, and form an assemblage different from Triassic
assemblages of other Newark Group basins. The Hartford Basin assemblage is largely Early Jurassic
in age; it is not restricted to this basin, but is also present in the Newark Basin of New Jersey and
the Culpeper Basin of Virginia. The recognition of Jurassic deposits in the Newark Group (Cornet et al.,
1973), which until recently has been considered entirely Triassic, is not new. As early as 1855, E.
Hitchcock, Jr., presented paleobotanical and geological evidence for a possible Rhaeto-Liassic age range
for the Hartford Basin sediments. Several other geologists and paleontologists of the mid-1800s speculated
that the deposits of the Hartford Basin extend into the Jurassic, but no convincing evidence was then
available. Newberry (1888) described a small megafossil flora from the Connecticut Valley but considered
it Rhaetian. Since then reptilean and fish remains have been interpreted as indicating only a Late Triassic
age (Reeside et al., 1957). Recently Reeve and Helsley (1972), in attempting to explain the paleomagnetic
discrepancy between the upper portion of the Chinle Formation, New Mexico, and the igneous rocks of the
upper Newark Group, suggested a post-middle Rhaetian, but pre-Pliensbachian age for much of the upper
portion of the Newark Group. The palynofloras from the Shuttle Meadow and Portland Formations of the
Hartford Basin largely confirm this suggestion, indicating that the Triassic-Jurassic boundary exists within
the strata of this basin.
TEXT-FIGURE 2. Hartford and Deerfield basins of Connecticut and Massachusetts. Numbers 1-13
indicate palynoflorule localities discussed in this paper. Middletown, Conn., is at Locality 13. North
Guilford is just south of Locality 1.
HARTFORD BASIN STRATIGRAPHY
The Hartford Basin sediments and igneous rocks are divided into four sedimentary and three volcanic
formations or units. The lowest formation, the New Haven Arkose, has been computed to have a
maximum stratigraphic thickness of 2600 m near the eastern edge of the basin, decreasing to around
1500 m near the western edge (Krynine, 1950) (Text-fig.3).
The Talcott Formation conformably overlies the New Haven Arkose in southern Connecticut, but
TEXT-FIGURE 3. Stratigraphic units of the Hartford Basin, showing their relative position and the
stratigraphic location of palynoflorule localities 1-13 (cf. Text-fig. 2). This is a restored composite section
from multiple sources, representing generalized North (left)-South (right) trends. The double line to the left
of Gaillard Graben indicates a fault, but because of restoration, no displacement is shown along the fault.
Locality 2 has the same position as Locality 1 and has therefore been omitted.
apparently it is missing in northern Connecticut and Massachusetts. In most of southern Connecticut,
the Talcott is a single basalt flow of variable thickness (30-75 m). Sanders (1970) has shown that within the
Gaillard Graben, which is restricted to the southeastern part of the basin, the Talcott Formation exists as
four lava-flow units interbedded with three sedimentary units, totaling about 305 m. The Gaillard Graben
has been interpreted as a post-depositional structure (Sanders, et al., 1963, p. 11). However, the existence
outside the Graben of only one (the lowest?) lava flow suggests that penecontemporaneous erosion
removed much of the once continuous Talcott Formation before deposition of the
overlying Shuttle Meadow Formation, and the Talcott was preserved in the graben as a result of downward
displacement of sediments in the graben before the erosional cycle began. On the other hand, the absence in
northern Connecticut and Massachusetts of basalt units referable to the Talcott might indicate only that the
basalts did not extend that far north. However, it seems reasonable that 300 m of superimposed volcanic
cones in Mt. Hitchcock in northern Massachusetts (Bain, 1941) may be related to the lava flows of the
Gaillard Graben, judging from stratigraphic position and thickness.
The Shuttle Meadow Formation, like the Talcott Formation, varies considerably in thickness and in
extent over the basin. In the southernmost part of the basin, within the Gaillard Graben, the Shuttle
Meadow Formation reaches its maximum thickness of about 270 m (Sanders, 1970). The formation steadily
decreases in thickness northward: 170 m in North Guilford, Conn.; 94 m near Middletown, Conn.; 94 m at
type section, New Britain, Conn.; 30 m in Farmington, Conn.; 15 m near Simsbury, Conn. (Krynine, 1950;
Lehmann, 1959; Schnabel, 1960). The upper part of the formation appears to be the most extensive. North
of Simsbury the Talcott basalt disappears, and the Shuttle Meadow Formation, if it continues north,
would be recognized as the uppermost part of the New Haven Arkose in northern Connecticut and
Massachusetts.
The Holyoke lava-flow unit conformably overlies the Shuttle Meadow Formation in Connecticut, and
the New Haven Arkose and Hitchcock volcanics in Massachusetts. This unit, consisting of at least two
lava flows, ranges in thickness from 120 to 170 m over most of the basin, but it has been recorded
under 92 m near the eastern and northern edges of the basin (Bain, 1941).
The East Berlin Formation conformably overlies the Holyoke lava-flow unit, and appears to thicken
both northward and southward from its minimum thickness near Avon, Conn.: 326-458 m in the Gaillard
Graben, Branford, Conn.; 170-180 m near Middletown, Conn.; 140-150 m near Avon, Conn.; 250 m near
Westfield, Mass. (Krynine, 1950; Lehmann, 1959; Schnabel, 1960; Colton and Hartshorn, 1966; Sanders,
1970). In the northern part of the basin, the upper part of the East Berlin Formation interfingers with the
Granby Tuff and local lava flows (Brophy et al., 1967). These volcanics and pyroclastics contain at least
one flow that apparently is continuous with the Hampden lava-flow unit (Bain, 1941).
The Hampden lava-flow unit conformably overlies the East Berlin Formation and ranges in thickness from
18 to 60 m. Much of this variation in thickness seems to be the result of differences in the number of
internal units or flows, which varies from two to eight (Chapman, 1965). In southeastern Connecticut a 12
m sedimentary unit has been found separating two flows (Sanders, 1970). The Portland Formation, which
conformably overlies the Hampden lava-flow unit and interfingers with some of the Granby Tuff (Bain,
1941), has a maximum thickness of more than 2000 m in Massachusetts. The "Chicopee shale" and "Longmeadow sandstone" are here considered as local members, at present recognized only in Massachusetts. In
Connecticut, where reported thickness rarely exceeds 920 m, post-depositional faulting and erosion are
responsible for removing much of this formation.
DESCRIPTIONS OF PALYNOFLORULE LOCALITIES
Of the 13 palyniferous localities thus far discovered in the Hartford Basin (Text-fig. 2, 3), 4 have been
selected as palynoflorule localities for purposes of formal description in this paper. Samples from the
9 remaining localities have yielded the same species of palynomorphs that are found at the formally
described localities.
Palynoflorule Locality No. 8
Formation: Upper New Haven Arkose.
Stratigraphic position: 116 ± 6 m below Holyoke lava-flow unit.
Geographic location: Massachusetts; Hampden County, 2.5 km south of Mt. Tom, just west of Holyoke;
roadcut on Southampton Road, 2.65 km east of junction with Country Road.
Palyniferous horizon: Tan to dark gray siltstone, .3-m-thick fossil plant bed situated within a 4.7+-m-thick
sequence of orange to tan coarse sand stone and conglomerate.
Remarks: Plant bed has produced dozens of well-preserved Clathropteris meniscoides Brongniart
fronds and Equisetites stems.
Palynoflorule Locality No. 1
Formation: Shuttle Meadow Formation.
Stratigraphic position: 100 ± 6 m below Holyoke lavaflow unit.
Geographic location: Connecticut; New Haven County, North Guilford; stream outcrop on east side of
Totoket Mountain, .825 km north of Bluff Head.
Palyniferous horizons: Thin layers of gray micaceous silty shale just below and .5 m above l-m-thick lake
bed containing abundant fossil fishes; additional layer 4.5 m above lake bed, consisting of variegated
greenish gray to red, poorly bedded silty claystone interbedded within a red siltstone and silty shale
sequence.
Remarks: See Cornet et al. (1973), for more detailed description of section.
Palynoflorule Locality No. 5
Formation: Lower Portland Formation.
Stratigraphic position: 257± 6 m above Hampden lava-flow unit.
Geographic location: Connecticut; Middlesex County, just west of Cromwell; neat top of ravine,
overlooking Chestnut Brook and Connecticut Route 9.
Palyniferous horizon: 8-cm-thick lens of greenish gray, silty shale, which is part of a micaceous, arkosic
sandstone sequence with numerous dinosaur foot-prints.
Remarks: Palyniferous horizon apparently thickens to 38 cm about 100 m upstream in Chestnut Brook.
Palynoflorule Locality No. 11
Formation: Middle Portland Formation.
Stratigraphic position: 1100 ± 6 m above Hampden lava-flow unit, provided no major faults exist.
Geographic location: Massachusetts; Hampden County, Agawam; cliff exposure on south bank of
Westfield River, next to Bridge Street bridge.
Palyniferous horizons: 6 beds of dark gray siltstone and shaly siltstone interbedded with brown-red
siltstone and sandstone.
Remarks: Cliff sequence measures 16 m thick, with an average interval of 2.3 m between palyniferous
horizons.
SYSTEMATIC PALEONTOLOGY
Samples were prepared using standard HF-maceration and ZnCl2 separation. Glycerin jelly was used as
mounting medium. Type specimens are deposited in the palynological collection at the Pennsylvania
State University. The assignment of specimens to species previously described in the literature is based on
5 or more specimens. If less than 5 specimens are described, and comparison with an established species
indicates affinity, our specimens are compared to that species. (The notation "cf." is used in such cases.
However, "cf." also is used in some instances where more specimens are available but doubt as to exact
affinity still exists.) A new species is proposed only if 10 or more specimens are described and if the
specimens constitute a coherent group distinct from any already in the literature. If less than 10 specimens
are described for a new form, it is given a reference number; for example, sp. 1 or Spore A. The median
rather than the mean is given for unimodal and relatively non-skewed site ranges. For skewed or very broad
size distributions, mean and mode are also given. Abundance of a species at a given locality is indicated by
the following terms: rare (less than 1%), scarce (1-3%), common (4-10%), abundant (11-40%), and
dominant (41-100%). A question mark indicates that rare, poorly preserved specimens are present, or that
at a certain cited locality there is some doubt as to affinity of the specimens concerned. Location of type
specimens is given with reference to an "X" scratched on the upper surface of the slide, per Traverse
(1958).
The terms intrastructure and intrastructural are used in descriptions for internal features of exines because
"infrasculpture" confuses structure (internal) and sculpture (external), and "infrastructure'' has been used in
other senses.
Genus Anapiculatisporites Potonié & Kremp 1954
Type species: Anapiculatisporites isselburgensis Potonié & Kremp 1954
Anapiculatisporites cf. A. dawsonensis
Reiser & Williams 1969
Plate 2, figure 9
Description: See Reiser and Williams, 1969, p. 3.
Remarks: One specimen measures 44 pm in diameter.
Locality: Portland Fm. 11 (rare).
Genus Converrucosisporites Potonié & Kremp 1954
Type species: Converrucosisporites triquetrus (Ibrahim 1933) Potonié & Kremp 1954
Basionym: Verrucosisporites triquetrus Ibrahim 1933
Converrucosisporites cameronii (de Jersey 1962)
Playford & Dettmann 1965
Plate 2, figures 3-8
Verrucosisporites cameronii de Jersey, 1962, p. 6, P1. 2, fig. 2, 3.
Triletes hungaricus Venkatachala & Góczán, 1964, p. 210, P1. 1, fig. 1~14.
Conbaculatisporites densus Mädler, 1964, p. 175, Pl. 1, fig. 9.
Converrucosisporites cameronii (de Jersey) Playford & Dettmann, 1965, p. 136, Pl. 12, fig. 11-13.
C. luebbenensis Schulz, 1967, P· 561, P1. 2, fig. 15-17; P1. 25, fig. 1.
C. minor Pocock, 1970, p. 47, P1. 8, fig. 4, 5, 7.
Emended description (more than 500 specimens): Isospores radial, trilete. Amb usually subaiangular
with broadly rounded apices and sides slightly concave to slightly convex; amb sometimes subspherical.
Laesurae extend almost to, but rarely reach, equaterial margin, and are usually bordered by narrow,
slightly elevated lips. Kyrtomes prominently to weakly developed, not always present. Laesurae frequently
bordered by arcuate folds concave toward proximal pole. Exine variably sculptured with a mixture of
rounded or blunt-topped verrucae, large and small bacula, broad and narrow-based coni, rate clavae, and
grana (grana comprising less than 50% of sculpture: compare description of Granulatisporites infirmus
(Balme) comb. nov.). Sculpture comprehensive, although usually more scattered and reduced
proximally. Distal and equatorial sculpture randomly spaced, distance between larger elements frequently
greater than their diameter. Bases of sculptural elements usually joined to form a low, imperfect ridge
system or incipient reticulum. When elements are large and closely spaced, many are fused into
elongate, irregular tugae. Sculpture usually dominated by 1 type: verrucae 0.8-1.8 um in diameter
and height, bacula 0.8-1.4 um high, coni 0.5-1.0 um high, and clavae about 1.0 um high. Exine 0.4-0.8 um
thick (exclusive of sculpture), single-layered. Equatorial diameter (60 specimens) ranges from
33 um to 51 um (median 42 um), 90% of grains between 36 um and 49 um·
Remarks: Specimens of this study compare very closely with C. cameronii from Australia, as well
as with the spore of Clathropteris meniscoides Brongniart figured by Harris (1931, Pl. 18, fig. 3).
The largest sculptural elements of the Australian form differ slightly from those of specimens from
the Hartford Basin in being taller and conate rather than blunt and verrucate. The list of synonyms
contains species that appear to be largely dominated by one type of sculpture: Triletes hungaricus with
broadly based coni, Conbaculatisporites densus with closely spaced bacula, Converrucosisporites
luebbenensis with verrucae and "truncae," and Converrucosissporites minor with broad verrucae.
Numerous specimens from locality 8 closely compare with these species. Rather than divide what appears
to be a natural species into 6 morpho-species, all of which would overlap in size and sculptural variation, 2
species are used that encompass the wide variation in sculpture with the least amount of overlap:
Granulatisporites infirmus (Balme) comb. nov. and Converrucosisporites cameronii.
Affinity: See discussion under affinity of G. infirmus.
Localities: Portland Fm., 5, 11 (rare to scarce); Shuttle Meadow Fm. 1, 3, 4, 12 (rare to common);
upper New Haven Arkose, 8 (abundant to dominant).
Genus Convolutispora Hoffmeister,
Staplin & Malloy 1955
Type species: Convolutispora florida Hoffmeister,
Staplin & Malloy 1955
Convolutispora klukiforma (Nilsson 1958)
Schulz 1967
Plate 1, figures 1-3
Description: Isospores radial, trilete. Amb usually subspherical, occasionally subuiangular. Distal and
equatorial sculpture composed of large irregular verrucae, basally joined to form a large reticulum;
lumina of reticulum 5-6 um in diameter. Verrucae generally aligned in 2 roughly perpendicular directions,
one axial, the other equatorial. Proximal sculpture similar but reduced in height and broader, forming a flat
contact area flanking the laesurae. Radius of laesurae 27-51% (median 34%) of equatorial diameter. Exine
thickness 1.7-5.l um. Equaterial diameter (53 specimens) 44-75 um (median 59 um), 90% of specimens
between 48 um and 71 um.
