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occasional papers of the 


Farlow 
Herbarium of cryptogamic botany 


No. 9 July, 1976 Harvard University, Cambridge, Massachusetts 


Donald H. Pfister | A Synopsis of the Genus Pulvinula 


A New Combination in the Genus Gymnomyces 


Norton G. Miller Studies on North American Quaternary Bryophyte Subfossils 
|. A New Moss Assemblage from the Two Creeks 


Forest Bed of Wisconsin 


Edited by: Reed C. Rollins 
Kathryn Roby 


occasional papers of the 


Farlow. 
Herbarium of cryptogamic botany 


No. 


No. 


No. 


No. 


No. 


No. 


No. 


No. 


No. 


1. Sylvia A. Earle: Hummbrella, a New Red Alga of Uncertain Tax- 


onomic Position from the Juan Fernandez Islands (June 1969). 


. |. Mackenzie Lamb: Stereocaulon arenarium (Sav.) M. Lamb, a 


Hitherto Overlooked Boreal-Arctic Lichen (June 1972). 


. Sylvia A. Earle and Joyce Redemsky Young: Siphonoclathrus, a 


New Genus of Chlorophyta (Siphonales: Codiaceae) from Panama 
(July 1972). 


. |. Mackenzie Lamb, William A. Weber, H. Martin Jahns, Siegfried 


Huneck: Calathaspis, a New Genus of the Lichen Family Cladonia- 
ceae (July 1972). 


. |. Mackenzie Lamb: Stereocaulon sterile (Sav.) M. Lamb and 


Stereocaulon groenlandicum (Dahl) M. Lamb, Two More Hitherto 
Overlooked Lichen Species (March 1973). 


. |. Mackenzie Lamb: Further Observations on Verrucaria serpu- 


loides M. Lamb, the Only Known Permanently Submerged Marine 
Lichen (April 1973). 


. Bruce H. Tiffney and Elso S. Barghoorn: The Fossil Record of the 


Fungi (June 1974). 


. Donald H. Pfister: The Genus Acervus (Ascomycetes, Pezizales). 


1. An Emendation. Il. The Apothecial Ontogeny of Acervus flavidus 
with Comments on A. epispartius (May 1975). 


. Donald H. Pfister: A Synopsis of the Genus Pulvinula. A New 


Combination in the Genus Gymnomyces. Norton G. Miller: Studies 
on North American Quaternary Bryophyte Subfossils. |. A New 
Moss Assemblage from the Two Creeks Forest Bed of Wisconsin 
(July 1976). 


A SYNOPSIS OF THE GENUS PULVINULA 


DonaLp H. PFIsTER! 


SUMMARY 

Seventeen species of the genus Pulvinula are discussed in this treatment. A synoptic 
key to these species is provided as are descriptions of most of them. A review of the 
pertinent literature is given and comments on the morphology of members of the genus 
are provided, One new species, P. neotropica Pfister is described and several new com- 
binations are made. Listed with the synonyms and excluded species of Pulvinula are a 
number of spherical-spored species of Pezizales which were examined in the course of 
this study and which are referred to other genera. 


INTRODUCTION 


Although Pulvinula Boudier is a well-defined member of the Pezizales, 
only Boudier (1907) previously attempted to treat all of the species of 
the genus together. The many species are scattered through the genera 
Crouania, Barlaea, Barlaeina, Detonia, and Lamprospora. To ferret them 
out, type studies have been necessary. In this preliminary report on the 
genus, I have brought together information gleaned from both the litera- 
ture and study of specimens. At present, single collections are heavily 
relied upon to provide details. In recognition of this, I have been very 
conservative in my treatment. I have preferred to cover collections which 
are at minor variance with described species by means of comments on 
those species rather than to describe new taxa prematurely. Therefore, the 
number of species treated is fewer than will finally be assigned to the 
genus. 

During the study of this genus for the Flora Neotropica project, it 
became apparent, as a practicality, that the temperate species required 
first attention. Nevertheless, many of the comments on species refer to 
collections from the Caribbean and will indirectly serve as a guide to the 
species of Pulvinula in that area. 


GENERAL COMMENTS 


The genus Pulvinula was originally mentioned by Boudier (1885) who 
listed Peziza convexella Karst., Peziza sanguinaria Cooke, and Peziza 
constellatio Berk. & Br. as species belonging to it. Later Boudier (1907) 
revised the nomenclature of two of the species and included a full generic 
description. At that time five species were included: Pulvinula cinnabarina 
(Fuck.) Boud., P. carbonaria (Fuck.) Boud., P. constellatio (Berk. & Br.) 
Boud., P. haemastigma (Wedw. ex Fr.) Boud. and P. subaurantia (Bomm. 
& Rouss.) Boud. The genus was not acknowledged until Le Gal's (1953) 
treatment of the genus from Madagascar. In the meantime, species of 


1Farlow Herbarium and Department of Biology, Harvard University, Cambridge, Massachusetts 
02138. 


2 DONALD H. PFISTER 


this genus were referred to Barlaea, Barlaeina, Crouania, Detonia, and 
Lamprospora. Le Gal (1953) selected Peziza convexella as type of the 
genus from the three original species mentioned by Boudier. Both she and 
Boudier considered this species a synonym of Pulvinula haemastigma, a 
view which I do not hold. For further comment on P. haemastigma see the 
discussion under P. convexella. 

In recent treatments, only Rifai (1968), Moravec (1969) and Pfister 
(1972) have attempted to describe and delimit species of Pulvinula. These 
treatments are unfortunately limited by the geographical areas which 
were covered. 

Taxonomic criteria which I consider of value in this genus are: the size 
of the ascospores; the size of the ascus; the presence or absence of croziers; 
the size of the apothecia; apothecial color; the type of substrate on which 
apothecia are produced. To some extent the number of ascospores per 
ascus is useful but there is variation among collections in this respect. 
The number of ascospores per ascus ranges from eight to four. All the asci 
of a single apothecium rarely contain eight ascospores. Pulvinula tetra- 
spora and P. lacteoalba are the only species where the asci consistently 
have four spores. Ascospores of all members of the genus are smooth. 

The construction of the excipular tissue is uniform among the species. 
However, though useful and diagnostic at the genetic level, it is of little 
use in distinguishing among species. The medullary excipulum is com- 
posed of textura intricata, the hyphae of which are thin, rarely exceeding 
4 »m in diam. The ectal excipulum is composed of textura globulosa, the 
cells of which are sometimes compressed, or of textura angularis. Figures 
1, 2A, and 2B show details of cross sections of apothecia. 

The asci of a majority, though not all, of the species here placed in 
Pulvinula arise from prominent croziers which form a characteristic 
horseshoe-shaped, two-pronged base. This particular type of pleurorhy- 
chous crozier (Chadefaud, 1943) is illustrated diagramatically in figure 3. 
Crozier formation has been observed in the majority, though not all, of 
the species of Pulvinula. There is variation from species to species in the 
attenuation of the ascus base and position of the basal septa; however, this 
type of ascus is always obvious no matter what variation might accompany 
it. In one species, P. miltina, there are generally no croziers but in several 
specimens I have examined there were occasional croziers. This is unusual, 
since in other species croziers are either always present or always absent. 

The genus has been placed in the Aleurieae (Dennis, 1968; Rifai, 1968; 
Korf, 1972), based on the occurrence of guttulate spores, of hyphoid hairs 
when present, and of carotenoid pigments. I do not feel this position is 
satisfactory. The species of the genus form a discrete unit which only 
superficially resemble other members of this tribe. The uniqueness of the 
genus is also evident in Arpin’s (1969) chemical analysis of pigments of 
Pulvinula constellatio in which he found a new monocyclic carotenoid 


4 DONALD H. PFISTER 


Fic. 2, A. Pulvinula albida. Portion of ectal excipulum. ru (isotype) & approx. 200. B. Pulvinula 
neotropica, Portion of ectal excipulum. ru (Pfister 811) X approx. 200. Scale equals 25 um. 


designated as P472. For the moment, however, rather than recognize 
yet another tribe in the heterogeneous family Pyronemataceae, I prefer to 
leave the genus in the Aleurieae. Since Pulvinula appears to be eymno- 
Pl g) 
hymenial, whereas at least some of the other Aleurieae are cleistohymenial, 
é / 
developmental studies may shed some light on its position. 


IDENTIFICATION OF SPECIES 


As an aid for the identification of species, I have constructed a synoptic 
key to the accepted species of Pulvinula using the method presented by 
Korf (1972). Using this key, the distinctive species can be easily identified. 
Certain species, however, cannot be separated without consulting descrip- 
tions for precise details of size, color, etc., which if included, would have 
made the synoptic key unwieldy. The numbers given in the key are those 
under which the name appears. The species are arranged in alphabetical 
order. If the fungus has more than one of the described characters, its 
numbers appear in lightface type. Those in boldface appear in only one 
category. 

