I based my identification and nomenclature on Frederiksen (1980a), and made no attempt to revise his taxonomy. In the following section, I have given not only standard taxon descriptions, but have also reviewed the available ecological information for each taxon.

Laevigatosporites haardtii (Potonié & Venitz) Thomson & Pflug 1953

Sporites haardtii Potonié and Venitz 1934, p. 13, pl. 1, fig. 13.

Laevigtosporites haardtii (Potonié and Venitz) Thomson and Pflug, 1953, p. 59, pl. 3, figs. 27-38.

Laevigatosporites sp., Tschudy and Van Lonen, 1970, pl. 1, fig. 1.

Description. 25-70 µm. Outline beanshaped. Shape of the dehiscence concave to straight. Dehiscence thick, on both sides of the equator nearly or entirely reached.

Affinity. According to Frederiksen (1980a), Laevigatosporites may represent the modern families Aspidiaceae, Aspleniaceae, Blechnaceae, Gleicheniaceae, Lomariopsidaceae, Polypodiaceae, and Pteridaceae.

Paleoenvironmental Notes. Laevigatosporites sp. was the most important palynomorph in Maastrichtian deposits described by Kroeger (1985). It was most common in abandoned channel and flood basin marshes, but was also found in point bar and temporary pond and swale deposits. Farley and Dilcher (1986), in a study of the Mid-Cretaceous Dakota formation, found L.ovatus to be more abundant in the "marshy lakeside" environment; floral differentiation among environments was not, however, strong. Laevigatosporites types were present in all three horizontal lignite samples tallied from the Eocene Gatuncillo Fm. of Panama by Graham (1985).

Graham (1987) indicated that he would have interpreted a high diversity and abundance of Laevigatosporites in the Miocene Uscari flora of Costa Rica as representative of local lowland marshes had other, Angiospermous, indicators been present. Nichols and Traverse (1971) reported a conspicuous abundance of Laevigatosporites in Paleocene Texas lignite samples characterized by palm pollen.

Farley (1990), in his study of the Early Eocene of Wyoming, found the greatest concentration of most spores, including Laevigatosporites, within the levee-crevasse splay environment. According to Farley, "...the abundance of Pteridiophyte spores reflects local stands of ferns whose rapid life style is well adapted to the unstable and damp substrate in the levee/crevasse splay environment.

Van der Hammen (1963) figured "Monolete spores" in his diagram from British Guiana. Almost all, he believed, were from the Polypodiaceae, which, he explained, "...occur frequently in freshwater swamps, but are also found under slightly brackish conditions." (p. 139)

The palynomorph flora of one Montana Paleocene coal described by Wilson and Webster (1946) contained 51.5% Laevigatosporites spp.

Extensive fern swamps were reported in the Orinoco Delta by Muller (1959). The distribution of Polypodiaceous types was highly dependant on proximity to parent populations. In some areas these spores dominated. In two of his cores, however, the presence of Polypodium was explained as exotic spores blown in from Trinidad and Tobago.

Occurrence. Up to 32% in 12 lignites and the clastic sample at Lake Somerville.

Range. Frederiksen (1980a) indicates that L. haardtii spores have been found from the Cretaceous to the Recent.

Genus Polypodiisporonites Potonié 1931

Polypodiisporonites favus Potonié 1931

Polypodii(?)-sporonites favus Potonié, 1931, p. 556, fig. 3.

Verrucatosporites favus (Potonié) Thomson and Pflug, 1953, p. 60, pl. 3, figs. 52-55; pl. 4, figs. 1-4 [misidentified].

Polypodiisporites favus (Potonié) Potonié, 1956, p. 78.

Reticuloidosporites favus (Potonié) Krutzsch, 1959, p. 215, pl. 42, figs. 467-470.

Description. 30 to 80 µm. Resembling verrucate. "Warts" at most 2 µm high, their shape corresponding to their position on a hemisphere. Sulcus unbroken. (Translated from Thomson & Pflug)

Affinity. Assigned as a "probable" Polypodiaceae by Frederiksen (1980a).

Paleoenvironmental Notes. Kroeger (1985), studying a Maastrichtian sequence from South Dakota, reported a similar taxon, Polypodiidites Ross, also assigned to the Polypodiaceae. This taxon was rare and found only in flood basin marsh and temporary pond/swale environments.

In the Orinoco Delta, Muller (1959) reported a strong coincidence of Polypodiaceous spores in surface samples and proximity of the mother plant. The heavily ornamented types, i.e. Polypodiisporonites type, settled most rapidly.

There are about 1000 living species in the Polypodiaceae and nearly all of them are epiphytes. Some, however, are not; Polypodium, for example, is terrestrial or ruprestral at the geographical and altitudinal limits of its range and epiphitic in the tropics. The genus can live from sea level to 4400 m., in a variety of moisture conditions. (Tryon and Tryon, 1982)

Occurrence. Up to 3% in 42 levels at San Miguel and up to 5% in 11 lignites and the clastic samples from Lake Somerville.

Range. Elsik (1968) found Polypodiisporonites afavus in the Paleocene Rockdale lignite and Frederiksen (1980a) found these grains from the Upper Claiborne to the Lower Vicksburg.

Microfoveolatosporis pseudodentatus Krutzsch, 1959, p. 212, pl. 41, figs. 463-466.

Microfoveolatosporis cf. M. pseudodentatus Krutzsch, 1959. Engelhardt, 1964, p. 69-70, pl. 1, fig. 6.

Microfoveolatosporis cf. M. pseudodentatus Engelhardt, 1964. Tschudy and Van Lonen, 1970, pl. 1, fig. 3.

Microfoveolatosporis pseudodentata Krutszch, 1959. Frederiksen, 1980, p. 29, pl. 1, fig. 4.

Description. The only known form species of the form genus possesses one ca. 1/2-2 1/2 µm thick wall, which is distinctly two-layered. V=1.5-2/1. The sculpture consists of a loose and sometimes delicate, netlike arrangement of roundish, foveolate cavities (z. T. of their small diameter) in the otherwise scabrate to smooth surface of the sporewall. The fovelolae are to approximately 1/2 of the outer/exterior layer deep. Sometimes, the outer wall layer shows beyond it a weak bar structure. The dehiscence is straight, + long, r=about 3/4 to 4/5, sometimes delicate and weak. Outline and figure slender-oval. Size ca. 65-80 µm. Affinity. Similar to Psilotum according to Kedves (1969) and to Schizaea pusilla Pursh, according to Engelhardt (1964).

Paleoenvironmental Notes. Psilotum grows on tree bases, logs, and hummocks in low wet woods, in the southeastern United States, as well as in the tropics. P. pusilla is a small, slender fern which lives in acid bogs in the northeastern part of North America (Mickel, 1979). Frederiksen (1980b) reported that Psilotum grows in humid subtropical, winter-dry tropical, and wet tropical regions.

Occurrence. Up to 6% in one lignite and the clastic sample at Lake Somerville.

Range. Found in the Paleocene Rockdale Lignite (Elsik, 1968)

Schizeaea tenuistriata (Pflanzl) Frederiksen, 1980a, p. 29, pl. 1, fig. 6.

Cicatricosisporites pseudodorogensis tenuistriatus Pflanzl, 1956, p. 239, pl. 16, fig. 5.

Affinity. According to Frederiksen (1981), similar to modern Schizaea laevigata Mett. and S. penicillata Kunth.

Occurrence. Found at 1% in one sample from Sequence B of San Miguel. Absent at Lake Somerville.

Range. Found from the Jackson (Moodys Branch) upwards in Mississippi (Frederiksen, 1980a).

Concavisporites discites Pflug In: Thomson & Pflug, 1953, p. 49, pl. 1, fig. 24.

Description. 15-35 µm. End- and Extexospore with concave equatorial outline. Torus indistinct, with disclike formation. Y-ledges dispersing in a straight line. (Translated from Thomson and Pflug)

Affinity. Possibly Gleicheniaceae, according to Frederiksen (1980a).

Occurrence. Less than 1% in two samples at San Miguel, and absent at Lake Somerville.

Range. Frederiksen (1980a) found one specimen in the Jackson age Yazoo Clay of Mississippi.

Lygodium? labratum Frederiksen, 1973, p. 69, pl. 1, figs. 5-10.

Lygodium labratum Frederiksen, 1980a, p. 30, pl. 1, figs. 10-11.

Description. Size 29-42 µm, mean 35 µm, holotype 34 µm. Outline triangular; sides slightly convex to slightly concave. Trilete; sutures closed, labra l.5-3 µm wide, rays straight, extending nearly full radius. Exine about 2 µm thick, sexine:nexine 2-3:1. Entire exine foveolate except labra; foveolae 0.5-1 µm in diameter, some of them anastomosing.

Occurrence. Less than 1% in one sample at San Miguel and absent at Lake Somerville.

Range. Frederiksen (1980a) found this taxon from the Upper Claiborne to the Lower Vicksburg of the Gulf Coast.

Punctati-sporites adriennis Potonié and Gelletich, 1933, p. 521, pl. 2, figs. 14-15.

Lygodiumsporites adriennis (Potonié and Gelletich) Potonié, 1956, p. 19.

Leiotriletes adriennis (Potonié and Gelletich) Krutzsch, 1959, p. 57

Description. Outline-convex-triangular, strongly ringed; side flat to scabrate, 1.5-2 µm thick, V=2/1; all-around homogenous thickness; r=only 1/3 - 2/3, in points running out, size ca. 70 µm.

Affinity. Spores of L.adriennis are similar to those removed by Manchester and Zavada (1987) from remains of the extinct Lygodium Kaulfussi Heer. The fossils, from the Upper Eocene Bridger Formation, consist of attached fertile and sterile fronds. The spores are trilete, psilate, and 55-70 µm in diameter. According to these authors, L. kaulfussi has been described from the Eocene of Alabama by Berry (1924). Similar spores were isolated from L. kaulfussi pinnules from Dorset, UK, by Chandler (1955). L. adriennis spores are common in Eocene Gulf Coast sediments. Frederiksen (1980a), citing the similarity to spores of A. aureum L., believed them to be produced by both Acrostichum, some species of which commonly occur in salt water today, and Lygodium. Citing Berry's identification of Acrostichum georgianum Berry leaves in the Claiborne and Jackson (Eocene) rocks of Georgia, Frederiksen stressed the resemblance to Acrostichum. Westgate and Gee (in press) found L. adriennis in association with Spinozonocolpites (thought to be synonymous with the mangrove Nypa) in Eocene sediments from Texas and consequently believed the affinity to be with brackish-water Acrostichum.

Paleoenvironmental Notes. The genus Lygodium contains Schizaceous ferns which, according to Manchester and Zavada (1987), are "...characterized by unusual leaves adapted for climbing. " (p. 392) Lygodium commonly lives along forest borders where it can get light. In tropical America, it may live in rain forests or more commonly, gallery forest, shrubby savannah or along borders of streams, from sea level to 1000 m. (Tryon and Tryon, 1982). Today, the genus Acrostichum is pantropical and contains ferns which inhabits a range of fresh to saltwater wetlands (Mickel, 1979).

Occurrence. Up to 2% in 18 samples from San Miguel and up to 1% in two samples at Lake Somerville.

Range. Dispersed spores attributed to Lygodium were present in Triassic sediments and macrofossils date from the Upper Cretaceous (Manchester and Zavada, 1987). Found in Upper Claiborne to Lower Vicksburg of the Gulf Coast by Frederiksen (1980a).

Ctenopteris? elsikii (Frederiksen) Frederiksen 1980

Undulatisporites sp., Elsik 1968, p. 294, pl. 8, fig. 4; pl. 10, fig. 6.

Undulatisporites elsikii Frederiksen, 1973, p. 69-70, pl. 1, figs. 11-12, 18.

Ctenopteris? elsikii (Frederiksen) Frederiksen, 1980a, p. 31, pl. 2, fig. 5.

Description. Size 22-47 µm, mean 30 µm, holotype 32 µm. Outline triangular, with convex to slightly concave sides and narrowly rounded corners. Trilete; sutures closed; labra wavy, o.5-1 µm wide and about 3 µm high, extending nearly to the outline. Exine 1-1.5 µm thick, psilate or rarely infrapunctate, sometimes with one or two coarse folds.

Affinity. Frederiksen (1980a) believed this grain to be very similar to several species of modern Ctenopteris.

Occurrence. Less than 1% in one sample from San Miguel and absent at Lake Somerville.

Range. Elsik (1968) found this spore in the Paleocene Rockdale lignite and Frederiksen (1980a) found it in samples from the Upper Claiborne to the Lower Vicksburg of the Gulf Coast.

Granulatisporites luteticus (Krutzsch) Frederiksen 1980

Punctatisporites luteticus Krutzsch, 1959, p. 61-62, pl. 1, figs. 3-7.

Granulatisporites luteticus (Krutzsch) Frederiksen, 1980a, p. 31, pl. 2, fig. 13.

Description. Shape roundish-triangular, surface completely dense with incomplete equally size punctate-granulate bordered sculptural elements. Outline lightly wavy. Spore wall three layered, V=1/1/1, only about 1-1.5 µm thick, from there frequently secondarily declined. Y-mark distinct, radiating in a straight course, r=1/2-3/4. Immediately on the rays, the sculptural elements are mostly fused. Size about 50-70 µm.

Affinity. According to Frederiksen (1980a), similar to spores of Acrostichum aureum L.

Paleoenvironmental Notes. Acrostichum aureum is a pantropical fern occurring in this country in peninsular Florida. It inhabits freshwater swamps and brackish to saltwater marshes (Mickel, 1979).

Occurrence. Less than 1% in one sample from San Miguel and 1% in the clastic sample at Lake Somerville.

Range. Frederiksen (1980a) found only one specimen from the Jackson age Yazoo Clay of Mississippi.

Cicatricosisporites dorogensis Potonié and Gelletich, 1933, p. 522, pl. 1, figs 1-5.

Description. Trilete spores, amb triangular, sides straight to slightly convex, apices rounded, equatorial diameter 50-56 µm. Laesurae reach full radius. Exine 3.0 to 4.0 µm thick, sculpture cicatricose, muri and vallae 2.0 to 2.5 µm wide, parallel to the sides of the spore proximally and distally. Exine may be radially thickened and the ends of the angles unsculptured.

Affinity. Frederiksen (1980a) attributed Cicatricosisporites to the Schizacean genera Anemia or Mohria. Graham (1985) believed that Cicatricosisporites resembled Ceratopteris in the Parkiaceae. Chandler (1955) illustrated similar spores isolated from the Early Tertiary megafossil Anemia colwellensis Chandler.

Paleoenvironmental Notes. Cicatricosisporites spores probably conspecific with C. dorogensis were rare in a Maastrichtian deposit described from North Dakota by Kroeger (1985). The genus occurred only in point bar deposits. Farley and Dilcher (1986) found Cicatricosisporites spp. to be more common in "marshy lakeside" environments of the Middle Cretaceous Dakota Formation.

Graham (1985) reported the taxon as common (15%) in one horizontal lignite sample but absent in two others from the middle(?) to upper Eocene Gatuncillo Fm. of Panama.

The very similar Cicatricosisporites intersectus was found in abundance in the Late Eocene to Early Eocene Kitsalano Fm. of British Columbia by Hopkins (1967). He indicated that it was restricted to the coastal lowlands.

Ceratopteris is a genus of tropical water ferns. Two species occur in this country and inhabit mud and quiet water in southern Florida and Louisiana. Anemia inhabits limestone substrates from Florida and the Caribbean to the Edwards Plateau of Texas and southern Mexico (Mickel, 1979). It is a terrestrial species and in general a genus of open habitats and well drained sites, with habitats from sea level to 3200 m. (Tryon and Tryon, 1982).

Occurrence. Up to 47% in 41 samples at San Miguel and to 59% in three lignite samples as well as in the clastic sample at Lake Somerville.

Range. Found in the Paleocene Rockdale Lignite by Elsik (1968) and in the Upper Claiborne to Lower Vicksburg by Frederiksen (1980a).

Lusatisporis perinatus Krutzsch et al., 1963, p. 98, pl. 30, figs. 10-11.

Selaginella sinutus Martin and Rouse, 1966, p. 185-186, pl. 1, figs. 7-8.

Selaginella perinatus (Krutzsch et al.) Frederiksen, 1980a, p. 33, pl. 3, figs. 14-15.

Description. Spore outline circular or only slightly triangular. Perispore decidedly coarsely granular, finely wrinkled, and extending from 2 to 5 µm beyond the limits of the endospore. Thickness of perispore ca. 1 µm. The laesurae of the trilete mark are wavy, and the margo is much wrinkled, thus giving the appearance of a sinuous ribbon. Small radial folds on the perispore occur on the interradials. Range of diameter: 28-48 µm. (Martin and Rouse, 1966)

Affinity. Martin and Rouse (1966) report that the spores are virtually identical to the extant species Selaginella wallacei and S. oregona.

Paleoenvironmental Notes. Most species of the fern ally Selaginella are found in tropical regions, but the genus is also found as far north as Alaska. Modern habitats range from wetland to dry pine woodlands and dunes (Mickel, 1979).

Occurrence. Less than 1% in one sample from San Miguel and absent at Lake Somerville

Range. Elsik (1968) found this species in the Paleocene Rockdale lignite. Frederiksen (1980a) found this species in samples ranging from the Upper Claiborne to Lower Vicksburg of the Gulf Coast.

Sphagnum stereoides (Potonié and Venitz) Martin and Rouse 1966

Sporites stereoides Potonié and Venitz, 1934, p. 11-12, pl. 1, figs. 4-5.

Stereisporites stereoides (Potonié and Venitz) Thomson and Pflug 1953, p. 53, pl. 1, figs. 64-73.

Sphagnumsporites steroides (Potonié and Venitz) Potonié, 1956, p. 17.

Sphagnum steroides (Potonié and Venitz) Martin and Rouse, 1966, p. 184, pl. 1, fig. 3.

Description. 15 to 30 µm. Exospore under 2 µm thick. Y-border reaching the equator.

Paleoenvironmental Notes. Nichols and Traverse (1971), in their palynological study of Late Paleocene Texas lignites, found Sphagnum percentages of a few to 10% indicative of a fluvial swamp environment.

Modern Sphagnum, or peat moss, lives in wetlands, most often in peat bogs. It creates an acid environment for itself and other plants. (Crum, 1976)

Occurrence. Less than 1% in two samples from San Miguel and 1% in one lignite sample from Lake Somerville.

Range. This species was found by Frederiksen (1981) from the Oligocene Vicksburg Group. Elsik (1968) found it in the Paleocene Rockdale Lignite.