Remarks: Contignisporites dunrobinensis (Couper 1958) Schult 1967 compares somewhat with this
species, but axial sculptural lineation weakly expressed, equatorial ribs well developed. C. klukiforma may
be conspecific with Striatella jurassica Mädler 1964. Species most prevalent in lake bed at Locality 1.
Affinity: Schizaeaceous fern?
Localities: Shuttle Meadow Fm., 1, 12 (rare).
Genus Deltoidospora Miner 1935
Type species: Deltoidospora hallii Miner 1935, designated by Potonié 1956.
Remarks: For generic synonymy, see Pocock, 1970, p. 27.
Deltoidospora hallii Miner 1935
Plate 3, figure 6
Deltoidospora hallii Miner, 1935, p. 618, P1. 24, fig. 7, 8.
D. cascadensis Miner, 1935, p. 618, Pl. 24, fig. 9-12.
Tripartina variabilis var. glabra subvar, gradata Malyavkina, 1949, p. 50, P1. 7, fig. 10, 12, 14.
Leiotriletes gradatus (Mal.) Bolkhovitina, 1953, p. 19, PI. 1, fig. 1~12; Pi. 7, fig. 10.
Deltoidospora gradata (Mal.) Pocock, 1970, p. 28, P1. 5, fig. 2.
Description: Isospores, trilete. Amb subtriangular in polar view, with sides nearly straight to moderately
convex. Laesurae extend to, sometimes ending short of, equatorial margins. Laesurae may or may not
be bordered by narrow raised lips; kyrtomes absent. Exine single layered, psilate, 0.8-1.0 um thick.
Size range (14 specimens): 20-39 um in diameter (median 31 um).
Remarks: The site range of our specimens completely overlaps the size ranges of species listed in the
synonymy. Miner (1935) separated D. hallii and D. cascadensis on the basis of shape, but our specimens
contain gradational types between spores with straight sides and ones with convex sides. The
presence or absence of narrow, raised lips is not a significant difference -- lips may be variable in
development as they are with the spore types referable to Clathropteris meniscoides Brongniart.
Locality: Portland Fm., 11 (rare).
Genus Dictyophyllidites Couper 1958
Type species: Dictyophyllidites harrisii Couper 1958
Dictyophyllidites paramuensteri n. sp.
Plate 3, figures 4, 5, 8
Holotype: Plate 3, fig. 5, size 63 um; Slide BS2, coordinates 23.2 x 116.2, ref 16.1 x 119.6.
Description: Isospores radial, trilete. Amb subtriangular with broadly rounded apices, and sides slightly
concave to slightly convex. Laesura extends almost to, rarely reaching, equatorial margin, sometimes
bordered by narrow, slightly elevated lips. Proximal side flattened, sometimes appearing slightly concave
in lateral compressions; distal side strongly convex. In lateral compressiqns depth of spore equal to or
greater than width. Kyrtomes usually weakly developed, sometimes strongly developed in polar
compressions, frequently not evident in lateral comptessions. Exine psilate to vaguely punctate, single
layered, 0.6-0.8 um thick. Equatorial diameter (252 specimens) ranges from 26 um to 110 um (median
68 um), 90% of grains between 41 um and 101 um.
Remarks: The distribution curve is very broad and basically unimodal. Three slight peaks occur at
60 um, 68 um, and 85 um, but are not distinct enough to warrant the creation of more than one organ
species. If this species were derived from one source, the extreme variation in size suggests some sort of
environmental stress or adaptive significance. The size range completely overlaps the ranges of
Dictyophyllidites harrisii Couper 1958 (36-56 um), D. mortonii (de Jersey) Playford & Dettmann 1965 (2845 um), Deltoidospora minor (Couper 1958) Pocock 1967 (26-56 um), D. australis (Couper 1358)
Pocock 1970 (58-90 um), and the spores (26-47 um) of the fern Dictyophyllum nilssonii (Brongniart)
Goeppert (quoted from Harris, 1944, by Couper, 1958). Dictyophyllum muensteri (Goeppert) Nathorst is
the only Rhaeto-Liassic fern known to produce spores of the Dictyophyllidites type of a size similar to D.
paramuensteri n. sp. The average size of these spores given by Harris (1931) is 65 um. Harris' illustration of
one such spore (Harris, 1931, Pl. 14, fig. 3) measures 76 um at the given magnification. These 2 figures
agree with the median of 68 um for D. paramuensteri n. sp.; hence, the implied relationship.
Comparison of this species with Deltoidospora Miner 1935 reveals some close similarities in shape. Many
of our specimens appear to lack kyrtomes, but cannot be separated by other characteristics from the
specimens that possess them. The thin, flexible exine of this species contrasts with the more rigid
exine generally found in Deltoidospora (synonym: Cyathidites).
This species is most prevalent in the lake bed at Locality 1.
Affinity: Probably Dictyophyllum muensteri, in part, but the size range could include some spores
of other ferns.
Localities: Portland Fm., 11?, 13 (rare); Shuttle Meadow Fm., 1, 12 (rare to scarce); upper New Haven
Arkose, 8 (rare).
Dictyophyllidites cymbatus
Venkatachala & Góczán 1964
Plate 1, figures 11, 12
Remarks: Specimens from the Hartford Basin agree well with this species, particularly in the strong
development of kyrtomes. Fifteen specimens range from 31 um-5l um (median 42 um) in equatorial
diameter. Exine 0.5-0.9 um thick.
Localities: Portland Fm., 5, 11 (rare).
Dictyophyllidites harrisii Couper 1958
Plate 1, figures 7, 8
Remarks: Eight specimens from the Hartford Basin range from 37 um to 51 um (median 43 um) in
equatorial diameter. This species is present in a mottled gray brown claystone, 5 stratigraphic m above the
lake bed at Locality 1, and appears to be separable from D. paramuensteri n. sp at this horizon by its lighter
color and more restricted size range. Dictyophyllidites spp. from the lake bed horizon do not show a similar
bimodal distribution in size.
Localities: Portland Fm., 5, 11 (rare); Shuttle Meadow Fm., 1, 12 (rare); upper New Haven Arkose, 8?.
Genus Dictyotriletes Naumova 1939
Type species: Dictyotriletes bireticulatus (Ibrahim 1932) Potonié & Kremp 1955 (designated by
Potonié & Kremp, 1955)
Basionym: Sporonites bireticulatus Ibrahim 1932 in Potonié, Ibrahim, and Loose, 1932.
Dictyotriletes Naumova 1939.
Ischyosporites Balme 1957.
Klukisporites Couper 1958.
Dictyotriletes sp. 1
Plate 1, figures 5, 6
Description: Isospores, trilete. Amb subtriangular in polar view, strongly convex distally. Proximal face
psilate; laesurae extend almost to equatorial margins. Distal side sculptured with a large, predominantly
complete reticulum, which extends slightly proximal to equatorial margin at apices, generally ending at
equatorial margin between apices. Muri of reticulum 2-7 um wide; lumina of reticulum 1.7-5.2 um in
overall diameter. Exine doublelayered, 1.7-3.0 um thick, increasing to 4.0 um at apices; endexine thinner
than ektexine, but not clearly delimited. Size (2 specimens): 44-48 um in polar-view diameter, one
specimen measuring 51 um in depth.
Remarks: This species is similar to Ischyosporites marburgensis de Jersey 1963 (recorded from the
Liassic of Australia), which differs in having a slightly thicker exine and more uniform distribution
and shape of lumina. Dictyotriletes sp. 1 has also been recovered from our Licking Run locality near
Midland, Va., where several distinct species of Dictyotriletes are present (Pi. 1, fig. 4).
Locality: upper New Haven Arkose, 8 (rare).
PLATE 1
1 Convolutispora klukiforma (Nilsson) Schulz. Locality 1, 53 um.
2 C. klukiforma. Locality 1, 56 um.
3 C. klukiforma. Locality 1, 63 um wide.
4 Dictyotriletes sp. Locality on Licking Run, near Midland, Va., 70 um wide.
5 Dictyotriletes sp. 1. Locality 8, 44 umwide.
6 Dictyotriletes sp. 1. Locality 8, 48 umwide.
7 Dictyophyllidites harrisii Couper. Locality 1, 44 um.
8 D. harrisii. Locality 5, 42 um.
9 Murospora sp. 1. Locality 11, 37 um.
10 Murospora sp. 1. Locality 11, 26 um.
11 Dictyophyllidites cymbatus Venkatachala & Góczán 1964. Locality 5, 39 um
12 D. cymbatus. Locality 5, 43 um.
13 Foveosporites agawamensis n. sp (Holotype). Locality 11, 46 um (under oil).
14 F. agawamensis n. sp. Locality 11, 25 um (under oil).
15 F. agawanzensis n. sp. Locality 11, 41 um.
16 Gleicheniidites cf. G. nilssonii Pocock. Locality 5, 41pm.
PLATE 2
1 Granulatisporites infirmus ( Balme) comb. nov. Locality 8; part of spore mass found in close association
with Clathropteris meniscoides Brongniart; grain on far left, 42 um.
2 G. infirmus.Locality 8; S.E.M., 41 um.
3 Converrucosisporites cameronii (de Jersey) Playford 8 ( Dettmann. Locality 8, 46 um.
4 C. cameronii. Locality 8, 42 um.
5 C. cameronii. Locality 8; S.E.M., 41 um wide.
6 C. cameronii. Locality 8; S.E.M.; part of spore mass probably derived from Clathropteris meniscoides
Brongniart; central grain, 37 um wide.
7 C. cameronii. Locality 8; S.E.M., 45 um.
8 C. cameronii. Locality 8, 56 um.
9 Anapiculatisporites cf. A. dawsonensis Reiser & Williams. Locality 11, 46 um.
10 Perotriletes cf. P. pseudoreticulatus Couper. Locality 11, 54 um.
11 Leptolepidites cf. L. major Couper. Locality 11; focus on sculpture to Fig. 12.
12 Leptolepidites cf. L. major. Locality 11, 54 um.
13 Todisporites rotundundiformis (Malyavkina) Pocock. Locality 11; 54 um.
14 T. rotundiformis. Locality 11; 46 um.
15 T. rotundiformis. Locality 11; 36 um.
Genus Foveosporites Balme 1957
Type species: Foveosporites canalis Balme 1957
Foveosporites agawamensis n. sp.
Plate 1, figures 13-15
Holotype: Plate 1, fig. 13, size 46 um; Slide AGA6-1, coordinates 52.8 x 111.2, ref 28.5 x 111.6.
Description: Miospores, trilete. Amb subspherical, rarely subtriangular. Laesurae margins slightly
thickened, sometimes as narrow conspicuous lips. Radius of laesura 35-46% of overall diameter of spore.
Exine 0.6-0.8 um thick, proximally covered with densely to sparsely spaced pits, 0.3-0.4 um in diameter.
Pits more closely spaced near laesura, decreasing in number equatorially; distal side generally lacking
pits. Size range (11 specimens): 25-52 um (median 41um).
Specific epithet refers to Agawam, Mass.
Locality: Portland Fm., 11 (rare).
Genus Gleicheniidites Ross 1949
Type species: Gleicheniidites senonicus Ross 1943
Gleicheniidites cf G. nilssonii Pocock 1970
Plate 1, figure 16
Description: See Pocock (1970, p. 32).
Remarks: The uniqueness of this species necessitates some description: our specimens compare very
closely with G. nilssonii, but have a slightly thicker exine (about 2.6 um thick between apices, decreasing
to about 2.1 um at apices). A distal triangular area of thickening is delimited by 3 arcuate ridges,
intersecting at the apices. Two specimens measure 41 um and 43 um in diameter. The small number of
specimens permits only tentative assignment at this time.
Locality: Portland Fm., 5 (rare).
Genus Granulatisporites Ibrahim 1933 emend.
Potonié & Kremp 1954
Type species: Granulatisporites granulatus Ibrahim 1933
Granulatisporites infirmus (Balme 1957) comb. nov.
Plate 2, figures 1, 2
Concavisporites infirmus Balme, 1957, p. 21, Pl. 2, fig. 32, 33 (Basionym).
Trachysporites asper Nilsson, 1358, p. 39, P1. 2, fig. 3.
Granulatisporites cf. G. asper (Nilsson) Playford & Dettman, 1965, p. 135, Pl. 12, fig. 10.
Emended description (more than 500 specimens): Isospores radial, trilete. Amb usually subtriangular
with broadly rounded apices, and sides slightly concave to slightly convex; amb sometimes subspherical.
Laesura extends almost to, rarely reaching, equatorial margin, usually bordered by narrow, slightly elevated
lips. Kyrtomes prominently to weakly developed, not always present. Laesurae frequently bordered by
arcuate folds concave toward proximal pole. Exine sparsely to densely sculptured with grana and small
verrucae, mixed with a few bacula and coni, all of which are frequently joined at their bases by a low,
imperfect ridge system or incipient reticulum. Sculpture is comprehensive, but usually
proximally scattered and more reduced; sculpture ranges from predominantly small grana and verrucae,
0.1-0.8 um in diameter (always more than 50% of sculpture), to a mixture of grana, verrucae, bacula, and
coni; occasionally either low coni or bacula dominate the larger sculptural elements. Exine 0.4-0.8 um thick
(exclusive of sculpture), single layered. Equatorial diameter (100 specimens) ranges from 22 um to 62 um
(median about 40 um), 30% of specimens between 33 um and 43 um. A few abnormal monolete and dilete
(chevron) grains observed.
Remarks: Playford and Dettmann (1965) considered Concavisporites infirmus Balme (equatorial
diameter 35-46 um, mean 40 um) to be distinct from Trachysporites asper Nilsson (size 38-41 um) and
their specimens, since C. infirmus possesses kyrtomes and a finely rugose sculpture. However, Balme's
illustrations bear remarkable resemblance to specimens of the present study, most of which have some
degree of kyrtome development. The illustrated specimen of T. asper (Nilsson, 1958, P1. 2, fig. 3) also has
raised arcuate folds flanking the laesurae. The finely rugose sculpture in the illustrations of C. infirmus is
not obvious (the sculpture appears to consist of discrete grana and verrucae); a basal incipient reticulum
could be described as finely rugose, because closely spaced sculptural elements would be united to give an
apparent lineation pattern (S.E.M. photomicrograph, P1. 2, fig. 2). The lack of obvious distinction between
C. infirmus, T. asper, and our specimens favors uniting these forms under the earliest name, C. infirmus.
This species conforms to the generic requirements of Granulatisporites ibrahim emend. Potonié & Kremp,
which has priority over Concavisporites Pflug and Trachysporites Nilsson.