1. ASCI 
1—1. Normal number of ascospores per ascus 
a. asci 4-spored 7, 16 
b. (asci 8-spored or number of spores 8 or fewer within a single apothecium ) 


A SYNOPSIS OF THE GENUS PULVINULA 5 


1—2. Form of ascus base 
a. (tapering toward the base ) 
b. ascus base abrupt 3, 9 
1—3. Presence of croziers 
a. (croziers present ) 
b. croziers absent 9, 13, 15 
1—4. Ascus length 
a. asci less than 150 um long 2, 6, 8, 17 
b. asci 150-250 um long, 3, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 16, 17 
c. asci greater than 250 um long 1, 4, 5, 13, 15, 16 
2. ASCOSPORES 
2—1. Ascospore size and shape 
a. ascospores elliptical 14 
b. ascospores globose, 9-11 um in diam 1, 2, 6, 10, 12 
c. ascospores globose, 11-16 um in diam 3, 6, 7, 8, 9, 11, 12, 13, 16, 17 
d, ascospores globose, 16 wm in diam or larger 3, 4, 5, 15, 16, 17 
3, PARAPHYSES 
3—1. Branching at the apices 
a. (apices mostly unbranched ) 
b. apices branched 2, 4, 5, 8, 9 
3—2. Apices of paraphyses 
a. (apices hooked or circinate ) 
b. apices straight 1, 6, 15 
3—3. Paraphyses width 
a. (less than 2 um in diam) 
b. greater than 2 um in diam 4, 5, 9, 11, 15 
4. SUBSTRATE 
4—1,. Composition and condition of substrate 
a. (apothecia on soil) 
b. apothecia on burned material, scorched soil, etc. 2, 3, 8, 11 
5. APOTHECIA 
5—1. Apothecial color 
a, apothecia white or light colored 1, 6, 7, 12, 16 
b. apothecia brightly colored 2, 3, 4, 5, 8, 9, 10, 11, 13, 14, 15, 17 
ACCEPTED SPECIES 
1. Pulvinula albida (Rick) Pfister, comb. nov. Fig. 2A 


= Detonia albida Rick, Brotéria 5: 29. 1906. 

Apothecia gregarious, 6-8 mm in diam. Disc white, margin somewhat 
convoluted at least when dry. Ectal excipulum up to 138-161 »m thick at 
the base of the apothecium, composed of globose to compressed globose 
cells 11-13 X 11-16 pm in diam, randomly arranged toward the outside 
and producing loose hyphoid hairs which form a tomentose outer layer. 


6 DONALD H. PFISTER 


Medullary excipulum composed of textura intricata of hyphae 1-2.2 »m in 
diam. Subhymenium not a distinct layer. Hymenium about 280 um thick. 
Asci 265-282 16-20 um, 8-spored. Bases of asci with prominent two- 
pronged croziers. Ascospores uniseriate, globose, hyaline, generally with 
no prominent oil globule, smooth-walled, 10.2-12 pm diam. Paraphyses 
gracile, 1-1.5 wm at base, 1.5-2 pm apically, straight or only slightly 
curved. 


SPECIMEN EXAMINED. Brazil. On soil, Sad Leopoldo, Rio Grande do Sul, 1932, det. 
Rick (FH). 

The specimen in the Farlow Herbarium general mycological collection, 
which appears to be part of the type collection, is the only specimen of 
this species I have seen. The species is close to the neotropical examples of 
Pulvinula globifera but differs in that the asci in the Rick specimen are 
much longer. It is also similar to P. convexella but differs in the form of 
the paraphyses which are straight in P. albida rather than curved as in 
P. convexella and the apothecia are white rather than orange. 


2. Pulvinula archeri ( Berk. in Hook.) Rifai 

= Pexiza archeri Berk. in Hook., The Botany of the Antarctic Voyage. 
II. Flora Tasmaniae 2:274. 1859. = Barlaea archeri ( Berk.) Sacc., Syll. 
Fung. 10:5. 1892. = Humaria archeri ( Berk.) Cooke, Handbook of Aus- 
tralian Fungi. p. 256. 1892. = Barlaeina archeri ( Berk.) Sacc. & Trav. in 
Sacc., Syll. Fung. 20:138. 1910. = Pulvinula archeri (Berk.) Rifai, Verh. 
K. ned. Akad. Wet. II 57(3) :213. 1968. 

= Pexiza gemmea Phil. in Cooke, Mycographia p. 236, fig. 398. 1879. = 
Barlaea gemmea (Phil. in Cooke) Sacc., Syll. Fung. 8:112. 1889. 

— Lamprospora pyrophila Snider, Mycologia 28:484. 1936. 

? = Barlaeina strasseri Bres. in Strasser, Verh. Zool.—Bot. Ges. Wein 
55:613. 1905. 

Apothecia gregarious to scattered up to 7 mm in diam. Disc orange, 
convex, margins more or less even. Ectal excipulum relatively small-celled, 
cells (10-) 13-15 am in diam, globose to subglobose. Medullary excipulum 
of textura intricata, hyphae 2 pm in diam. Hymenium less than 150 pm 
thick. Asci 135-145 x 10-11 »m with or without prominent croziers, taper- 
ing toward the base. Ascospores globose, generally with a single large oil 
droplet, smooth-walled, 9-10.5(-11.5) »m in diam. Eight or fewer asco- 
spores per ascus. Paraphyses slender, 1-2 »m in diam, circinate and pig- 
ment filled, mostly branched apically. 

SPECIMENS EXAMINED. Austria. Holotype of Barlaeina strasseri, Sonntagberg 26. IX. 
1904 (s). U.S.A. Holotype of Peziza gemmea, on fallen leaves of Sequoia sempervirens 
(Harkness 876) (x); on soil in burn area, Benton Co., Mary's Peak, Philomath, Ore., 
H. J. Larsen (no. 247), 11. IX. 71 (osu); on burned material, Polk Co., Camp 
Kilowan and vicinity 5 mi. S. of Falls City, Ore., H. J. Larsen, 27. V. 1972 (osu); on 


soil in fire spot, Benton Co., Mary’s Peak, Philomath, Ore., H. J. Larsen (no, 252), 17. 
IX. 1971 (osu). 


A SYNOPSIS OF THE GENUS PULVINULA 7 


| 


Fic. 3. A-H. Diagrams illustrating the formation and development of asci in most species of 


Pulvinula. 


A collection from Oregon (Benton Co., Mary’s Peak Campground, 27. V. 
1972) sent for examination by Harold Larsen agrees with this species 
except in the form of the paraphyses, where they are practically straight 
and mostly unbranched. 

Based on collections I have seen from the western United States, I pre- 
sume this is the most common species found on burned wood and sur- 
rounding soil in that part of the country. Both Peziza gemmea and 


8 DONALD H. PFISTER 


Lamprospora pyrophila were described from material from the western 
states. The holotype of L. pyrophila in the mycological herbarium at the 
University of Washington was not available for study. 


3. Pulvinula carbonaria (Fuck. ) Boud. 

= Crouania carbonaria Fuck., Jahrb. Nassauischen Vereins Naturk. 
27-28 :64. 1873. = Peziza sanguinaria Cooke, Mycographia p. 14, fig. 19. 
1879 (a name change). = Pulvinula carbonaria (Fuck.) Boud., Hist. 
Class. Discom. d’Eur. p. 70. 1907. 

Apothecia gregarious, 1-3 mm in diam. Dise at first subconcave then 
flat to repand, orange-scarlet. Receptacle smooth, lighter than the disc. 
Ectal excipulum about 115 »m thick, composed of textura globosa to com- 
pressed angular cells, 9-11 16-23 um. Ectal excipulum of textura in- 
tricata, hyphae about 2 pm in diam. Subhymenium not a distinct layer. 
Hymenium about 215 nm thick. Asci 196-230 »m long, 4-8 spored. Ascus 
base moderately broad with a prominent crozier. Ascospores uniseriate, 
globose, hyaline, with a single large guttule and some smaller satellite 
guttules, smooth-walled, 15-17 ym diam. Paraphyses gracile, up to 1.5 pm 
in diam throughout; mostly strongly bent or curved apically with abun- 
dant carotenoids. 


SPECIMENS EXAMINED. Fuckel, Fungi rhenani no. 2482 (ru). U.S.A.: Burnt ground 
in woods, Burbank, E. Tenn., R. Thaxter (7944), Aug. 20-Sept. 5, 1887 (rH). Burnt 
spot in hemlock woods, Purgatory Swamp, Norwood, Mass., D. H. Linder, 27. IX. 
1936 (FH). 


4. Pulvinula cinnabarina ( Fuck.) Boud. 

= Crouania cinnabarina Fuck., Jahrb. Nassauischen Vereins Naturk, 
27-28:64. 1873. = Peziza laeterubra Cooke, Mycographia p. 14, fig. 20. 
1879 (a name change, non Peziza cinnabarina Schw.). = Lamprospora 
laeterubra (Cooke) Lagarde, Ann. Mycol. 4:217. 1906. = Pulvinula cin- 
nabarina (Fuck.) Boud., Hist. Class. Discom. d’Eur. p. 70. 1907. [= 
Lamprospora cinnabarina (Fuck.) Moser in J. Gams, Kleine Kryptogamen- 
flora 2a:112. 1963. ( basionym not cited) ]. 

Apothecia usually gregarious, up to 7 mm in diam. Disc vermillion, 
drying red, margin wavy. Receptacle smooth where exposed. Ectal ex- 
cipulum up to 115 »m thick at the base of the apothecium; composed of 
radially arranged elongate cells, 7 < 11 um in diam. Medullary excipulum 
composed of textura intricata of hyphae up to 4 um in diam. Subhymenium 
not a distinct layer. Hymenium about 280-320 ym thick. Asci 260-275 « 
20-23 wm, 4-8-spored (8 predominating in type collection), The ascus 
tapers toward the base with a definite crozier. Ascospores uniseriate, 
hyaline with one or a few guttules, smooth-walled, 16-20 pm in diam. 
Paraphyses thin, up to 2.2-4 »m in diam throughout, curved or only 
slightly curved apically, sometimes branched. 


A SYNOPSIS OF THE GENUS PULVINULA 9 


SPECIMENS EXAMINED. Fuckel, Fungi rhenani no. 2481 (FH). U.S.A.: On soil among 
mosses, Deep Creek, Great Smoky Mountains National Park, Tennessee, 11. XI. 1970. 
P. E. Powell and D. H. Pfister (349) (FH). 


5. Pulvinula convexella (Karst. ) Pfister, comb. nov. 

= Peziza convexella Karst., Not. Sallsk. Fauna Fl. Fenn. Forh. 10:123. 
1869. == Leucoloma convexellum ( Karst.) Rehm, Ascomycetes Lojkani 
p. 8. 1882. == Barlaeina convexella ( Karst.) Sacc., Syll. Fung. 18:24. 1906. 