Taxodium hiatipites Wodehouse, 1933, p. 493, fig. 17.

Taxodiaceaepollenites hiatus (R. Potonié, 1931) Kremp, 1949 [misidentified]. Engelhardt, 1964, p. 71, pl.1, fig. 10.

Inaperturopollenites cf. I. hiatus (R. Potonié) Thomson and Pflug, 1953. Tschudy and Van Lonen, 1970, pl. 2, figs. 5-6.

Cupressacites hiatipites (Wodehouse) Krutzsch, 1971, p. 41.

Description. 25-50 µm. Outline spherical, less often decayed. Swollen and burst open, thereby a characteristic bill-like aperture (Hiatus). Exine over 1 µm thick, psilate to scabrate, distinctly two-layered. Sometimes one can discern an indistinct ligula and an exit pore.

Affinity. Probably the Taxodiaceous genera Taxodium or Glyptostrobus, according to Frederiksen (1980a).

Paleoenvironmental Notes. C. hiatipites pollen is extremely rare in Eocene Gulf Coast lignites (Elsik, 1978; Frederiksen, 1981). The taxon is present in larger percentages in Late Paleocene sediments (Elsik, 1978; Gennett, unpublished data from Louisiana; Kroeger, 1985). Elsik mentioned that "Taxodium is abundant in many marine and non-marine clastic sediments of Paleocene-Lower Eocene [sic] of the Gulf Coast. (p. 29)" and suggested that distribution is tied to water transport.

Presently distributed along the Atlantic and Gulf Coastal Plains and in the Mississippi Valley, Taxodium (baldcypress) most often grows in permanent swamps (Harrar and Harrar, 1962). Coverage of T. distichum ranges up to 40% in the Mississippi delta and to 60% in Florida; Taxodium comprises 60% of arboreal pollen in some parts of the Mississippi delta (Delcourt et al., 1984). Most Cupressaceae species shed huge amounts of wind dispersed pollen.

Occurrence. Less than 1% in 5 samples from San Miguel and 1% in three lignite samples and the clastic sample at Lake Somerville.

Range. Frederiksen (1980a) found this taxon infrequently to occasionally in the Upper Claiborne to Lower Vicksburg of the Gulf Coast; Elsik (1968) found more abundant grains in the Paleocene Rockdale lignite.

Because of present uncertainties in the classification of Tertiary bisaccate pollen (T. Timmcke, oral commun., 1989), I followed Kroeger (1985) and combined the few grains into a descriptive group.

Affinity. Frederiksen (1980a) classified all bisaccate pollen present in his Gulf Coast Eocene as members of the Pinaceae. Individuals present in the San Miguel samples resemble members of the modern genera Picea or Pinus as illustrated by McAndrews et al (1978).

Paleoenvironmental Notes. Species of the Pinaceae were reported as infrequent to rare by Frederiksen (1980a) in Gulf Coast Jackson Group samples. Potter reported only rare occurrences in the Tennessee Claiborne and Kroeger (1985) described the group as uncommon in the Paleocene of South Dakota. On the other hand, Mancini (1981) listed Pinaceae pollen as characteristic of Alabama marine shales interbedded with Middle Paleocene Naheola lignites. An increasingly coniferous Cordilleran flora (Leopold and McGinitie, 1972) may have provided a long distance source for bisaccates; Bartley (1967) for example, reported a site in Quebec almost 500 km from a pine source receiving 24% Pinus pollen. Dilcher (1973) noted, for the Middle Eocene of Tennessee, "The Appalachian highland is often used to explain the presence of temperate forms in the Mississippi embayment and may be a valid explanation for the presence of some pollen such as Pinus which is not yet reported as a megafossil and might have come from upland plants rather distant from areas of deposition." (p. 56)

Occurrence. Less than 1% in one sample from San Miguel and 1% in one lignite at Lake Somerville.

Ephedra claricristata Shakhmundes, 1965, p. 226-227, fig. 10.

Ephedra eocenica Shakhmundes, 1965, p. 219-220, figs. 2- 3.

Ephedripites (Distachyapites) tertiarius Krutzsch, 1970, p. 156, 158, fig. 20; pl. 44, figs. 1-21.

Gnetaceaepollenites eocenipites (Wodehouse, 1933) R. Potonié, 1958 [misidentified]. Engelhardt, 1964, p. 70, pl. 1, fig. 8.

Ephedra sp. (distachya-type), Fairchild and Elsik, 1969, p. 83 pl. 37, fig. 2.

Ephedra sp. (type A of Steeves and Barghoorn 1959), Tschudy and Van Lonen, 1970, pl. 1, fig. 1.

Ephedra type A of Stevens and Barghoorn, 1959. Tschudy, 1973, p. B17, pl. 4, figs. 22-23.

Occurrence. Less than 1% in 9 samples at San Miguel and absent at Lake Somerville.

Range. Found by Frederiksen (1980a) in the Upper Claiborne to Lower Jackson.

Ephedripites hungaricus Nagy, 1963, p. 278, figs. 1-3, 12A

Ephedra hungarica (Nagy) Frederiksen, 1980a, p. 37, pl. 7, fig. 4.

Description. Pollen grain perprolate (47 X 19 µm) tapering towards the supposed poles. Walls thin, provided with about 10 to 11 ridges. Ridges narrowing and converging towards the poles, 1 to 1.5 µm wide. Grooves between the ridges not wider than 0.5 µm. In the center of the ridges is a dark stripe. At lower focus, a white stripe (about 0.5 µm wide) becomes visible between the dark stripes on both margins of the ridges. Towards the ends of the pollen grain the ridges converge and appear to be somewhat twisted.

Affinity. Ephedra is an extant plant. Specimens from the San Miguel lignites are similar to modern grains pictured in McAndrews et al. (1978) and in the TAMU pollen reference collection.

Paleoenvironmental Notes. Extant forms are common in arid climates. Gray (1960), Hopkins (1967) and Frederiksen (1985) believed that Ephedra may have had a broader ecological range in the Eocene than at present. These authors have suggested that, because Ephedra has been found mostly in brackish or marine sediments, that it may have part of the Eocene beach flora. Hopkins and Sweet (1976) found Ephedra pollen in black shale and coal in the Eocene/ Oligocene Kishenehn Formation of southeastern British Columbia. They suggested that the plants were growing in dry locations at the lee of mountains. Frederiksen (1981) suggested that Ephedra was part of the shrub layer because of its appearance in low quantities in many samples. This, according to Frederiksen indicated abundant, widespread plants with low pollen productivity.

Ephedra has been found in Quaternary cores from Lake Superior, making up an average of .03% of the pre-1890 pollen sum. Janssen (1967) also pictured Ephedra in his study of pollen in Quaternary sediments at Stevens Pond, Minnesota. Ephedra is not included in Lakela's (1965) Flora of Northeastern Minnesota, nor is it described in Gleason and Cronquist's (1963) flora of the eastern United States. Long distance aerial transport is inferred for these sites.

Occurrence. Less than 1% in two samples from San Miguel and absent at Lake Somerville.

Range. Frederiksen (1988) reported Ephedra hungarica from the lower part of the Lisbon Formation (Middle Eocene) into the Oligocene. Ephedra first appeared in the Gulf Coast in the Claiborne Formation (Fairchild and Elsik, 1969); it was established in the continental interior by the end of the Cretaceous (Frederiksen, 1985).

Monoporopollenites gramineoides Meyer, 1956, p. 111, pl. 4, fig. 29.

Graminidites gramineoides (Meyer) Krutzsch, 1970, p. 15.

Graminidites spp. Tschudy, 1973, p. B17, pl. 4, figs. 34-35.

Description. Mean of long and short dimensions, 19-36 µm, mean 30 µm. Exine, 0.3-0.5 µm thick, considerably folded, usually crushed to an oval shape; nearly psilate but fairly punctate, granulate or verrucate; outline nearly smooth. Diameter of pore 1.7-2.5 µm; width of annulus 2.5-3 µm.

Affinity. Gramineae, according to Frederiksen (1980a)

Occurrence. Less than 2% in two levels from San Miguel and absent from Lake Somerville.

Range. Frederiksen (1988) reported first occurrences of Graminidites just below the base of the Gosport Sand.

Milfordia Erdtman, 1960, p. 46.

Monulicipollenites Fairchild in Stover et al., 1966, p. 2-3.

Restioniidites Elsik, 1968, p. 313.

Milfordia Erdtman emend. Krutzsch, 1970, p. 18.

Affinity. Restoniaceae, Flagellariaceae, or Centolepidaceae, according to Frederiksen (1980a).

Paleoenvironmental Notes. According to Frederiksen (1981), "Milfordia pollen is very abundant in some Eocene and lower Oligocene deposits in Europe, and the Centrolepidaceae-Flagellariaceae-Restoniaceae may have been important marsh plants during this time interval...However... the producers of this pollen do not seem to have been important in the peat-forming swamps. (p. 531-532)" Milfordia was rare in the Gulf Coast coals studied by that author.

Occurrence. Up to 2% in four levels from San Miguel and absent from Lake Somerville.

Range. The genus was reported from the Claiborne Group by Fairchild and Elsik (1969) and from the Jackson Group by Frederiksen (1980a).

Description. Pollen grains monoporate, bilaterally symmetrical with the poriferous or non-poriferous usually flattened. Spheroidal to elliptical, the longest axis of the pore usually at angles to the longest axis of the grain. Pore margin uneven with thickened endexine. Exine reticulate with coarsely and finely reticulate areas clearly demarcated in a distinctive pattern on the surface of the grain.

Affinity. Frederiksen (1980a) believed that this genus was Monocotyledonous "...possibly Ruppiaceae or Potomogetonaceae according to Machin (1971, p. 856)." (p. 38.)

Paleoenvironmental Notes. Fowler (1971) believed these pollen grains to have come from brackish to fresh water hydrophytes. The genus is rare in samples examined by Frederiksen (1980a, 1981).

Occurrence. Less than 1% in one sample from San Miguel and absent from Lake Somerville.

Range. Found from Upper Middle Eocene to Oligocene on the Gulf Coast (Frederiksen, 1980a).

Description. Grains spheroidal or oblately flattened and somewhat triangular in outline. Pores three on the equator with their apertures broadly elliptical and meridionally oriented, only slightly protruding above the surface, and with the exine immediately surrounding them slightly thickened corresponding to the Corylus pattern. Texture smooth.

Affinity. As can be inferred from the synonym Engelhardtia, Momipites is very similar to pollen of the modern Juglandaceous genus Engelhardtia. Momipites, the form-genus, was isolated by Crepet et al. (1975, 1980) from fossil catkins similar to the present-day Engelhardtia-Oromunnea-Alfaroa complex. According to Nichols (1973), morphological differentiation between Alfaroa and Engelhardtia pollen is not possible, and some fossil grains are not comparable to either genus, resembling more closely the other Juglandaceous genera. Leaves from the Claiborne Group of Tennessee, more similar to Alfaroa and Oreomunnea than Engelhardia, were described by Dilcher and Manchester (1986) and designated as Oreoroa claibornensis. Manchester (1983) also described fossil wood from the tribe Engelhardieae (undifferentiated) occurring in the Yegua formation. "Momipites...is abundant in the same formation," he said.

The name Momipites was originally given to the genus by Wodehouse (1933) and signified a resemblance to Momisia, a member of the Ulmaceae (elm family).

Paleoenvironmental Notes. According to Frederiksen (1985), "...during the past 30,000 years in the highlands of Papua, New Guinea, the relative frequency of Engelhardtia has fluctuated independently of...other genera with which it is now associated...even during short time periods Engelhardtia seems to change its associates as climatic and ecological conditions change."

Momipites has, in fact, been correlated with changing environmental conditions. Pocknall (1987) used the ratio between Momipites spp. and spp. to define biozones in the Paleocene and Late Eocene of Wyoming. He found that the change between assemblages dominated by the two genera coincided with sedimentological change between fluvial and fluvial lacustrine due to increased tectonism. The increase in relief due to tectonism also initiated a more seasonal climatic regime.

Thoughts on Momipites are numerous. Nichols (1971) found that in the Paleocene of Texas, one species of "Engelhardia" was more common in clays and another in lignites. Traverse (1955) discovered that Momipites in the Oligocene Brandon Lignite sequence was more common in lignitic sand. Graham and Jarzen (1969) described an Oligocene sequence in Puerto Rico in which Momipites-rich clastics alternate with Rhizophora-rich peat. In his study of the Tennessee Claiborne, Potter (1976) noted that both important Momipites species (probably equivalent to M. microfoveolatus and M. coryloides) were common both in clastic sediments and at the base of his lignite seam. Momipites grains were rare in most of the lignite. He believed this taxon to be part of the "Background" rain.

Some authors have used Momipites pollen as a marsh indicator. Elsik (1986) suggested that this taxon was blown into a Yegua-Jackson marsh to form the Momipites dominated assemblage in Zapata county. Elsik (1978) assigned a "regional pollen" status to Momipites spp. because modern Engelhardia grows in upland areas, and because Manning lignites exhibit a low diversity of arboreal pollen. Mukhopadhyay (1989) well used Momipites as a marsh indicator in cursory examinations of San Miguel and Lake Somerville lignites. Conversely, Elsik (1978) found abundant Momipites in a fluvial Yegua Fm. lignite from Madison Co., TX.

In his study of clays and brown coals from the Eocene of Germany, Pflug (1957) grouped Momipites as a genus more likely to be abundant in clays and inferred that it came from lakeshore vegetation. Kvacek (1972), reviewing the occurrence of Engelhardia leaf fossils in Europe, emphasized that, because it is not represented in the Glyptostrobus-Taxodium-Alnus-Nyssa community of the Miocene brown coal swamp forest, Engelhardia was not adapted to lowland swampy conditions during the Tertiary. He believed the fossil Engelhardia of Europe to be "...a typical subtropical component with a great demand on high atmospheric humidity." (p. 30)

In addition to pollen, peltate leaf hairs from the Engelhardieae appear occasionally in palynomorph preparations (Elsik and Dilcher, 1974).

Modern forms are restricted to Asia, where they may live in swampy areas in buttressed form, and the highlands of Central America, where they require summer rainfall, constant high humidity, warm dry winters and an almost unseasonal climate (Leopold and McGintie, 1972). Engelhardtia and Alfaroa both grow today in Costa Rica. Holderidge et al (1971) recorded E. mexicana as a canopy tree, and the two species of Alfaroa as an intermediate size tree. Life zones included wet tropical, moist tropical, premontain wet, and premontane rain forest. Fredericksen (1980c) listed the climatic requirements of the Engelhardtia-Oromunnea-Alfaroa complex as: 1) humid subtropical with precipitation in all seasons and more than three cm rain in the driest month in summer, or 2) winter dry tropical, with more than two months dry, or wet tropical, with ten to twelve months wet.

Flenley (1979) described Engelhardia as basically a montane genus, with some species growing in lowland forests. One species, E. spicata, is found in mountain swamps.

Range. A common Tertiary palynomorph, Momipites is common throughout the Late Eocene and Oligocene of the United States; Nichols (1973) gave the range of the M. coryloides group, which contains both M. coryloides and M. microfoveolatus, as Paleocene to Miocene. The genus persisted into the Miocene in Idaho, and Alaska, but not the Gulf Coast.

Momipites coryloides Wodehouse, 1933, p. 511, fig. 43.

Engelhardtia sp., Fairchild and Elsik, 1969, p. 83, pl. 37, figs. 8-9.

?Momipites sp., Tschudy and Van Loenen, 1970, pl. 2, fig. 15.

Triatriopollenites sp., Tschudy and Van Loenen, 1970, pl. 3, figs. 1-2.

Triatriopollenites sp. of the T. coryphaeus type (20-30 µm.), Tschudy, 1973, p. B16, pl. 4, figs. 12-13.

Description. Oblately flattened and triangular in outline, 21-33.1 µm in diameter.

Affinity. Crepet et al. (1975) removed similar pollen, described as "...triporate, occasionally tetraporate, very finely scabrate exine, mean equatorial diameter is 19.6 µm", from a Claiborne-age macrofossil, Eokachyra. Perianth parts, peltate scales, and stomata referable to the extant Juglandaceous Engelhardtia-Oromunnea-Alfaroa complex were present as macrofossils.

Paleoenvironmental Notes. Both the floral and pollen structure of Eokachyra suggested to Crepet et al. (1975) that pollination was anemophilous. These characteristics included small perianth, exposed anthers, and smooth pollen grains. Pollination mechanisms in the extant E-O-A complex are not well known, but according to these authors the floral structure also suggests anemophily.

Occurrence. Present in all samples from 1 to 69% from San Miguel and Lake Somerville, at levels between 2 and 69%.

Range. Frederiksen (1980a) gives the range of M. coryloides in South Carolina as Late Paleocene to Upper Eocene, and for the Gulf Coast as Latest Paleocene to Lowest Oligocene.

Engelhardtioidites cf. E. microcoryphaeus (R. Potonié, 1931) Potonié, Thomson, and Thiergart, 1950. Engelhardtia microfoveolata Stanley, 1965, p. 300-301, pl. 45, figs. 8-13.

Triatriopollenites sp. of the T. coryphaeus type (13µm- 18µm), Tschudy, 1973, p. B16, pl. 4, figs. 1-3.

Mompites microfovveolatus (Stanley) Nichols, 1973, p. 107.

Description. Oblate triporate pollen grains, outline in polar view triangular with straight to slightly convex sides and rounded apexes; equatorial diameter 15-22 µm. Endexine about 0.3 µm thick; ektexine thickness on the order of 0.6 µm making a total exine thickness of about 1 µm. Sculpture a fine punctation with lumina approximately 0.3 µm wide. Pores meridionally elongated; dimensions 1 X 1.5 µm. Some specimens have arclike area that surrounds the pore region; this feature does not appear to be caused by the thickening of the exine but rather appears to be the result of more stain being accepted in this arclike region due perhaps to differences in exine chemistry.

Affinity. Pollen grains similar to but not formally identified as M. microfoveolatus were removed by Crepet et al. (1980) from a Juglandaceous catkin, Eoengelhardtia, preserved in the Claiborne Formation of Tennessee. They described these grains as small in size, averaging 14.8 µm in diameter, straight sided and triangular. Because of the floral and pollen morphologies, as well as the presence of large peltate scales, the authors believed this microfossil taxon to be closely allied with the modern Engelhardia-Oromunnea-Alfaroa complex. They further suggested that the genus, partially because of the small size of the pollen, was related to the Old World section Psilocarpeae.

Range. Frederiksen (1980a) gave the range for M. microfoveolatus as Cretaceous through Jackson in South Carolina and Uppermost Paleocene to Jackson on the Gulf Coast. Muller (1981) listed worldwide range of the M. fragilis type, including M. microfoveolatus, as Lower Campanian up to the Recent.