Granulatisporites infirmus grades into Converrucosisporites cameronii (de Jersey) Playford & Dettmann in
our samples, but for practical reasons the two forms are separated by gross sculptural differences.
Granulatisporites subgranulosus (Couper, 1958) Fisher 1972 differs from G. infirmus by having a much
thicker, double-layered exine (l.5-3.0 um thick).
Affinity: Granulatisporites infirmus has been recovered by the first author from sporangia of Clathropteris
meniscoides Brongniart at Locality 8. Associated spore masses in sporangial preparations contain C.
cameronii. Some dispersed spore preparations from the megafossil plant layers are dominated by C.
cameronii, while others are dominated by G. infirmus, although both forms are invariably present in the
same preparation. Specimens of Equisetites spp. are the only other megafossils present in the plant bed.
Probable roots and rhizomes of C. meniscoides and Equisetites are preserved in situ. Only one species of
Clathropteris appears to be present, although it exhibits a wide range in frond site.
Localities: Portland Fm., 5, 11 (rare to scarce); Shuttle Meadow Fm., 1, 3, 4, 12 (rare to common);
upper New Haven Arkose, 8 (abundant to dominant).
Genus Harrisispora Pocock 1970
Type species: Harrisispora mollis (Bolkhovitina 1956) Pocock 1970
Basionym: Adiantum mollis Bolkhovitina 1956
Remarks: See Pocock (1370, p. 38, 39) for generic description and generic synonymy.
Harrisispora sp. 1
Plate 3, figure 3
Description: Miospores, trilete. Amb subtriangular with slightly convex sides. Radius of laesura about
30% of diameter of spore; laesura occupies for nearly its entire length a narrow zone or groove (1.6-3.0 um
wide) of exinal thinning where ektexine and endexine apparently separate. Both endexine and ektexine
participate in forming the laesura. Narrow, raised lips present on surface of endexine in groove. Adjacent to
the narrow zone of thinning, a band of exine (67 um wide) is inaapunctate and thickened on each side of
laesura. Exine double layered, psilate, about 1.4-1.6 um thick (exclusive of proximal area of thickening);
ektexine 1.0 um thick, endexine 0.4-0.6 um thick. Size (1 specimen): 80 um in diameter.
Affinity: Possibly ferns related distantly to Lygodium (Pocock, 1970).
Locality: Portland Fm., 11 (rare).
Genus Leptolepidites Couper 1953
Type species: Leptolepidites verrucatus Couper 1953
Leptolepidites cf. L. major Couper 1958
Plate 2, figures 11, 12
Remarks: Two specimens, measuring 46 um and 54 um, agree with L. major in size, and in having
verrucae frequently in contact. However, the verrucae
of our specimens are much smaller (1.0-2.5 um high; 1.0-3.0 um in diameter), agreeing in size and in
number around the edge with Leptolepidites bossus (Couper) Schulz. Our specimens appear to be
intermediate between L. major and L. bossus.
Locality: Portland Fm., 11 (rare).
Genus Murospora Somers 1952
Type species: Murospora kosankei Somers 1952
Murospora sp. 1
Plate 1, figures 9, 10
Description: Miospores, trilete. Amb subcircular to subtriangular in polar view; equatorial outline of
cingulum generally parallels that of central body. Laesura extends almost to, sometimes reaching,
equatorial margin of central body. Radial apices occasionally protrude beyond general outline of central
body, producing a bulge on the cingulum. Laesura usually bordered by narrow, raised lips. Exine double
layered, psilate. Ektexine extends over whole surface, including cingulum. Endexine generally restricted to
central body, occasionally extending as lobes into, sometimes to outer margin of, cingulum. Exine 0.9-1.0
um thick, ektexine about 0.4 um thick, endexine about 0.6 um thick. Size range (7 specimens): 26-53 um in
overall diameter (median 37 um); diameter of central body 22-41 um; width of
cingulum 3-7.0 um.
Locality: Portland Fm., 11 (rare).
Genus Perotriletes Couper 1953
Type species: Perotriletes granulatus Couper 1953
Perotriletes cf. P. pseudoreticulatus Couper 1953
Plate 2, figure 10
Description: Miospores radial, trilete. Amb subtriangular in polar view. Laesura extends almost to
equatorial margin, flanked by narrow, raised lips. Exine (exclusive of perispore) about 0.6 um thick;
both proximal and distal faces sculptured with irregular, occasionally anastomosing, ridges, 1-2 um
high. Perispore closely fitting, densely covered with small grana, 0.1-0.4 um high; perispore attached
only at laesura margins. Overall site (2 specimens) 55-56 um, about 52-53 um in diameter, excluding
perispore.
Remarks: The presence of grana on the perispore might be sufficiently different from the type descrip-
tion to warrant the creation of a new species. However, because of the small number of specimens, a
tentative assignment is made until further information can be obtained. Our specimens differ significantly
from other described species of Perotriletes.
Locality: Portland Fm., 11 (rare).
Genus Todisporites Couper 1958
Type species: Todisporites major Couper 1958
Todisporites rotundiformis (Malyavkina 1949) Pocock 1970
Plate 2, figures 13-15
Cyclina pseudolimbata var. rotundiformis Malyavkina, 1949, p. 53, P1. 9, fig. 13.
Leiotriletes rotundiformis (Mal.) Bolkhovitina, 1956, p. 30, P1. 1, fig. 42b.
Todisporites minor Couper, 1958, p. 135, P1. 16, fig. 9, 10.
T. rotundiformis (Mal.) Pocock, 1970, p. 30, P1. 5, fig. 15.
Description: See Couper (1958, p. 135).
Remarks: Our specimens are very similar to T. minor, but have a thinner exine (0.4-0.7 um) and a
distribution curve skewed toward smaller size. Measurement of 32 specimens from Locality 11 gives a size
range of 29 um to 54 um (median 39 um, mean 40 um, mode 35 um). However, these differences are not
significant. Pocock (1970) gives a size range of 38 um to 46 um, Couper (1958) gives 32 um to 50 um
(mode 45 um), Malyavkina (1949) gives 30 um to 40 um. Some of our larger specimens are subtriangular
in shape, but invariably these have major folds.
Affinity: Couper (1958) compared T. minor with the spores of Todites princeps (Presl) Gothan,
which are smaller (20-40 um) and have a mean (32 um) close to our mode.
Localities: Portland Fm., 10, 11 (rare).
Genus Verrucosisporites Ibrahim 1933 emend.
Potonié & Kremp 1954
Type species: Verrucosisporites verrucosus Ibrahim 1933
Verrucosisporites cheneyi n. sp.
Plate 3, figures 1, 2
Holotype: Pl. 3, fig. 1, size 73 um; Slide BS3, coordinates 40.5 x 122.7, ref. 56.2 x 126.2.
Description: Isospores, radial, trilete. Amb circular to subcircular, proximal face less convex than distal
side. Exine densely sculptured with verrucae and a few grana, 0.9-3.4 um high, 1.5-2.8 um wide.
Verrucae sometimes in contact with one another, occasionally fused into larger elements. Radius of
laesurae variable, 26-78% of equatorial diameter. Exine (exclusive of sculpture) approximately 0.8 um
thick, single layered. Equatorial diameter (41 specimens) 58-83 um (median 70 um), 90% of grains
between 59 um and 79 um.
Remarks: This species compares closely with the Permian Verrucosisporites trisectus Balme &
Hennelly 1956b, but differs somewhat in size range (V. trisecatus: 65-135 um), and in the degree of
suture development (laesurae of V. trisecatus frequently extend onto distal surface of spore). The
sculpture of both species is very similar.
Species most prevalent in lake bed at Locality 1.
Species named after Jane Cheney, director emeritus of the Children's Museum of West Hartford, Conn.,
in recognition of her efforts to preserve the natural history of the Connecticut Valley.
Affinity: Osmundaceous fern?
Localities: Portland Fm., 6, 11, 13 (rare); Shuttle Meadow Fm., 1 (rare); upper New Haven Arkose,
8 (rare).
Spore A
Plate 3, figure 7
Description: Miospore, trilete. Amb triangular in polar view, apices rounded. Radius of laesura about 30%
of diameter of spore. Laesura open, bordered by a kyrtome consisting of narrow bands of exinal thickening.
Distai side extended, equatorially strongly concave, forming triradiate fold centered at distal pole. Exine
double layered; endexine 0.7 um thick; ektexine 1.0 um thick, loosely attached along triradiate fold. Size:
98 um in diameter.
Locality: Portland Fm., 11 (rare).
Genus Pilasporites Balme & Hennelly 1956a
Type species: Pilasporites calculus Balme & Hennelly 1956a
Pilasporites allenii Batten 1968
Plate 4, figure 11
Description: See Batten (1868, p. 638).
Remarks: Our specimens agree closely in all respects with P. allenii Batten from the Lower Cretaceous of
England, except in exine thickness, which is not a significant difference for the creation of a new species.
Measurement of 72 specimens from Locality 8 gives a size range of 30 um to 54 um (median 39 um, mean
39 um), indicating a slightly larger average size than that of the type description (mean 36.7 um; Batten,
1968). Exine of our specimens is 0.5-0.8 um thick as compared with 1.25-1.75 um given by Batten. Slightly
irregular distribution in the size-range curve of our specimens suggests more than one source plant, as is
also indicated by megafossils.
Affinity: Equisetites cf. E. sarranii Zeiller (abundant in layers containing P. allenii), as well as possibly two
other species of Equisetites, present at Locality 8.
Localities: Portland Fm., 5, 2 1 (rare); Shuttle Meadow Fm., 1 (rare in lake bed); upper New
Haven Arkose, 8 (rare to scarce).
Genus Araucariacites Cookson 1947
Type species: Araucariacites australis Cookson 1947
Remarks: Cookson (1947), following the suggestions of Erdtman (1947), created the "sporotype" names
Granulonapites (± = form-generic name) and Araucariacites (± = organ-generic name), as alternative names
to be combined with the epithet, australis. The 1972 I.C.B.N. restriction on the use of alternative names is
retroactive only to January 1, 1953. Consequently, we consider subsequent attempts to validate one of the
two generic names (Couper, 1953, Nilsson, 1958) unnecessary, since Cookson's monotypic species was
validly published. Her sporotypes and sporomorphs are handled normally in all respects as genera and
species, respectively. Our choice of the generic name, Araucariacites, is based on the fact that Cookson
does not accept the name Granulonapites in her publication: wherever the generic concept is given a single
name, that name is always Araucariacites.
Araucariacites punctatus (Nilsson 1958) comb. nov.
Plate 6, figures 14, 15
Granulonapites punctatus Nilsson, 1958, p. 70, Pl. 5, fig. 22.
Description: Pollen grains inaperturate, spherical to oval, frequently folded. Exine 0.5-0.7 I~m thick,
densely sculptured with small grana, 0.2 um or less in diameter. Size range (50 specimens): 28-63 um
(median 44 um) in maximum width or diameter, 90% of specimens between 34.0 um and 56.8 um.
Remarks: This species agrees in all morphological respects with Araucariacites australis Cookson, but
has a smaller mean and smaller site distribution than generally recognized for A. australis. Couper (1953)
reported grains of similar size under the name A. australis from Jurassic-lower Oligocene strata of New
Zealand. Nilsson (1318) described one specimen (upon which A. punctatus is based) from the middle
Keuper of Scania.
This species is generally much more common than A. australis in our samples.
Localities: Portland Fm., 5, 6, 7, 9, 10, 11, 13 (rare to scarce); Shuttle Meadow Fm., 1, 12 (rare).
Araucariacites australis Cookson 1947
Plate 6, figure 13
Remarks: This species is distinguishable from A. punctatus (Nilsson) comb. nov. by its larger size and
usually darker color. Range in size for 26 specimens: 54-94 um (median 73 um). Our specimens compare
well in shape and in manner of folding to grains of Brachyphyllum mamillare Brongniart as illustrated by
Couper (1958, Pl. 27, fig. 1, 2). Species is more frequently encountered in the Portland Fm. than in older
strata. Some specimens assigned to this species may be unsplit examples of A. fissus Reiser & Williams.
Localities: Portland Fm., 5, 11 (rare); Shuttle Meadow Fm., 1 (rare).
Araucariacites fissus Reiser & Williams 1969
Plate 7, figures 6, 7
Description: See Reiser & Williams (1969, p. 17).
Remarks: The size distribution (17 specimens) indicates a larger average size than that given by
Reiser & Williams (1969), being 58-110 um (median 85 um) rather than 34-95 um (?mean 57 um) for
the Australian forms. The indicated Early Jurassic age for A. fissus Reiser & Williams agrees well with
the proposed age for the Portland Fm. This species becomes more frequent in younger strata. The largest
forms are more common at Locality 5.
Discussion of relationship of A. fissus to Triangulopsis: Our specimens (esp. P1. 7, fig. 7) from Locality
11 compare well in overall shape, manner of splitting, and size range to Triangulopsis discoidalis Döring
1961, recorded from the Upper Jurassic of Germany and western Canada. However, T. discoidalis
apparently has a much thicker, double-layered exine, separable into an outer envelope and an internal
body (Pocock, 1970, p. 74). Döring considered T. discoidalis to be microplanktonic, while Pocock's study
of a large population of this species indicated morphological affinity with vascular plants. Döring
considered Zonalapollenites to be synonymous with Triangulopsis, suggesting a similar basic structure.
It is possible that A. fissus is related to Triangulopsis discoidalis, lacking only the development of a
distinct endexine.
Localities: Portland Fm., 5, 10, and 11 (rare).
Genus Perinopollenites Couper 1958
Type species: Perinopollenites elatoides Couper 1958
Perinopollenites elatoides Couper 1958
Plate 4, figures 12, 13
Description: See Couper (1958, p. 152).
Remarks: One specimen from~locality 11 clearly shows a pore (Pl. 4, fig. 12). Other specimens possess
a highly folded "perine," which may simulate a crude reticulum (Pl. 4, fig. 13). Specimens from Locality 5
generally have a slightly smaller body, giving the appearance of a relatively larger "perine." Six specimens
range in maximum diameter from 34 um to 52 um; diameter of central body 28 um-43 um.
Localities: Portland Fm., 5, 11 (rare).
Genus Callialasporites Sukh Dev 1961
Type species: Callialasporites trilobatus (Balme 1957) Sukh Dev 1961 (designated by Sukh Dev, 1961)
Basionym: Zonalapollenites trilobatus Balme 1957
Zonalapollenites (in part) Pflug in Thomson and Pflug, 1953, p. 66, 67.
Callialasporites Sukh Dev, 1961.
Applanopsis Döring, 1961.
Triangulopsis (in part) Doring, 1961.
Pflugipollenites Pocock, 1962.
Applanopsipollenites Levet-Carette, 1964.
Description: See Sukh Dev, 1961, p. 48; Balme, 1957, p. 32.
Remarks: Sukh Dev (1961) and Pocock (1970) discuss the synonymy of Zonalapollenites and
Tsugaepollenites, and differences between these two genera and species now placed in Callialasporites.