= Peziza constellatio Berk. & Br., Ann. Mag. Nat. Hist. Ser. 4. 17:142. 
1876. = Leucoloma constellatio (Berk. & Br.) Rehm, Ber. Naturhist. 
Vereins Augsburg 26:5. 1881. = Aleuria constellatio (Berk. & Br.) Gill., 
Champ. France, Discom. p. 207. 1888. = Plicariella constellatio Lindau in 
Engler & Prantl, Nat. Pflanzenfam. 1(1):180. 1897. = Pulvinula con- 
stellatio (Berk. & Br.) Boud., Hist. Class. Discom. d’Eur. p. 70. 1907. = 
Barlaeina constellatio (Berk. & Br.) Rehm in Dodge, Trans. Wisconsin 
Acad. Sci. 7:1037. 1914. 

= Crouania livida Rehm in Hazsl., Math. Naturwiss. Ber. Ungarn 21: 
262. 1886. = Barlaeina livida (Rehm in Hazsl.) Sacc. & Trott. in Sacc., 
Syll. Fung. 22:621. 1913. 

Misapplication: Pulvinula haemastigma (Hedw. ex Fr.) Boud. sensu 
Boud., Hist. Class. Discom. d’Eur. p. 70. 1907. 

Apothecia less than 8 mm in diam. Disc yellow-orange. Ectal excipulum 
of globose, subglobose to angular cells. Medullary excipulum of narrow 
diameter hyphae forming textura intricata. Asci 230-255(-270) pm X 
16-20 pm. Eight spores or sometimes fewer per ascus. The base of the 
asci with prominent two-pronged croziers. Ascospores smooth-walled, 
uniseriate, globose, hyaline with one or more oil droplets, (16—) 18-20 pm 
in diam. Paraphyses up to 2.3 pm in diam, moderately curved and reg- 
ularly branched apically. 

This is the type species of the genus Pulvinula selected by Le Gal 
(1953). Boudier (1907) synonymized this species with P. haemastigma 
(Hedw. ex Fr.) Boud., a synonymy which has been since followed by most 
workers, including Le Gal. I do not follow this position and consider P. 
haemastigma a nomen confusum. My reason is that the illustration upon 
which the species concept must now be based, in the absence of a speci- 
men, is not adequately diagnostic in several critical features, such as the 
form of the paraphyses, the form of the apothecium, and the configuration 
of the crozier at the ascus base. In addition, there is no information on 
ascus or ascospore size. Boudier (1907) pointed out that Saccardo (1889) 
had interpreted Peziza haemastigma differently than he had. Neither Le 
Gal nor subsequent workers have designated a lectotype from among 
the eligible specimens in the Karsten Herbarium. There are two specimens 
bearing practically the same information which might have been referred 
to by Karsten. Having examined the specimens, I designate number 1273 
lectotype of P. convexella. 


10 DONALD H. PFISTER 


There have been five varieties and one subspecies described and at- 
tributed to Pulvinula convexella or to specific names listed in its syn- 
onymy. These are listed below. Such a proliferation of infraspecific taxa 
suggests something of the diversity within this complex species. 

Peziza constellatio var. fuckelii Cooke, Mycographia p. 45, fig. 82. 1879. 

Barlaea constellatio var. minuta F. E. Clements, Bot. Surv. Nebraska 4: 10. 1896. 

Pulvinula convexella subsp. tjibodensis Penz. & Sacc., Malpigia 16: 202. 1901) 

Detonia constellatio var. aurantiaca F, E. Clements & E. §. Clements, Cryptogamae 
Formationum Coloradensium no. 114. 1906. 

Lamprospora haemotostigma [sic] var. gigantea Thind & Batra, Jour. Indian Bot. 
Soc. 38: 221. 1959. 

Pulvinula haemastigma var. luteoflava Moravec, Ceska Mykol. 23: 25. 1969. 


CRITICAL SPECIMENS EXAMINED. Finland, Lectotype: Tavastia australis, Tammela, 
Mustiala, ad terram nudam ad rivuli marginem in horto, 30. VIII. 1867, leg. et det. 
P. A. Karsten (Herb. Karsten 1273); Travastia australis, Tammela, Saloris (=Soloinen), 
in via, 6. VII. 1878, leg. & det. P. A. Karsten (u, Herb. Karsten 1275); Tavastia 
australis, Tammela, Mustiala, 30. VIII. 1867, leg. et det. P. A. Karsten (u, Herb. 
Karsten 1272). U.S.A. On moist rotted log in shade in spruce-fir forest, Gothic, 
Colorado, leg. Mary Alt, 8. VII. 1963 (osu) and eight additional specimens from 
North America in FH general herbarium. 


A number of specimens from several exsiccati examined in the course of 
this study were issued under one of the synonymous names. Those which 
are unquestionably referrable to Pulvinula convexella are as follows: H. 
Sydow and P. Sydow, Mycotheca germanica, nos. 496 (sub Barlaea con- 
vexella), 1620, 1621 (sub Barlaea constellatio); P. Sydow, Mycotheca 
Marchica, no. 2862 (sub Crouania constellatio); H. Rehm, Ascomyceten, 
no. 406 (sub Detonia constellatio); E. Bartholomew, Fungi Columbiani, 
no. 2909 (sub Barlaea subaurantia ); J. B. Ellis and B. M. Everhart, North 
American Fungi, no. 2306 (sub Barlaea constellatio); F. E. and E. S. 
Clements, Cryptogamae Formation Coloradensium, no. 114 (sub Detonia 
constellatio aurantiaca); A. Allescher and J. N. Schnal, Fungi Bavarici 
Exsiccati, no. 942 (sub Barlaea constellatio ); W. Krieger, Fungi Saxonici, 
no. 1038 (sub Barlaea constellatio ). 


6. Pulvinula globifera ( Berk. & Curt.) Le Gal 

= Peziza globifera Berk. & Curt., J. Linn. Soc., Bot. 10: 366. 1868. == 
Barlaea globifera (Berk. & Curt.) Sacc., Syll. Fung. 8: 114. 1889. = Hum- 
aria globifera (Berk. & Curt.) Cooke, Handbook of Australian Fungi 
p. 256. 1892. = Barlaeina globifera (Berk. & Curt.) Sacc. & Trav. in Sace., 
Syll. Fung. 19: 139. 1910. == Pulvinula globifera (Berk. & Curt.) Le Gal, 
Prodr. Flore Mycol. Madagascar 4:94. 1953. 

My Caribbean collections of Pulvinula globifera have all been white, 
as described earlier (Pfister, 1972). Rifai (1968) also gives a full descrip- 
tion of Pulvinula globifera (Berk. & Curt.) Le Gal. The geographical 
range and the degree of intraspecific variation remain to be clarified. 
Rifai pointed out that Le Gal (1953) misinterpreted this species, and 


A SYNOPSIS OF THE GENUS PULVINULA 11 


considers the species description by Le Gal to be P. orichalcea (Cooke ) 
Rifai. The collection studied by Gamundi (1966) is also probably P. 
orichalcea although I have not seen her specimen. The morphology of the 
paraphyses of P. globifera is variable; occasionally they are curved and at 
times become inflated apically and below the septa. This is especially evi- 
dent at the margin. 

W. C. Denison’s collection from Arizona (osu 23,991) might also be 
referred to Pulvinula globifera. | have no data on the color or size of the 
apothecia when fresh. 

Two varieties of Pulvinula globifera have been described. They are 
listed below, but no attempt has been made to reevaluate their status. 
Both have recently been treated. Rifai (1968) reported that Peziza globi- 
fera var. etiolata was not significantly different from the holotype of 
Peziza globifera to consider it distinct. I have not reexamined Cooke's 
specimen. 

Barlaea globifera var. sphaeroplea (Berk. & Curt.) Sace., Syll. Fung. 
8:114. 1889. = Sphaerosporella brunnea (Alb. & Schw. ex Fr.) Svréek & 
Kubicka fide Rifai (1968). 

Peziza globifera var. etiolata Cooke, Mycographia p. 236, fig. 399. 1879. 
= Pulvinula etiolata (Cooke) Le Gal, Prodr. Flore Mycol. Madagascar 
4:91. 1953, see Rifai (1968). 


SPECIMENS EXAMINED: eleven specimens from Puerto Rico and Guadeloupe (FH). 


7. Pulvinula lacteoalba J. Moravec, Ceska Mykol. 23: 231. 1969. 

This is probably a 4-spored form of Pulvinula globifera and should not, 
perhaps, be considered a distinct species. For a description see the original 
publication by Moravec. 


SPECIMENS EXAMINED. Holotype: Ad terram nudam in olla cum Sparmania africana 
in domo-Liban, districtus Jiecin, 22. V. 1966, leg. Jiri Moravec (PR 674713). 


8. Pulvinula laeterubra (Rehm) Pfister, comb. nov. 

= Barlaea laeterubra Rehm, Ann. Mycol. 5: 516. 1905. = Barlaeina 
laeterubra (Rehm) Sacc. & Trott. in Sacc., Syll. Fung. 22: 622. 1913, == 
Detonia laeterubra (Rehm) Dodge, Trans. Wisconsin Acad. Sci. 17: 1037. 
1914, = Lamprospora wisconsinensis Seaver, North American Cup-Fungi 
(Operculates ) p. 69. 1928 (a name change). 

Apothecia gregarious, 14 mm diam. Disc salmon-red (sometimes 
yellow ). Ectal excipulum 90-115 pm thick composed of textura angularis, 
the cells of which are about 11 17 pm in diam, cells more or less radially 
arranged. Medullary excipulum composed of textura intricata of hyphae 
2 wm in diam. Subhymenium not a distinct layer. Hymenium about 180 »m 
thick. Asci (140-) 161-184 (-200) um, 1-8 ascospores per ascus, ascus base 


has a prominent two-pronged crozier. Ascospores uniseriate, globose, hy- 


1 DONALD H. PFISTER 


aline generally with indiscrete oil droplets, smooth-walled, 11-13.8 »m 
in diam. Paraphyses thin (about 1 um), curved and commonly branched. 