Occurrence. To 13% in 83 samples from San Miguel and in quantities between 1% and 11%, in all samples from Lake Somerville.

Triatriopollenites cf. T. coryphaeus (R. Potonié, 1931) Thomson and Pflug, 1953. Engelhardt 1964, p. 78, pl. 4, fig. 47.

Platycarya spp. Tschudy 1973, p. B14, pl. 2, figs. 30- 31.

Description. 10 to 25 µm, rarely somewhat larger. Exine smooth to scabrate, rigid, only with insignificant secondary folds. Contour convex-triangular. Pores never prominent. Exine cross section ribbon shaped, on all places equally thick. Ektexine always over 3X thicker than the endexine. Large atrium without intrapunctate structure. The Early Tertiary representatives sometimes appear endoplicate. (Translated from Thomson and Pflug)

Affinity. Elsik and Dilcher (1974) commented that the exines of their Platycarya grains are folded and thinned or split in a manner similar to extant Platycarya. According to Nichols (1973), "Fossil pollen exhibiting the true Platycarya group morphology can be assigned to the modern genus." (p. 105) Wing and Hickey (1984) showed that high Platycarya pollen percentages coincided with an abundance of Platycarya macrofossils in the sediment record at their site in Wyoming. They also figured Platycara pollen removed from anthers of staminate inflorescences of P. americana obtained from the Golden Valley Fm.

Paleoenvironmental Notes. Frederiksen (1985) summarized occurrences of Platycarya in the Early Tertiary and concluded that "...Platycara (or a close relative)..." may have lived near the shores of tropical seas... and quite possibly even in peat swamps." (p. 12). Other authors, transporting modern ecological tolerances back into time, attributed very high percentages in Tertiary lignite and lignitic detritus to long distance transport, assuming that Platycarya lived on dry hilly substrates (Gruas-Cavagnetto, 1976; Auffret and Grus-Cavagnetto, 1975). Wing and Hickey (1984), taking into account the abundance of Platycarya megafossils in the uppermost levels of their oxbow lake infill as well as modern ecological information, typified Early Tertiary Platycara as "...small, probably shrubby trees that rapidly colonized open or unstable ground, were fast growing and maturing plants...and may have formed thickets of dense, almost monospecific growth (p. 407)."

Occurrence. In values to 2% in three samples from San Miguel and absent from Lake Somerville.

Range. Platycarya appeared in the earliest Eocene and is one of the characteristic indicators of the Early Eocene throughout most of the Northern Hemisphere (Wolfe, 1973). Frederiksen (1980a) noted only one probable occurrence from the Upper Middle Eocene of Mississippi. Leopold and McGinitie (1972) citing Tschudy (1973) reported that Platycarya's last appearance in the Mississippi Embayment was in the Lower Jackson Group, "...just before a Late Eocene cool period. (p. 179)." Fairchild and Elsik (1969) noted that Platycarya is common in Claiborne sections.

Carya veripites Wilson and Webster 1946, p. 276, fig. 14.

Caryapollenites cf. C. simplex (R. Potonié, 1931) Raatz, 1937. Engelhardt, 1964, p. 78, pl. 5, fig. 51.

Carya sp. or Caryapollenites sp. of C. simplex (Potonié) Raatz 1937, Tshudy and Van Loenen, 1970, pl. 3, fig. 11.

Carya sp. or Caryapollenites sp., Tschudy and Van Loenen, 1970, pl. 3, figs. 12a-b.

Description. Grains circular to slightly triangular in equatorial view, oblately flattened; diameter 27-33 µm; germ pores three, round to broadly elliptical in shape, 3-4.5 µm long, all in one hemisphere, placed 3-4.5 µm from the equator; exine smooth, translucent, approximately 1.5 µm thick near equator; polar areas apparently thinner and sometimes corroded or broken, resulting in an irregularly modified area near the center of the grain.

Affinity. Carya (modern hickory).

Paleoenvironmental Notes. Frederiksen (1980a) reported these grains as infrequent to occasional in his Late Eocene samples. In the Oligocene Brandon lignites of Vermont, Carya was most abundant in the clastic samples beneath the lignite (Traverse, 1955).

There are 20 species of hickory now growing in the United States. Most species prefer the uplands, but one, Carya aquatica (water hickory) lives in the deep swamps of the southern coastal plains. (Harrar and Harrar, 1962) Modern-day Carya is wind pollinated, but according to Lewis et al. (1983), is large and not well adapted to anemophily.

Occurrence. Less than 1% in 19 samples from San Miguel and up to 2% in seven lignites from Lake Somerville.

Planera thompsoniana Traverse 1955

Planera thomsoniana Traverse, 1955, p. 52, fig. 10 (53)

Ulmus cf. Zelkova Gray, 1960, table 1 and fig. 1f.

Ulmipollenites cf. U. undulosus Wolff, 1934. Engelhardt, 1964, p. 79, pl. 5, fig. 58.

Ulmipollis sp., Tschudy and Van Lonen, 1970, pl. 3, figs. 16, 22, 25.

Description. Ca. 5-stephanoporate pollen grain with the pores strictly equatorial or nearly so, the pores with thickened rims, both of ektexine (annuli) and endexine (costae). Both hemispheres may have linear thickening in the exine, that is, arci. The sculpture is a low reticulum with wide muri. Sculpture much less pronounced in one hemisphere, this hemisphere often collapsed into the grain. Size: ca. 35 µm by 41 µm (Much flattened, disc-shaped.) Thickness of exine: ca. 2 µm.

Affinity. Frederiksen (1980a) noticed that these grains have arci which are typical of Planera and Zelkova.

Occurrence. Found at 1% from one sample from Sequence E of the San Miguel lignite. Absent from Lake Somerville.

Range. Found from the Upper Claiborne to Lower Vicksburg by Frederiksen (1980a).

Pollenites stellatus Potonié, 1931, p. 4, fig. 20.

Polyporopollenites stellatus (Potonié) Thomson and Pflug, 1953, 91-92, pl. 10, figs. 85-94.

Polyatrio-pollenites stellatus (Potonié) Pflug, 1953, p. 115, p. 24, fig. 47.

Pterocarya vermontensis Traverse, 1955, p. 45, fig. 9 (29).

Pterocaryapollenites stellatus (Potonié) Potonié, 1960, p. 132.

Pterocaryapollenites vermontensis (Traverse) Potonié, 1960, p. 132.

Pterocarya stellatus Martin and Rouse, 1966, p. 196, pl. 8, figs. 79-80.

Multiporopollenites sp., Tschudy and Van Lonen, 1970, pl. 3, fig. 33.

Description. Ca. 6-stephanoporate pollen grain, with the pores showing typical Juglandaceous tendency to have the pores more or less displaced off the equator into one hemisphere. Pores with thickened rims, that is, annuli. Exine sculpturing scabrate. Size: ca. 37 µm. Thickness of exine: ca. 1.6 µm.

Affinity. Pterocarya

Paleoenvironmental Notes. Frederiksen (1980a) found this taxon infrequently in his Jackson Group sediments. Leopold and McGinitie (1972) reported Pterocarya as "...common in the southeast and midcontinent. (p. 182)" by the Eocene. They noted that the modern Pterocarya "...requires summer rainfall, but can tolerate low annual rainfall and subhumid conditions. (p. 182)"

Occurrence. Less than 1% in one sample from San Miguel and absent from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg from the Gulf Coast (Frederiksen, 1980a).

Pollenites anulus (Potonié, 1931) Potonié and Venitz, 1934 [misidentified]. Engelhardt, 1964, p. 79, pl. 5, figs. 60-61.

Juglanspollenites sp., Tschudy and Van Loenen, 1970, pl. 3, figs. 29-30.

Multiporopollenites spp. Tschudy, 1973, p. B15, pl.3, figs. 20-22.

Nothofagus tschudyi Elsik, 1974, p. 290, 292-294, pl. 1, figs. 1-5; pl. 2, figs. 1-9.

Cf. Nothofagus Dombeyi Type. Elsik, 1974b, p. 2, fig. 44.

Celtis tschudyi (Elsik) Frederiksen, 1980a, p. 43, pl. 8, figs. 23-25.

Taxonomic notes. C tschudyii was later placed in the larger C. tschudyi group by Frederiksen (1988). The group encompasses a continuum of four species. He felt that "...the species seem to intergrade so much that they are impossible to differentiate consistently." (p. 51)

Affinity. Thought by Frederiksen (1980a) to be equivalent to modern Celtis.

Paleoenvironmental Notes. Grains of Triporopollenites pliktosus Anderson 1960, probably equivalent to Celtis, were more common in the clay layer of Potter (1976) from the Claiborne of Tennessee.

Celtis laevigata (Texas sugarberry) is an common component of sweetgum-oak floodplains in parts of the Texas Big Thicket (Ajilvsgi, 1979). Flenley (1979) says of the occurrence of Celtis in the East African Quaternary, "...common in moist and dry lowland forests and its pollen exhibits very high relative export. Abundance of this pollen type therefore indicates forest in the lowlands, but not necessarily near the pollen site." (p. 134)

Occurrence. Less than 1% in one sample from San Miguel and less than 1% in one lignite and the clastic sample from Lake Somerville.

Range. Frederiksen (1988) gives the range of the triporate members of the C. tschudyi "group" as Middle Upper Sabinian to Upper Claiborne.

Genus Malvacipollis Harris 1965 emend. Krutzsch 1966

Malvacipollis tschudyi (Frederiksen) Frederiksen 1980

?Aff. Nothofagus sp., Tshudy and Van Loenen, 1970, pl. 3, figs. 23, 27-28.

Echiperiporites spp. Tschudy, 1973, p. B15, pl. 3. figs. 13-14.

Echiperiporites tschudyi Frederiksen, 1973, p. 75, 78, pl. 2, figs. 19-22.

Malvacipollis tschudyi Frederiksen, 1980a, p. 44, pl. 8, fig. 27.

Description. Size 28-29 µm, mean 33 µm, holotype 35 µm. Originally spheroidal, outline round. Exine 1.5 µm thick, sexine:nexine 4:1; tegillate; ectosexine and endosexine equally thick. Sexine granulate and conate, the coni about 1 µm in diameter at the base, tapering evenly toward the point, and 1-2 µm high. Pores arranged only around the equator, although this is not immediately obvious because the grains were spheroidal and became compressed in many different orientations. Pores may be somewhat difficult to see; the grains are apparently 3- to 8-porate; holotype has 6 or 7 pores. Pores 1-2, µm in diameter; slight lanbra and (or) slight annuli present; vestibula 1-1.5µm deep; nexine thickened (to about 0.5 µm) under vestibula; no endospores evident.

Affinity. Frederiksen (1980a) originally hypothesized a Malvaceous sporophyte for M. tschudyi, but later (Frederiksen, 1988) noted that, because the sexine is not thickened under the spines, M. tschudyi probably belonged to a member of the Euphorbiaceae; he cites Tschudy (1973) as noting a resemblance to the family Picrodendraceae. Zamaloa and Romero (1990) and Martin (1974), referring to the southern hemisphere species of Malvacipollis, emphasize the difficulties of distinguishing between pollen of the families Malvaceae and Euphorbiaceae. Zamaloa and Romero felt comfortable, however, with, an assignment of their M. argentina grains to the Malvaceae.

Paleoenvironmental Notes. Frederiksen (1980a) found this taxon infrequently in his Claiborne and Jackson samples.

Modern Malvaceaean species in the Big Thicket of Texas range from herbs to shrubs. Hibiscus militaris is common in open area of levees. Kosteletzkya virginica (salt marsh mallow) is typical of coastal prairies and can be found in areas of seawater incursion and natural salt outcroppings. Callirhoe papaver (winecup) is a common wildflower in uplands. (Ajilvsgi, 1979) Malvaceous plants in British Guiana grow in swamps and marsh forests, along rivers, in strand scrub, and in mangrove along rivers. (Van der Hammen, 1963)

Occurrence. Less than 1% in five samples from San Miguel and absent from Lake Somerville.

Range. Frederiksen (1988) gave the range of Malvacipollis cf M. tschudyi as Lower to Middle Claibornian.

Description. Grains small, subtriangular in polar view with straight to slightly concave sides; equatorial diameter 19-24 µm. Pores usually close to the angles Exine 1.5 to 2.5 µm; sexine as thick as nexine; sculpture ill-defined, ?finely reticulate.

Affinity. Anacolosa in the Oleaceae according to Frederiksen (1981).

Occurrence. 1% from one sample from Sequence E at San Miguel. Absent at Lake Somerville.

Range. Frederiksen (1980a) found only one specimen from the Jackson of Alabama; Engelhardt (1964) reported examples of a species from the Claiborne. Cookson and Pike (1954) gave the genus range as Eocene.

Monosulcites asymmetricus Frederiksen 1973, p. 79, pl. 2, figs. 23, 28-29, 34-35.

Affinity. According to Frederiksen (1981), probably Palmae.

Paleoenvironmental Notes. Berry (1924) described numerous occurrences of palm pollen in the Texas Eocene, mostly in the Jackson age Fayette sandstone. These included Phoenicites?, a fragment of a pinnate palm leaf which could have belonged to a number of palm genera, from Fayette Sandstone localities near Millican and Wellborn, Brazos Co., TX; Palmoxylon lacunosum from numerous Texas locations; palm nut or Palmocarpon from the Yegua Fm. of Brazos County; and Palmocarpon sessile, a chunk of inflorescence from near Millican, Brazos County.

On their pollen diagram from Recent Borneoan peats, Anderson and Muller (1975) charted several palm genera derived from sporophytes commonly growing in peat swamps. These included Calamus, which grows along the margins of swamps; Cytostachus, which is abundant on shallow peat along the transition zone from mangrove and on the perimeter of swamps, and a species of Oncosperma which is common along inland margins of the mangrove belt and in riparian fringes.

Van der Hammen (1963) described a palm marsh forest in coastal deltas of British Guiana. Palms were found growing in swamp forest, rain forest, and on sand ridges. He pointed out that the genus Mauritia occurs in the low parts of savannas, in swamps, and on low river banks.

In his study of the Orinoco Delta, Muller (1959) described an open, bush-like palm swamp, "...with emerging clusters of palms, mainly Mauritia sp., Manicaria sacchifera, and Euterpe sp." This type of swamp is located in the central delta area, where the soil is permanently waterlogged, few fluctuations occur in water level, and pure peat is formed. Some palms also grow in the upper deltaic swamp forest, where there are more marked seasonal variations in water level. Mauritia also occurs as scattered, stunted individuals in the central deltaic, permanently inundated, "herbaceous swamp." This also is a peat forming environment. Muller suggested that the distribution of these vegetation types in the central deltaic back swamp represents a succession controlled by decreasing nutrient availability. The latter two vegetation types also grow on peat in the outer delta, where tidal influences are most important. According to Muller, the pollen percentages of Mauritia, which is a spiny, medium heavy grain, fluctuate strongly in relation to proximity of Mauritia populations, and reach 30% in the source area. He noticed a similar pattern with psilate palm types.

Occurrence. Less than 1% in five samples from San Miguel and absent at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg on the Gulf Coast (Frederiksen, 1980a).

Genus Monocolpopollenites Pflug & Thomson in Thomson & Pflug 1953 emend. Nichols et al. 1973

Monocolpopollenites tranquillus (Potonié) Thomson and Pflug 1953

Pollenites tranquillus Potonié, 1934, p. 51, pl. 2, figs. 3, 8.

Monocolpopollenites tranquillus (Potonié) Thomson and Pflug, 1953, p. 62-63, pl. 4, figs. 24-37, 39-47.

Palmaepollenites tranquillus (Potonié) Potonié, 1958, p. 97, pl. 11, fig. 138.

Monosulcites sp., Tschudy and Van Loenen, 1970, pl. 1, fig. 15.

Description. 20-45 µm. Exine to 2 µm thick, always scabrate or intrapunctate. The equator has a characteristic asymmetry four to six true, loop formed contour, with rounded-off corners. Two corners are situated on the intersection with the colpus-plane, one close (or shut) by that an acute one as the other. The colpus lies not exactly symmetrically. Colpus bulge about 1 µm wide, itself towards the end scarcely tapered. Colpus seldom gaping. Figure tabular. (Translated from Thomson and Pflug)

Affinity. Palmae, according to Frederiksen (1980a).

Occurrence. To 4% in 24 samples from San Miguel and up to 4% in seven lignites at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg of the Gulf Coast (Frederiksen, 1980a).

Genus Sabal Adanson

Sabal cf. S. granopollenites Rouse 1962

Sabal granopollenites Rouse, 1962, p. 202, pl. 1, figs. 3-4.

Sabal cf. S. granopollenites Rouse. Frederiksen 1980a, p. 45, pl. 9, figs. 6-8.

Description. Pollen monocolpate, fusiform in outline, coarsely granulate to weakly reticulate. The single colpa is long and narrow with weak margins. Size range 28-32 µm.

Affinity. Engelhardt (1964) noted a strong resemblance of his Sabalpollenites to modern Sabal sp. (Palmae), or palmetto.

Paleoenvironmental Notes. Frederiksen (1985) considered Sabal to be a widespread member of the shrub layer during the Eocene. Daghlian (l978) reported Sabal dortchii Daghlian megafossils, identified by cuticle to the genus Sabal, from the Middle Eocene of Tennessee. Palm leaf fossils were closely associated with backswamp or flood plain deposits and abandoned channel sediments. A. Raymond (oral commun., 1993) suggested that Sabal-type palm macrofossils may be contained in channel sediments from Welch Park at Lake Somerville. Potter (1976), in his investigation of a oxbow lake in the Middle Eocene of Tennessee, interpreted the increase of Sabalpollenites as an invasion of palms into a lowland community.

Dilcher (1973) remarked, "There are several modern species of Sabal, nearly all of which have been grown in Florida and observed to recover very well from rather severe winter frosts. Thus the use of palms as an indication of a tropical or subtropical climate is open to question." (p. 43)

Berry (1924) identified macrofossils found in the Fayette Sandstone (Jackson Group) of Webb County, Texas as Sabalites vicksburgensis. He noted that it resembled in habitat the modern Sabal palmetto by which he meant it preferred environments near the coast.