Callialasporites cf. C. dampieri (Balme 1957) Sukh Dev 1961
Plate 4, figure 10
Description: See Balme (1957, p. 32).
Remarks: Three specimens range in overall diameter from 60 um to 77 um; body 56 um-59 um in
diameter. Sculpture of saccus is finely granulate. Endexine of body is about 1.3 um thick. A slightly
darkened area is present on the central part of the body. One specimen has saccus folded back on body,
resembling a similar condition found in C. segmentatus (Balme) comb. nov.
Affinity: Gamerro (1965) found grains of C. dampieri and C. trilobatus (Balme) Sukh Dev in male cones in
organic connection with leaves of the gymnosperm Apterocladus lanceolatus Archangelsky.
Locality: Portland Fm., 11 (rare).
Callialasporites segmentatus (Balme 1957) comb. nov.
Basionym: Zonalapollenites segmentatus Balme 1957, p. 33, Pl. 9, fig. 93, 94.
Remarks: A formal transfer is made of this species here, although no specimens definitely referable to it
were found.
Callialasporites trilobatus (Balme 1357) Sukh Dev 1961
Plate 4, figure 4
Description: See Balme (1957, p. 33).
Remarks: This species is larger than our specimens of Callialasporites dampieri, 7 specimens measuring
78 um to 104 um in overall diameter; diameter of body 46 um-84 um. Exine of body is 0.8-1.1 um thick.
Saccus is densely covered with small grana, 0.2-0.8 um in diameter. The shape of the body ranges from
oval to subtriangular, with 3 lobes well developed on forms with subtriangular bodies. Forms with oval
bodies tend to possess sacci without indentations. Sukh Dev (1961) described a new species,
Callialasporites monoalasporus, that closely resembles our forms with oval bodies. More specimens are
needed to determine if a significant distinction exists between our forms resembling C. monoalasporus
Sukh Dev, and C. trilobatus. In addition, C. monoalasporus may be conspecific with C. turbatus (Balme)
Schulz 1967.
Affnity: Gamerro (1965) illustrates grains of C. trilobatus and C. dampieri, as well as forms resembling C.
monoalasporus, which he found associated with Apterocladus lanceolatus Archangelsky. Reiser &
Williams (1963) found in their study forms of Callialasporites gradational between different species of this
genus, suggesting that many of them are derived from one source plant.
Locality: Portland Fm., 11 (rare).
Genus Alisporites Daugherty 1941 emend. Nilsson 1958
Type species: Alisporites opii Daugherty 1941
Alisporites thomasii (Couper 1958) Nilsson 1958
Plate 4, figure 5-7
Pteruchipollenites thomasii Couper, 1958, p. 150, P1. 26, fig. 10, 11.
Alisporites thomasii (Couper) Nilsson, 1958, p. 83, P1. 8, fig. 1.
Description: See Couper (1958, p. 150).
Remarks: Our specimens compare closely with A. thomasii in overall size and shape, as well as exine
structure and sculpture. Twenty-one specimens range from 48 um to 77 um in overall breadth (median
69 um); breadth of body 20 um-46 um (median 36 um); length of saccus 36 um-57 um (median 40 um).
Some specimens, not included in the above measurements, have bodies compressed more or less obliquely
with pendantly attached sacci, producing forms suggestive of pollen of extant Pinus. More study is needed
to determine whether these unique forms should be included in A. thomasii.
Localities: Portland Fm., 5, 11 (rate).
Genus Pityopollenites Reissinger 1950 emend. Jansonius 1962
Type species: Pityopollenites pallidus Reissinger 1950
Vitreisporites Leschik 1956 (imprint date, 1955).
Caytonipollenites Couper 1958.
Vitreisporites Leschik emend. Jansonius 1962.
Remarks: Pityopollenites was validly published by Reissinger in 1950, and was monotypic at the time of
publication. Vitreisporites, as originally described by Leschik in 1956, required a faint laesura and a size of
the central body under 20 um. Jansonius (1962) removed these characteristics from the generic description,
considering them either not convincingly demonstrated or too restrictive. For further discussion, see Dunay
(1972).
Pityopollenites pallidus Reissinger 1950 emend. Nilsson 1958
Plate 4, figures 2, 3
Pityopollenites pallidus Reissinger, 1950, p. 105), P1. 15, fig. 1-5.
Vitreisporites signatus Leschik, 1956 (imprint date, 1955), p. 53, P1. 8, fig. 10.
Caytonipollenites pallidus (Reissinger) Nilsson, 1958, p. 78, P1. 7, fig. 12-14.
Description: See Nilsson (1958, p. 54).
Remarks: Twenty-one specimens of this study range fiom 22 um to 33 um in overall breadth (median 27
um); breadth of body 7 um-15 um (median 27 um); length of saccus 14 um-22 um (median 17 um).
Specimens from Locality 5 are generally smaller than those from Locality 11.
Locatities: Portland Fm., 5, 11 (rare).
Genus Platysaccus Naumova 1939 emend.
Potonié & Klaus 1954
Type species: Platysaccus papilionensis Potonié & Klaus 1954
Remarks: Platysaccus was validly published by Naumova in 1939 (p. 355, fig. 1), although no
type species was designated, which is not required by the I.C.B.N. before 1358. Potonié and Klaus (1954)
designated their new species, P. papilionensis, as type, and emended the genus.
Platysaccus cf. P. lopsiensis (Malyavkina 1961) Pocock 1970
Plate 4, figure 1
Podocarpus lopsiensis Malyavkina in Samoilovich et al., 1961, p. 130, Pl. 35, fig. 9a,b.
Platysaccsus lopsiensis (Mal.) Pocock, 1970, p. 85, Pl. 18, fig. 6, 7.
Description: See Pocock (1370, p. 85).
Remarks: Two specimens compare closely with P. lopsiensis, measuring 56 um to 59 um in overall
breadth (grains not completely expanded); breadth of body 26 um-27 um ; length of saccus 26 um-46 um.
Locality: Portland Fm., 11 (rare).
Genus Podocarpidites Cookson 1947
Type species: Podocarpidites elliptica Cookson 1947 (designated by Potonie', 1958)
Remarks: This generic name was validly published by Cookson (1947) under circumstances similar to
those described above under Araucariacites Cookson.
Podocarpidites sp. 1
Plate 4, figure 14
Description: Pollen grains, bisaccate. Body of grain generally longer than broad; sacci strongly distally
pendant, rarely expanded, usually highly folded. Distal surface traversed by an elongate sulcus, about 20
um long, flanked by basal attachment of sacci. When expanded, saccus measures about 36 um in length,
50 um in breadth. Proximal surface of body covered by a closely adhering cappa, 1.4-2.6 um thick, tightly
folded to form a verrucose and rugose surface. Exine of body about 0.9 um thick. Sacci possess fine reticulum with lumina 0.2-0.3 um in diameter. Size of body (5 specimens): 40-46 um long; 31-34 um broad.
Sacci generally about 30 um in diameter (folded). Overall breadth approximately 50-60 um, but highly
dependent on shape of sacci.
Remarks: Bisaccates are gehetally rare in the Shuttle Meadow Fm. However, they increase slightly
in abundance in the lake bed and underclay at Locality 1. A similar florule from Licking Run near
Midland, Va. (see Discussion), contains as much as 15% bisaccates in certain lacustrine horizons. Most
bisaccates from floras dominated by Corollina meyeriana compare best with Podocarpidites and
Pityosporites, although examples of Alisporites have been found. Generally poor preservation of most of
the Shuttle Meadow bisaccates prevents description of several other types present.
Localities: Shuttle Meadow Fm., 1, 12 (rare); upper New Haven Arkose, 8?
Circulina-Corollina-Classopollis
Nomenclatural problem: Circulina and Corollina were validly published as generic names by Malyavkina
(1949), even though her generic and specific descriptions have been regarded as vague and inadequate for
validation (Pocock and Jansonius, 1961; Reyre, 1969; Traverse, Ames and Spackman [CCFSP, 1974, in
press: 38-103, 38-128]). Classopollis is generally regarded as the acceptable generic name, but was
originally described as tricolpate by Pflug (1953). Subsequent emendation of Classopollis by Pocock and
Jansonius (1961) and Couper (1958) largely corrected the description of this pollen type, but Chaloner
(1962) and Pettitt and Chaloner
(1964) provided the most accurate description, clarifying the structure of the exine with electron
micrographs of well-preserved grains. Klaus (1960) emended the description of Circulina and Corollina,
providing Hilfstypus -- better called neotype, which is admissible under 1372 I.C.B.N., Guide for
Determination of Types, p. 76, for establishing taxonomic validity. However, Klaus chose to distinguish
Circulina from Corollina, making Classopollis a junior synonym of Corollina.
S. Samoilovich (Institute of Petroleum, Leningrad, USSR; pers. comm. to A. Traverse, 1973) has indicated
that Malyavkina's type material was not preserved. She provided photographs (Pl. 4, fig. 9; Pl. 5, fig. 10)
of "neotypes" (chosen by Zauer, supplied to Klaus, and chosen by him as Hilfstypus) for species of
Circulina and Corollina, derived fiom the same Lower Jurassic stratigraphic horizon as studied by
Malyavkina (from the same locality, a borehole 80 km to the west of the settlement of Sagiz). Even
though the assignment of these "neotypes" by Zauer is subjective, it is perhaps the only way to obtain
stability of nomenclature and to avoid personal preference for the name Classopollis.
It is interesting to note that the "neotype" of Corollina compacta Malyavkina (Pl. 5, fig. 10) compares well
with Classopollis in exinal structure, even though a well-developed distal pseudopore is not present, and
equatorial striations are questionably present. Thus Klaus was apparently correct in making Classopollis
synonymous with Corollina rather than with Circulina, It is further significant that the "neotype" of
Circulina simplex Mal. closely resembles Malyavkina's illustrations, lacks a well-developed ring tenuitas
(rimula) and distal pseudopore, and resembles specimens of Spheripollenites Couper and endoexinal bodies
of Classopollis more than it does Circulina meyeriana (Mal.) Klaus, raising the question of the correctness
of Klaus's emendation of Circulina.
Reyre (1969) considers Circulina Mal. emend. Klaus as "an exceptional case within the genus
Classopollis, corresponding to the coincidence of the limiting cases of the two characters, equatorial
thickening and ornamentation of the intrastructure." He describes two species, Classopollis quezeli and C.
kieseri, with a typical Classopollis intrastructure, but without equatorial striations or with only vaguely
defined striations (pseudostriations). These two species closely resemble grains that have been erroneously
assigned to Corollina meyeriana (Klaus) Venkatachala & Goçzán (Schulz, 1967, P1. 22, fig. 6) or
Circulina meyeriana Klaus (Geiger and Hopping, 1968, P1. 4, fig. 32). In fact this confusion suggests
that there is really no clear generic distinction between grains with a massive intrastructure and no
equatorial thickening or striations, and grains with a structured exine, equatorial thickening, and striations.
Consequently the only valid genetic characteristics are those that Circulina emend. Klaus and Classopollis
have in common: frequent occurrence as tetrads, spherical shape, size range (18-60 um), proximal trilete
mark, distal pseudopore, and subequatorial furrow or rimula. Since the intrastructure of the exine ranges
from massive through various types (Reyre, 1969), equatorial striations vary in number and completeness,
particularly in some species, for example, Classopollis torosus (Reissinger) Balme and C. kieseri Reyre,
and because the amount of equatorial thickening is variable, even in a single species, these characteristics
are best reserved for specific distinction.
It is therefore clear that Circulina (as emended by Klaus) and Corollina should be combined into a single
genus, with Classopollis as a junior synonym. Circulina Malyavkina may not be identical with Circulina
Mal. emend. Klaus (the status of which is also confused). Corollina was monotypic when published,
whereas Circulina was not. Thus, Corollina does not present problems of generic typification and is
therefore chosen to embody all of the common generic characters.
Genus Corollina Malyavkina, 1949 emend.
Corollina Malyavkina 1949.
Circulina Malyavkina (in part), 1949.
PLATE 3
1
2
3
4
5
6
7
8
Verrucosisporites cheneyi n. sp. (Holotype). Locality 1, 73 um.
V. cheneyi n. sp. Locality 1, 85 um wide.
Harrisispora sp. 1. Locality 11, 80 um.
Dictyophyllidites paramuensteri n. sp. Locality 1, 65 um wide.
D. paramuensteri n. sp. (Holotype). Locality 1, 63 um.
Deltoidospora hallii Miner. Locality 11, 27 um.
Spore A. Locality 11, 98 um.
Dicryophyllidites paramuensteri n. sp. Locality 1, 53 um.
PLATE 4
1 Platysaccus ccf. P. lopsiensis (Malyavkina) Pocock. Locality 11, 58 um.
2 Pityopollenites pallidus Reissinger. Locality 5, 22 um.
3 P. pallidus. Locality 5, 26 um.
4 Callialasporites trilobatus ( Balme) Sukh Dev. Locality 11, 90 um.
5 Alisporites thomasii (Couper) Nilsson. Locality 5, 58 um.
6 A. thomasii. Locality 11, 63 um.
7 A. thomasii, Locality 11, 73 um.
8 Circulina simplex Malyavkina. Locality I, 25 um.
9 C. simplex Malyavkina (Neotype). Locality indicated under description, about 22 um.
10 Callialasporites cf. C. dampieri ( Balme) Sukh Dev. Locality 11, 75 um.
11 Pilasporites allenii Batten. Locality 8, 44 um.
12 Perinopollenites elatoides Couper. Locality 11, 41 um.
13 P. elatoides. Locality 11, 44 um.
14 Podocarpidites sp. 1. Locality 1, 58 um.
Classopollis Pflug 1953.
Classopollis Pflug emend. Couper 1958.
Corollina Mal. emend. Klaus 1960.
Circulina Mel. emend. Klaus 1960.
Classopollis Pflug emend. Pocock & Jansonius 1962.
Monilapollis Chang 1963.
Pagiophyllumpollenites Chang 1963.
Classopollis Pflug emend. Reyre 1969. [Also the species listed below referred to "Pollenites" by Thiergart
and by Reissinger.]
Generic type: Corollina compacta Malyavkina, 1949, p. 124, Pl. 46, fig. 10. (Liassic).
Holotype: Malyavkina, 1949, P1. 46, fig. 10, 11.
Neotype: Plate 5, fig. 10. Designated by Zauer, photograph supplied by Samoilovich, 1973.
Stratigraphic position and locality of neotype: Borehole Kozhegali K36, 466.8 m-471.9 m, Lower
Jurassic, 80 km west of Sagiz near Caspian Sea.
Species assigned to genus: Circulina funifera Malyavkina, 1949, p. 123, P1. 46, fig. 6; BennettiteaePollenites reclusus Thiergart, 1949, p. 11, P1. 2, fig. 14, 15, P1. 3, fig. 6; Pollenites torosus Reissinger,
1950, p. 115, P1. 14, fig. 20.