SPECIMENS EXAMINED. U.S.A. Holotype of Barlaea laeterubra, [on soil], Madison, 
Wisconsin, 7. V. 1904, Harper (412) (s, Herb. Rehm), isotype (FH). Also six speci- 
mens in FH general herbarium. 

There are two Puerto Rican collections (DHP 1320 and 1329) which 
differ slightly from Pulvinula laeterubra: they are yellow, and the para- 
physes are broader. 

Seaver (1928) proposed the name Lamprospora wisconsinensis, when 
he treated Barlaea laeterubra in Lamprospora, since he also treated 
Lamprospora laeterubra (Cooke) Lagarde. See the list of excluded species 
for comments on this fungus. He gave the spore size as smaller than 10 pm, 
which is not the case in the type collection where the ascospores are 12- 
14 »m in diam. Red-orange North American collections with spores from 
12-14 um in diam, have been erroneously referred to Pulvinula constellatio 
and P. convexella. They should be referred to Pulvinula laeterubra. 


9. Pulvinula miltina ( Berk. in Hook.) Rifai 

= Peziza miltina Berk. in Hook., Fl. Nov. Zealand. 2: 199. 1855, = 
Barlaea miltina ( Berk.) Sacc., Syll. Fung. 8: 113. 1889. = Humaria miltina 
(Berk.) Cooke, Handb. Austral. Fungi. p. 256. 1892. = Barlaeina mil- 
tina ( Berk.) Sacc. & Trav. in Sacc., Syll. Fung. 19: 139, 1910. = Pulvinula 
miltina ( Berk.) Rifai, Verh. K. Ned. Akad. Wet. II 57(3): 204. 1968. 

For a description of this species of Pulvinula see Rifai (1968). 

Several Caribbean collections are similar to Pulvinula miltina but with 
spores 9-11.5 ym in diam, and asci 140-165 pm long. These measurements 
are smaller than those given for P. miltina by Rifai. The Caribbean col- 
lections are all on charred wood. They have the same broad ascus base as 
is present in P. miltina, but might well represent an undescribed species. 
These collections are listed below. 

SPECIMENS EXAMINED. Guadeloupe, F.W.I. On charred wood and surrounding soil, 
Le Mamelles, Guadeloupe National Park, January 5, 1974 (DHP 813) Fx. Puerto Rico. 


On burned bamboo, El Verde, Luguillo National Forest, April 8, 1974. (DHP 1331) 
Fu; on burned branches, as above (DHP 1325) Fu. 


10. Pulvinula mussooriensis (Thind, Cash & Singh) Batra & Batra 

= Lamprospora mussooriensis Thind, Cash & Singh, Mycologia 51: 
457. = Pulvinula mussooriensis (Thind, Cash & Singh) L. R. Batra & 
S. W. T. Batra, Kansas Univ. Sci. Bull. 44: 167. 1963. 

Pulvinula mussooriensis is very near P. niveoalba except in color and in 
ascus length. In P. mussooriensis the asci exceed 200 »m and the hymen- 
ium is yellow, whereas in P. niveoalba the asci are 160-180 wm and the 
hymenium is white. Both have variable spore numbers with 8 spores per 
ascus predominating. Whether the two species should be distinguished 


A SYNOPSIS OF THE GENUS PULVINULA 13 


from each other is a question which can be resolved only by additional 
field collections and by additional distributional data. Both are known only 
from the type collections. 


SPECIMEN EXAMINED. India. Isotype of P. mussooriensis, (Brewery Road, Mussoorie), 
on soil amid mosses, Aug. 12, 1956 (spr). 


11. Pulvinula neotropica Pfister, sp. nov. Fig. 2B 

Apothecium ad 4 mm diam, convexum, flavovirens, in sicco luridum. 
Asci 165-177 11-14 pm, octospori. Ascospori globosi laevigati (12—) 13— 
14(-15) pm. Paraphyses filiformae ad 4 »m diam ad apices. In ligno 
ustulato, Holotypus: on burned wood, El Yunque, Luquillo National 
Forest, Puerto Rico, D. H. Pfister (1342) and J. D. Rogers, 9. IV. 1974 in 
FH. 

Apothecia gregarious, 24 mm diam. Disc pale yellowish-greenish, con- 
vex, drying buff, margin more or less even. Receptacle pulvinate, smooth 
or slightly furfuraceous. Ectal excipulum 30-45 um thick, of globose cells 
and/or laterally compressed cells, cells 10-15 > 25-32 ym arranged 
radially, 3-5 cells deep. Medullary excipulum composed of more or less 
tightly interwoven textura intricata of hyphae, about 2 »m in diam. Sub- 
hymenium not clearly differentiated from the medullary excipulum. Hy- 
menium about 200 pm thick. Asci 165-177 « 11-14 um, arising from 
prominent croziers, not broad at base, 8-spored. Ascospores uniseriate, 
globose, hyaline, with a single large oil globule, smooth-walled (12) 13- 
14(-15) pm. Paraphyses thin, 2-3 pm below, 3-4 »m toward the apex, 
mostly curved in the upper portion but not strongly. 

SPECIMENS EXAMINED. Puerto Rico. Holotype: on burned wood, El Yunque, Luquillo 
National Forest, D. H. Pfister (1342) and J. D. Rogers, 9.1V.1974 (FH); on charcoal 
in fire spot, El Toro Trail, El Yunque, D. H. Pfister et al. (639), 9.VII.1973 (FH). 
Guadeloupe, F.W.I. On soil and charred wood, Les Mamelles, Guadeloupe National 
Forest, 5.1.1974, D. H. Pfister (811), Martha Sherwood, and Steve Carpenter (FH); 


burn site, Parc Tropicale, D. H. Pfister (1249), S. Carpenter, M. Sherwood, 10.1.1974 
(FH). 


12. Pulvinula niveoalba J. Moravec 

= Pulvinula niveoalba J. Moravec, Ceska Mykol. 23:231. 1969. 

Pulvinula niveoalba can be distinguished by its relatively small apothe- 
cia (0.6-3 mm in diam), white hymenium, ascus size (160-180 »m) and 
ascospore dimensions (9.5-12.2 ym ). 

SPECIMEN EXAMINED. Czechoslovakia. Holotype, ad terram humidam nudam in 


societe Pulvinulae haemastigmae, viae cavae in piceto prope Drhleny, districtus Mlada 
Boleslav, 23. VI. 1969, leg. Jiri Moravec (PR 674714). 


13. Pulvinula orichalcea (Cooke ) Rifai 
= Peziza orichalcea Cooke, Mycographia p. 235, fig. 397. 1879. = 
Barlaea orichalcea (Cooke) Sacc., Syll. Fung. 8:114. 1889. = Barlaeina 


14 DONALD H. PFISTER 


orichalcea (Cooke) Sacc. & Trav. in Sacc., Syll. Fung. 19: 140. 1910. = 
Pulvinula orichalcea (Cooke) Rifai, Verh. K. Ned. Akad. Wet. II. 57(3): 
213. 1968. 

Le Gal's (1953) description of Pulvinula globifera is, according to Rifai, 
based on P. orichalcea. I have seen two collections of this species from the 
Caribbean. 

SPECIMENS EXAMINED. Jamaica. On soil, trail from Whitfield Hall to Portland Gap, 
to Blue Mountain Border of St. Thomas and Portland Parishes, R. P. Korf et al., 17. I. 


1971 (CUP-MJ-593 ); on soil, Traveller’s Rest, Silver Hill Gap on the border of Port- 
land and St. Andrew Parish, R. P. Korf et al., 8. I. 1971 (CUP-MJ-103). 


14. Pulvinula ovalispora Boud. Fig. 1 
= Pulvinula ovalispora Boud., Bull. Soc. Mycol. France 33: 16. 1917. 
This is the only species of the genus reported to have ellipsoidal spores. 

When describing Pulvinula ovalispora, Boudier indicated that, although 

the spores were oval, all other characteristics agree with those of the 

genus. The ascus bases are forked as is typical of the genus. The type 
collection consists of a single, small, partially dissected apothecium, thus 
details of the excipulum were not studied. 


SPECIMEN EXAMINED. Algeria, Ad terram nudam, Februario, leg. Rene Mairé (pc). 


15. Pulvinula salmonicolor (Seav. ) Pfister 

= Lamprospora salmonicolor Seaver, Mycologia 17: 47. 1925. = Pulvi- 
nula salmonicolor ( Seay.) Pfister, Phytologia 24: 211. 1972. 

Apothecia usually solitary, 5-7 wm diam. Disc yellow-orange to orange, 
convex, drying buff-salmon, margin even. Receptacle pulvinate, smooth 
where exposed. Ectal excipulum up to 65 »m thick at the base of the 
apothecium, composed of globose to slightly laterally compressed cells, 
30-35 x 40-45 pm, becoming somewhat radially arranged at the margin, 
random at the base of the apothecium. Medullary excipulum composed 
of tightly interwoven hyphae (textura intricata), 2-3 wm in diam. Sub- 
hymenium not clearly differentiated from the medullary excipulum. 
Hymenium about 375-400 ym thick. Asci 242-286 > 22-23 ym, 8-spored, 
ascus base lacking croziers. Ascospores uniseriate, globose, hyaline, gen- 
erally with a single oil globule, smooth-walled though the cytoplasm 
sometimes gives them a granular appearance, 20-23 um diam. Para- 
physes stout, 4 wm at the base, 7-10 pm apically, straight. 