Fan palms, like Sabal palmetto, are more common in subtropical zones of Central America, but can occur in tropical zones (Holeridge et al., 1971). The most common habitat is sandy or swampy coastal regions (Corner, 1966). They are most abundant where the mean annual temperature exceeds 24 degrees C, but a few grow in cooler forests. Frederiksen (1980c) reports that Sabal grows in summer-dry and/or arid subtropical, humid subtropical, winter-dry tropical, and wet tropical climates. Ajilvsgi (1979) described palmetto oak flats in the Big Thicket of Texas, which occupy the lowest, flattest terrain of the Big Thicket including broad swamplands and tortuous bayous. Dwarf palmetto is the most distinctive plant in this association, and may cover the forest floor with little or no vegetation between. In this environment, water stands for fairly long periods of time, and in the summer, evaporation causes large areas of cracked clayey soils be exposed.

Sabal secretes nectar freely, and is visited by bees (Lovell, 1926).

Occurrence. To 8% in 32 samples from San Miguel and 1% in one lignite sample from Lake Somerville.

Range. Claiborne to Lower Jackson of the Gulf Coast (Frederiksen, 1980a).

Genus Arecipites Wodehouse 1933 emend. Nichols et al. 1976

Arecipites columellus Leffingwell 1971

Sabalpollenites cf. S. convexus Thiergart, 1938. Engelhardt, 1964, p. 71. pl. 2, fig. 14.

Monosulcites sp., Tschudy and Van Loenen, 1970, pl. 1, figs. 10, 14.

Arecipites columellus Leffingwell, 1971, p. 40-41, pl. 7, figs. 1-2.

Description. Monosulcate, sulcus extending the entire length of the grain, never expanding at the ends; shape elongate ellipsoidal; exine 1.2 to 1.5 µm thick, and thinnest adjacent to the sulcus; exine partially tectate, forming distinct ectosexinous and columella layers that are best seen in optical section, sexine thicker than nexine; reticulate, lumina about 0.5 µm wide and of uniform size; grain size (15 specimens) 30 to 40u X 20 to 25 µm.

Affinity. According to Frederiksen (1980a), "this species is identical with modern pollen of Serenoa serrulata (Michx.)(Palmae)(p. 45)". Arecipites grains in the San Miguel and Lake Somerville deposits are fairly similar to S. serrulata pollen pictured by Kedves (1980, p. 363).

Paleoenvironmental Notes. Elsik (1978) interpreted a preponderance of Calamuspollenites-Arecipites pollen as indicating an herbaceous community inhabiting Calvert Bluff (Paleocene) and Yegua-Jackson marshes in the absence of the still-evolving forbs. Potter (1976) hypothesized that the increase of Arecipites in the upper part of a lignite seam represented an invasion of palm trees during the final stage of a Middle Eocene oxbow lake. Farley (1990) noted a low of Arecipites in the levee-crevasse splay environment of the Early Eocene of Wyoming.

Serenoa, or saw palmetto, is a branching palm closely resembling and with habitats similar to that of Sabal (Corner, 1966). Frederiksen (1980c) reported Serenoa as growing in humid subtropical and winter-dry tropical regions.

Lovell (1926) mentions Serenoa repens as a honey producer.

Occurrence. Appears in 114 levels in values to 39% at San Miguel; to 14% in 15 lignites and the clastic sample at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg of the Gulf Coast (Frederiksen, 1981).

Calamuspollenites eocenicus Elsik and Dilcher, 1974, p. 74, pl. 28, figs. 66-67.

Description. Bilateral pollen grains with one furrow which reaches the apices; furrow is closed to open. Furrow margin when split is thin and recurved. Exine of three layers; outer layer punctate with scabrate-microrulgulate ornamentation. Dimensions about 20 X 30 µm. Furrow width depends on whether or not the exine is split and recurved and degree of compression and folding of grain. Exine slightly thicker than 1 µm (but less in floor of furrow). Foot layer or endexine and columellate layer about equal thickness; less than 0.5 µm each. Tectum very thin, punctae very small, microrugulate 0.5 µm or less wide and generally very short.

Affinity. Probably Palmae (Frederiksen, 1980a)

Occurrence. Less than 1% in one sample from Sequence C of the San Miguel Lignite. Less than 1% in two lignites from Lake Somerville.

Range. Claiborne to Jackson (Frederiksen, 1980a)

Affinity. According to Frederiksen (1983), the genus Liliacidites represents "Probably mainly Palmae" (p. 46), but he added the possible choices Liliaceae, Amaryllidaceae, and Iridaceae. Frederiksen believed that it was impossible to determine on the basis of pollen alone which genera of Palmae produced individual species of Middle Eocene Liliacidites. Engelhardt (1964) noted that Couper (1953) erected this genus for specimens which could not be placed more accurately in other genera, and that pollen with similar morphology occurs in the Palmae.

Paleoenvironmental Notes. Elsik (1978) used sparse Liliacidites occurrence in the Calvert Bluff (Paleocene) and common Liliacidites occurrence in the Yegua-Jackson to indicate a marsh environment. The "marsh assemblage" was only a short episode in the beginning of peat development. Two species of Liliacidites were found by Potter (1976) to be conspicuously present at the top of a Middle Eocene lignite. This was interpreted as an invasion of palms surrounding a lowland community surrounding a freshwater pond, a late stage of the infilling of an oxbow lake. Farley (1990) found the highest amounts of Liliacidites in swamp-facies carbonaceous shales in his study of the Early Eocene of Wyoming.

Range. From near the top of the Tallahatta Fm. (Middle Eocene) into the Oligocene (Frederiksen, 1988).

Liliacidites variegatus Couper, 1953 [misidentified]. Engelhardt, 1964, p. 71, pl. 2, figs. 13-16.

Liliacidites sp., Tschudy and Van Loenen 1970, pl. 1, fig. 16.

Liliacidites tritus Frederiksen, 1973, p. 80-81, pl.3,

figs. 13-16.

Liliacidites tritus Frederiksen, 1980a, p. 46, pl. 9, figs. 14-15.

Description. Size 32-39 µm, mean 34 µm, holotype 33 µm. Outline more or less oval; may be slightly polygonal or asymmetrical;ends rounded or slightly pointed; length /width 1.3-1.9. Exine about 0.3 µm thick excluding ornamentation; distinctly reticulate, the muri clavate in optical section and simpli- to duplibaculate in design. Muri 1 µm high and 0.7-1 µm wide; lumina round to polygonal to oval, 0.5 x 0.5 2-0.5 x 1 µm in size on proximal side; lumina may be slightly smaller on distal side near sulcus. Sulcus extends full length or nearly full length of grain; sulcus gaping, slightly open all along its length or edges overlapping; edges of sulcus turned up so that the edges look thickened.

Affinity. L. tritus is "Very similar to modern pollen of Pseudophoenix sp.," according to Frederiksen (1980a, p. 46.)

Occurrence. To 2% in 18 samples from San Miguel. To 7% in two lignites at Lake Somerville.

Liliacidites sp., Tschudy and Van Loenen, 1970, pl. 1, figs. 14-15.

Liliacidites vittatus Frederiksen, 1973, p. 80, pl. 3, figs. 1-5.

Liliacidites yeguaensis Elsik, 1974b, p. 98, pl. 1, figs. 14-15.

Liliacidites vittatus Frederiksen. Frederiksen, 1980a, p. 46, pl. 9, figs. 16-17.

Description. Size 28-38 µm, mean 34 µm, holotype 35 µm. Broadly lenticular in outline, the ends slightly pointed; length/width 1.1-1.5. Reticulate; muri in optical section ribbon-like, discontinuous and clavate, with the clavae 0.5-3 µm apart; between clavae, muri are supported by thin bacula. Exine including reticulum 2 µm thick; exine proper 0.5-.07 µm thick;exine proper 0.5-0.7 µm thick, baculate interval 0.3-0.5 µm high, muri 0.7-1 µm high. Muri 1 µm wide and duplibaculate in design; lumina polygonal, about 2-3 µm in diameter on proximal face, 10-12 lumina present down length of grain; lumina may be smaller near sulcus. Sulcus extends full length or nearly full length of grain; usually slightly open, without thickened margins.

Affinity. Not known.

Occurrence. To 19% in 97 samples from San Miguel. To 15% in 15 lignites and the clastic sample at Lake Somerville.

Description. Oval, flattened along the polar axis, generally separated by a continuous equatorial aperture in two more or less equal parts; borders of apertures somewhat irregular and with a torn appearance, with reduced ectexine thickness; wall + 1 µm thickness, surface bearing scattered spines; endexine thin, <1 µm thick; columellae fused into a finely reticulate pattern; muri < 1 µm; lumina rounded, even in size, < 1 µm across; spines 5-7 µm long, uniformly spaced, conical, with a fairly blunt, rounded apex and an often slightly expanded base, flat underneath, apparently resting on separated columellae. 33 (35) 43 µm (8 specimens).

Affinity. According to Muller (1968), these grains have "...identity with the pollen of the palm genus Nypa without reasonable doubt." (p. 12)

Paleoenvironmental Notes. Because S. echinatus is almost certainly the pollen of Nypa, a mangrove palm, it has been used as an indicator of a marine-influenced, saline environment. Srivastava and Binda (1991), in a study of the early Tertiary of Saudi Arabia, stressed the association of S. echinata with open-marine dinoflagellates. They believed their S. echinata mangrove to be part of a continuum of tropical estuarine vegetation. It was included in the most marine "sea-proximal zone," an "...area under the influence of the highest and lowest tides..." and "...supporting mangrove vegetation." (p 57). Nypa was the "most distinctive" palynomorph in a lignite from the Claiborne Group of Webb Co., TX (Westgate and Gee, in press). The presence of Nypa suggested to these authors tropical to subtropical conditions, as well as an intertidal area with water temperatures greater than 24 degrees C. S. echinatus was also an important constituent of an early to Middle Eocene (?) intertidal sequence from Morocco (Fechner, 1988). Again, the presence of S. echinatus indicated a mangrove swamp environment.

Anderson and Muller (1975) found Nypa pollen only in clays in the basal portion of their profile from a modern swamp in Sarawak. Flenley (1979) notes that Nypa grows in brackish water, usually at the landward side of mangrove swamps.

Flenley (1979) also gave the present distribution of Nypa as mainly Ceylon, Assam, Burma, Malesia, and China.

Occurrence. Absent at San Miguel and 1% in the clastic sample at Lake Somerville.

Range. Nypa has been recorded from the Paleocene/Eocene of West Africa, France, Britain, Poland, Southern U.S.S.R. [sic], Egypt, Central India, Borneo (pollen), Brazil, and Texas, according to Flenley (1979). Citing Muller (1970), he charted the first appearance of Nypa pollen in the Senonian.

Pollenites liblarensis Thomson in Potonié et al., 1950, p. 55, pl. B, figs. 26-27.

Tricolpopollenites liblarensis (Thomson) Thomson and Pflug, 1953, p. 96, pl. 11, figs. 111-132.

Cupuliferoidaepollenites liblarensis (Thomson) Potonié, 1960, p. 92, pl. 6, fig. 94.

Tricolpopollenites liblarensis (Thomson) Thomson and Pflug, 1953. Tschudy, 1973, p. B18, pl. 4, figs. 31- 33.

Description. 20-30 µm. Outline spindleshaped or ellipsoidal. Width-length index between 0.5 and 0.8. Exine nearly smooth to vaguely interrugulate.

Affinity. Frederiksen (1980a) suspected that Cupuliferoipollenites pollen was Fagaceous, also Leguminosae in part. Crepet and Daghlian (1980) removed a very similar pollen type from a Castaneoid (Fagaceae) inflorescence from the Claiborne Group of Tennessee. These grains were prolate with an average polar diameter of 15.0 µm and an average equatorial diameter of 9.0 µm (P/E=1.67). SEM photography showed a surface consisting of striate, anastomosing fused ridges characteristic of modern Castaneoids. Leaves of the extinct Fagaceous genus Dryophyllum and castaneoid fruits were found in the same sediments.

Paleoenvironmental Notes. Cupuliferoidaepollenites grains dominated Potter's (1976) clay zone from the Claiborne of Tennessee, with percentages of over 60% in this zone. He believed that this type represented "...waterborne and windborne background and local pollen derivation" (p. 83) One upper Eocene lignite seam from Hungary was co-dominated was co-dominated by Cupuliferoidaepollenites liblarensis and Cupuliferoipollenites cingulum. (Kedves, 1965). Pflug (1952) found large proportions of C. liblarensis associated with the brown coal seams in Germany. Frederiksen (1985) described Pflug's Interpretation: "Rapid subsidence caused flooding of the swamp. The water was too deep for trees to grow. The main sporomorph type in the marsh or shallow lake deposits was the Cupuliferoidaepollenites liblarensis type, produced by Fagaceous trees in the closed forest outside the swamp." (p. 58)

Anderson and Muller (1975) figured up to 30% Lithocarpus/Castanopsis in a profile from a modern Sarawak peat swamp. The high percentages were found in a zone dominated by Shorea albida, which represents roughly a midpoint in the plant succession. Pollen of the Fagaceous taxa are found in percentages to about 10% in surface samples representing the lower half of the successional sequence.

Many modern species of the Castaneoidea are thought to be insect pollinated (Crepet and Daghlian, 1980); Lithocarpus is primarily entomophilous (Lewis et al, 1980; Flenley, 1979). According to Flenley, both Lithocarpus and Castanopsis are common in Malesian lower montane forests. Castanopsis is wind-pollinated and exports large amounts of pollen.

Occurrence. To 81% in 134 levels at San Miguel and to 18% in 13 lignites and the clastic sample at Lake Somerville.

Range. Frederiksen (1981) found this species in samples from the Upper Claiborne to the Lower Vicksburg.

Foveotricolpites prolatus Frederiksen, 1973, p. 81, 84, pl. 3, figs. 17-22

Affinity. According to Frederiksen (1981), similar to pollen of modern Spartium junceum L. (Leguminoseae).

Occurrence. 1% in four samples from San Miguel. Absent at Lake Somerville.

Range. Claiborne to Lower Vicksburg (Frederiksen, 1981)

Paleoenvironmental Notes. The Quercoideae were well-developed by the beginning of the Oligocene, when, because of a climatic deterioration and a sudden drop in sea level, they began to dominate the flora. At one Catahoula Fm. (Oligocene) locality, the oak pollen

comprised 85% of the palynoflora (Daghlian and Crepet, 1983). The authors believed that this evolutionary success was due to hybridizations caused by Eocene climatic change.

Modern-day temperate oaks are wind pollinated, but tropical oaks are primarily insect pollinated (Lewis et al., 1983). The various temperate oak species may live anywhere from dry, sandy soils to swamps, as with Quercus bicolor. According to Flenley (1979), Quercus grains have "high relative export."

Quercoidites sp. was characteristic of the clay zone in Potter's 1976 study of the Tennessee Claiborne.

Quercoidites inamoenus (Takahashi) Frederiksen 1980

Tricolpopollenites inamoenus Takahashi, 1961, p. 313, pl. 22, figs. 42-49.

Quercoidites cf. Q. henrici (R. Potonié, 1931) Potonié, Thomson, and Thiergart. Engelhardt 1964, p. 71, pl. 2, fig. 15.

Quercus sp., Fairchild and Elsik, 1969, p. 84, pl. 37, fig. 22.

Tricolpopollenites sp., Tschudy and Van Loenen, 1970, pl. 4, fig. 2.

Quercus sp., Elsik, 1974b, pl. 4, fig. 2.

Quercoidites inamoenus (Takahashi) Frederiksen, 1980a, p. 47, pl. 10, figs. 3-8.

Affinity. Frederiksen (1980a) assigned this highly variable species to Quercus or to the extinct Fagaceous genus Dryophyllum. He included a large range of colpus-pore forms within this species, including both tricolpate forms with simple colpi or with geniculi, and tricolporate forms, with ora present as slits. These tricolporate forms have been isolated by Daghlian and Crepet (1983) from oak catkins from the Oligocene Catahoula Formation near Huntsville, Texas. C. Ferguson (oral commun., 1988) removed similar pollen from shales at the same site. This tricolporate "oak" pollen is not readily recognizable as oak either to myself nor to other palynologists experienced with Quaternary oak pollen (J. Jones, 1988, oral commun.; S. Hall, 1988, oral commun.), nor does it resemble modern oak pollen pictured by Lieux (1980).

Occurrence. To 10% in 96 levels at San Miguel and to 3% in four lignites and the clastic sample at Lake Somerville.

Range. From just above the base of the Lisbon Fm. (Middle Claiborne) into the Oligocene (Frederiksen, 1988).

Pollenites microhenricii Potonié, 1931, p. 26, pl. 1, fig. V19c.

Pollenites henrici microhenrici (Potonié) Potonié and Venitz, 1934, p. 27.

Tricolporopollenites microhenrici (Potonié) Thomson and Pflug, 1953, p. 96, pl. 11, figs. 62-110.

Quercoidites microhenrici (Potonié) Potonié, 1960, p. 93.

Description. 20-30 µm. Outline spindleshaped. Width-length index ca. 0.5, but also somewhat more or less. Very complex, here and there likely a larger quantity of botanical units are embraced. lets separate from one another scarcely. Viewed beside size and shape is the intergranulate or intrabaculate structure. Ektexine and endexine equally strong. Several have geniculi.

Affinity. Frederiksen (1980a) speculated that Q. microhenricii was derived from a Fagaceous sporophyte.

Occurrence. To 57% in 101 levels from San Miguel and to 2% in two lignites and the clastic layer at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1981). Elsik (1968a) reported this taxon from the Paleocene Rockdale lignite.

BT-68, ??Rubiaceae?? Traverse, 1955, p. 75, fig. 13 24 (138).

Tricolpites sp. 3 Engelhardt, 1964, p. 72, pl. 2, fig. 19.

Tricolpopollenites sp., Tschudy and Van Lonen, 1970, pl. 4, figs. 9, 14.

Fraxinoipollenites spp., Frederiksen, 1980a, pl. 10, figs. 14-17.

Fraxinoipollenites medius Frederiksen, 1980a, pl. 10, figs. 11-12.

Description. Tricolporate pollen grain with long furrows quite unmodified except for slight margos. Reticulate sculpture with wide muri and large lumina. Reticulum has columellate structure.

Affinity. Probably not Fraxinus, according to Frederiksen (1980a).

Paleoenvironmental Notes. Fraxinus megafossils are common in Claiborne Group sediments, but dispersed pollen grains in the same beds assigned to Fraxinus by Taylor (1987) are not of this type (Call and Dilcher, 1992).

Small Species Group

Paleoenvironmental Notes. Modern Fraxinus is wind pollinated (Lewis et al., 1983)

Occurrence. To 12% in 85 samples from San Miguel and to 4% in 11 lignites and the clastic sample at Lake Somerville.

Large Species Group

Occurrence. Up to 5% in 26 samples from San Miguel. 1% in one lignite sample and 2% in the clastic sample at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1981)

Tricolpites sp. 2. Engelhardt, 1964, p. 72, pl. 2, fig. 18.