Emended generic description: Pollen grains distally pseudoporate, proximally trilete, spherical to ovoid,
or compressed, acorn shaped; occur as tetrads or isolated grains; exine two layered, with distinct endexine
and ektexine. Endexine comprises a spherical, internally separate body, which frequently may be
poorly developed and difficult to distinguish, or well developed, sometimes occurring as separate bodies
from rupnued grains. Ektexine (modified from Reyre, 1969) composition variable on different parts of the
grain; composed of inner structured layer, comprising inuastructure, and outer tegillum, which is
continuous over entire grain and covered with outer sculpture uniformly distributed over whole surface. Intrastructure massive, alveolate, punctate, vermiculate or pseudovermiculate, reticulate or
pseudoreticulate, but can be heterogeneously organized: reduced, or thickened, in parts of the grain;
reduced at distal pole (pseudopore) or only along circular line surrounding it, reduced along a subequatorial
furrow (ring tenuitas or rimula) on distal side of equator, and sometimes reduced at proximal pole
(triangular area). Intrastructure generally thidcened in equatorial zone on proximal side of furrow or
rimula, but never thinner than ektexine proximai to equatorial zone or belt; intrastructurai elements
within this equatorial zone may or may not be organized into more or less continuous circumequatorial
striations; they are absent if intrastructure of this zone is massive.
Remarks: Endexinal body may or may not possess a poorly developed trilete mark; it infrequently
possesses a distal area of thinning or tenuitas. The neotype of Circulina simplex Malyavkina (our P1. 4,
fig. 9) and illustrations by Malyavkina (Pl. 46, fig. 7-3) strongly resemble separated endexinal bodies
of Corollina, and therefore do not possess sufficient characteristics to place C. simplex in Corollina.
Circulina funifera Mal. (Pl. 46, fig. 6), on the other hand, appears to possess a subequatorial furrow
(border sharply differentiated from the body, narrow, double), and exinal intrastructure (border covered
with fine, radially directed, straight lines), which possibly place it in Corollina.
Corollina torosus (Reissinger 1950) Klaus 1960 emend.
Plate 5, figures 2-3, 14
Pollenites torosus Reissinger, 1950, p. 115, Pl. 14, fig. 20.
Classopollis classoides Pflug, 1953, Fig. 4, J-M, P1. 16, fig. 20-25, 229-37; emend. Pocock & Jansonius,
1961, p. 443, P1. 1, fig. 1-3.
Classopollis cf. C. torosus (Reissinger) Balme, 1957, p. 37, Pl. 11, fig. 114-119.
C. torosus (Reissinger) Couper, 1958, p. 156, Pl. 28, fig. 2-7; Nilsson, 1958, p. 74, Pl. 7, fig. 68.
C. torosus (Reissinger) Balme; Chaloner, 1962, p. 13, fig. 2, Pl. 11, fig. 1, 2; emend. Burger, 1965,
p. 65, P1. 1, fig. 1, 2.
Monilapollis hsui Chang, 1963, p. 435, 439, P1. 1, fig. la-lc, 2a-2c.
"Classopollis torosus" Pettitt & Chaloner, 1964, P1. 1, fig. 1-5.
Emended description: Isolated pollen grains or occurring as atrads; spherical to ovoid, or acorn-shaped;
diameter in equatorial view (100 grains): 18-41 um (median 32 um), 90% of grains between 24 and 37 um
(Couper, 1958, notes range 24-46 um, mode 32 um). Diameter in polar view 24-37 um (diameter of
subequatorial furrow 83-88% of overall diameter). Distal pseudopore variable in size, even for grains of
same size: 4.3-8.5 um in diameter, frequently distorted and widened in equatorial view. Proximal
triangular area of thinning, which sometimes contains trilete mark, 6.8-13.6 um high. Exine two
layered; ektexine divisible into outer tegillum, continuous over whole surface of grain, and inner structured layer, which is thickened in equatorial region to form a band or belt. Endexine variably distinct
to indistinct, spherical, frequently scabrate, occasionally with a small proximal trilete mark. Structure
of equatorial band ranges from pseudoreticulate, to vaguely striate with aligned intrastructural columellae
(pseudostriations), to distinctly striate, with columellae fused to form more or less parallel bands,
the number of striations frequently varying from one part of band to another. Sometimes combinations of
grains with all degrees of striation development occur in same tetrad. When striations are well developed,
maximum number per grain ranges from 6 to 10. Width of equatorial band, which frequently ranges from 5
um to 13 um at its maximum, usually decreases in one area to 4-10 um, and striations decrease in number
to 4 or 5 through anastomosis or disintegration of more proximal striations into discrete intrastructural
elements. Exine thickness varies with grain size, and presumably also with preservation; it is thickest in
equatorial band: 1.3-2.4 um (Couper, 1958, records 3.0 um), decreasing gradually toward proximal pole:
for example, 2.l um down to 0.9 um; thickness on distal side of equaterial furrow (range 0.7-1.7um)
generally slightly less than that proximal to equatorial band. Intrastructure of ektexine mostly
pseudoreticulate, with positive elements largest on proximal side of equatorial furrow, particularly if
equatorial band is pseudoreticulate; pseudoreticulum on distal side of grain usually finer in pattern,
occasionally becoming punctate. Sculpture of tegillum variable (perhaps owing to preservation); light
microscope indicates roughened surface; S.E.M. indicates microscabrate sculpture, mixed with
microverrucae and microconi (Pl. 5, fig. 14; see also Pettitt and Chaloner, 1964, p. 614); sculpture
distinctly less well developed than in species figured by Reyre (1969), perhaps less well developed than in
forms described under Classopollis torosus (Reissinger) Couper emend. Burger 1965.
Remarks: The emended description of Classopollis classoides Pflug by Pocock and Jansonius (1961) is not
adequate and conflicts with our present knowledge of the exine structure of this genus. We find no
distinction between C. classoides and C. torosus, these species agreeing in exinal structure and variation in
striation development (Pocock and Jansonius, 1961, Pl. 1, fig. 1-9). The size ranges overlap widely, but the
mean equatorial width for C. classoides (29 um) is somewhat smaller than that of C. torosus (32 um).
However, Harris (1957) noted a still smaller mean in his samples (25 um) for grains placed by Pettitt and
Chaloner (1964) in C. torosus. These differences may result in part from preservational states and
preparation techniques. The mean diameter for C. torosus from Locality 11 is 32 um, while it is
25 um from Locality 5. Table 1 gives data for relative abundance of C. torosus that suggest the
existence of two separate "Circumpolles" populations, intermixing at Locality 5. The morphology of these
two populations does not permit their distinction in a mixed sample, as the ranges of variation widely
overlap. Therefore, only one form-species is described, although two different source plants may be
indicated.
Affinity: Hirmerella muensteri (Schenk) Jung 1968, possibly Pagiophyllum connivens Kendall 1948.
Localities: Portland Fm. 5, 6, 7, 9, 10, 11 (dominant); Shuttle Meadow Fm. 1, 3, 4, 12 (rare to scarce);
upper New Haven Arkose, 8 (scarce).
Corollina itunensis (Pocock 1962) comb. nov.
Plate 5, figure 1
Classopollis itunensis Pocock, 1962, p. 71, Pl. 11, fig. 176, 177; P1. 12, fig. 178 (Basionym).
Description: See Pocock (1970, p. 104).
Table 1. Approximate percentages of major Corollina species in Hartford Basin (Based on Corollina counts
only. Most of palynoflorules counted were at least 90% Corollina.)
Localities arranged in stratigraphic order, youngest at right: N.H.A. = New Haven Arkose; S.M.F. = Shuttle
Meadow Formation; P.F. = Portland Formation; + = present, but less than 1%.
STRATIGRAPHIC POSITION
INTRASTRUCTURAL
TYPE
Corollina
SPECIES
N.H.A.
95%
2.5%
2.5%
S.M.F.
97%
1%
+
No. 12
97%
+
3%
No. 5
2%
4%
58%
No. 11
1%
+
23%
36%
76%
C. meyeriana
C. murphyi n. sp.
C. torosus
C. torosus
massive to micropunctate
vermiculate (non-striate)
pseudoreticulate to
punctate (non-striate)
pseudoreticulate to
punctate (pseudostriatestriate)
Remarks: Our specimens compare very closely in structure to C. torosus, but differ significantly in size
and color or transparency. They are very light in color, pale yellow. Equatorial-view diameter (16
specimens) from 44 um-57 um (median 48 um); depth from pole to pole 36-53 um. These specimens
appear distinct from C. torosus, because they have been found as tetrads and in a fragment of a pollen mass
that contained only grains of the above description. The lighter color is not the result of thinning of the
exine: endexine 1.3 um thick; ektexine 0.3-0.4 um proximally, 1.3 um equatorially, and 0.9 um distally.
This species is common to the Canadian Jurassic, first occurring in zone J1 3 of Pocock (1970), which is
equivalent to the upper part of the Lower Jurassic.
Locality: Portland Fm., 11 (rare).
Corollina meyeriana (Klaus 1960) Venkatachala & Goçzán 1964
Plate 6, figures 3-9
Circulina meyeriana Klaus, 1960, p, 165, P1. 36, fig. 57-60.
Corollina meyeriana (Klaus) Venkatachala & Goçzán, 1964, p. 217, Pl. 3, fig. 1-15.
Gliscopollis meyeriana (Klaus) Venkatachala, 1966, p. 93.
Description: Isolated pollen grains or occurring as tetrads; spherical to ovoid, or acorn shaped,
occasionally subtriangular in polar view; diameter in polar view (50 specimens): 20-31 um (median 26 um)
(Klaus recorded "33 um (35 um) 40 um"); diameter in equatorial view (12 specimens): 22-28 um (median
26 um); distal pseudopore diameter 7-12 um; subequatorial furrow present; height of proximal triangular
area of thinning 7-12 um; trilete radii each about 1.7 um long. Exine usually single layered, although
occasionally an endexine body is present; exine 0.4-1.6 um thick, sometimes thickened in equatorial region
(1.0-1.6 um thick), gradually thinning toward proximal pole; intrasaucture massive to micropunctate,
frequently punctate if equatorial region thickened; sculpture roughened to microscabrate.
Remarks: The size range given by Klaus is much larger than and does not overlap with, the
grains from the Hartford Basin. However, de Jersey (1971) noted a size range for this species in the
Ripley Road Sandstone and Helidon Sandstone of Australia of "21 um to 39 um in equatorial diameter."
Occasionally proximal or distal parts are found separately, suggesting separation along the subequatorial
furrow, but this does not necessarily indicate the operculate condition proposed by Venkatachala and
Goçzán (1966). Kendall (1943) gives size range for pollen associated with Brachyphyllum scotii
Kendall as 30-40 um in diameter.
Affinity: Brachyphyllum scotii Kendall 1949. See Barnard, 1968, p. 175, P1. 1, fig. 3, 6, for further
information on pollen associated with B. scotii.
Localities: Portland Fm., 5, 6, 7, 9, 10, 11 (rare to common); Shuttle Meadow Fm., 1, 3, 4, 12 (dominant);
upper New Haven Arkose, 8 (dominant).
Corollina murphyi n. sp.
Plate 5, figure 11-13, 1T, 16
Holotype: Plate 5, fig. 11, size 30 um; Slide PL9, coordinates 37.7 x 109.6, ref. 52.4 x 111.3.
Description: Pollen grains, spherical to ovoid, occasionally subtriangular in polar view; diameter in
polar view (30 specimens) 28-42 um (median 34 um); diameter in equatorial view (6 specimens): 30-37
um; distal pseudopore diameter 6.5-9.5 um; subequatorial furrow present. Width of proximal area of
thinning 5.2-10.6 um: area not always observable, rarely triangular, frequently irregular in shape. Exine two
layered, 0.9-3.4 um thick; endexine distinct to indistinct, spherical, psilate, thin (0.3 um thick), frequently
folded or missing in ruptured grains; ektexine 1.8-3.4 um thick in equatorial region, without striations to
possessing 1 or 2 irregularly widened, imperfect circumequatorial bands forming a belt about 4.5 um wide;
intrastructure vermiculate to imperfectly pseudoreticulate, formed by broad irregular islands of structural
thickening, which are separated by narrow anastomosing lines of exinal thinning; sculpture indistinct to
microscabrate.
Remarks: Intrastructural elements broader and mote massive than in C. torosus. Species named after
Sister Mary Ellen Murphy, Department of Chemistry, Saint Joseph College, West Hartford, Conn., who
is contributing important organic geochemical evidence for sediments of the Hartford Basin.
Localities: Portland Fm., 5, 11 (scarce to common); Shuttle Meadow Fm., 1, 12 (scarce); upper
New Haven Arkose, 8 (scarce).
Corollina simplex (Danzé-Corsin & Laveine 1963) comb. nov.
Plate 6, figure 10
Basionym: Classopollenites simplex Danzé-Corsin & Laveine 1963
Classopollenites simplex Danzé -Corsin & Laveine, 1963, p. 106, P1. 11, fig. 7, 8.
Classopollis simplex de Jersey & Paten, 1964, p. 12, P1. 7, fig. 4-6, non Danzé-Corsin & Laveine 1963.
C. simplex (Danzé -Corsin & Laveine) Reiser & Williams, 1969, p. 16, P1. 6, fig. 15.
Excluded (homonymous) species: Classopollis simplex Reyre, 1969, p. 312, P1. 54, fig. 3-5.
Description: See Reiser and Williams( 1969, p. 16).
Remarks: Specimens from the Hartford Basin range in polar view diameter from 17 um-28 um, which
agrees well with the range given by de Jersey and Paten (1964). However, Danzé-Corsin and Laveine
(1963) reported a range of 20-40 um. The generic name Classopollenites Danzé-Corsin & Laveine 1363
is a superfluous substitute for Classopollis Pflug 1953, and is, hence, illegitimate. Furthermore,
Classopollis simplex de Jersey & Paten 1964, although validly published, is a homonym, and is also
conspecific with C. simplex Danzé-Corsin & Laveine. Classopollis simplex Reyre is distinctly different and
is therefore an illegitimate homonym.
It should be noted that Corollina simplex is probably not a natural species, since it has been found as
anomalous grains in tetrads of both C. meyeriana and C. torosus. C. simplex grades into forms of C.
meyeriana with a slightly thickened equatorial band, as well as into small compact forms of C. torosus.
However, this does not preclude use of the concept of C. simplex.
Localities: Portland Fm., 5, 7, 9, 11 (rare); Shuttle Meadow Fm., 1, 3, 12 (rare to scarce); upper New
Haven Arkose, 8 (rare).
Corollinoid? pollen
Plate 6, figures 11, 12
Description: Two unusual palynomorphs resembling Corollina possess a circumequatorial saccus or
pseudosaccus. Both grains have p'roximal triangular area of exinal thinning, equatorial furrow of exinal
thinning, and distal pseudopore (although not obvious on specimen in P1. 6, fig. 12). One specimen (Pl. 6,
fig. 11) has massive intrastructure and narrow flange or saccus, the other (Pl. 6, fig. 12) an intrapunctate
structure and expanded, well-developed saccus. These differences suggest that they may not be conspecific.