SPECIMENS EXAMINED. Guadeloupe, F.W.L. On soil among mosses, La Soufriére, July 
20, 1973, D. H. Pfister (594) and W. Sarriera; on soil, Les Mamelles, Guadeloupe 
National Forest, 5.1.1974, D. H. Pfister (887), Martha Sherwood, and Steven Carpenter 
(FH); on soil at base of a banana plant, Camp Jacob, Saint Claude, 500-550 m, 7.1. 
1974, D. H. Pfister (1063), Martha Sherwood, and Steven Carpenter (FH); on soil, 
Pare Tropicale, Basse Terre, 10.1.1974, D. H. Pfister (1197), Martha Sherwood, and 
Steven Carpenter (FH), Puerto Rico. On soil among mosses, El Yunque, 9.IV.1974, 
D. H. Pfister (1340 & 1343). 


A SYNOPSIS OF THE GENUS PULVINULA 15 


The species was discussed earlier (Pfister, 1972). Recent collections 
have permitted this more complete description. 


16. Pulvinula tetraspora (Hansf.) Rifai 

= Lamprospora tetraspora Hansf., Proc. Linn. Soc. New South Wales 
79:126. 1954. == Pulvinula tetraspora (Hansf.) Rifai, Verh. K. ned. Akad. 
Wet. II. 57(3) :207. 1968. 

Misapplication: Pulvinula etiolata (Cooke) Le Gal, Prodr. Flore Mycol. 
Madagascar 4:91. 1953. = Peziza globifera var. etiolata Cooke, Myco- 
graphia p. 236, fig. 399. 1879. 

Rifai (1968) provides a complete description of this species, which, in 
addition to Pulvinula lacteoalba, are the only two species thus far known 
to have exclusively 4-spored asci. 


17. Pulvinula sp. 

There is a distinct species of Pulvinula which resembles P. convexella 
and P. carbonaria but which differs in that the apothecia are on soil rather 
than on charcoal and the asci are smaller than in P. convexella. The only 
specimen of this species I have seen is in Fuckel, Fungi rhenani no. 2290) 
(FH). This specimen was given the name Crouania humosa (Fr.) Fuckel. 
However, Fuckel’s description and specimen do not agree with Fries 
concept of the species, thus that epithet is not available. Future type 
studies may provide an available name for this species. 


SYNONYMS, EXCLUDED SPECIES AND COMMENTS ON SPHERICAL-SPORED 
SPECIES OF PEZIZALES 


In searching for species of Pulvinula, a number of species were exam- 
ined which were referrable to other genera of the Pezizales or were 
synonyms of accepted species of Pulvinula. They are listed alphabetically 
below under the accepted name if one could be determined. 


LAMPROSPORA ASPERELLA (Rehm) Boud., Hist. Class. Discom. d’Eur. p. 69. 
1907. = Crouania asperella Rehm, Hedwigia 24:226. 1885. = Barlaea 
asperella (Rehm) Sacc., Syll. Fung. 8:113. 

The placement of this species in Lamprospora by Boudier (1907) seems 
to be correct. The structure of the excipulum agrees with that found in 
Lamprospora sensu Rifai (1968). Seaver (1928) synonymized it with L. 
crec hqueraulti. 


LAMPROSPORA ASTROIDEA (Hazsl. in Cooke) Boud. Hist. Class. Discom. 
d'Eur. p. 68. 1907. = Peziza astroidea Hazsl. in Cooke, Mycographia p. 29, 
fig. 49. 1879. = Barlaea astroidea (Haszl. in Cook) Sace., Syll. Fung. 8: 
111. 1889. 

Peziza astroidea was treated by Boudier (1907) as Lamprospora. I 
have not seen a specimen of this species but follow Boudier’s placement 


16 DONALD H. PFISTER 


of it. Cooke’s plate shows the apothecia with fimbriate margins which 
would certainly substantiate Boudier’s deposition of it. 


BARLAEINA CENTROSPORA Kirschst., Ann. Mycol. 33:206. 1935. 

This is a species of Lamprospora sensu stricto and is close or identical 
to Lamprospora crechqueraulti. The specimen should be reexamined 
when a critical revision of the genus Lamprospora is done. Holotype in B 
examined. 


PLICARIA CHAIGNONI Pat., Bull. Soc. Hist. Nat. Autun. 17:154. 1904. = 
Pulparia planchonis (Dun. ex Boud.) Korf, Pfister, and Rogers, Phytologia 
21:206. 1971. Holotype in rH examined. 


LAMPROSPORA CHOPRAIANA Batra, Mycologia 52:665. 1960. 
This is not a Pulvinula but type material is too scanty to make an identi- 
fication. Rifai (1968) excluded it from Lamprospora. 


PULVINULA CONSTELLATIO (Berk. & Br.) Boud., Hist. Class. Discom. d’Eur. 
p- 70. 1907. 

Pulvinula constellatio has been accepted as a distinct taxon by most 
workers. I feel it intergrades with P. convexella and tentatively regard it 
as a synonym as did von Hohnel (1917). Pulvinula constellatio is part of a 
complex of species, including P. cinnabarina, P. convexella, and P. laeteru- 
bra, which is yet to be adequately defined. Complete synonymy of this 
species is given under P. convexella. 


BARLAEA DISCOIDEA P. Henn. & E. Nym., Monsunia 1:33. 1900. = Barlaeina 
discoidea (P. Henn. & E. Nym.) Sacc. & Syd., Syll. Fung. 16:710. 1902. = 
Pulvinula discoidea (P. Henn. & E. Nym.) Batra, Univ. Kansas Sci. Bull. 
44:143. 1963. 

The type material of this taxon was not located. In this group, where 
microscopic features are critical for proper and accurate identification, no 
attempt is made to place it. 


PULVINULA ETIOLATA (Cooke) Le Gal, Prodr. Flore Mycol. Madagascar 
4:91. 1953. = Peziza globifera Berk. var. etiolata Cooke, Mycographia 
p. 236, fig. 399. 1879. 

Rifai (1965) considered Le Gal’s use of this name a misapplication. 
According to him, the specimen described is referrable to Pulvinula tetra- 
spora. Peziza globifera var. etiolata, according to Rifai, is indistinguishable 
from Pulvinula globifera. I have not studied Cooke's specimens. 


BARLAEINA FEURICHIANA Kirschst., Ann. Mycol. 33:205. 1935. 
This is a species of Lamprospora sensu stricto. Holotype in B examined. 


PEZIZA GEMMEA Phil. in Cooke, Mycographia p. 236, fig. 398. 1879. 
This is a synonym of Pulvinula archeri (Berk. in Hook.) Rifai. 


A SYNOPSIS OF THE GENUS PULVINULA 17 


BARLAEINA HENNINGSI Kirschst., Notizbl. Bot. Gart. Berlin-Dahlem 15:830. 
1943. 

This is a species of Lamprospora, close or identical to L. crec hqueraulti. 
Holotype in B examined. 


CROUANIA KNAJASCHENSIS Karst., Hedwigia 23:37. 1884. = Barlaea knajas- 
chensis (Karst.) Sacc., Syll. Fung. 8:113. 1889. = Lamprospora knajaschen- 
sis (Karst. ) Boud., Hist. Class. Discom. d’Eur. p. 68. 1907. 

No specimen of this species could be located in the Karsten Herbarium 
at Helsinki. From the original description, it appears to be a species of 
Pulvinula since it is said to have smooth spores, curved paraphyses which 
are relatively narrow, and asci within the proper size range. However, it 
was treated by Boudier (1907) as a Lamprospora. Without a specimen 
the question is irresolvable. 


PEZIZA LAETERUBRA Cooke, Mycographia p. 14, f. 20. 1879. = Lamprospora 
laeterubra (Cooke) Lagarde, Ann. Mycol. 4:213. 1906, 

This name was proposed as a substitute for the epithet cinnabarina 
when Cooke wished to transfer Crouania cinnabarina Fuck. to Peziza. 
Both Cooke (1879) and Lagarde (1906) described and illustrated the 
globose ascospores as being finely reticulate. I have not examined their 
specimens, but Fuckel’s specimen in Fungi rhenani no. 2481 is definitely a 
Pulvinula and has smooth ascospores. The fungus with ornamented spores 
illustrated and discussed by Cooke and Lagarde is apparently without a 
legitimate name. 


CROUANIA Livipa Rehm ex Hazsl., Math. Naturwiss. Ber. Ungarn. 21:262. 
1886. 

This is a synonym of Pulvinula convexella, which see for complete syn- 
onymy. Apparently Rehm did not intend to publish this as a new species 
since the packets are annotated as Peziza convexella Karst. and the 
species was published as Leucoloma convexellum (Rehm 1882). Specimen 
examined: in excavatione terrae callis “Riu mare” infra alpen Retyezat 
Transsylvaniae 8/1872, Lojka sub 1870. 


BARLAEINA PLATENSIS Speg., Anales Mus. Nac. Hist. Buenos Aires ser: 3. 
1:70. 1902. 

The ascospores of this species are not spherical but rather are broad 
ovoid-ellipsoid, 10-12 % 9-10 »m in diam and smooth. The hairs are 
flexuous and blunt tipped and arise from = globose cells of the outer 
layer of the ectal excipulum. The cells of the ectal excipulum are brown 
colored. The paraphyses are characteristic in that they are swollen apically 
and many times swollen again below the septa giving the paraphyses a slim 
moniliform appearance. This may represent a species of Leucoscypha. 
Holotype in Lps examined. 


18 DONALD H. PFISTER 


LAMPROSPORA PYROPHILA Sydner, Mycologia 28:484. 1936. 

The holotype is deposited at the University of Washington herbarium 
but for some reason is not available for study. I wish to thank Dr. Amy Y. 
Rossman of Oregon State University for locating the specimen in the Uni- 
versity of Washington herbarium. Since a number of specimens from the 
northwest were examined and found to agree with both the original 
description of Lamprospora pyrophila and with Rifai’s (1968) description 
of Pulvinula archeri, Lamprospora pyrophila is here considered to be a 
synonym of Pulvinula archeri. 