Tricolpopollenites sp., Tschudy and Van Loenen, 1970, pl. 4, figs. 3-6, 10, ?12.

Tricolpites n. sp. A (microreticulate) Tschudy, 1973, B13, pl. 2, figs. 11-12 only.

Platanus occidentalis Frederiksen, 1980a, pp. 48-49, pl. 10, fig. 19.

Affinity. According to Frederiksen (1980a), P. occidentaloides is "Very similar to Platanus occidentalis L. except that the fossils are slightly larger and the colpi are slightly deeper than in the modern grains." (p 49). My few specimens, as well as the one illustrated by Frederiksen (1980a) vary markedly from the modern specimen pictured by McAndrews et al. (1978), especially in relation to the depth of the colpus.

Description. Polar axis 26-35 µm, equatorial axis (in polar view) 22-35 µm, holotype 31 µm. Tricopate. Spheroidal to subprolate; broadly rounded at the poles. Exine 1.25-1.5 µm thick including reticulum; "nexine":"endosexine":"ectosexine" ratio about 1:15:1. Lumina a little less than 0.5 µm in diameter. Colpi extend 2/3-3/4 length of grain and are moderately deeply incised; colpi appear narrow in equatorial view but gape widely in polar view; edges of colpi very rough and sometimes beaded; margins thickened little if any.

Paleoenvironmental Notes. Macrofossils of Platanus were found in the Eocene Clarno Nut Beds of Oregon. The wood and fruit morphology were similar to those of modern specimens, but the leaves were primitive (Manchester, 1981). P. neptuni, found in the Eocene and Oligocene of Central Europe, co-occurred with Dryphyllum and Laurophyllum and was often accompanied by evergreen arborescents and drought resistant species (Hably, 1980).

In Frederiksen's (1981) Eocene samples from the southeastern United States, Platanus pollen was more common in clastics and was thought more likely to have come from trees outside the peat-forming environment.

Platanus (sycamore) trees grow in most of eastern north America, in moist or wet alluvial sites (Gleason and Cronquist, 1963). It is "...one of the commonest of streambank and bottomland trees...(p. 197)", according to Harlow (1957). P. occidentalis is a pioneer plant growing below levees and on low banks in the Big Thicket of Texas (Ajilvsgi, 1979). Frederiksen (1980b) reported that Platanus grows in summer-dry subtropical and or semi arid, humid subtropical, and winter-dry tropical regions. Members of the Platanaceae are wind pollinated and release large amounts of pollen (Lewis et al., 1983).

Occurrence. Less than 1% in 20 samples from San Miguel and 1% in one sample from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a). Not found below the uppermost Claiborne by Tschudy (1973).

Salixipollenites parvus Frederiksen, 1980a, p. 49, pl. 10, figs. 20-27.

Description. Size 15-24 µm, mean 20 µm, holotype 16 µm. Tricolpate. Subprolate to prolate; broadly rounded at the poles. Exine about 1 µm thick including ornamentation. Reticulum medium coarse in relation to small size of the grain--lumina are 0.5-1 µm in diameter. Muri slightly less than 1 µm high and about 0.5 µm wide, clavate in optical section and distinctly simplibaculate in design. Colpi extend 2/3 to 3/4 length of grain, inner edges of colpi appearing thickened.

Occurrence. To 4% in 47 levels at San Miguel and to 6% in 13 lignites from Lake Somerville.

Range. Near the top of the Lisbon Fm. into the Oligocene (1988).

Fraxinus? pielii Frederiksen, 1980a, p. 49, pl. 10, figs. 28-32.

Description. Size 24-33 µm, mean 28 µm, holotype 23 X 26 µm. Oblate; outline square, with sides slightly to moderately convex. Tetracolpate, colpi 1.5-3 µm deep, margins lacking. Exine 1 µm thick; tegillate; ectosexine:endosexine:nexine ratio about 1:1:1, finely infrareticulate to finely infragranulate, the lumina or grana 0.3-0.5 µm in diameter.

Affinity. Frederiksen (1980a) observed that this species has a finer reticulum than modern Fraxinus.

Occurrence. To 1% in four samples from San Miguel. Absent at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Albertipollenites? aranosea Frederiksen 1973, p. 84, pl. 3, figs. 30-34.

Rousea araneaosa (Frederiksen) Frederiksen 1980, p. 49, pl. 10, figs. 33-34.

Description. Size 21-37 µm, mean 29 µm, holotype 33 µm. Prolate, outline oval, poles broadly to narrowly rounded. Tricolpate, colpi extending nearly full length of grain, ends of colpi typically broadly rounded, colpi margins slightly thickened. Bases of colpi sometimes torn or ragged-some specimens appear to be raggedly tricolporate. Exine excluding ornamentation 1 µm thick; sexine-nexine 3:1. Reticulate, the muri clavate in optical section, clavae 0.7-1 µm high, muri 0.3 µm wide, lumina 1.5-2.5 µm in diameter.

Affinity. Frederiksen gave the affinity of R. araneosa as "...probably Bignoniaceae. (p. 49)".

Paleoenvironmental Notes. One member of the Bignoniaceae, Catalpa bignonioides, lives on riverbanks and swamp margins in the southeastern United States. Enallagma latifolia (black-calabash) inhabits rich, moist soil near salt water in Florida and the Caribbean. The family also includes several vines. (Harrar and Harrar, 1962)

Occurrence. Less than 1% in 11 samples from San Miguel and 1% in one lignite from Lake Somerville.

Range. Middle of the Lisbon Fm. (Middle Eocene) into the Oligocene (Frederiksen, 1988).

Tricolpopollenites sp., Tschudy and Van Lonen 1970, pl. 4, figs. 20 a-b.

Rousea monilifera Frederiksen 1981, pl. 10, figs. 20 a- b.

Description. Size 36-45 µm, mean 40 µm, holotype 41 µm. Tricolpate. Subprolate to prolate, rarely spheroidal; broadly rounded at poles. Exine 0.5-1 µm thick excluding ornamentation. Coarsely reticulate; muri 2-3 µm high, clavate in optical section, heads of clavae rounded or more often radially elongate; muri 0.5-0.8 µm wide and coarsely beaded in design (simplibaculate), the beads 0.7-1 µm in diameter, that is, of greater diameter than the width of the muri. Muri may be somewhat discontinuous. Lumina about 2-3 µm in diameter, except those near the colpi, which are only about 1 µm; lumina polygonal to rectangular. Colpi deeply invaginated, extending nearly full length of grain, 0.5-2 µm wide, with edges not thickened.

Affinity. According to Frederiksen (1981), very similar to Armeria (Plumbaginaceae); also similar to Amanoa (Euphorbiaceae), according to Elsik and Dilcher (1974).

Occurrence. 1% in four samples from San Miguel; absent from Lake Somerville.

Striatopollenites terasmaei Rouse, 1962, p. 212, pl. 4, figs. 30, 35.

Striatopollis terasmei (Rouse) Frederiksen, 1980a, p. 50, pl. 11, fig. 6.

Description. Pollen grains elliptical in outline. Tricolpate, with the colpas narrow and simple. Exine striate, with some 20-30 longitudinal ridges covering the surface. The ridges are about 2µm apart and curve inward at the poles, as in species of Schizaea and Ephedra. Size range 24 to 26 µm.

Occurrence. Less than 1% in four levels from San Miguel. Absent from Lake Somerville.

Range. Bashi-Hatchetigbee Fm. (Late Sabininan, Lower Eocene) into the Oligocene (Frederiksen, 1988)

Cupuliferoipollenites cf. C. insleyanus (Traverse, 1955) R. Potonié, 1960. Engelhardt, 1964, p. 72-73, pl. 2, fig. 23.

Castanea sp., Fairchild and Elsik, 1969, p. 83, pl. 37, fig. 6.

Description. 10-30 µm, most not over 28 µm, Outline spindle-shaped to cylindrical, also egg-shaped. Not spherical or biconical. Cavernae + parallel up to the poles extending. Exine smooth or weakly interrugulate.

Affinity. Frederiksen (1981), citing his own unpublished data, stated that Cupuliferoipollenites came from "Mainly Dryophyllum; (an extinct genus of Fagaceae)...perhaps few of these grains were produced by Castanea or and (or) Castanopsis." (p. 50) Recently, Jones (1988) pointed out that the type specimen of Dryophyllum should be assigned to the Juglandaceae, however, he added that most of the species included in the genus Dryophyllum now need to be reassigned to an undesignated Fagaceous genus. Nichols (1970), who examined modern grains of Castanea, felt that his C. cingulum grains (probably equivalent to Frederiksen's Cupuliferoipollenites spp.) were without a doubt referable to that modern genus. Engelhardt (1964) also mentioned the resemblance of this taxon to Castanea. For a further discussion of Cupuliferoipollenites, see the section on "Scenario 3: Mangroves" in the chapter on "Paleoecology."

Taxonomic Notes. Frederiksen (1984, written communication) refers to the taxa Cupuliferoipollenites, Siltaria, and Araliaceoipollenites as "...problematic." They form a sculptural continuum, and have very similar morphology. Cupuliferoipollenites grains are, by his definition, psilate.

Paleoenvironmental Notes. Pflug (1952) believed that "Castanea" pollen from German brown coals came from low trees or shrubs because of its limited dispersal in the sediments. Cupuliferoipollenites made up generally 69 or more percent of lignite assemblages in Potter's study of the Claiborne of Tennessee. It was also present at levels up to 45% in the clay layer. He thought that Cupuliferoipollenites was locally derived. Dilcher (1973) discussed the taxon in Tennessee clay samples, offering the opinion that "The abundance of Castanea pollen in all of the clay samples studied suggests that this pollen came from wind-pollinated trees. According to Faegri and Van der Pijl (1966), tropical species of Castanea are insect-pollinated and would not be expected to yield abundant pollen. Thus the abundance of Castanea pollen in these sediments is an indirect indication that a somewhat temperate climate was present in the Mississippi embayment area during the Middle Eocene." (p. 55-56)

Farley (1990), in a study of palynomorph distribution in Early Eocene rocks of Wyoming, found the highest occurrence of Cupuliferoipollenites in the pond facies. He considered Cupuliferoipollenites to have been allochthonous, but possibly of local origin.

Pflug (1952) found an abundance of Cupuliferoipollenites cingulum in a German Early Eocene lignite. His interpretation is reported by Frederiksen (1985): "Subsidence slowed, the level of the peat surface rose relative to ground-water level, and a low open forest formed a swamp vegetation that produced mostly fagaceous pollen of the Cupuliferoipollenites cingulum type plus some triporate, palm, and other pollen" (p. 58).

In the Late Oligocene Brandon lignites of Vermont, Castanea was found only in the lignites and not in the silt and clay layers (Traverse, 1955). Percentages were low, only to 4.6%.

Berry (1924) described leaf macrofossils of Dryophyllum brevipetiolatum as exceedingly common in the Fayette sandstone of East Texas. Some of his specimens were collected in Trinity and Grimes Counties, Texas.

Frederiksen (1980c) reports modern Castanea/Castanopsis from summer dry subtropical and/or semiarid, humid subtropical, winter-dry tropical, and wet tropical regions.

Occurrence. Present in all samples from San Miguel, with percentages to 75% and to 63% in all samples from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Genus Chrysophyllum Linnaeus

Chrysophyllum brevisulcatum (Frederiksen) Frederiksen 1980

Cupuliferoipollenites brevisulcatus Frederiksen, 1973, p. 85, pl. 3, figs. 28-29.

Chrysophyllum brevisulcatum Fredericksen, 1980a, p. 51, pl. 11, fig. 12.

Description. Size 14-21 µm, mean 17 µm, holotype 19 µm. Prolate; sides straight and poles broadly rounded. Tricolporate; colpi about 0.5 µm wide, extending only 1/2-3/5 length of grain, margins not thickened; ora lalongate, typically 1.5-2 x 4-4.5 µm. Exine 0.5-0.7 @ thick at poles and about 1 µm thick at the equator; thickening produces darkened equatorial band 6-7 µm wide. Nexine everywhere very thin. Exine psilate.

Affinity. Frederiksen (1980a) believed this pollen type to be morphologically identical to Chrysophyllum.

Paleoenvironmental Notes. Sapotaceous pollen was common in parts of the German Brown coals (Frederiksen, 1985). Frederiksen (1980a) found Chrysophyllum to be "infrequent" or "occasional" in a few of his Late Paleogene samples from the Gulf Coast; up to 40% were found in some Jackson lignites. He hypothesized that the grains had poor regional dispersal and may have been low plants.

Van der Burgh (1967) found wood and pollen of the Sapotaceae to be equally well represented in the Miocene Rhenish brown coals. Berry (1924) identified leaf fossils from the Fayette Sandstone from near Wellborn, Brazos Co., TX as Chrysophyllum preoliviforme, resembling the extant C. oliviforme of South Florida.

Chrysophyllum is a member of the family Sapotaceae, which contains 50 genera of trees and shrubs, eight of which are found in the subtropical forests of the United States. Chrysophyllum, or satinwood is a tropical genus with 65 species of evergreen trees and shrubs. The genus currently grows in southern Florida, where favored habitats are hammocks, flatwoods, and sandy plains (Harrar and Harrar, 1962). Of the three species of Chrysophyllum found in Costa Rica, all are canopy trees. Life zones include tropical moist, tropical wet, premontain rainforest, and premontane wet (Holdridge et al., 1971). Frederiksen (1980c) reported the genus in humid subtropical, winter-dry tropical, and wet tropical regions. Pennington (1990) reported 43 species in the Neotropics. Three of these species grow in periodically flooded habitats in the Amazon Basin. A few others live in wet forests, but the majority are dry land plants.

Pollination is entirely zoophilous (Frederiksen, 1981).

Occurrence. To 7% in 93 levels from San Miguel, excluding Sequence B and to 21% in 12 lignites from Lake Somerville.

Range. From near the base of the Lisbon Fm. (Middle Eocene) into the Oligocene (Frederiksen, 1988).

Genus Cyrillaceaepollenites Potonié 1960

Occurrence. "Unclassified" Cyrillaceaepollenites occurs to 6% in 32 samples from San Miguel (including Sequence B) in quantities to 14% in nine lignites and the clastic sample from Lake Somerville.

Cyrillaceaepollenites kedvesii Frederiksen 1980

Cyrillaceaepollenites kedvesii Frederiksen 1980a. p. 51, pl. 11, figs. 13-18.

Description. Length of polar axis 18 to 24 µm, mean 24 µm, holotype 25 µm. Spheroidal or nearly so. Tricolporate; colpi geniculate, narrow, extending nearly full length of grain, with thickened margines 0.5 to 1µm wide; ora lalongate, 1 to 2 µm wide and as long as 6 µm. Exine 1 µm thick; ectosexine:endosexine: nexine ratio 1:1:5:1 but collumellae are not visible or are only faintly visible; design punctate to nearly psilate.

Affinity. Unknown.

Occurrence. To 11% in 60 samples (excluding Sequence B) from San Miguel and to 1% in two lignites at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Cyrillaceaepollenites megaexactus (Potonié)

Potonié 1960

Pollenites megaexactus Potonié 1931, p. 26, pl. 1, fig. V42b.

Pollenites cingulum bruehlensis Thomson in Potonié et al., 1950, p. 56, pl. B, figs. 31-33.

Tricolporopollenites megaexactus bruehlensis (Thomson) Thomson and Pflug, 1953, p. 101, pl. 12, figs. 50-57.

Cyrillaceaepollenites megaexactus (Potonié) Potonié, 1960, p. 102.

Cyrillaceaepollenites cf. C. megaexactus. Tschudy, 1973, p. B17, pl. 4, figs. 14-17.

Cyrillaceaepollenites megaexactus (Potonié) Potonié. Frederiksen, p. 51, pl. 11, figs. 19-22.

Description. 10-24 µm. Outline globular. Width-length index about 1-0.9. Smooth to hyaline smooth. Frequently creased in the polar areas. Cavernae reaching the poles. Without Cavium. [Translated from Thomson and Pfulg]

Affinity. Cyrillaceae, Cyrilla, and (or) Cliftonia according to Frederiksen (1980a).

Paleoenvironmental Notes. Frederiksen (1981) believed pollen from Cyrillaceaepollenites n. sp. to have come from plants common on the coastal plain exclusive of peat-forming areas. Fechner (1988) reported Cyrillaceae from the Middle Eocene of North Africa, believing it to be a plant indicative of salt marshes because of its ability to endure physiological drought. The salt marshes at her site hypothetically stretched behind a belt of mangroves.

Traverse (1955) reported reciprocal pollen percentages of the dominants Cyrilla, found preferentially in the lignites, and Quercus. Traverse theorized that the Cyrilla plants grew around the margin of the swamp, and the Quercus pollen was exotic and indicative of deeper water.

Cyrilla racemiflora, the only modern American species, often forms nearly impenetrable thickets along the margins of swamps and in rich river bottoms, as well as on exposed sandy ridges (Harrar and Harrar, 1969). Ajilvsgi (1979) listed black titi or swamp cyrilla (Cyrilla racemiflora) as an understory element in the drier outer edges of baygall wetlands in the Big Thicket of Texas. She described it as a tall shrub or small tree which usually forms large colonies. C. racemiflora is commonly found in the understory of Taxodium wetlands in the Okefenokee Swamp and in the pocosins of North Carolina (Brown, 1990). I have seen this species as a large shrubby pioneer migrating onto the marsh plants on a floating mat in the Mississippi delta.

The monotypic Cliftonia monophylla is a bushy shrub or small understory tree found in alluvial swamps, in sandy or peaty soils, and on river bottoms subject to short periods of inundation (Harrar and Harrar, 1969). Both Cliftonia and Cyrilla are visited by bees and are important honey producers (Lovell, 1926; Elias, 1980)

Occurrence. To 7% in 48 levels (excluding Sequence B) at San Miguel; to 5% in three lignites from Lake Somerville.

Range. Middle Talahatta Fm. (Middle Eocene) into the Oligocene (Frederiksen, 1988).

Cyrillaceaepollenites? ventosus (Potonié)

Frederiksen 1980

Pollenites ventosus Potonié, 1931, p. 555, fig. 15.

Pollenites ventosus Potonié. Engelhardt, 1964, p. 79, pl. 5, fig. 59.

Pollenites pseudolaesius Potonié 1931. Fairchild and Elsik, p. 84, pl. 37, fig. 23.

Tricolporites sp., (cf. Pollenites ventosus Potonié 1934). Tschudy and Van Loenen, 1970, pl. 4, fig. 30.