Saccus or pseudosaccus on each grain appears to be attached at equatorial furrow. Exine equatorially
thickened (about 1.7 um thick), appearing massive in optical section. Size (2 specimens): 42 um in overall
diameter; body 31-34 um in diameter; diameter of pseudopore 9-11 um; height of triangular area of
thinning about 14 um; saccus or pseudosaccus extends 4-6 um beyond perimeter of body.
Remarks: These grains appear to have a close affinity with Corollina.They show different degrees
in development of the saccus or pseudosaccus, suggesting a developmental trend. The presence of this
circumequatorial feature is striking if these grains were derived from the "Circumpolles." More specimens of these forms in these or other sediments are needed to permit a more complete study.
Locality: Portland Fm., 11 (rare).
Genus Circulina Malyavkina 1943 emend.
Type species: Circulina simplex Malyavkina 1949
Circulina Malyavkina (in part), 1949.
Species assigned to genus: Circulina simplex Malyavkina, 1949, p. 224, P1. 46, fig. 7-9.
Excluded generic concept: Circulina Malyavkina emend. Klaus, 1960.
Excluded species: Circulina funifera Malyavkina, 1949, p. 123, 124, P1. 46, fig. 6.
Emended description: Pollen grains inaperturate, oval to circular, scabrate, punctate, or smooth. Exine
single layered, lacking internal structure of Corollina.
Remarks: This genus is intended for small inapertutate grains that do not have the germinal structures
observed in Spheripollenites Couper 1958 or Exesipollenites Balme 1957, and that may be isolated
endexinal bodies of Corollina Malyavkina 1943 emend.
Circulina simplex Malyavkina 1943 emend.
Plate 4, figures 8, 9
Holotype: Malyavkina (1949, P1. 46, fig. 7-9).
Neotype: Plate 4, fig. 3, in this paper, a specimen earlier designated by Zauer, photograph supplied by
S. Samoilovich, 1973 (for geographic and stratigraphic location, see Corollina compacta neotype, in
earlier discussion of "Circulina-Corollina-Classopollis Nomenclatural Problem").
Emended description: Pollen grains inaperturate, oval to circular, punctate to smooth, occasionally
scabrate. Exine single layered, sometimes folded to give the impression of an eccentric double wall. Exine
about 0.7 um thick. Diameter of grains about 30 um.
Localities: Portland Fm., 5, 7, 9, 10, 11, 13 (rare); Shuttle Meadow Fm., 1, 3, 12 (rare); upper New Haven
Arkose, 8 (rare).
Genus Camerosporites Leschik 1956 emend. Scheuring 1970
Type species: Camerosporites secatus Leschik 1956
Camerosporites reductiverrucatus n. sp.
Plate 7, figures 1-5
Holotype: Plate 7, fig. 1, size 54 um; Slide AGA1-22, coordinates 12.6 x 111.4, ref. 56.7 x 110.8.
Description: Pollen grains, spherical to ovoid, or acorn shaped, occasionally present as tetrads.
Triradiate fold sometimes present at proximal pole. Equatorial furrow or ring tenuitas occasionally well
developed; proximal part of grain often found without distal part. No distal tenuitas observed. Grains
frequently folded. Exine of proximal part of grain slightly thicker than distal part;·proximal part generally
sculptured with low, broad verrucae, more closely spaced around equator, forming equaterial band 4-12 um
wide (mode 8.5 um). Distal part of grain usually psilate, but occasionally possesses a few scattered low
vetrucae arranged around distal pole. Sculpture of proximal part of grain varies considerably from an
abundance of low verrucae, simulating an equatorial band, to almost no sculpture, with a few verrucae
restricted to equatorial region. Exine single-layered?; exine of distal part of grain 0.8-0.9 um thick,
occasionally intrascabrate; exine of proximal part of grain 0.8-1.2 um thick. Verrucae 2.5-4.5 um wide,
irregular in shape, about 0.4 um in height. Size range (50 specimens): 39-70 um in overall diameter
(median 56 um); 90% of grains between 48 um and 65 um.
Remarks: This species is larger than other species of this genus reported from the Keuper. Our highly
sculptured forms resemble Camerosporites secatus Leschik, but lack the thick verrucae of this species.
Other sparsely sculptured forms show a tendency of our species to lose the characteristic sculpture of
the genus, and these would probably not be placed in Camerosporites if they did not intergrade with
highly sculptured forms. This species suggests that at least one source plant for Camerosporites may have
persisted into the Jurassic.
Locality: Portland Fm., 11 (rare).
Genus Cycadopites Wodehouse 1933 ex Wilson & Webster 1946
Type species: Cycadopites follicularis Wilson & Webster 1946
Cycadopites Wodehouse 1933.
Entylissa Naumova 1933.
Cycadopites Wodehouse ex Wilson & Webster 1946.
Monosulcites Cookson 1947.
Ginkgocycadophytus Samoilovich 1953.
Ginkgoretectina Malyavkina 1953.
Monosulcites Cookson ex Couper 1953.
Entylissa Naumova ex Potonié & Kremp 1954.
Cycadopollenites Danzé-Corsin & Laveine 1963.
Ginkoretectinapollenites Danzé-Corsin & Laveine 1963.
Remarks: For further synonymy, see Jansonius (1962, p. 80). Potonié (1958) separated many of the genera
in the proposed synonymy by the shape of the sulcus or by the degree of overlap of sulcus
margins. Subsequent authors have placed many of these genera in synonymy, but they have continued
to separate Monosulcites and Cycadopites on the basis of sulcus shape. However, variation in sulcus shape
and margin overlap probably reflect differences in orientation and compression during fossilization, as
well as variation in the original shape of the grains. Features of the sulcus should therefore be used only
as an aid in distinguishing species, but caution should be exercised in assigning grains to different genera
on this basis alone.
Cycadopites andrewsii n, sp.
Text-figure 4; Plate 8, figures 17-21
Holotype: Plate 8, fig. 20; size 37 um long, 25 um wide; Slide CLRZ-1, coordinates 34.4 x 120.9, ref.
26.1 x 121.5.
Description: Fusiform to ovoid pollen grains, monosulcate; longitudinal ends rounded to bluntly pointed;
sulcus narrowest at distai pole, usually widening toward longitudinal extremities. Exine distally smooth,
proximally sculptured with closely spaced verrucae, which form a very distinct pseudoreticulum. Verrucae
0.3 um x 0,3 um to 1.7 um x 2.6um wide, broadest at proximal pole, decreasing in diameter, to a lesser
degree in height, toward extremities and equatorial margin. "Lumina" of pseudoreticulum decrease in size
away from proximal pole, usually
TEXT-FIGURE 4. Diagram of polar cross-section of pollen grains of Cycadopites andrewsii n. sp.,
showing relationship of ektexine and endexine and distribution of sculpture.
disappearing at, or slightly distal to, equatorial margin. Exine double layered, ektexine (exclusive of
sculpture) about 0.3-0.4 um thick, sculpture 0.1-0.4 um high, endexine about 0.2 um thick. Endexine
apparently loosely attached except at sulcus margins, sometimes separating, in part or completely, as a
distinct inner envelope. Lips of sulcus conform to area of apparent ektexinal and endexinal fusion; lips
widest at distal pole (1.7-2.6 um wide), tapering to extinction at longitudinal extremities. Size range (30
specimens): length 34-46 um (median 38 um); width 20-30 um (median 22 um). Ratio of width to length
1:1.2-1:2.4 (median 1:1.7).
Remarks: Several monosulcate species described in the literature compare with C. andrewsii n. sp., but
apparently have an intrareticulum rather than an external pseudoreticulum; for example, Monosulcites
rhaetoliassicus Madler 1964 (Rhaeto-Liassic), Cycadopites reticulatus (Nilsson 1958) comb. nov. (Lower
Jurassic), Ginkgoretectina couperi (Pocock 1962) Pocock 1970 (Upper Jurassic), and G. ferrei Pocock 1970
(Upper Jurassic).
Species named after Professor Henry N. Andrews, paleobotanist, University of Connecticut, Storrs.
Locality: upper New Haven Arkose, 8 (rare to scarce).
Cycadopites deterius (Balme 1957) Pocock 1970
Plate 8, figures 10-12
Entylissa deterius Balme, 1957, P· 29, Pl. 6, fig. 75-77.
Cycadopites gracilis Sah & Jain, 1965, p. 282, P1. 7, fig. 146-149.
Description: Pollen grains monosulcate, elliptical. Sulcus frequently widest at extremities, narrower at
distal pole. Sulcus margins sometimes parallel one another for much of grain length, occasionally overlap
to a lesser or greater degree. Lips generally absent. Exine single layered, 0.5-0.9 um thick, frequently
slightly thicker at longitudinal extremities. Sculpture usually psilate distally, scabrate to psilate proximally,
with larger grana frequently concentrated on proximal side at extremities. Grain color ranges from dark to
light, depending on state of preservation. Size range (23 specimens): 30-60 um long (median 50 um); 14-35
um wide (median 24 um). Ratio of width to length 1:1.5-1:3.0 (median 1:2.2).
Remarks: The size range of our specimens is slightly larger than that given by Balme (1957) for Entylissa
deterius (41-60 um long), but the average size is the same. The sculpture on E. deterius is psilate, ranging to
scabrate on some of our specimens.
Cycadopites gracilis is included in the synonymy because it contains a combination of features found
in our specimens: scabrate sculpture, overlapping sulcus margins, and a similar, but slightly smaller,
size range (length 32-44 um). Together, both E. deterius and C. gracilis cover the range of variations
found in our specimens, but cannot be distinguished in out samples as separate organ species.
Cycadopites carpentieri (Delcourt & Sprumont) Singh 1964, per the description in Couper (1958),
compares very well to this species in overall shape, exine thickness, and features of the sulcus. Although
the size range of C. carpentieri overlaps that of C. deterius, the median size of C. carpentieri is about 10 um
larger, suggesting that Couper's specimens are a distinct form. The grains illustrated by Delcourt &
Sprumont (1955) show that the specimens upon which C. carpentieri was based may have had a
significantly thicker exine than our specimens.
Specimens from the Shuttle Meadow Fm. Are slightly more fusiform (median ratio of width to length
1:2.6) than those from the Portland Fm. and fall into the lower part of the size range of specimens
from the Portland: length 31-46 um (median 40 um); width 14-19 um (median 17.0 um). The sulcus
of Shuttle Meadow specimens trends to be more linear in shape, sometimes gaping only at one end, or
rarely at the distal pole.
Localities: Portland Fm., 5, 7, 10, 11, 13 (rare to scarce); Shuttle Meadow Fm., 1 (rare).
Cycadopites durhamensis n. sp.
Plate 8, figures 13-15
Holotype: Plate 8, fig. 13, size 26 um long, 12 um wide; Slide LTG-1, coordinates 40.0 x 124.0, ref.
50.7 x 125.7.
Description: Pollen grains monosulcate, fusiform or elliptical. In polar comptessions longitudinal extremities generally bluntly pointed to rounded; in lateral compressions ends sometimes pointed, nipple
like. Sulcus generally straight, margins frequently overlapping for much of the length of grain.
Occasionally one end of sulcus expanded and open; rarely both ends of sulcus expanded on same grain.
Infrequently grains ate compressed, with sulcus gaping widely at distal pole: such grains frequently show
rents. Lips generally not developed, but occasionally sulcus margins at distal pole thickened. Exine single
layered, 0.5-0.7 um thick, possibly slightly intrascabrate. Sculpture psilate. Size range (21 specimens):
22-31 um long (median 27 um); 9-16 um wide (median 13 um). Ratio of width to length 1:1.8-1:3.0
(median 1:2.1).
Remarks: This species ranks among the smallest of monosulcates reported from the Mesozoic. It
differs from Cycadopites minimus (Cookson) Pocock 1970 per description of Couper (1958, p. 157, Pl.
26, fig. 23-25), in being slightly smaller (smaller average size), and in having a more fusiform outline
(ratio of width to length 1:2.1 rather than 1:1.4 as in Couper's specimens). The tendency of the sulcus
margins to overlap in our specimens also serves to distinguish C. durhamensis from C. minimus.
Our specimens compare well in shape and size with the Rhaeto-Liassic Ginkoretectinapollenites
punctata (Malyavkina) Danzé-Corsin & Laveine (1963, p. 109, Pl. 11, fig. 17) but apparently differ
significantly in exine thickness,
Species named after the geographic locality, Durham, Conn.
Affinity: Perhaps Otozamites latior Saporta, which is the most abundant species and only genus of
cycadophyte found in the black, microlaminated micritic limestone at Locality 1 (recovered by first
author) and Locality 1 (Newberry, 1888, p. 90). C. durhamensis n. sp. is by far the most abundant
monosulcate pollen grain at Locality 1.
Localities: Shuttle Meadow Fm., 1, 12 (rare).
Cycadopites cf. C. jansonii Pocock 1370
Plate 8, figures 7-9
Monosulcites "sp. K" Jansonius, 1962, p. 79, P1. 16, fig. 28-30.
Cycadopites jansonii Pocock, 1970, p. 109, P1. 26, fig. 10, 15.
C. paruvus Bolkhovitina 1953, p. 63, P1. 10, fig. 5, 6 (in part) ex Pocock, 1970, p. 109, P1. 26, fig. 11.
Description: Pollen grains monosulcate, elliptical or fusiform. Sulcus frequently widest at extremities,
narrower at distal pole. Sulcus bordered by a pair of longitudinal lips extending full length of the grain.
Lips widest (2.5-7.8 um wide) at distal pole. One lip of a pair frequently wider than the other. Exine double
layered, 0.5-0.7 um thick; ektexine about 0.4 um thick; endexine about 0.2 um thick, occasionally
separating in part as an internal envelope. Length (6 specimens) 37-48 um; width 27-29 um. Ratio of width
to length 1:1.6-1:2.7.
Remarks: Our specimens agree with the type description in grain size (length 33-54 um; width
23-36 um, Pocock, 1970) and in the form of the lips. Our specimens are referred to C. jansonii with "cf."
because of difference in exine thickness (the exine of C. jansonii is about 1.5 um thick). Ginkgo parva
Bolkhovitina 1953, part of which species is an apparently identical form, was not validly published because
of the listing of an alternative name. Pocock (1970), intending to make a new combination, Cycadopites
parvus, for part of Bolkhovitina's species, actually validated the name C. parvus.
Locality: Portland Fm., 5 (rare).
Cycadopites reticulatus (Nilsson 1958) comb. nov.
Plate 8, figures 1-3
Basionym: Entylissa reticulata Nilsson, 1958, p. 62, Pl. 5, fig. 14.