BARLAEINA ROSEA Kirschstein in Schieferdecker, Die Schlauchpilze der 
Flora von Hildescheim p. 97. 1954. 

This is a species of Lamprospora. It appears to be very similar to L. 
dictydiola Boud. Holotype in B examined. 


PEZIZA SANGUINARIA Cooke, Mycographia p. 14, fig. 19. 1879. 

The name was introduced by Cooke as a nomen novum to replace the 
epithet carbonaria when he transferred Crouania carbonaria to Peziza. 
See Pulvinula carbonaria for complete notes on this species. 


BARLAEINA STRASSERI Bres. in Strasser, Verh. Zool.—Bot. Ges. Wein 55: 613. 
1905. 

This is a Pulvinula close, if not identical, to Pulvinula archeri. The small 
ascospores, 8-10 wm in diam, and the apically branched paraphyses are 
very characteristic. Holotype in B examined. 


PEZIZA SUBAURANTIA Bomm. & Rouss., Bull. Soc. Roy. Bot. Belgique 23: 
134. 1884, = Barlaea subaurantia (Bomm. & Rouss.) Sacc., Syll. Fung. 8: 
114. 1889. = Pulvinula subaurantia (Bomm. & Rouss.) Boud., Hist. Class. 
Discom. d’Eur. p. 70. 1907. 

I was unable to locate the holotype of this species in either BR or in PC, 
thus its placement within the genus is unresolved. The specimen in Fungi 
Colombiani no. 2909 issued under this name is Pulvinula convexella. 


ACKNOWLEDGEMENTS 


I wish to thank the curators and directors of the following herbaria for searching for 
and forwarding specimens to me for study: The Herbarium and Library, Royal Botanic 
Gardens, Kew; Instituto de Botanica C. Spegazzini, La Plata; Section for Botany, 
Swedish Museum of Natural History (Naturhistoriska Riksmuseet), Stockholm; Botani- 
cal Museum, University of Helsinki, Helsinki; Muséum National d’Histoire Naturelle, 
Laboratoire de Cryptogamie, Paris; National Fungus Collections, Agricultural Research 
Center-West, Beltsville, Maryland; Plant Pathology Herbarium, Cornell University, 
Ithaca, New York; Botanischer Garten und Botanisches Museum, Berlin-Dahlem; 
Herbarium, Department of Botany and Plant Pathology, Oregon State University, 
Corvallis, Oregon. 

I also wish to thank Dr. Richard P. Korf for reading the manuscript and making 
valuable suggestions. 


A SYNOPSIS OF THE GENUS PULVINULA 19 


LITERATURE CITED 


Bouvier, E. 1885. Nouvelle classification naturelle des Discomycétes charnus. Bull. 
Soc. Mycol. France. 1: 91-120. 

—————— . 1907. Histoire et Classification des Discomycétes d’Europe. Klinchsieck, 
Paris. 221 pp. 

Cooke, M. C. 1879. Mycographia, seu icones fungorum. I. Discomycétes. Williams and 
Norgate, London. 267 pp. 

Dennis, R. W. G. 1968. British Ascomycetes. J. Cramer. Lehre. 455 pp. 

Gamunpi, I. J. 1966. Nota sobre Pezizales bonaerenses con comentarios sobre el 
“status” de algunos géneros. Revista Mus. La Plata. Secc. Bot. 10: 47-68. 

Houne., F. X. R. von. 1917. Mycologische Fragmente, CXX—CXC. Ann. Mycol. 15: 
293-383. 

Korr, R. P. 1972. Synoptic key to the genera of the Pezizales. Mycologia 64: 937-994. 

LacarpE, J. 1906. Contribution a l’études de Discomycétes charnus. Ann. Mycol. 4: 
125-256. 

Lr Gat, M. 1953. Les Discomycétes de Madagascar. Prodr. Flore Mycol. Madagascar 
4: 1-465. 

Moravec, J. 1969. Nékteré operkulatni diskomycety nalezené v okresech Mlada 
Boleslav a Jitin. Some operculates found in the districts of Mlada Boleslav and 
Jitin. Ceska Mykol. 23(4) : 222-235. 

Puiuies, W. 1887. A Manual of the British Discomycetes. Kegan Paul, Trench and 
Co., London. 461 pp. 

Prister, D. H. 1972. Notes on Caribbean Discomycetes. II. Two species of Pulvinula 
from Puerto Rico. Phytologia 24; 211-215. 

ReuM, H. 1882. Ascomycetes Lojkani lecti in Hungaria, Transsilvania et Galicia. 
Friedlander & Sohn, Berlin. 70 pp. 

Rirar, M. A. 1968. The Australasian Pezizales in the Herbarium of the Royal Botanic 
Gardens, Kew. Verh. K. ned Akad. Wet. II 57(3): 1-295. 

Saccarpo, P. A. 1889. Sylloge Fungorum hucusque cognitorum. 8. Patavii. 1143 pp. 

SEAVER, F. J. 1928. North American cup-fungi (Operculates). Published by the author. 
New York. 284 pp. 


STUDIES ON NORTH AMERICAN QUATERNARY 
BRYOPHYTE SUBFOSSILS 
I. A NEW MOSS ASSEMBLAGE FROM THE TWO CREEKS 
FOREST BED OF WISCONSIN! 


Norton G. MILLER? 


SUMMARY 


Plant fossils in an exposure of Two Creeks Forest Bed peat about 11,850 years old, 
near Green Bay, Wisconsin, include 32 species of mosses representative of a variety of 
forest and nonforest habitats. The assemblage, which contains few aquatics and many 
calciphiles, establishes that a diverse flora of temperate, boreal, and arctic mosses 
occurred in northwestern Wisconsin just prior to and during active glaciation. Most 
of the mosses identified presently grow in Wisconsin or elsewhere in the upper Great 
Lakes area, but two, Aulacomnium turgidum and Hypnum bambergeri, now rarely 
occur farther south than arctic and subarctic regions. Present-day occurrences of some 
of these species in the Great Lakes region may, in part, date from the period when 
forest bed sediments accumulated. The represented vegetation, an open, more or less 
dry Picea glauca woodland with rich fens and dry sites, perhaps on dune sand, is 
inferred from pollen spectra, mosses, cones, seeds and twigs obtained from the peat. 


The Two Creeks Forest Bed is a buried organic deposit first described 
in detail from exposures along the shore of Lake Michigan near the base 
of the Door Peninsula in northeastern Wisconsin. The forest bed contains 
a record of vegetation and soil that developed during an ice-free period 
between two glacial advances late in Wisconsinan time. Both glaciations 
overrode northeastern Wisconsin leaving behind tills which under- and 
overlie the forest bed. These deposits, the buried forest bed, and associ- 
ated sediments contain a record of environmental conditions that may 
extend back 14,000 radiocarbon yrs B.P. The forest bed is generally con- 
sidered to represent part of the interval between 11,000 and 12,500 yrs 
B.P. ( Black, 1970). 

Paleobotanical and paleoenvironmental studies of the buried forest 
were first carried out by Wilson (1932, 1936) who provided a compre- 
hensive analysis of the deposits and their contained fossils. Foremost 
among plant materials recovered were mosses, 19 of which were identified 
in forest bed peats and clays from two sites (Cheney, 1930, 1931; Wilson, 
1936). The orientation and manner of growth of certain mosses in the 


1 The first in a series aimed at providing a better understanding of the development of distribution 
patterns of North American bryophytes during Pleistocene and Holocene times. The usual American 
usage of Holocene, i.e., postglacial time, which extends from about 10,000 yrs Before Present 
(B.P.) to now, is being followed, even though some feel that the “postglacial” is best treated as a 
subdivision of the Pleistocene. Because the geologic setting and general paleobotany of deposits 
containing fossil Bryophyta help to determine what environmental conditions prevailed when burial 
occurred, these topics are treated to some extent also. To document the sometimes fragmentary 
remains of bryophytes encountered in Quaternary sediments, illustrations or citations of published 
drawings of fossil material are generally provided. The term subfossil indicates that specimens, 
though found in a fossil context, are not petrified or coalified, but consist of essentially unaltered 
tissues that almost always lack cellular contents. 

2 Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138 


21 


22 NORTON G. MILLER 


deposits, distribution of species within the sediments, and the preponder- 
ance of aquatic and subaquatic mosses facilitated reconstruction of the 
plant communities represented and geological events associated with 
deposition of the upper till. Culberson (1955) also studied mosses in the 
forest bed and reported species not previously found. In addition, mosses 
are mentioned by Schweger (1969) as occurring in a deposit of Two 
Creeks peat about 50 km northwest of exposures along the Lake Michigan 
shore. The vegetation of northeastern Wisconsin during Two Creeks time, 
based on pollen analysis, has been treated by West (1961) and Schweger 
(1969). 


LOCATION AND GEOLOGIC SETTING 


The forest bed exposure under consideration occurs in Scott, Wisconsin, 
mostly in the southwest quarter of Sect. 23, T. 24 N. / R. 21 E., on the 
Norbert F. Peters farm, about 6.5 km northeast of the city of Green Bay. 
According to unpublished data assembled by the late F. T. Thwaites 
when he visited the pit on 30 June and 1 July 1958, the sediments were 
exposed on a north-south trending wall of a borrow pit, then about 75 m 
long and 10 m deep (Fig. 1 & 2). The topmost deposit was a pale red 
glacial till, the base of which was fairly level and not over 2.5 m below the 
top of the exposure. Sediments below the till, however, were mixed and 
were not organized into horizontal strata to any great extent. At certain 
places dark red, obscurely stratified clay occurred immediately beneath 
the till, but at others sand was present. Thwaites’ manuscript notes record 
that peat, associated logs, and other woody materials (Fig. 2) occurred 
intermixed with the clay at an isolated position along the wall. However, 
in 1957, Pfleger (Rubin & Alexander, 1960, p. 153) noted an organic rich 
silt and sand layer about 2 m thick with tree trunks and forest litter below 
the till. Since active digging was going on at this time, these observations 
indicate the organic bed and associated sediments were variable laterally. 
Red clay also underlies the peat, but sediments beneath this clay were 
covered or remained unexcavated, and therefore are not described in 
information available to me. 