Tricolporites sp., Tschudy and Van Loenen, 1970, pl. 4, figs. 31-32.

Cyrillaceaepollenites of the Pollenites ventosus type. Tschudy, 1973, p. B17, pl. 4, figs. 20, 21.

Pollenites laesius type. Elsik 1974, pl. 4, fig. 115.

Cyrillaceaepollenites ventosus (Potonié) Frederiksen, 1980a, p. 51-52, pl. 11, figs. 23-24.

Description. Size 16-22 µm, mean 19 µm. Oblate. Probably tricolporate, but the ora are obscure. Colpi extend 1/2 to 2/3 (generally about 1/2) the distance to the poles. Exine 0.5 - 0.8 µm thick, sexine: nexine ratio is 3:1, integillate to indistinctly tegillate, weakly punctate to coarsely granulate; outer and inner surface smooth or rough. Most specimens have a compression fold that forms a dark, circular to rounded ring cut be the tips of the colpi.

Affinity. Unknown.

Paleoenvironmental Notes. Frederiksen (1981) noted that Pseudolaesopollis ventosa is widely distributed in Eocene detrital samples from the Mississippi embayment, but is rare in coals. He believed "...the species must have been at least partly wind pollinated; or if entirely zoophilous, the producing plants must have been very abundant and widespread on the coastal plain...these plants may have been an important plant constituent of deltaic swamp communities only during the Jackson. Even then, they were probably members of only some of those communities. (p. 524)." Elsik (1978) found an occasional sample of Manning Fm. lignite with an abundance of P. ventosa.

Occurrence. Up to 4% in 56 samples from San Miguel and to 8% from 13 lignites and the clastic sample from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Genus Siltaria Traverse 1955

Siltaria cf. S. scabriextima Traverse 1955

Siltaria scabriextima Traverse, 1955, p. 51, fig. 10 (50).

Cupuliferoipollenites sp., Tschudy and Van Loenen, 1970, pl. 7, figs. 15-16.

Siltaria cf. scabriextima Traverse. Frederiksen, 1970, p. 52, pl. 11, fig. 25.

Description. Tricolporate pollen grain with longitudinal furrows extending nearly from pole to pole. Large and clearly differentiated elliptical transverse furrows ca. 4µm by 6µm. Both furrows have lightly developed margos or thickened rims. Longitudinal furrows underlain by longitudinal costae of the exine, which are cut nearly through under the transverse furrows. Scabrate sculpture. Columellate tectum. Size: ca. 24µm by 32µm. Thickness of exine: ca. 2.5µm.

Taxonomic note. Siltaria, according to N. Frederiksen (written commun., 1984) forms a continuum with Cupuliferoipollenites and Araliaceoipollenites (see discussion with Cupuliferoipollenites). He typified Siltaria as "finely columellate and ... punctate in plain view (that is, having pits or bumps that are 1/2 micrometer or smaller)."

Paleoecological Notes. Siltaria was found only in silts, and not in the lignites of the Oligocene Brandon Lignite (Traverse, 1955).

Occurrence. Present in 123 levels to 36% at San Miguel and to 12% in 11 lignites and the clastic sample from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Siltaria pacata (Pflug) Frederiksen 1980

Siltaria pacata (Pflug) Frederiksen, 1980a, p. 52, pl. 11, fig. 25.

Tricolporopollenites pacatus Pflug in Thomson and Pflug, 1953, p. 99, pl. 12, figs. 118-121.

Ailanthipites pacatus (Pflug) Potonié, 1960, p. 96.

Description. 25-35 µm. Outline widely ellipsoidal, to almost spherical. Width-length index between 0.7 and 1. Exine so elegantly intrabaculate, that it in the inspection appears scabrate, even shiny. Cavernae converge polewards and fall into a Cavium. Germinal aperture similar to Tricolpites dolium. Cavernae are essentially thin, equatorial ring extremely long.

Affinity. Kedves (1969) saw a resemblance with the Simaroubaceae and Cornaceae. Frederiksen (1980a) suggested Diospyros in the Ebanaceae.

Occurrence. One sample from sequence C from San Miguel. Absent at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Genus Araliaceoipollenites Potonié 1960

Occurrence. Undifferentiated large Araliaceoipollenites grains (that is, not including A. granulatus) were present in quantities to 3% in 12 samples from San Miguel.

Paleoenvironmental Notes. The Araliaceae is primarily a tropical family, but some members can be found north as far as Canada. Using leaf structure, Dilcher and Dolph (1970) identified a Araliaceaen macrofossil, which they named Dendropanax, in the Claiborne Group of Tennessee.

Araliaceoipollenites granulatus (Potonié) Frederiksen

1980

Pollenites pseudocingulum granulatum Potonié, 1931, p. 32, pl. 1, figs. 2, 6, 19, 24, 26-27.

Araliacoipollenites granulatus (Potonié) Frederiksen, 1980a, p. 52, pl. 11, figs. 29-30.

Taxonomic note. A. granulatus, according to N. Frederiksen (written commun., 1984), forms a continuum with Siltaria and Cupuliferoipollenites (see discussion with Cupuliferoipollenites). Araliaceoipollenites was described as being columellate and granulate.

Occurrence. To 47% in 115 levels at San Miguel and to 3% in 8 lignites and the clastic sample from Lake Somerville.

Range. Lower part of the Middle to Upper Eocene Lisbon Fm. through the Eocene-Oligocene boundary in Alabama and Mississippi (Frederiksen, 1988).

Araliaceoipollenites profundus Frederiksen 1980

Tricolpopollenites spp. of the T. henrici type. Tschudy, 1973, p. B16, pl. 4, figs. 10-11.

Araliaceoipollenites profundus Frederiksen, 1980a, p. 53, pl. 12, figs. 2-4.

Description. Size 33-58 µm, mean 45 µm, holotype 41 µm. Tricolporate. Subprolate to perprolate, mostly prolate; outline lenticular with pointed to slightly flattened ends. Exine 1.5-2.5 µm thick, tegillate; ectosexine 0.5 µm thick, endosexine 0.5 -1.5 µm thick, nexine about 0.25-0.5 µm thick. Design distinctly granulate to coarsely punctate or finely reticulate. Colpi extending nearly from pole to pole, very narrow and deeply invaginated almost to the polar axis; ora round to lolongate, 4-6 µm long, often indistinct.

Occurrence. To 2% in 24 levels at San Miguel and 1% in one lignite from Lake Somerville.

Range. Upper Claibornian (Upper Lisbon Fm.) into the Oligocene (Frederiksen, 1988).

Araliaceoipollenites megaporifer Frederiksen 1980

Tricolporites sp. (?Araliaceoipollenites) Tschudy and Van Loenen, 1970, pl. 4, figs. 22a-b.

Araliaceoipollenites megaporifer Frederiksen, 1980a, pp. 52-53, pl. 11, figs. 31-32; pl. 12, fig. 1.

Description. Size 14-29 µm, mean 23 µm, holotype 26 µm. Tricolporate. Subprolate to prolate, mostly prolate; outline oval with rounded ends. Exine about 1 µm thick, columellate; sexine: nexine ratio 2:1. In some specimens the exine thickens from slightly less than 1 µm at the equator to more than 1 µm at the poles because of a thickening of the endosexine. Design granulate, surface rough. Colpi very narrow and extending from three-fourths of the grain to the full length; thickenings of the colpi margines 0.3 to 1 µm wide. Ora round, 2.5 - 4 µm in diameter, extending beyond the colpi margines.

Occurrence. To 3% in 21 samples from San Miguel and 1% in three lignites from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a). Rouse and Mathews (1988) commented that the Araliaceoipollenites species "... are very widespread..." in Middle Eocene to Early Oligocene samples, "...extending into the Beaufort Sea in the north and offshore Labrador in the east. They are so characteristic of the Upper-Eocene interval off Labrador that Williams and Brideaux (1975) established the Araliaceoipollenites megaporifer Zone for the interval." (p. 1272)

Genus Foveotricolporites Pierce 1961

Foveotricolporites sp.

Description. Size 46-54 µm. Prolate; outline elliptical. Tricolporate; colpi narrow, extending nearly full length of grain; ora lolongate, 0.5-1.5 µm wide; 0.5-3 µm deep, and 5-8 µm long. Exine 2 µm thick, tegillate, ectosexine:endosexine:nexine ratio 2:1:2. Foveolate, the fovelolae about 0.3 µm in diameter.

Affinity. Possibly Cornaceae, according to Frederiksen (1980a).

Occurrence. Less than 1% in three levels from San Miguel. Absent at Lake Somerville.

Range. Frederiksen (1980a) found this taxon only in the Oligocene Forest Hill Sand.

Affinity. My grains of Ilex are very similar to modern species pictured in McAndrews et. al. (1978) and in the TAMU pollen reference collection. D. Taylor (written commun., 1988) believes that Ilexpollenites from the Claiborne "...appears to have affinities to Aquifoliaceae as previously suggested."

Paleoenvironmental Notes. Pflug (1957) found Ilex, in his study of German Upper Eocene brown coals, to be more common in clastics, and more likely to have come from lakeshore trees. Bartlett and Barghoorn (1973) noted that Ilex is not a prolific producer, and concluded that the presence of Ilex in Holocene sediments from Panama indicated proximity in the vicinity of the vegetational site. Furthermore, they felt that the occurrence of Ilex, a genus not presently growing in the sample area, signified a more seasonal climate than is present today on the Atlantic coast of the Canal Zone.

Van der Hammen (1963) found significant number of Ilex grains in his samples from British Guiana, hypothetically coming from species in swamp forest and marshwood.

Modern Ilex is a tree or shrub genus which contains 200 species in both hemispheres (Harrar and Harrar, 1962) and is widely distributed in both temperate and tropical areas (Li, 1972). Habitats are extremely variable.

One American species, Ilex decidua (winterberry) grows both on moist, rich upland soils and in swamps (Harrar and Harrar, 1962). In the Big Thicket of Texas, I. opaca (American holly) is found on well-drained ridges, whereas I. coriacea is an inhabitant of acid bogs (Ajilvsgi, 1979). Because of its ability to grow in wetlands, and its appearance in lignites (Hopkins, 1967; Rachele, 1976; Traverse, 1955), Ilex can be considered a probable indicator of wetland depositional environment.

Members of the Aquifoliaceae are largely insect pollinated, (Lewis et al., 1983; Lieux, 1983). Flenley (1979) points out that Ilex grains are of "medium/low relative export." Large concentrations of Ilex pollen in many samples from a site is unlikely. Pollen of this taxon is found in quantities to 50% of the pollen sum in upper levels of "treehouse" stratigraphies in the Okefenokee; surface samples show greatly reduced percentages in the adjacent marsh (Rich and Spackman, 1979).

Occurrence. "Undifferentiated" Ilex was tallied to 2% in 36 levels from San Miguel, and to 3% in seven lignite samples from Lake Somerville.

Range. According to Muller (1981), the first appearance of Ilex pollen is in the Turonian of Australia; Ilex first appears in the United States in the Cenomanian of California. During the Tertiary, Ilex distribution is cosmopolitan.

Ilex infissa Frederiksen, 1980a, p. 53, pl. 12,

figs. 10-14.

Description. Size 19-28 µm, mean 24 µm, holotype, 28 µm. Prolate spheroidal to subprolate. Tricolporate; colpi narrow (0.5-1 µm wide), rather deeply invaginated, extending nearly full length of grain, bordered on each side by thickenings 2 µm wide; ora distinct, lalongate, slitlike, 0.5 µm wide and 3.5-5 µm long, cutting through marginal thickenings of the colpi. Exine 1.5-2 µm thick, sexine:nexine ratio 2:1, densely clavate, the clavae 1.3-2 µm long.

Occurrence. To 2% in 12 samples from San Miguel and 1% in one lignite from Lake Somerville.

Range. Found in the Jackson Group by Frederiksen (1980a).

Tricolporopollenites iliacus medius Pflug and Thomson in Thomson and Pflug, 1953, p. 106, pl. 14, figs. 46-60.

Ilexpollenites cf. I. iliacus (R. Potonié, 1931) Thiergart, 1937. Engelhardt, 1964, p. 73, pl. 2, fig. 22.

Ilexpollenites sp., Tschudy and Van Loenen, 1970, pl. 4, figs. 19, 18?

Ilex media (Pfleug and Thomson) Frederiksen, 1980a, pp. 53-54, pl. 12, figs. 15-16.

Description. 25-45 µm. Height of the "little clubs" (clavae) normally not under 1/12 of the maximum diameter of the grain, clavae smallest partly globeheaded. Diameter of globular heads mostly not under 2 µm. Clavae height mostly over 3 µm. Outline globular to ellipsoidal. Exine regular with clavae sitting on it. [Translated from Thomson & Pflug]

Range. Members of the family Aquifoliaceae appeared during the Turonian. During the Tertiary, the family became widespread (Muller, 1981). Found by Frederiksen (1980a) in the Upper Claiborne to Lower Vicksburg.

Occurrence. Present in two samples from the San Miguel Sequence E at 3% and 10%. Absent at Lake Somerville.

Paleoenvironmental Notes. Because of its distribution, Frederiksen (1981) believed pollen of Verrutricolpoites n. sp. to have come from plants more common outside of the peat-forming communities. One species, Verrutricolporites rotundiporis was compared by Germeraad et al. (1968) to the modern pollen of Crenea, which grows along water courses and swampy areas, and in the mangrove vegetation of tropical regions.

Occurrence. Undifferentiated Verrutricolporites is present to 2% in 10 samples from San Miguel and 1% in three lignites and the clastic sample from Lake Somerville.

Tricolporopollenites spp. Tschudy, 1973, p. B15, pl. 3, figs. 17-19?

Verrutricolporites cruciatus Frederiksen, 1980a, p. 54, pl. 12, figs. 17-19.

Description. Size 26-34 µm, mean 29 µm, holotype 31 µm. Prolate: outline oval with rounded to slightly pointed ends. Tricolporate; colpi narrow, extending about four-fifths the edge of the grain, edges thickened little or not at all; ora distinct, lalongate, 1-3 µm X 3-8 µm. Exine 1.5-2 µm thick including ornamentation; sexine:nexine ratio 1:1; verrucate, the elements irregular in design, about 0.5-1 µm in diameter and 0.3-0.5 µm high; negative reticulum rather well developed.

Affinity. Unknown

Occurrence. To 2% in 12 samples from San Miguel; in sequence H, where only Verrutricolpites spp. was tallied, there were two levels with less than 1%. At Lake Somerville, to 1% in three lignites and the clastic sample.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Verrutricolporites ovalis (Potonié) Frederiksen 1980, p. 54, pl. 12, figs. 20-21.

Pollenites cingulum ovalis Potonié, 1934, p. 83, pl. 4, fig. 8.

Tricolporopollenites sp. 5. Engelhardt, 1964, p. 74, pl. 3, fig. 30.

Tricolpate rugulose-verrucose pollen. Fairchild and Elsik, 1969, pl. 37, fig. 18.

Unidentified tricolpate pollen having relatively coarse verrucos-rugulose ornament. Elsik, 1974, pl. 4, fig. 119.

Occurrence. 1% in one sample from San Miguel Sequence G. Absent at Lake Somerville.

Range. A few grains were found by Frederiksen (1980a) in Jackson and Vicksburg sediments.

Verrutricolporites tenuicrassus Frederiksen, 1980a, p. 54, pl. 12, figs. 22-25.

Description. Size 23-34 µm (five specimens), holotype 31 µm. Prolate, outline oval. Tricolporate; colpi 0.5-1µm wide, extending nearly full length of the grain; ora lalongate about 3 X 6 µm. Exine 0.7-1.2 µm thick at equator and 2-3 µm thick at poles; sexine: nexine ratio 2-3.1; verrucate, the elements irregular in design. 0.5-1 µm in diameter and 0.2 -0.5 µm high; negative reticulum present; exine may be indistinctly tegillate.

Affinity. Possibly Fagaceae, according to Frederiksen (1980a)

Occurrence. To 2% in 12 samples from San Miguel.

Range. Found by Frederiksen (1980a) in the Lower Jackson.

Pollenites kruschi Potonié, 1931, p. 4, fig. 11.

Tricolporopollenites kruschii (Potonié) Thomson and Pflug, 1953, p. 103, pl. 13, figs. 14-63.

Nyssapollenites cf. Nyssa accessorius (R. Potonié, 1934) R. Potonié 1950, Engelhardt, 1964, p. 74, pl. 3, fig. 33.

Tetracolporites sp., Engelhardt, 1964, p. 76, pl. 4, fig. 50.

Nyssa sp., Fairchild and Elsik, 1969, p. 84, pl. 37, fig. 16.

Nyssa kruschii (Potonié) Frederiksen, 1980a, pp. 54-55, pl. 13, fig. 1.

Description. 15-50 µm. Outline + globular or secondary pearshaped. Unsculptured. Intrarugulate structure. All pores circular and circumscribed by the Caverna. [Translated from Thomson and Pflug].

Paleoenvironmental Notes. Megafossil evidence for Nyssa has often been in the form of endocarps (fruits) because, according to Eyde and Barghoorn (1963), most Nyssa foliage is without conspicuous identifying features. The presence of these fruits in the Brandon and Wilcox lignites, as well as in European brown coals, suggests that Nyssa was a member of the Paleogene peat forming swamp community. In the Brandon lignites, some fruit species were found only in the silt and not the lignite. Dilcher and McQuade (1967) described the endocarp N. eolignitica from Middle Eocene silts in Tennessee; they believed it to be similar both to the modern N. ogeche and the Oligocene N. lescurii from the Brandon silts.

A study of wood and pollen from the Miocene Rhenish brown coal by Van der Burgh (1967) yielded relatively equal proportions of pollen and fruits of Nyssa.

Nyssa pollen was common in a Yequa Fm. lignite located in fluvial strata in Madison County, Texas (Elsik, 1978) where it was considered "...indicative of a swamp environment. (p. 28)." A Yegua Fm. lignite from Brazos County, Texas which I have examined also yielded abundant Nyssa pollen.

Nyssa occurred preferentially in the lignite layers of the Oligocene Brandon Lignite deposit of Vermont. Values were to 5%, as opposed to 1.2% in the silts and clays (Traverse, 1955). Clark (1986), in his study of the modern barrier beach vegetation of Long Island, used a minimum of 3% of Nyssa pollen to indicate a local presence in the form of a maritime forest community.

Frederiksen (1980c) reported modern Nyssa growing in humid subtropical, winter-dry tropical, and wet tropical(?) regions. Nyssa is, according to Harlow (1957), "...a typical swamp-forest tree, found as well along lake shores, but growing on drier sites when planted." (p. 259) In the North, it grows in wet or moist soils (Curtis and Bausor, 1943).

