PALYNOLOGICAL CONTRIBUTIONS TO THE CHRONOZOGY AND STRATIGRAPHY OF THE HARTFORD BASIN IN CONNECTICUT AND MASSACHUSETTS

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.. Remarks:

PALYNOLOGICAL CONTRIBUTIONS TO THE

CHRONOZOGY AND STRATIGRAPHY OF THE

The Pennsylvania State University

University Park, Pennsylvania 16802

ABSTRACT

Recent discoveries of palynoflorules at numerous localities in the Newark Group basins of the easternUnited States provide new evidence for correlation of deposits in these basins Floras from the ShuttleMeadow and Portland Formations of the Newark Group rocks in the Hartford Basin of Connecticut andMassachusetts indicate that the Triassic-Jurassic boundary is located within the rocks of this basin: ashift 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 theShuttle 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 forthis 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 ofCarnian-Norian age from New Jersey Paleobotanical and geological evidence is summarized, showing thatthe predominantly Corollina (Hirmerella) palynofloras of the Hartford Basin were associated with awarm, seasonally wet and dry climate, and casting doubt on the idea that Corollina-producing plantscomposed 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 (Danzé-Corsin & 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

Recent discoveries of palynomorphs in most of the major Newark Group basins of the eastern UnitedStates (Text-fig 1) provide new evidence for correlating Newark Group strata with European Triassic-Jurassic 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 thenavailable 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 theHartford 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 volcanicformations 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 theGaillard 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 innorthern 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 steadilydecreases 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, andthe New Haven Arkose and Hitchcock volcanics in Massachusetts This unit, consisting of at least twolava 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 thickenboth 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 "Long-meadow 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 beenselected 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 PennsylvaniaState 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 broadsize distributions, mean and mode are also given Abundance of a species at a given locality is indicated bythe 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 frequentlybordered 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, andgrana (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 frequentlygreater than their diameter Bases of sculptural elements usually joined to form a low, imperfect ridgesystem 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 fromthe 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 withbroadly 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, 2species 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, singlelayered, 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 organspecies If this species were derived from one source, the extreme variation in size suggests some sort ofenvironmental 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

(28-45 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 ofone 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

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 equatorialdiameter 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 similarbimodal 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, predominantlycomplete reticulum, which extends slightly proximal to equatorial margin at apices, generally ending atequatorial margin between apices Muri of reticulum 2-7 um wide; lumina of reticulum 1.7-5.2 um inoverall diameter Exine doublelayered, 1.7-3.0 um thick, increasing to 4.0 um at apices; endexine thinnerthan 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 distributionand shape of lumina Dictyotriletes sp 1 has also been recovered from our Licking Run locality nearMidland, 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

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 ofspecimens 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 elevatedlips 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) andtheir specimens, since C infirmus possesses kyrtomes and a finely rugose sculpture However, Balme'sillustrations 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 tothe 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 innumber 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 inoverall 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 umhigh Perispore closely fitting, densely covered with small grana, 0.1-0.4 um high; perispore attachedonly at laesura margins Overall site (2 specimens) 55-56 um, about 52-53 um in diameter, excludingperispore

Remarks: The presence of grana on the perispore might be sufficiently different from the type

descrip-tion to warrant the creadescrip-tion 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 oflaesurae variable, 26-78% of equatorial diameter Exine (exclusive of sculpture) approximately 0.8 umthick, 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 ofsuture 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 Slightlyirregular 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) forthe Australian forms The indicated Early Jurassic age for A fissus Reiser & Williams agrees well withthe proposed age for the Portland Fm This species becomes more frequent in younger strata The largestforms 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 adistinct 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 5generally 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 slightlydarkened 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 areneeded to determine if a significant distinction exists between our forms resembling C monoalasporusSukh 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, tightlyfolded to form a verrucose and rugose surface Exine of body about 0.9 um thick Sacci possess fine reticu-lum 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 Mostbisaccates 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 indicatedthat 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) Eventhough the assignment of these "neotypes" by Zauer is subjective, it is perhaps the only way to obtainstability 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 vaguelydefined 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 suggeststhat 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 Verrucosisporites cheneyi n sp (Holotype) Locality 1, 73 um

2 V cheneyi n sp Locality 1, 85 um wide

3 Harrisispora sp 1 Locality 11, 80 um

4 Dictyophyllidites paramuensteri n sp Locality 1, 65 um wide

5 D paramuensteri n sp (Holotype) Locality 1, 63 um

6 Deltoidospora hallii Miner Locality 11, 27 um

7 Spore A Locality 11, 98 um

8 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;

Bennettiteae-Pollenites reclusus Thiergart, 1949, p 11, P1 2, fig 14, 15, P1 3, fig 6; Bennettiteae-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 bodiesfrom 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 face Intrastructure massive, alveolate, punctate, vermiculate or pseudovermiculate, reticulate or

sur-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 subequatorialfurrow (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 orrimula, 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 coveredwith 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 tured layer, which is thickened in equatorial region to form a band or belt Endexine variably distinct

struc-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 thesetwo populations does not permit their distinction in a mixed sample, as the ranges of variation widelyoverlap 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)