Stratigraphic relationships at the Peters’ borrow pit are less clear than 
at comparable exposures along the western shore of Lake Michigan, but 
Thwaites (ms. notes) interpreted deposits in the two areas similarly. He 
considered sediments at the pit to have been laid down in a lake im- 
pounded in front of the ice that produced the upper till. In his opinion, 
the peat-log mass was rafted in because of the absence of roots in the 
position of growth. R. F. Black, who studied the site several times in 
the 1960's, interprets (in litt.) the organic matter as having been incor- 
porated into lake sand when the rising lake level drowned the forest in 
front of the advancing ice. He notes that none of the trees were in growth 


QUATERNARY BRYOPHYTE SUBFOSSILS 23 


Fic. 1. Sediments exposed on west-facing north-south wall of borrow pit near Green Bay, Wis- 
consin, summer 1958. Organic deposit about midway up the wall. (Photograph courtesy of H. H. 
lltis. ) 


Fic. 2 Close-up of most of the organic deposit showing logs and other woody material; 


a 


deformed peat layer visible at left. Summer 1958. (Photographs courtesy of H. H. Iltis. ) 


24 NORTON G. MILLER 


position and that the organic bed was folded and disturbed greatly by the 
overriding ice. Although these workers indicate possible redeposition of 
organic materials, some of the blocks of sediment I studied have an un- 
mixed vertical succession of species that is consistent with a shift from 
more or less dry and open forest communities to those that are wetter. 
At least part of the forest bed, therefore, appears to have been preserved 
as it had accumulated. 

The following age determinations have been published for wood as- 
sociated with peat in the borrow pit: 11,940 + 390 yrs B.P. (Y-147X, 
Preston et al., 1955, p. 958) and 11,140 + 300 yrs B.P. (W-590, Rubin & 
Alexander, 1960, p. 153). These determinations agree favorably with the 
generally accepted age of the Two Creeks Forest Bed, 11,850 + 100 yrs 
B.P. (Broecker & Farrand, 1963), which has been taken by some to mark 
the end of Twocreekan time (Black & Rubin, 1968 ). 

To summarize, the Two Creeks forest developed on lacustrine sediments 
deposited out of a lake that stood higher than the present level of Lake 
Michigan. The lake beds accumulated on till whose precise age is not 
known but is often correlated with an advance of the late Woodfordian 
Cary ice or a somewhat more recent readvance. This lower till was not 
recorded at the Peters’ borrow pit but can be seen along the shore of Lake 
Michigan and elsewhere in the immediate area. Forest became established 
on the lake sediments following a lowering of the lake level, and soil 
development and peat accumulation occurred. Glacier ice readvanced into 
the region about 11,850 yrs B.P. causing the lake level to rise and de- 
stroy the forest. Lacustrine sediments accumulated on top of the soil for 
some time following this date. Ice later spread over northeastern Wiscon- 
sin and then receded, leaving behind a deposit of reddish drift generally 
considered to correlate with the Valders till, described from somewhat 
farther south in Wisconsin. Recent data published by Evenson (1973) 
indicate that till above the forest bed is younger than the Valders, al- 
though this interpretation has been contested by Black (1974; see also 
Evenson et al., 1974). Evenson (1973) has proposed the name Two Rivers 
till for that present above the Two Creeks forest bed. 


METHODS 


Ten large, dry blocks of sediment (ca. 20 x 10 & 10 cm) were placed 
individually in beakers, covered with a solution of 0.5% trisodium phos- 
phate and heated gently to loosen clays and silts and rewet the plant 
materials (Benninghoff, 1947). Inorganic materials were washed from the 
organic fraction through a small mesh sieve with distilled water after 
about 3 hr of treatment. Residues were soaked for several hours in distilled 
water to leach out all trisodium phosphate and were bottled in dis- 
tilled water or 70% ethyl alcohol until studied microscopically. 


QUATERNARY BRYOPHYTE SUBFOSSILS 25 


To determine the variability of residues, subsamples removed from a 
well-mixed slurry of the wet fossil material were separated into their 
component species. The area occupied by individual taxa was determined 
by measuring a tightly packed layer of fragments, one plant deep, using 
a grid divided into square centimeters. This procedure was repeated twice 
for two residues to establish the reproducibility of the sampling method, 
and data from three additional sediment blocks were gathered. Data for 
individual samples were calculated in percentages. 

A sample of the organic material cut from the center of a dry sediment 
block was analyzed for pollen following procedures outlined in Faegri 
and Iversen (1964). Pollen counts from inorganic sediments were not 
made because of possible contamination and uncertainty about the orig- 
inal orientation of sediment blocks in the exposure. 


RESULTS 


A compact moss layer, resting directly on an accumulation of forest 
litter or soil, occurred in each sediment block. The combined thickness of 
the organic material varied from sample to sample but the maximum noted 
was 5 cm. Intermixed clays and silts were absent except for a small amount 
of inorganic sediment found with plant fragments along the upper and 
lower limits of the organic layer. No plant fossils were found embedded 
in the silts and clays which under- and overlie the organic horizon. 

Pollen analysis and vascular plant megafossils. Pollen counts obtained 
from samples of the thin (ca. 3 cm thick) moss layer in two sediment 
blocks are given in Table 1. The principal trees represented are spruce 
(Picea spp.) and tamarack (Larix laricina) which together account for 
about 80% of the percentage base. Pine pollen is nearly absent. About 15% 
of the sum consists of various nonarboreal (NAP) taxa, especially mem- 
bers of the Cyperaceae, which comprise about one-half of the NAP totals. 
These pollen assemblages agree with results obtained by West (1961) 
from samples of Two Creeks peat exposed along the shore of Lake Michi- 
gan, with the exception of somewhat higher spruce (80%) and lower 
Cyperaceae (usually < 5%) representation in most of his spectra. A pollen 
spectrum in Schweger (1969) from sediments at the borrow pit contains 
much less spruce (ca. 40%) and more Cyperaceae and Gramineae (ca. 
45%) than either spectra reported here. Though differing quantitatively, 
the counts are similar in terms of pollen types represented, and the differ- 
ences probably relate to the mixed nature of the “composite sample” 
Schweger processed (i.e., peat and organic silt mixed). My analyses were 
of single, relatively thin, strata. 

The organic bed contains abundant cones, twigs and needles of spruce 
and tamarack (Fig. 3-9). Seeds of the two genera were also found, but 
no other seeds (or fruits) were noted. Although some spruce cones are 


26 NORTON G. MILLER 


TABLE 1. POLLEN CONTENT OF MOSS LAYER AT THE PETERS’ BORROW PIT, 


BROWN CO., WISCONSIN 


AP? Sample 1 
n % 

Pinus 2 0.6 
Picea 241 68.5 
Larix 31 8.8 
Betula -- — 
Carpinus-Ostrya — _ 
Quercus 4d 1.1 
Carya 1 0.3 
Ulmus 1 0.3 
Populus 9 2.6 
AP totals 289 82.1 

NAP* 
Alnus — — 
Myrica - - 
Salix 7 2.0 
Cyperaceae 23 6.5 
Gramineae 8 2,2 
Ambrosia vi 2.0 
Artemisia 7 2.0 
Xanthium il 0.3 
High-spine Compositae 10 2.8 
NAP totals 63 17.9 

MISC” 
Polypodiaceae 3 0.8 
Moss spores _ _ 
Unidentifiable 24 6.4 


Sample 2 
% 


n 
2 0.5 
271 71.5 
35 9.2 
1 0.3 
2 0.5 
1 0.3 
1 0.3 
1 0.3 
10 2.6 
324 85.5 
1 0.3 
1 0.3 
2 0.5 
24 6.3 
11 2.9 
‘i 1.8 
5 1.3 
4 1.1 
55 14.5 
7 1.7 
1 0.2 
17 4.2 


“Percentage base = sum arboreal pollen (AP) + nonarboreal pollen (NAP) 


. Percentage base = sum AP + NAP + MISC 


QUATERNARY BRYOPHYTE SUBFOSSILS af 


Fics. 3-9. Cones and twigs from Two Creeks peat at Peters’ borrow pit.—3—5. Cones of Picea 
glauca (Moench) Voss.—6. Probable cone of Picea mariana (Mill.) B.S.P.—7—8. Cones of Larix 
laricina (DuRoi) K. Koch.—9. Twig of Larix laricina. 


small for the species, most were identified as white spruce (Picea glauca) 
because the cones are at least two times longer than broad and their scales 
have entire margins. Because cones beneath living white spruce trecs are 
somewhat variable in size and may be decayed to the point that the scale 
margins are no longer entire, identifications were made using only well- 
preserved specimens. About nine of every ten cones in the sediment blocks 
were of P. glauca, the remainder being assignable to P. mariana. 

Bryophyte remains. Mosses in some sediment blocks were preserved 
mostly as unfragmented plants, indicating that water transport was mini- 
mal and that in situ burial probably occurred. Other sample residues 
contain mixtures of species from different habitats, and in these cases it 
seems likely that the assemblage was brought together by moving water, 
No liverworts were found. 