In the present-day Okefenokee Swamp, Nyssa occupies peat substrates which are more mineral-rich than those in which Taxodium flourishes (Fair-Page and Cohen, 1990). In the dismal swamp, N. sylvatica inhabits the full range of swamp environments, including the rarely flooded forest community and the most extensively flooded forest community. It dominates in the maple-gum community, where water levels fluctuate for 6 months out of the year. (Brown, 1990) N. aquatica rarely germinates when submerged, according to Huenneke and Sharitz (1990).

Nyssa is entomophilous (Lewis et al, 1983), but incidental shedding of pollen occurs.

Occurrence. To 44% in 102 levels from San Miguel and to 15% in 13 lignites and the clastic sample from Lake Somerville.

Range. Pollen of the Nyssaceae first appeared in the Paleocene (Tiffney, 1985).

Tricolporopollenites dolium (R. Potonié, 1931) Thomson and Pflug, 1953 [misidentified]. Engelhardt, 1964, pl. 73, pl. 2, fig. 21.

Tricolporopollenites kruschii (Potonié, 1934) Thomson and Pflug 1953. Elsik, 1968, p. 628, pl. 34, figs 3a-b only.

Rhoipites angustus Frederiksen, 1980a, p. 55, pl. 13, figs. 2-8.

Description. Size 24 to 32 µm, mean 29 µm, holotype 32 µm. Prolate spheroidal to prolate; outline oval to diamond shaped, poles rounded to somewhat flattened. Tricolporate; colpi about three-fourths the length of the grain and very narrow (sides of the colpi may be pressed together), exine not thinned along colpi so that colpi walls appear very thick; ora distinct, round, 2-2.5 µm in diameter, endannuli apparently lacking. Exine 1 µm thick, minutely reticulate.

Affinity. Frederiksen (1980a) listed Mastixia, Nyssa, and Rhus barclayi as possible related taxa. Wodehouse (1933), in defining the genus, noted that the species R. bradleyi was a perfect match with the modern species Rhus typhina (staghorn sumac). The five megafossil species of Rhus present in his Green River Formation were in agreement with this match.

Paleoenvironmental Notes. N. Frederiksen is less than secure with his identification of Rhoipites latus as Rhus. He (written commun., 1988) suggested that "...The [R. angustus] parent plant...lived in a low-stress environment (??)...From the ornamentation one would guess that it was insect pollinated, which means the plants must have been abundant if the grains are plentiful...this is all speculation." His thoughts on pollination are in general agreement with those of Wodehouse (1933), who believed R. bradleyi to be insect pollinated and living in great abundance close to the site of deposition.

The controversial Rhus (sumac) is a shrub or small tree. Some species live in moist, rich soil; others in dry, rocky areas. (Vines, 1982) Species of Rhus pioneer in larger clearings of the uplands of the Texas Big Thicket. (Ajilvsgi, 1979).

Frederiksen (1980c) reported Rhus as living in summer-dry subtropical and/or semiarid, humid subtropical, winter-dry tropical, and wet tropical regions.

Occurrence. To 40% in 130 samples from San Miguel. To 23% in 12 lignites and the clastic sample from Lake Somerville.

Range. R. angustus was found in the Upper Claibornian to Lower Vicksburgian of the Gulf Coast by Frederiksen (1980a); Elsik describes this species as Tricolporopollenites kruschii from the Sabinian Rockdale Lignite of Texas. Matthewes and Rouse (1984) found R. angustus from Early to Early Middle Eocene of the Fraser River area, British Columbia. Similar Rhoipites species were described from the Middle Eocene of California (Frederiksen, 1983).

Rhoipites latus Frederiksen 1980

Tricolporopollenites sp 4 Engelhardt, 1964, p. 74, pl. 3, fig. 29.

Tricolpopollenites sp., Tschudy and Van Loenen, 1970, pl. 5, figs. 13a-b.

Tricolporites sp., Tschudy and Van Loenen, p. 70, pl. 5, fig. 1.

Tricoloporopollenites n. sp. B (Parthenocissus type). Tschudy, 1973, p. B17, pl. 4, figs. 18-19.

Rhoipites latus Frederiksen, 1980a, pp. 55-56, pl. 13, figs. 9-13.

Description. Size 34-44 µm, mean 39 µm, holotype 42 µm. Prolate; outline oval. Tricolporate; colpi deep, narrow, extending nearly full length of grain, bordered by thickenings 1.-2 µm wide,; ora distinct and round, slightly lalongate, 2.5-3.5 µm in the greatest dimension, wider than colpi and creating depressions in marginal thickenings. Exine exclusive of ornamentation 0.5-0.7 µm thick. Exine reticulate; muri coarsely clavate in cross section, clavae 1.5 µm high, thin baculae present between clavae; muri duplibaculate, 0.5-0.8 µm thick and wide; luminae polygonal to longitudinally elongate, 0.5-1.5 µm X 1-2.5 µm.

Affinity. Rouse (1977) noticed a resemblance to the Tileaceous tropical genus Triumfetta; Barnett (1989) used this name for her R. latus specimens. Tschudy (1973) believed that R. latus may have been related to the modern genus Parthenocissus.

Paleoenvironmental Notes. The Rhoipites latus pictured by Matthewes and Rouse (1984) from Canada is much more finely reticulate than Frederiksen's Gulf Coast specimens. These authors interpreted the climate of British Columbia in the Early Eocene to be warmer and more humid than at present.

No significant environmental data is available for Rhoipites latus. Because of its distribution in clastics, Frederiksen (1981) believed pollen of "Parthenocissus?" to have come from plants common in non-peat forming coastal areas.

Frederiksen (1980c) reported Parthenocissus as growing in summer-dry subtropical and/or semiarid, humid subtropical, and winter-dry tropical areas. Virginia creeper, common in eastern North America, is a woody vine. Two species of Parthenocissus are found in moist soils throughout most of the eastern part of the United States (Gleason and Cronquist, 1963); P. heptaphylla is found in rocky canyons and on sandy soil in the Edwards Plateau (Powell, 1988).

Occurrence. To 4% in 64 samples from San Miguel and in 16 lignite and the clastic sample from Lake Somerville.

Range. Middle of the Early Claiborne Tallahatta Fm. through the Eocene-Oligocene boundary in Alabama and Mississippi (Frederiksen, 1988). Frederiksen (1980a) found R. latus in Gulf Coast samples of Upper Claibornian to Vicksburgian age. R. latus was present in rocks of Early to Late Eocene in South Central British Columbia and in Early to Middle Eocene in Arctic rocks (Rouse, 1977). Similar Rhoipites grains are also found from the Middle Eocene of Nevada (Wingate, 1983).

Description. Tricolporate pollen with long, wide longitudinal furrows that are as if they were lateral extensions of the longitudinal furrows. Reticulate sculpture with very high muri.

Affinity. Frederiksen (1983) noted that "Specimens with rather straight colpi have some resemblance to pollen of Xylosma (Flacourtiaceae), whereas specimens with arched colpi are similar to pollen of Styrax (Styracaceae)."

Large Species Group

Occurrence. To 3% in 53 samples from San Miguel. To 10% in four lignites at Lake Somerville.

Small Species Group

Occurrence. To 7% in 73 samples from San Miguel. To 2% in seven lignites from Lake Somerville.

Range. Frederiksen found specimens of the genus in samples ranging from Late Claiborne to Early Vicksburg.

Genus Caprifoliipites Wodehouse 1933

Caprifoliipites tantulus Frederiksen, 1980a, p. 57, pl. 14, figs. 1-2.

Description. These grains match the description of Caprifoliipites micrireticulatus (Pflug and Thomson in Thomson and Pflug, 1953) Potonié, 1960, but they are only 14-19 µm in greatest dimension (holotype 14 µm) whereas the size of C. microreticulatus was given as 18-30 µm.

Affinity. Fredericksen (1980a) suggested that Caprifoliipites tantulus pollen is similar to that of Viburnum (Caprifoliaceae).

Paleoenvironmental Notes. C. tantulus was common to abundant in Frederiksen's (1981) Jackson age lignites; it was less abundant in his clastics. C. tantulus pollen was also found in one of Frederiksen's Late Claiborne samples; Jones and Gennett (1991) found a few C. tantulus grains in marine sediments from the Claiborne Stone City Fm.

Frederiksen (1981) believed these grains to appear zoophilous, but conceded that they could be transitional from zoophilous to anemophilous. He also believed that, due to their abundance in coal and patchy but wide distribution in clastics that the C. tantulus plant was a heavy producer, in part anemophilous, and possibly shrubby or herbaceous.

Crepet (1985) mentions Viburnum-type pollen described from the Middle Eocene of France by Gruas-Cavagnetto (1978), and suggests "...Hymenoptera (wasp), Myophily (rodent), Ornitophily (bird)..." as possible pollination mechanisms for that taxon.

Harrar and Harrar (1967) reported about 100 species of Viburnum in the United States, occurring as both trees and shrubs. There are six species in the southeastern United States, all small trees. One species, V. rufidulum, grows in a variety of conditions, but attains its best growth on moist, rich alluvium. Another, V. prunifolium, is found more often at forest borders and is not so fond of moist conditions.

Frederiksen (1980c) reported Viburnum as growing in summer-dry subtropical or semiarid, humid subtropical, winter-dry tropical, and wet tropical regions.

Occurrence. To 12% in 104 samples from San Miguel and to 33% in 18 lignites from Lake Somerville.

Range. Middle of the Early Claiborne Tallahatta Fm. to the top of the Jackson Fm. in Alabama and Mississippii (Frederiksen, 1988).

Ailanthipites berryi Wodehouse, 1933, p. 512, fig. 44.

Tricolporopollenites sp. 1. Engelhardt, 1964, p. 73, pl. 3, fig. 25.

Description. Generally ellipsoidal, but somewhat various in shape according to the degree of their expansion, tricolporate with furrows long, almost reaching from pole to pole, furrow rim and pore rim conspicuous, projecting deeply inward. Exine reticulate, pitted with the pits elongate and linearly arranged, forming a sort of thumb print. Grains 20-25 µm. 1 µm broad and 26-30 µm long. Holotype 36-19.8-61.5.

Affinity. Frederiksen (1980a) suggests possible affinities with Anacardiaceae, Burseraceae, Leguminoseae, Sapindaceae, and Simaroubaceae. Wodehouse (1933) believed "...with a fair degree of confidence... (p. 512)" that it should be assigned to the modern genus Ailanthus, megafossils of which he reported in the Green River Flora.

Paleoenvironmental Notes. Wodehouse (1933) theorized that "...they are obviously insect borne, yet the occur in the shales in large numbers. Therefore, they should belong to a plant which grew abundantly near the place of deposition (p. 512)."

The only species of Ailanthus present in America today, the "tree of Heaven" is introduced from China. It is an extremely opportunistic, fast growing species which can live anywhere. Members of the Simaroubaceae are primarily entimophilous, but are also facultative anemophilous (Lewis et al., 1983).

Occurrence. To 3% in 50 samples from San Miguel. To 1% in seven lignites and the clastic sample at Lake Somerville.

Range. The known range on the Gulf Coast of Ailanthipites berryi is Upper Eocene to Lower Oligocene (Frederiksen, 1980a). Various species of Ailanthipites were also found by Frederiksen (1983) in Middle Eocene clastics in California.

Description. Size 42-46 µm. Oblate spheroidal to suboblate; outline more or less round. Tricolporate; colpi extend about two-thirds the distance to the poles, bordered by thickenings 1-2.5 µm wide; ora round, 4-10 µm in diameter. Exine excluding ornamentation 1 µm thick. Exine reticulate; muri clavate in optical section, clavae 1.5 µm high, muri 0.5 µm wide and duplibaculate; lumina 1-2 µm in diameter.

Range. Found in Jackson Group samples by Frederiksen (1980a).

Affinity. According to Frederiksen (1980a), Alangiaceae, probably Alangium.

Occurrence. To 2% in 23 samples at San Miguel. Absent at Lake Somerville.

Myraceidites parvus Cookson and Pike, 1954, p. 206, pl. 1, figs. 27-31.

Myrtaceidites parvus nesus Cookson and Pike, 1954, p. 206, pl. 1, figs. 29-31.

Myrtaceidites parvus anesus Cookson and Pike, 1954, p. 206, pl. 1, figs. 27-28.

Cupanieidites sp., Tschudy and Van Loenen, 1970, pl. 4, figs. 23-24.

Description. Grains small, 9-14 µm in equatorial diameter; amb subtriangular with convex sides; arci present which may or may not enclose polar islands. Exine psilate, indistinctly patterned, or granular.

Affinity. Muller (1968) believed palynomorphs of this genus to be "in all probability pollen from Myrtaceae." (p. 22) Frederiksen (1983) noted that M. parvus is a heterogenous form species representing mainly genera such as Myrtus, Eugenia, and Calyptranthes as well as perhaps Ardisia in the Myrsinaceaea.

Paleoenvironmental Notes. Frederiksen (1981) noted that Myrteacidites is slightly more common in Eocene clastics than in lignites and that the palynomorph may represent a shoreline plant.

Along with Cupaniedites orthoteichus, Myrtaceidites parvus is one of the few palynomorphs described both from the Gulf Coast and from Late Eocene lignites from Australia (Milne, 1988). Although M. parvus is present in quantities up to 1.6% in the Australian deposit, four other species are present, and the genus Myrtaceidites totals over 10% of the palynomorphs in one sample. These lignites formed in a "predominantly non-marine" setting (p 286).

Crepet (1985) cited the occurrence of Myrtaceidites sp. (i.e. Eugenia-type pollen) in the Middle Eocene Lawrence Clay Pit of Tennessee, using this genus as an example of "Chiropterophily, Ornithophily, Melittophily." In other words, he believed the Myrtaceidites plant to have been pollinated by bats, birds, or bees.

One Myrtaceous plant common in the southeastern United States is Eugenia axillaris, which is found on sandy soils near salt water. Another species, Anamomis simpsonii is found on hammocks. Calyptranthese pallens grows in Florida on hammocks and coastal soils of coral origin (Harrar and Harrar, 1969).

Pollination of modern Myrtaceae species is largely zoophilous and only incidentally anemophilous (Lewis et al, 1983). Flenley (1979) noted that Myrtaceous grains exhibit "some relative export."

Occurrence. Less than 1% in 28 samples from San Miguel. Present to 2% in four lignites from Lake Somerville.

Range. Upper part of the Middle to Upper Claiborne Lisbon Fm. through the Eocene-Oligocene boundary in Alabama and Mississippi (Frederiksen, 1988). Grains of the family Myrtaceae have been found since the Santonian. The earliest occurrence in North America of Myrteacidites sp. is in the Mid-Eocene of Tennessee (Muller, 1968).

Genus Cupanieidites Cookson & Pike 1954 emend.

Chmura 1973

Cupanieidites orthoteichus Cookson and Pike, 1954, p. 213, pl. 2, figs. 73-78.

Duplopollis orthoteichus (Cookson and Pike) Krutzsch, 1959, p. 145.

Duplopollis myrtoides Krutszch, 1959b, p. 145, pl. 34, figs. 25-44; text-fig. 13.

Cupanieidites orthteichus Cookson and Pike, 1954. Engelhardt, 1964, p. 74-75, pl. 3, fig. 34.

Duplopollis sp., Fairchild and Elsik, 1969, p. 84, pl. 37, fig. 19.

Duplopollis sp., Tschudy and Van Loenen, 1979, pl. 4, figs. 25-27.

Description. Grains isopolar to subisopolar; amb sharply triangular with straight sides; arci prominent, forming distinct polar island, equatorial diameter 20-30 µm. Sexine reticulate, mesh fine, somewhat indistinct.

Affinity. According to SEM studies by Taylor (in press), C. orthoteichus has affinities to the Cupanieae group of the Sapindaceae.

Paleoenvironmental Notes. Cupanieidites orthoteichus is one of the few taxa found both in the San Miguel and in the Late Eocene of Western Australia, where it was found in frequencies of less than 1% by Milne (1988). Berry (1924) assigned leaf macrofossils from the Jackson Group of Georgia to Cupanites.

Crepet (1985) cited the occurrence of Sapindaceaous Diplopeltis hueglii-type flowers from the Middle Eocene of Central America by Kemp (1976), and hypothesized "Melittophily" (bees) as the form of pollination. In Costa Rica, the three species of Cupania live in habitats including dry tropical, where C. guatemalensis lives as a shrub, wet tropical, moist tropical, moist premontane, wet premontain; where C. macrophylla and C. glabra are intermediate trees; and premontain rainforest, where C. macrophylla occupies a canopy niche (Holdridge et al).

According to Lewis et al. (1983), although modern-day Sapinaceae are insect pollinated, the anthers are exposed and some pollen grains may become wind-borne.

Occurrence. Present to 4% in 36 samples from San Miguel. To 1% in five lignites from Lake Somerville.

Range. Base of the Middle to Upper Claiborne Lisbon Group to the top of the Jackson Group in Alabama and Mississippi (Frederiksen, 1988). The genus Cupanieidites has been found as early as the Conacian (Muller, 1981).

Gothanipollis sp. 1. Engelhardt, 1964, p. 75, pl. 3, figs. 35-37.

Gothanipollis cockfieldensis Engelhardt, 1964, p. 598- 600, pl. 1, figs. 1-4.

Gothanipollis sp., Fairchild and Elsik, 1969, p. 84, pl. 37, fig. 20.

Gothanipollis sp., Tschudy, 1973, p. B16, pl. 4, fig. 4 only.

Gothanipollis cockfieldensis Engelhardt, 1964b. Frederiksen, 1980, fig. 59, pl. 14, fig. 16.

Description. Pollen grains, syncolporate; polar view triangular with concave sides and truncated recurved apices; equatorial view, lens-shaped; three colpi straight, distinct, united at poles with vestibulate ora located at apices; exine 1 µm thick, sculpture granulate, psilate near colpi. Size range from 18 to 25 µm.

Affinity. Frederiksen (1980, 1985) hypothesized Gothanipollis to be Loranthaceous. Taylor (in press) believed G. cockfieldensis to have similarities with large flowered Neotropical Loranthaceous grains. Jarzen (1978), working in the Paleocene, linked Gothanipollis with the loranthaceous genus Aetanthus. Habitat was cited as "northern Andes temperate forests at elevations of 2000 feet or more". It also resembles the Chilean genus Psitacanthus cuneifolius as pictured by Heusser (1971).