Description: Pollen grains monosulcate, elliptical. Sulcus may or may not be expanded at longitudinal
extremities, but never at distal pole. Frequently grains collapse, with one margin ofsulcus overlapping
the other. Sulcus sometimes bordered by very narrow lips. Longitudinal ends of grain generally rounded,
sometimes pointed. Exine structured with an intrareticulum that has its greatest development proximally,
decreasing in size equatorially, distally, and toward the extremities, Proximai surface of grain frequently
has an irregular surface, reflecting thickening of the exine in the reticulum. Lumina of reticulum 0.2-2.2 um
in diameter. Exine 0.6-l.0 um thick, thicker proximally. Sculpture psilate distally, generally irregular
proximally because of reticulum. Size range (6 specimens): 47-60 um long, 22-32 um wide. Ratio of width
to length 1:1.7-1:2.3.
Localities: Portland Fm., 5, 11, (rare).
Cycadopites westfieldicus n. sp.
Plate 8, figures 4-6
Holotype: Plate 8, fig. 4, size 60 um long, 29 um wide; Slide AGA1-23, coordinates 48.7 x 124.2,
ref. 56.8 x 126.4.
Description: Pollen grains monosulcate, elliptical. Sulcus frequently widest at extremities, narrow at
distal pole. Sulcus margins occasionally overlap at distal pole. Longitudinal ends of grain rounded, rarely pointed. Lips well developed, thin, occasionally difficult to observe on corroded grains. Lips widest
at distal pole (5-11 um wide), tapering toward longitudinal extremities. Lips of a pair frequently unequal
in width. Exine double layered, about 1.3 um thick; ektexine 0.8-0.9 um thick; endexine 0.4-0.5 um thick.
Sculpture psilate. Size range (21 specimens): 46-63 um long (median 56 um); 22-41 um wide (median 29
um). Ratio of width to length 1:1.4-1:2.4 (median 1:1.8).
Remarks: This species is larger than Cycadopites jansonii Pocock, but has an exine of similar thickness.
Species named after the Westfield River, site of Locality 11.
Locality: Portland Fm., 11 (rare).
Cycadopites sp. 1
Plate 8, figure 16
Description: Pollen grains monosulcate, elliptical. Sulcus open at longitudinal extremities of grain,
margins narrower but basically straight between extremities. Lips absent. Exine (exclusive of sculpture)
about 2.6 um at equatorial margin, single layered. Sculpture consists of scattered large verrucae and coni,
with some bacula, primarily concentrated around equatorial sides of grain and, to a very minor extent,
on proximal and distal faces. Sculptural elements 0.7-1.2 um high, 0.8-2.6 um wide. Size (2 specimens):
45-46 um long, 26-29 um wide. Ratio of width to length 1:1.6-1:1.7.
Locality: upper New Haven Arkose, 8 (rare).
PL ATE 5
1
2
3
4
5
6
Corollina itunensis (Pocock) comb. nov. Locality 11, 53 um.
Corollina torosus (Reissinger) Klaus, emend. Locality 11, 34 um-36 um.
C. torosus (Reissinger) Klaus, emend. Locality 11, 32 um-38 um.
C. torosus (Reissinger> Klaus, emend. Locality 11, 31um-34 um.
C. torosus (Reissinger) Klaus, emend. Locality 5, 28 um-30 um.
C. torosus (Reissinger) Klaus, emend. Locality 11, 22 um-25 um.
7
8
9
10
11
12
13
14
15
16
C. torosus (Reissinger) Klaus, emend. Locality 11, 30 um.
C. torosus (Reissinger) Klaus, emend. Locality 11, 37 um.
C. torosus (Reissinger) Klaus, emend. Locality 11, endexinal body, 34 um (under oil).
Corollina compacta Malyavkina (Neotype). Locality indicated under description, about 33 um.
Corollina murphyi n. sp, (Holotype). Locality 5, 30 um.
C. murphyi n. sp. Locality 8, 34 um.
C. murphyi n. sp. Locality 8, 30 um.
Corollina torosus (Reissinger) Klaus, emend. Locality 11; S.E.M., 28 um-33 um.
Corollina murphyi n. sp. Locality 8, 32 um.
C. murphyi n. sp. Locality 11; polar view with endexinal body, 42 um.
PLATE 6
1 Brachyphyllum Icotii Kendall. Locality on Licking
Run, near Midland, Va.; cuticle of leaf showing stomata; field about 250~Lm across.
2 B. stotii Kendall. Locality on Licking Run, near
Midland, Va.; cuticle of leaf; field about 850CLn
aCfOSS.
3 Corollina meyeriana (Klaus) Venkatachala & Gdcza'n.
Locality 8, 26cLm.
4 C. nzeyeriana. Locality 8, 29CLm.
5· C, meyeriana. Locality 8; S.E.M., 23CLm.
6
3
10
11
12
13
14
15
C. meyeriana. Locality on Licking Run, near Midland,
Va.; each grain about 54CLm.
7 C. meyeriana. Locality 1, 26~Lm.
8 C. meyeriana. Locality 1, 24~rrr.
C. meyeriana. Locality 1; S.E.M., I~um.
Corollz`na st'nrplex (Danz~-Corsin & Laveine) comb.
nov. Locality 11, 24CLm.
Corollinoid! pollen. Locality 11, 42~Lm.
Corollinoid! pollen. Locality 11, 42c~m.
Ar-aucariaL-ires australiJ Cookson. Locality 11, 71Clm.
At·alccc?ri*cites prrtzc-tatrrs (Nilsson) comb. nov. Locality
11, S~CLm·
A. punctattls (Nilsson) comb. nov. Locality 5, 48CLm·
DISCUSSION
Stratigraphic Correlatio~n
Text-figure 5 summarites the stratigraphy of the
Hartford Basin as correlated, by this study and
other information, with the upper part of the Newark
Basin sediments.
PalynoAoras dominated by Corollina (exceeding
70% of assemblage) have been reported from the
Rhaeto-Liassic and Upper Jurassic (Kendall, 1343;
Harris, 1357; Couper, 1918; Lantz, 1358a,b; Pocock
and Jansonius, 1961; Chaloner, 1962; Venkatachala
and Gdna'n, 1964; Wall, 1965; Burger, 1965; Reyre,
1969; Volkheimer, 1371). Floras composed of more
than 30% Corollina meyeriana have thus far been
reported only from the basal Liassic of Larachbeg,
Morvern, Scotland (Kendali, 1943). However, floras
composed of a complex of Corollina species, totaling
more than 300Jo, have been reported from the
Rhaetian Ktissen facies, Hungary (Venkatachala and
Gdcza'n, 1964). This complex contains both equaterially striate and non-striate forms, C. classoides and
C. meyeria·t;ta. Palynofloras composed of 700/0 or
more Corollina torosus are common in the Liassic
(Harris, 1957; Couper, 1958; Wall, 1965; Volkheimer, 1371), but have also been recorded in undifferentiated Rhaeto-Liassic strata (Chaloner, 1362;
Jung, 15)68). In general, non-striate to pseudostriate
species are most common in the Rhaetian and early
Liassic, while equatorially striate species predominate
in the Liassic (Geiger and Hopping, 1968; Reyre,
1363).
The palynofloras of the Hartford Basin show an
upward change in Corollina from more than 30%
Corollina meyeriana in the Shuttle Meadow Formation to more than 30% Corollina torostls in the
Portland Formation. The age of this shift would lie
somewhere between the oldest reported record of
a flora dominated by Corollina torosus and the
youngest record of a flora dominated by Corollina
meyeriana. Although there is admittedly limited
evidence, probably the shift occurred between the
limits of Rhaetian and basal Liassic. Paleomagnetic
data and isotopic dates (171-136 m.y.) for the
Hartford Basin lava-flow units agree with a RhaetoLiassic age for this shift (Reesman et al., 1373; Reeve
and Helsley, 1372).
The saatigraphic position of the Triassic-Jurassic
boundary in the Hartford Basin might naturally be
placed in that format~on where the greatest change
in Corollina species occurs, thus relegating strata
containing the flora dominated by Corollina meyeriana to the Triassic, and strata dominated by Cnrollina
torosus to the jurassic. It might further be argued
that the Corollina-dominated K~issen facies supports
a Rhaetian age for the Shuttle Meadow Formation.
However, evidence of a basal Liassic Age for the
formation comes from the composition of the nonCorollina portion of the assemblage. The Shuttle
TEXT-FIGURE 5· Suggested time-stratigraphic correlation of
Hardord Basin and upper sediments of the Newark
Basin. A = Grllidlas~orites trilobat~s (Balme) Sukh
Dev; D = Callialasporites cf. dampieri; H = Hirmerella
muenlteri (Schenk)Jung; K = Convolzltis~ora RluRiforma
(Nilsson) Schuiz; M = flora dominated by Corolli~na
meyeriana (Klaus) Venkatachala & Gciczain; S = Brachyphyllzlm scottii Kendall; T = Aora dominated by Corollina torosus (Reissinger) Klaus. RedFeldius is a subholostean fish with a restricted zone in the Newark
Group. Numbers shown for the Newark and Hardord
basins refer to our palyniferous localities in these basins.
The letters defined above indicate presence of the indicared form. M and T generally indicate dominance. Parentheses indicate forms present in variable but minor
amounts.
Meadow Formation contains Conuoluti~pma &IzlRiforma, which is characteristic of the European Liassic,
rarely being found in the Rhaetian, and absent from
the K~issen facies. Conversely, the K~ssen facies contains three species of Oval~olliJ in relatively large
numbers (5-16%). This pollen genus is frequently
found in the Rhaetian, but is very rare to absent in
Liassic paiynofloras. In lacking Oual~olliJ, the Shuttle
Meadow flora is similar to basal Liassic floras.
Kendall (2949) provided only a brief description
of the Larachbeg, Scotland, palyno~ora. However,
she did provide a very detailed description of cuticle
from Brachyphyllum scotii Kendall, associated with
abundant pollen (more than 90%) supposedly derived from that plant. At a locality 2.2 km north of
Midland, Virginia (Culpeper Basin), reported by
Baer and Martin (1349), a similar association of
CorolLina and well-preserved cuticle, including
whole leaves, of B. scotii was recently discovered in
a gray brown shale-siltstone sequence exposed along
a brook, Licking Run. Many palyniferous layers also
occur in the Licking Run section. Fish-bearing black
calcareous shale outcrops suatigraphically about 2.2
m below the megafossil plant bed~ The fish fauna
from this locality is remarkably similar to the one
from the Shuttle Meadow Formation at Locality 1
(Comet, et al., 137~; Schaeffer, 1967; P. E. Olsen,
pers. comm.). A more detailed study of the Midland
flora is in progress.
The Midland flora is essentially identical to that
from the Shuttle Meadow Formation, including the
presence of Conuolzltispora RI~R~forma. The abundant
pollen, including tetrads and pollen masses, associated
with the B. scotii cuticle at Midland, is identical with
Corollina meyeriana (Klaus) Venkatachala and
Gdcza'n. Several distinct species of Dictyot:·'!etes
(a genus characteristic of the Jurassic) are also present. One of these species (Dictyotriletes sp. 1) is
present at Locality 8 in the upper New Haven
Arkose of Massachusetts: (Locality 8 is considered
contemporaneous with part of the Shuttle Meadow
Fm.; Text-fig. 3). Moderately corri~ded but identifiable cuticle, including whole leaves, of B. scotii has
also been recovered from Shuctle Meadow Locality 1
of the Hartford Basin and from lake beds above
the lowest lava-flow unit (First Watchung Mt.),
just east of Round Top Mountain, near Oldwick, New
Jersey (Locality 4, Newark Basin, Text-fig. 5).
The Shuttle Meadow, Midland, and Larachbeg
palynofloras appear to have much in common. The
identification of B. scotii merely indicates that a
basal Liassic age is possible. Only one age horizon
is known for this conifer – thus it is of little timestratigraphic value, especially if it were the only
source for C. meyeriana, which is doubtful. However,
the presence of B. scotii, C. klukiforma, Dictyotriletes
sp. 1, the dominance of C. meyeriana, and the absence
of any palynomorph known to be characteristic of
only the Triassic, favors an early Liassic age for the
Shuttle Meadow and Midland floras.
Todites pinceps (Presl) Gothan was discovered
by the first author in the youngest sediments of
the Culpeper Basin, near Leesburg, Va. This fern
has been found mainly in late Rhaetian to Middle
Jurassic age strata (Harris, 1948, p. 182, 183j.
ClathropteriJ meniscoides Brongniart has been found
at Localities 2 and 8 in the Hardord Basin, and
Locality 4 in the Newark Basin. This fern may
range from Carnian through the Liassic, butis
particularly common in the Rhaetian and lower
Liassic (Harris, 1931)· Both of these ferns were
found in association with a flora dominated by
C. meyeriana, and this strongly indicates a RhaetoLiassic age for this type of association.
In order to examine palynofloras that are slightly
older than those of the Shuttle Meadow Fm. of
Gonnecticut, one must ref~r to strata in the Newark
Basin of New Jersey, because palynomorphs have
not been found in the older Talcott Fm. of Connecticut, nor in strata of the New Haven Arkose known
to be older than the Talcott (Text-fig. ~). Correlation of footprint and fish faunas (P. E. Olsen, pers.
comm.), and of palynofforules and lava flows for the
Hartford and Newark basins (Text-fig. 5), indicates
that some strata in New Jersey containing a typical
Shuttle Meadow palynoflora may be older than the
Shuttle Meadow Formation of Connecticut. The
seventeen fish localities so far investigated by Olsen
from sediments between the First (lowest) and
Third (highest) Watchung lava-flow units lack the
subhoiostean, Redfi~ldizls spp., which is frequently
found in younger strata. Therefore it is apparent
that the flora dominated by C. meyeriana (occurring
with B. scotii) has considerable snatigtaphic range,
calculated in the Newark Basin to be greater than
600 m (not including thickness oflava-flow units).
The presence of Upper Triassic reptiles, such as
StegomuJ, Hypsognathus, and the phytosaur, cf.
Rutiodon (P. E. Olsen and J. H. Osuom, pers.
comm.), in the middle part of the New Haven Arkose (Text-tig. 5) strongly favors placing the TriassicJurassic boundary just above the Talcott Formation
(which at present would not conflict with paleomagnetic data). The boundary might correlate with
the erosional unconformity in the Hartford Basin.
Recently a well-preserved Carno-Norian palynoflorule (Locality 1, Newark Basin, Text-fig. 5) was
obtained from gray shales of the lower Brunswick
Formation at Milford, New Jersey (gray members
L-M of McLaughlin, 1946). The florule is dominated
by bisaccates: AliJporites, Kla~sipolleniteJ, Pityosporites, Protodiploxypinzls, Platysacczls, Triadispora,
Chordasporites?, and PodosporiteJ? There are also
moderate percentages of Camerosporites verrucosus
M~dler, Patinasporites densz~s Leschik, VaLlasporites
ignacii Leschik, and tetrads of spherical psilate grains
("Genus A") only before described by Dunay
(1972) from the Upper Triassic Dodtum Group of
northwest Texas. The stratigraphic position of this
middle Keuper florule from New Jersey is about
920 m above "gray member B" of McLaughlin
(1344), which is the top of the underlying Lockatong
Formation. However, the relationship of the florule
to the overlying First Watchung Mountain basalt is
less certain, as most of the Brunswick Formation,
including the lava flows, is not preserved around
Milford, N.J., and very few "unfaulted" sections of
the entire lower Brunswick have been measured.