Paleoenvironmental Notes. Modern members of the Loranthaceae (mistletoe family) are commonly parasitic. Quaternary pollen studies, for example that of Watts and Bradbury (1982), show that Loranthaceous pollen, usually that of the pine-dependent Arceuthobium, is found in sediments near the place in which the host was growing. In their study of an Oligocene pollen flora from British Columbia, Rouse and Matthews (1979) found pollen from an apparently parasitic Gothanipollis plant associated with Taxodium, Metasequoia, and Juglans.

Crepet (1985) cited Loranthaceous pollen found in North-central Europe and Germany by Krutzsch (1970). He hypothesized "Melittophily (bees), Myophily (flies), Ornithophily (birds), Phalaenophily (marsupial flying squirrels)" as pollination mechanisms.

Occurrence. Present to 1% in four samples from San Miguel. Absent from Lake Somerville.

Range. Base of the Uppermost Claiborne Gosport Fm. to the top of the Jackson Group in Alabama and Mississippi (Frederiksen, 1988). Gothanipollis appears in both Gulf Coast and Central European sediments from the Middle Eocene (Fairchild and Elsik, 1969); Elsik (1978) found the taxon from the base of the Lower Eocene to the top of the Upper Eocene in Gulf Coast sediments, but the range extended into the Paleocene and Oligocene in the Pacific Northwest. Gothanipollis is also reported from the Middle Eocene Mission Valley Formation of California (Frederiksen, 1983), as well from the Middle Eocene of south central British Columbia (Rouse, 1977). The genus is missing from Wingate's (1983) Nevada flora, and has not, according to Rouse (1977), been found in any Arctic horizons.

Bombacacidites nacimientoensis Anderson 1960, p. 23, pl. 8, fig. 15.

Bombacacidites nacimientoensis (Anderson 1960) Elsik, 1968, p. 620, p. 22, figs. 1-2, 4.

Bombacacidites sp., Tschudy and Van Loenen, 1970, p. 5, figs. 17-19.

Description. Oblate, tricolpate pollen with short wide furrows bordered by a wide margo; furrows reaching about one-half the distance to the pole; shape intersemiangular in polar view, the poles slightly protruding; sculpture reticlulate, with narrow and probably clavate, muri in the polar region; reticulum transitional into clavate sculpturing at the periphery.

Affinity. Probably Bombacaceae, according to Frederiksen (1981). Westgate and Gee (in press) suggested an affinity to the Bombacaceae-Sterculiaceae-Tiliaceae complex for the similar B. paulus. Elsik (1968) listed the Bombacaceae or Tiliaceae as possible affinities. Taylor (1988) noted a close resemblance to the Bombax clade.

Paleoenvironmental Notes. Westgate and Gee (in press) proposed that the Bombacacidites paulus in their South Texas Middle Eocene lignite might be from a plant similar to the back mangrove and forest tree Brownlowia or to several genera of the Bombacaceae which are pantropical rain forest trees.

Berry described leaf fossils of Bombacites jacksoniana from the Jackson Group near Somerville, Texas and from the Fayette Sandstone in Webb Co., Texas.

Bartlett and Barghorn (1973) illustrated a similar palynomorph from the extant neotropical plant Pseudobombax septenatum, which occurs on forested hills, in pastures, and along roads, usually on sandy soil. Pseudobombax pollen was present in only one of four modern surface sediment samples from Gatun Lake in Panama, and the authors believed the pollen "...undoubtedly came from a large tree of this genus growing in the vicinity of the sampling site." (p. 227) Van der Hammen (1963) mentioned that the Bombacaceae may occur in rain and marsh forests, in swamp woodlands along river banks, and also in savannas.

Occurrence. Absent at San Miguel. 1% in one lignite from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Tiliaepollenites sp., Engelhardt, 1964, p. 77, pl. 4, fig. 48.

Tiliaepollenites sp., Tschudy and Van Loenen, 1970, 1970, pl. 5, figs. 13, ?11a-b, ?14-15.

Description. Size 16-31 µm, mean 25 µm, holotype 24 µm. Tricolporate, rarely tetracolporate. Peroblate; outline rounded triangular, occasionally nearly round. Exine 1 µm thick including ornamentation; sexine: nexine ratio 2-3.1. Sexine reticulate, the muri about 0.3 µm wide and slightly clavate in optical section, rising 0.5 µm or less above exine surface. Lumina polygonal, about 1 µm in diameter, with a small granum in the center of each lumen. Sexine slightly overhangs apertures; colpi and ora 1-2 µm wide at equator; reticulum extends to edges of apertures. Nexine at the apertures thickens perpendicular to sexine; nexine (endannulus) 1.5-2.5 µm thick around vestibula, thinning slightly toward bases of vestibula.

Affinity. Bombacaeae-Sterculiaceae-Tiliaceae complex (Frederiksen, 1973).

Paleoenvironmental Notes. From its distribution, Frederiksen (1981) believed Intratriporopollenites n. sp. to have come from plants abundant on the Gulf Coastal Plain, but living mainly or entirely outside the "deltaic" swamp. Fechner (1988) and Mohr and Fechner (1986) hypothesized the similar I. ceciensis from the Middle Eocene of North Africa to represent the Tiliaceous mangrove Brownlowia, which is not restricted to tropical coastal environments.

Berry (1924) described macrofossils of Tilia jacksoniana from the Fayette sandstone, near Wellborn, Brazos Co., TX.

The Tiliaceae is described by Lewis et al. (1983) as primarily a tropical woody family with some native North American species. The flowers often secrete a sweet scent and nectar, with entomophily by bees, flies, and moths. Tilia pollen is produced in large quantities and is found in the pollen rain at substantial distances from the source.

Occurrence. Present at less than 1% in five samples from San Miguel and at 6% in one lignite from Lake Somerville.

Range. Lower Claiborne Tallahatta Fm. through the Eocene-Oligocene boundary in Alabama and Mississippi (Frederiksen, 1988).

Reticulataepollis reticlavata Frederiksen, 1980a, p. 60, pl. 14, figs. 23-26.

Description. Size 16-30 µm, mean 23 µm, holotype 23 µm. Tricolporate. Oblate; outline rounded triangular. Exine 0.05 µm thick; the muri coarsely clavate in optical section, muri 0.5-1 µm thick; clavae 1.5-2 µm high, projecting slightly above muri; muri 1 µm wide, lumina 1.5-2.5 µm in diameter. Colpi boat shaped, 5.5 µm long and 1 µm wide; ora 1-1.5 µm in diameter, endannulus 3-5.5 µm in diameter.

Affinity. According to Frederiksen (1980a) similar to Kirkia and Ligustrum.

Occurrence. 1% from one sample from Sequence A from San Miguel. Absent at Lake Somerville.

Range. Upper Claiborne to Lower Jackson (Frederiksen, 1980a).

Occurrence. Undifferentiated Symplocos sp. is present at less than 1% in 6 samples from San Miguel and in 2 samples from Lake Somerville.

Symplocoipollenites sp. 1. Engelhardt, 1964, p. 75, pl. 4, fig.39.

Triporopollenites sp., Tschudy and Van Loenen, 1970, pl. 3, fig. 13.

Porocolpopollenites spp., Tschudy, 1973, p. B15, pl. 3, figs. 5-6.

Symplocos contracta Frederiksen, 1980a, p. 61, pl. 15, figs. 6-9.

Description. Size 22-34 µm, mean 28 µm, holotype 32 µm. Peroblate; outline triangular with straight to slightly convex. Tricolporate. Exine finely foveolate and tegillate; midway between apertures it is 1.5-2 µm thick, Nexine 0.5 µm, endosexine 0.71 µm, ectosexine 0.7-1 µm, ectosexine 0.3 to 0.5 µm; sexine thins gradually towards apertures, where it is 0.3-0.5 µm thick. Colpi 1-2.5 µm long lacking marginal thickenings; ora obscure; vestibula 0.5 µm or less deep, typically slitlike in optical section.

Affinity. Frederiksen (1980a) cites the opinion of many authors that no modern genera other than Symplocos have pollen of the Porocolpopollenites-Symplocoipollenites type.

Paleoenvironmental Notes. Fossil Symplocos pollen has been found in the Brandon lignite (Traverse, 1955) and the Eocene German brown coals, where the plant apparently lived in coastal swamps (Frederiksen, 1981).

The modern genus Symplocos, or sweetleaf, is usually seen as a shrub, but occasionally as a tree. It is found in the West Indes, Asia, and Australia, and in the United States ranges from Delaware to Florida west to eastern Texas and Arkansas (Harrar and Harrar, 1962). Symplocos habitats range from coastal plains to mountains; it lives in swamps and stream margins, in sandy thickets, in alluvial woods, in rich, moist forest soils, and in upland areas (Frederiksen, 1981; Harrar and Harrar 1962). Frederiksen (1980c) classified the climatic range of Symplocos as humid subtropical, winter dry tropical, and wet tropical. The family is insect pollinated (Machin, 1971).

Occurrence. Present at less than 1% in 6 samples from San Miguel. Absent from Lake Somerville.

Range. Upper Claiborne and Lower Vicksburg (Frederiksen, 1980a).

Symplocoipollenites sp., Tschudy and Van Loenen, 1970, pl. 5, figs. 6 a-b.

Porocolpopollenites spp., Tschudy, 1973, p. B16, pl. 4, figs. 8-9.

Symplocos gemmata Frederiksen, 1980a, p. 61-62, pl. 15, figs. 10-14.

Description. Size 19-31 µm, mean 26 µm, holotype 29 µm. Tricolporate. Oblate; outline triangular with strongly convex to nearly straight sides. Exine 1 µm thick, sexine:nexine ratio 2:1 except at apertures. Sexine indistinctly tegillate and rather sparsely to densely gemmate to granulate, the elements typically varying in size on each specimen, from 0.3 to 1.5 µm in diameter and to as much as 1 µm in height. Ornamentation covers entire exine up to edge of apertures. Colpi 0.5-1 µm wide at the equator, narrowing rapidly away from the equator; colpi very short, usually not extending past the endannulus, often barely visible so that some grains look triporate; colpi may be bordered by narrow (0.5-µm-wide), smooth margins which wrap around the ends of colpi. Shallow vestibula present. Endannuli 2-3 µm thick, with ora about 2.5 µm in diameter; sexine does not thicken at apertures.

Occurrence. Present at less than 1% in four samples from San Miguel. Absent at Lake Somerville.

Range. Middle Claiborne Lisbon formation to the top of the Jacksonian in Mississippii and Alabama (Frederiksen, 1988).

Symplocos jacksoniana Traverse 1955, p. 73, fig. 13.

Symplocoipollenites jacksonius (Traverse) Potonié, 1960, p. 107.

Proteacidites sp., Engelhardt, 1964, p. 75, pl. 4, fig. 41.

Symplocoipollenites sp., Tschudy and Van Loenen, 1970, pl. 5, fig. 9.

Description. Tricolporate pollen grain of distinctive triangular shape as seen in polar view. Longitudinal furrows very short, scarcely visible in the polar views. Transverse furrows also short and narrow. Conspicuously distinct ektexine and endexine. Sculpture pitted, similar to that of S. paniculata Wall. Columellate tectum. Costae, mostly transverse, under furrow. Size: ca. 24 µm. (Grains have very short polar axes-always present in polar views.) Thickness of exine: ca. 2 µm.

Occurrence. Present in two samples from San Miguel; absent at Lake Somerville.

Range. Symplocos jacksoniana appears in the Middle Claiborne Lisbon Fm. on the Eastern Gulf Coast and occurrences continue through the Eocene-Oligocene boundary (Frederiksen, 1988).

Tricolporopollenites sp. 7. Engelhardt, 1964, p. 74, pl. 3, fig. 32.

Symplocos tecta Frederiksen, 1980a, p. 62, pl. 15, figs. 17-20.

Description. Size 24-36 µm, mean 31 µm, holotype 34µm. Oblate or peroblate; outline rounded triangular. Tricolporate; colpi extend one-third to one-half the distance to poles, not bordered by thickenings; vestibula shallow, often slitlike in optical section, sometimes covered by folds; ora obscure in polar view. Exine 3-4 µm thick midway between apertures; tegillate, ectosexine:endosexine:nexine ratio about 2-4:1:1; sexine thins toward apertures. Exine infraverrucate, elements 0.5-1 µm wide; very fine negative reticulum present.

Affinity. Frederiksen (1980a) believed this taxon to be similar to modern pollen of Symplocos glauca.

Occurrence. One specimen from Sequence E of San Miguel; absent at Lake Somerville.

Range. S. tecta ranges from the base of the Jackson into the Oligocene in Mississippi and Alabama. The range top is not known (Frederiksen, 1988).

Symplocoipollenites sp., Fairchild and Elsik, 1969, p. 84, pl. 37, fig. 15.

Symplocos arcuata Frederiksen, 1980a, p. 60-61, pl. 15, figs. 1-4.

Description. Size 26-30 µm (six specimens), mean 28 µm. Oblate or peroblate; outline triangular with convex sides. Tricolporate, colpi extending about one-third the distance to the poles, not bordered by thickenings; ora obscure in polar view; vestibulum split shaped in optical section because both nexine and sexine are arched outward at the apertures. Exine 1 µm thick excluding ornamentation, weakly tegillate, sexine:nexine ratio 1.5:1, at the apertures, sexine is about 1.3 µm thick and nexine 1 µm thick; thickening of exine at apertures (tumescence) produces darker exine color in aperture region. Exine rugulate to verrucate, elements 0.5 µm wide and 0.2-0.5 µm high; no negative reticulum present.

Occurrence. One grain from San Miguel in Sequence F; absent at Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a)

Description. Normally four pores (in rare cases, also three). These sit symmetrically in relation to each other. Pores with meridional colpi and equatorial pores (?). Polar axis is squarish (in rare cases triangular), symmetrical.

Affinity. Based on a study of the Sapotaceae by Harley (1986), Taylor (1989) decided that the grains of Tetracolporopollenites brevis from the Claiborne Group of Tennessee belonged to Group B of this family. According to Taylor, "The fossil pollen is similar to this extant pollen group in which are found the subfamilies Mimusopsioideae and Madhucoideae. The fossil grains are not identical to the pollen grains of any extent genus and may be from an extant lineage or taxon within the family." (no page)

Paleoenvironmental Notes. Anderson and Muller (1975) mentioned palynomorphs designated as "Palaquium-complex" in their Quaternary pollen flora from Borneo. These grains include sapotaceous genera from both coastal areas and peat swamp forest.

Sapotaceous megafossils were described by Berry (1924) from the Fayette Sandstone of Webb, Grimes, Fayette, and Brazos Counties, Texas. The specimen from the last location was compared to the modern genus Mimusops.

Mimusopsis is a Paleotropical genus. The closely related genus Manilkara comprises tree and shrubs; there are thirty Neotropical species. M. inundata is found on swampy ground below 200 m. in the lowland rainforests of Amazonia; it is particularly frequent on periodically or permanently flooded sites. Two species grow on periodically flooded ground. Most species, however, prefer dry land. One species, M. jaimiqui, grows in coastal scrublands and thickets in Florida, often in association with mangroves. (Pennington, 1990).

Very little is known about the pollination mechanisms of the Sapotaceae. Some are pollinated by bats. Pennington (1990) assumed that individuals would be visited by bees, insects and larger animals because of the floral structure and the tasty nature of the corolla.

Occurrence. Present to 10% in 41 samples from San Miguel and to 2% from eight lignites from Lake Somerville.

Range. Upper Claiborne to Lower Vicksburg (Frederiksen, 1980a).

Genus Ericipites Wodehouse 1933

Ericipites redbluffensis Frederiksen 1980

Description. Size of tetrad 27-32 µm, mean 29 µm, holotype 29 µm. Distinct notches present in outline of tetrad where grains meet. Individual grains more or less spheroidal; outline of grain in polar view triangular with concave to convex sides. Tricolpate with definite geniculi and probably no ora, colpi extending near full length of grain or sometimes syncolpate. Colpi of adjacent grains probably meet fundamentally according to Fischer's rule, but because the grains are syncolpate or nearly so,, colpi of all four grains meet at the center of the tetrad. Exine 1 µm thick, tegillate, ectosexine: endosexine: nexine ratio 1:2:1; sharply infragranulate to finely infraverrucate; outline rough.

Affinity. Frederiksen (1980a) gave the affinity of Ericipites as "Ericaceae?" (p. 65). The tetrads are also very similar to pollen extracted from Eomimosoidea plumosa Crepet and Dilcher, a fossil Leguminosae (Mimoisoideae) flower described by Crepet and Dilcher (1977) from the Middle Eocene of Tennessee. These palynomorphs, placed in the form genus Ericipites, consist of tetrads about 32 µm in diameter. Herendeen and Dilcher (1990) added that the flower most closely resembles the genera Dinizia and Fillaeopsis belonging to the tribe Mimoseae. According to Herendeen and Dilcher, leaves and fruits which co-occur with the inflorescence also resemble Dinizia in many respects.

Paleoenvironmental Notes. Van der Burgh (1967) found pollen of Ericaceae but no wood in his Rhenish brown coals. In the same deposit, wood but no pollen of Leguminosae was present.

Wolfe (1978) pointed out that the Late Claiborne assemblages from Tennessee contains diverse Leguminoseae (Fabaceae) macrofossils. It was significant to him that the family is common in dry tropical vegetation today. Additionally, Berry (1924) described fossils of the genus Mimosites in the Jackson age Fayette sandstone from near Wellborn, Brazos Co., TX.

Crepet (1985) suggested "Chiropterophily (bats), Melittophily (bees), Ornithophily (birds)" as possible pollination mechanisms for Eomimosoidea plumosa, based on the brush type flowers of both that species and modern Mimosoids.

Occurrence. Present to 1% in 25 samples from San Miguel. Absent at Lake Somerville.

Range. Frederiksen (1980a) found E. redbluffensis from the upper part of the Claiborne into the Oligocene.

Aesculiidites circumstriatus (Fairchild) Elsik, 1978, p. 632, pl. 27, fig. 10-18; pl. 28, fig. 1-7.

Tricolpites circumstriatus Fairchild in Stover, Elsik, and Fairchild, 1966, p. 5, pl. 2, fig. 3-7.

Description. Pollen isopolar, slightly prolate, spheroidal. Colpi long, extending almost to the poles. Exine less than 1 µm thick, layering not distinguishable. Colpi margins slightly thickened in the equatorial area, in some with rudimentary transverse furrow. Sculpture consists of minute irregular striations more or less parallel to equatorial plane. In polar view, striations appear roughly concentric around pole. Size range; equatorial diameter, 14 to 16 µm; length 18 to 22 µm based on 21 specimens.

Occurrence. 1% in one sample from Sequence B at San Miguel; absent from Lake Somerville.

Range. Elsik (1968) found this taxon in the Paleocene Rockdale lignite.