Kummel (1836, p. 58) measured one such section
"between the mouth of Lawrence Brook, east of
New Brunswidc, and the base of the first trap ridge
back of Bound Brook" as being 3050 m thid(. Because of the relative proximity of this measured section to Milford, N.J. (in relationship to the rest of
the basin), this figure can be safely used to calculate
the approximate distance of the~Milford florule beneath the First Watchung Mountain lava-flow unit,
a distance of critical importance in the following
argument (6. Text-fig. 5).
Van Houten (1363) estimated the rate of sedimentation fbr the Brunswick Formation at 30~ +mml1000 yr. The Milford florule lies approximately
2100 m below the First Watchung Mountain basalt
(= 3050 m-320 m). The Milford Carno-Norian
florule would then be separated in time from the
oldest known occurrence of a florule dominated by
Corollilza meyerzana (occurring with Brachyphyd
Illm SGOtii) in the Newark Basin by at least 6.3 m.y.
This amount of time would accommodate much of
the Norian and Rhaetian stages and agrees well
with the paleomagnetic normal polarity of the
lava flows, which can readily be explained if most of
the Rhaetian occurred before extrusion of the lavaflow units (Creer, 1371; Reeve and Helsiey, 1972).
For comparison, the maximum thickness of Upper
Triassic strata in England and Germany appears to
be 1400 m, but it is usually less than 500 m
(Geiger and Hopping, 1968; Wills, 1970).
A probable upper Norian palynoflorule (samples
provided by Donald Baird, Princeton Univ.; coll. by
N. K. Resch) occurs in a gray siltstone with reptilean
footprints, exposed in the construction of U.S. Rt. 280
behind the old Second Precinct Police Station at
Newark, N.J. (Newark Basin Locality 11, Textfig. 5). The florule is significant in that it possesses
equatorially striate grains of Corollina torosus
(Reissinger) Klaus, as well as C. meyeriana (Klaus)
Venkatachala 8r Gdcza'n, Patinasporites densuJ
Leschik, possibly Enzonalasporites vigens Leschik and
TriadiJpora sp. C. torosus, generally not reported
below the Rhaetian, and Triadispora sp., rarely reported above the middle Norian, are seldom found
in the same assemblage. In addition the presence
of P. densus suggests an age no younger than
Norian because, while this species occurs as high
as the lower Rhaetian, it is rare above the Norian.
Corollina spp. represent about ~O% of the assemblage. Bisaccates of the Pityosporites and Alisporites types are common, and Converrucosisporites
cameronii (de Jersey) Playford 8r Dettmann and
Granulatisporites infirmus (Balme) comb. nov. are
much more common than in older floruies of the
Newark Group. This palynoflorule is calculated to
be about 1100 m below the First Watchung
Mountain basalt.
The rather low stratigraphic position of middle
and upper Keuper florules in the Brunswick Formation supports the paleomagnetic evidence that at
least part of the Brunswick below the First
Watchung Mountain basalt is Rhaetian, but does
not rule out the possibility that strata containing
the generally Liassic C. ~neyeriana-B. Jcotii flora
may extend downward into the upper Rhaetian.
For this reason this floral assemblage is here designated Rhaeto-Liassic wherever it occurs.
The Portland Formation florules of the Hartford
Basin are generally dominated by C. toroszls (Table 1)
and its presumed megafossil sources, Hirmerella sp.
(Locality 7, Comet et al., 1373) and Hirmerella
nzzlenJteri (Schenk) Jung 1368 (Localities 10, 11,
and 13). Lower Portland flotules (Localities S, 6, 7,
and 13) appear to vary laterally and to fluctuate vertically in Corollirza spp. percentages, with C. meyeraana sometimes comprising much more than 2~0, but
rarely being dominant. poor preservation of most of
these fforules makes specific identification difficult,
but cuticle fragments show that the ratio of
Brachy~hylltlm scotaz to Hirmerella spp. varies in
proportion to the apparent ratio of C. meyeriana
to C. toroJus.
The minor constituents of the Portland Formation assemblage include several good Liassic or
Jurassic indicators, such as Carollina itzlnensir, Callialasporites 6. C. dampieri, CaLLzalasporites trilobatt~s, Araz~cariacites fissz~s, Cycadopites reticulatzls,
Perotrilete~ 6. P. pselldoreticlllatzls, LRptole~iditeJ
cf. L. major, and Gleicheniidites cf. G. ·nilssonii.
The overall composition of the flora compares well
with that of the Liassic (Balme, 1857; Couper,
1358; Nilsson, 1958; Wall, 1365; Schulz, 1967;
Reiser and 'Williams, 1363; Pocock, 1370; Volkheimer, 1971; Volkheimer, 1972; Antonescu, 1373),
particularly in the dominance of Corollina torosus
and the presence of Corollina itzlnensiJ, C. meyeriana,
Araucariacites australiJ, A,Fssus, Cycadopites deterizlJ,
C. reticsclattcJ,'C. janJonii, AliJporites thomasii,
Pityopollenites pallidus, Platysaccus 6. P. lopJiensis,
PerinopoLleszites elatoides, Callialasporites spp.,
GranulatiJporiteJ infirmus, ConveP·PucosiJporiteJ cameronii, Todisporites rotundiformis, Gleicheniidites cf.
G. rrzilssonii, LRptolepidktes sp., and HarrisiJpo~a sp.
Even though many of these palynomorphs are long
ranging and not good index fossils, together they
make up a flora unlike any flora reported from the
Triassic.
In contrast to those of the Upper Triassic, palynofloral assemblages of the Liassic are relatively uniform, both geographically and chronologically.
Although new species appear throughout the Liassic,
floral change is gradual, with a. number of plant
types forming dominant associations over wide geographic areas. Minor floral constituents seem to be
the most variable. Several good index spores and
pollen have been recognized, such as certain species
of Dictyotriletes, ContzgniJporiteJ, and Callialasporites. When present in significant amounts (generally
greater than 1%), these forms can be used to distinguish the upper part of the Liassic from the lower
and middle parts.
Antonescu (1973) reported the presence in very
small amounts of several characteristic upper Liassic
palynomorphs in the middle Liassic of Rumania:
Callialasporites and Leptolepidites. The rare occurrence at our Locality 11 of CallialaJporiteJ 6. C.
dampieri, CallialasporiteJ trilobatzls, Leptolepidites
PL ATE 7
1
2
3
4
5
6
7
Carmerosporites reductiverrucatus n. sp. (Holotype).
Locality II, fj4Clm.
C. reductiverrucatus n. sp. Locality 11; composite
photograph of two focal levels, 46Ccm-561~m.
C. reductiverrucatus n. sp. Locality 1 1, 5 ~um.
C. reductiverrucatus n. sp. Locality 11, 58tcm.
C. reductiverrucatus n. sp. Locality 11, uiradiate fold
present; distal part of grain missing, 33CLal·
Araucariacites fissus Reiser & Williams. Locality 11,
30CLm·
A, fissus. Locality 11, 76tGm.
PLATE 8
1
2
Cycadopites reticukr~us (Nilsson~ comb. nov. Locality
C. retict~latzls~ (Nilsson) comb. nov. Locality 11, 48~m
(under oil).
C. retictilatzis (Nilsson) comb. nov. Locality 5, 32CLm
wide (under oil).
4 C~L-crdopites z~~e.rtfEelclic/rs n. sp. (Holotype). Locality
11, GOE~Lm.
5. u~esif~elrlicus n. sp. Localit)r 11, ~O~m (under oil).
6 C. rceJrf;rldicNs n. sp. Localit)r 1 1, (,0 CLm(under oil).
7 Cycadopites 6. C. janJonii Pocock. Locality ~, 36~m.
8 C~~cadopitss 6. C.iansonii. Localir)· ~, 35~Lm.
9 Cycado~iles 6. C.iansonii. Locality ~, 46CLm.
10 C3,cadopites deterius (Balme) Pocuck. Locality 5,
11
12
13
14
15
16
17
18
19
20
21
48~m.
C. deteri~s. Locality 11, 4 I~m.
C. dett·ri~s. Locality ~, Tl~m~
Cycadopites durhamensis n. sp. (Holotype). Locality
1, 26~Lm.
C. dllrhamenJis n. sp. Locality 1, 28~m.
C. dtirhamensis n. sp. Locality 1, 29Ccm.
C~lcadopites sp. 1. Locality 8, 4SjCLm·
Cycado~ites andrewsii n. sp. Locality 8, S.E.M.; 34CLm.
C. annrewsii n. sp..locality 8, 37ctm.
C. andrewsii n. sp. Locality 8, 341~Lm.
C. andrewsii n. sp. (Holotype). Locality 8, ~7cLm.
C. andrewsii n. sp. Locality 8, 42CLm.
cf. L. niirlor, and C~rOllind zt~nensis, generally not
reported below the upper Liassic, suggests that the
middle part of the Portland Formation is perhaps
Pliensbachian or youn~er in aae. Dictyotriletes spp.
(Pi. 1, fig. 4-6) and Convolz~tispora ~luRiforma are
present in the Hartford and Culpeper basins in
strata presumed to be older than the Portland Formation, Such an occurrence strongly favors an early
to middle Liassic age for much of the Portland
Formation (Text-fig. 5). The upper part of the
Portland is as yet unsampled, but based on stratigraphic thickness, this part of the formation could
well be upper Liassic.
Galton (1971) compared the prosauropods Anchisaurus and AmmosaurEls, and the primitive crocodile StegomoJuchzcs of the upper Portland Formation
with closely related reptiles of the Navajo Sandstone
of northeastern Arizona, suggesting that these formations were closely contemporaneous. Galton, on
the basis of previous authors, presumed the Portland
Formation to be Upper Triassic, and therefore suggested that the Navajo Sandstone might also be
Triassic. However, most previous investigators have
assigned a probable Early to Middle Jurassic age
to the Navajo (Galton, 1971, p. 791, Text-fig. 13).
If the sedimentation rate mentioned above for the
lower Brunswick Formation can be applied to the
2000+m of Portland Formation, a minimum 6.25
m.y. duration would be indicated. This would be
less than the extent of time indicated by palynological
evidence, suggesting that the average rate of sedimentation for the Portland Formation was about
half that given by Van Houten (1363) for the
Brunswick Formation. Comparison of similar types
of sediment were made for sections of the lower
Brunswick Formation along the Delaware River,
N.J., and the Portland Formation along the Connecticut and Wesfield rivers, Mass.
Pdleoecology
Several types of environment for source plants
have been proposed for assemblages dominated by
Corollina pollen, particularly C. torosus: (1) coastal
swamp habitat to account for Corollina pollen in
offshore marine deposits and poor Corollina representation in back delta environments (Hughes and
Moody-Smart, 1367); (2) coastal or lagoonal environment under dry climatic conditions to account
for sediments rich in Corollina, laterally associated
with strata containing marine microplankton assemblages in one direction, laterally grading to strata
rich in Monosulcites-type pollen in the other direction
(Pocock and Jansonius, 1961; Venkatachala and
Gocza'n, 1964); (3) upland sandy slope environment with source plant tolerance of drought conditions (Vakhrameev, 1970). Occasional association of
megafossils of Corollina- producers with Equisetites,
or of Corollina pollen with Equisetites and articulate spores, is not significant enough to suggest any
particular environmental relationship (Batten, 1973).
Similar associations occur in the Hartford Basin,
and we suggest that Corollina pollen, because of
relatively high density, probably reached some of the
same sites of deposition largely through river drainage
systems, as did Equisetites. A similar course of
transportation could easily account for the dominance
of small, broken fragments of Hirmerella and
Brachyphyllum at many megafossil localities.
The sediments of the central area of the basin are
predominantly fine grained. Many of these sediments
were probably deposited in variable, shallow to
deepwater lakes, as evidenced by turbidites, slumps,
and dark, gray black shales and siltstones within
reddish brown shale and siltstone sequences (Sanders,
1368), locally containing horizons with reptilean footprints. Since no known evidence of paleosols or
root penetration zones exists in these finer grained
deposits, and a large footprint excavation (Rocky Hill
Dinosaur State Park, Conn.) indicates extensive mudflat conditions, it is safe to assume that Corollina
producers did not frequently grow there,
Predominantly coarse-grained sediments in the
marginal parts of the basin support fluvial origin
(Krynine, 1950). It is possible that Corollina producers grew on these fans and deltas, thus supporting, in
part, Vakhrameev's sandy slope environment. It is
interesting that this fluvial environment also suggests
an association with small basin swamps, lakeside
habitats, and lagoons. However, it is difficult to
envision a relatively pure population of Corollina
producers restricted to the basin. Corollina accounts
for more than 30% of the palynoflora in sediments
from large areas (in absolute terms, non-Corollina
palynomorphs are usually less than 350 per gram
of sediment), and there do not seem to be sufficient
areas of potential Corollina- producer environments
in the basin proper to account for the masses of pollen
found.
The absence of true coals and scarcity of coniferous
megafossils in the basin argue against extensive
swamps such as those dominated by Taxodium or
Rhizophora today. The dominance of Corollina
producers in the younger sediments of the Culpeper,
Newark, and Hartford basins suggests that a large
geographic area was covered by this assemblage.
Others have suggested that dominance of Corollina
pollen probably indicates close proximity of marine
or coastal environments (Pocock and Jansonius, 1961;
Wall, 1965; Batten, 1373).
Some previous authors (Pocock and Jansonius,
1361; Venkatachaia and Gdcza'n, 1964; Vakhrameev, 1970) have claimed that Corollina pollenproducers occurred in regions with generally arid to
semiarid climatic conditions, but have not given
convincing evidence to support this opinion. However,
the source plants show no convincing xeromorphic
adaptations -- microphyIly by itself is not necessarily
such an adaptation. Furthermore, Krynine (2850)
proposed a warm, humid savanna climate, witha
heavy but seasonally distributed rainfall and a dry
season of two or three months for the Hartford
Basin. Recent recognition of microlaminated (varved)
micritic lake beds in the Shuttle Meadow Fm.,
probably produced in a meromictic lake under the
influence of a seasonal climate (Comet, et al., 1373),
might support Krynine's wet-dry climatic interpretation. It is highly improbable that the Hartford Basin
flora, which includes many kinds of cryptogams
(based on spore diversity), including large-leaf forms
of Clathropteris, and has few known representatives
with well-developed xeromorphic cuticular adaptations, could tolerate long periods of aridity.
Acknowledgments
The authors acknowledge the support of The National Science Foundation (Earth Sciences Section,
grant number GA-36870) for all phases of this project.
We also acknowledge the generous advice on field
problems and stratigraphic correlation, as well as in
location of palyniferous zones, of P. E. Olsen, Yale
University. Professor Donald Baird, Princeton University, has provided us with the sample designated No. 11, from the Newark Basin. Professor
F. G. Van Houten, Princeton University, read portions
of the manuscript and suggested useful improvements.
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