In order to determine species composition at various levels in the 
organic bed, two sediment blocks showing clay—moss/ forest litter—clay 
layering were cut in two vertically and the cut edges were soaked in water 


28 NORTON G. MILLER 


to facilitate examination. A compact, peaty forest litter, 3 to 4 cm thick 
was present in both blocks. Materials of Drepanocladus uncinatus and 
Thuidium abietinum (subordinate) occurred near the top of the litter, 
together with cones and needles of white spruce and tamarack. While 
both of these mosses occur in a variety of nonforest habitats, D. uncinatus 
is a typical species of moist to dry conifer stands, and T. abietinum is 
sometimes found in open white spruce forests on dry soil. Downward in 
the litter, the mosses drop out and the amount of degraded plant material 
increases. Above the forest litter and on top of the mosses was a dense 
mat of Tomenthypnum nitens, which generally occurs in wet, calcareous 
habitats that can be forested or open. Relatively few conifer needles and 
no cones were found with this moss, indicating a decline in the abundance 
of trees at the site. A shift from more or less dry to wetter conditions 
indicates a rise in the water table possibly relating to flooding associated 
with the advance of glacier ice or to periglacial activity. 

Variability in the composition of moss assemblages obtained from five 
sediment blocks is given in Table 2. Three of the samples yielded fifteen 
or more species while two had eight or nine. Some species occur in all 
samples but others are present in just one or a few. Four samples are 
dominated by Tomenthypnum nitens and from one to seven subdominant 
species occur with it. About half of the identified species are rare either in 
a given sample or in all examined materials. It is tempting to conclude 
that numerical relationships between species listed in the table also held 
in nature. However, the percentage values, which though similar to field 
determined estimates of percent cover, are not strictly comparable to 
measurements of abundance in nature for at least two reasons: sample 
residues are in part mixtures of species from distinct communities that may 
have existed at different times, and it is impossible to reconstruct the 
orientation of the mosses as they actually grew, therefore making measure- 
ments of area only an approximation of the original situation. 

Mosses identified in the forest bed are given below, with an indication 
of their abundance in the residues. The following scale is used: rare—10 or 
fewer fragments; sparse—11 to 20; rather abundant—21 to 40; abundant—41 
to 60; and very abundant— > 60. A fragment varied from isolated leaves to 
leafy branches to essentially complete plants. Notes are also given on the 
present distribution of identified taxa (particularly in the Great Lakes 
region), diagnostic characteristics and the occurrence of structures associ- 
ated with the sexual reproductive cycle (e.g., archegonia, antheridia, 
sporophytes, etc.). Unless otherwise specified, a record documenting a 
recent collection of the species in Wisconsin has been found (see Cheney 
& Evans, 1944; Forman, 1967). Species indicated by an asterisk have not 
been previously reported from the forest bed. Crum’s recent book (1973) 
on the mosses of northern Michigan contains helpful notes on the habitat 
preferences of many of the species found in the forest bed. 


QUATERNARY BRYOPHYTE SUBFOSSILS 


TABLE 2, AREAS (PERCENT COVER) OCCUPIED BY MOSS SUBFOSSILS 
IN SAMPLES ONE TO FIVE 


Sample—Subsample 
Designation 

Species 
Tomenthypnum nitens 
Bryum pseudotriquetrum 
Campylium polygamum 
C. stellatum 
Brachythecium turgidum 
Ditrichum flexicaule 
Drepanocladus uncinatus 
Thuidium abietinum 
Hylocomium splendens 
Scorpidium turgescens 
Hypnum bambergeri 
Distichium capillaceum 
Drepanocladus aduncus var. polycarpus 
Eurhynchium pulchellum 
Amblystegium serpens 
Tortella fragilis 
Drepanocladus sp. 
i IN recurvirostrum 
Dicranella heteromalla 
Mnium marginatum 
Tortella sp. 
T. tortuosa 
Myurella julacea 
Encalypta procera 
Aulacomnium palustre 
A. turgidum 
unknown 


TOTALS 


*From samples analyzed for pollen 


1 2 3 4 
a® b? a® b* 

67.9 67.2 54.7 52.8 70.3 83.8 
7.7 7.3 101 122 08 £03 
88 64 91 112 63 £76 
1.6 18 8.1 6.1 6.3 trace 

—- O02 5.1 4.1 _ — 
1.1 18 4.1 4.1 9.3 — 
44 9.1 a — 3.9 1.3 
5.5 3.6 _ — — 6.7 
1.1 0.5 — —- 2.3 — 

— — = 0.8 — 

_ — 4) 4.6 — _ 

trace trace 3.0 4.] -_ = 
= ae 15 m0) — trace 
0.7 0.9 - — — _ 
0.5 0.4 trace — — 03 

— 0.5 trace — — — 

—_ — 02 — — — 
0.2 0.2 — — — _ 
0.2 0.2 — — — trace 
0.2 — trace trace = — 

— — trace trace — _ 

_— — trace trace _ — 


99.9 100.1 100.0 100.2 100.0 100.0 


29 


trace 

0.2 
trace 
trace 
trace 
trace 
trace 
100.0 


30 NORTON G. MILLER 


14 


Fics. 10-16. Subfossil mosses from Two Creeks peat at Peters’ borrow pit.—l10. Sphagnum 
papillosum Lindb., cross section of branch leaf.—11. same, hyaline cell and pores, outer face of 
branch leaf.—12. Dicranella heteromalla (Hedw.) Schimp., plant with apical antheridial cluster.—13. 
same, two leaves.—14. Encalypta procera Bruch., leaf.—15. same, cells and papillae from upper part 
of leaf.—16. Tortella fragilis (Drumm.) Limpr., plant with leaf tips mostly broken off. 


*Sphagnum papillosum Lindb.—Rare, one branch leaf only. Identified 
on the basis of trapezoidal chlorophyllose cells, which are broader on the 
inner than outer face, presence of a resorption furrow, and hyaline cells 
with relatively few pores. No papillae occur on the inner lateral walls of 
hyaline cells, making the material var. laeve Warnst. A fairly common, 
widespread species in the Great Lakes region. Fig. 10 & 11. 

Ditrichum flexicaule (Schwaegr.) Hampe—Abundant, leafy plants 
matted together or solitary, in one case intermingled with Bryum pseudo- 
triquetrum, Campylium stellatum and Tortella fragilis. Not known to 


QUATERNARY BRYOPHYTE SUBFOSSILS 31 


occur in Wisconsin but present in the Straits of Mackinac region, Michi- 
gan. Fossil material from New York State is illustrated in Miller (1973). 

Distichium capillaceum (Hedw.) B.S.G.—Abundant, gametophytic ma- 
terials only. Referred here with fair certainty, but the related D. inclina- 
tum (Hedw.) B.S.G., which differs primarily on the basis of sporophyte 
characters, may also be present. Forman (1967) cites collections of D. 
capillaceum from rotting logs and soil at forested sites on the Door Penin- 
sula. Fossil material illustrated in Miller (1973). 

*Dicranella heteromalla (Hedw.) Schimp.—Rather abundant. Not re- 
ported before from North American Quaternary deposits. Some specimens 
bear antheridia. Fig. 12 & 13. 

Encalypta procera Bruch—Rather abundant, occurring mainly as iso- 
lated leaves or as stem fragments with few leaves, only one large leafy 
plant found. Preservation fair; upper leaf cells with forked papillae; lower 
hyaline, nonpapillose cells mostly decayed. All leaves lack any indication 
of an awn; filiform brood bodies are absent. No doubt still a member of 
the Wisconsin flora, since records of Encalypta streptocarpa Hedw. in 
Cheney and Evans (1944) probably refer to E. procera. Apparently not 
recognized previously as a fossil in North America. Fig. 14 & 15. 

*Trichostomum tenuirostre (Hook. & Tayl.) Lindb.—Rare, eight well- 
preserved leafy plants. The specimens agree well with most herbarium 
collections so-named, but North American materials of Trichostomum 
need revision. Leaves of the fossil specimens clearly show the typical 
region of basal hyaline cells that do not extend up the margin as in 
Tortella. Apparently unknown in the present flora of Wisconsin but oc- 
curring in the Upper Peninsula of Michigan. No records of its fossil 
occurrence in North America are known to me. 

Tortella fragilis (Drumm.) Limpr.—Sparse, mostly leafy plants, a few 
with setae, and several isolated leaves. An infrequent moss of calcareous 
situations in the Great Lakes region. Fig. 16. 

°T. inclinata (R. Hedw.) Limpr.—Rare, three plants, somewhat poorly 
preserved but with certain leaves showing the characteristic concave, + 
cucullate apex. A rare moss in the Great Lakes area where it has been 
collected most often on dune sand. Unknown from Wisconsin but reported 
from several counties bordering the Straits of Mackinac, Michigan. A 
previous fossil record exists from northwestern New York State in a deposit 
a few hundred years older than the Two Creeks Forest Bed (Miller, 1973, 
q-v. for illustrations of fossil material ). 

T. tortuosa (Hedw.) Limpr.—Rather abundant, leafy plants, preserva- 
tion generally good. This species and T. fragilis are known from recent 
collections made in Door County, Wisconsin, at stations near the fossil 
bed. Illustrations given in Miller (1973). 

*Bryoerythrophyllum recurvirostrum (Hedw.) Chen—Sparse, somewhat 
degraded leafy plants. Fossil material illustrated in Miller (1973). 


32 NORTON G. MILLER 


Fics. 17—25. Subfossil mosses (cont.).-17. Mnium marginatum (With.) Brid. ex P. Beauv., 
plant.—18. same, three leaves from same plant.—19. same, leaf margin with a paired teeth.—20. 
same, upper leaf cells.—21. Myurella julacea (Schwaegr.) B.S.G., fragment of plant.—22. same, 
edge of leaf with teeth and papillose cells.—23. Thuidium recognitum (Hedw.) Lindb., nearly com- 
plete plant.—24. same, stem leaves from same plant.—25. same, paraphyllia. 


QUATERNARY BRYOPHYTE SUBFOSSILS 33 


*Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb.—Rare, two leaves. 
Identification probable but the characteristic roughened awn is poorly 
preserved. A common calciphile usually of open, + dry situations through- 
out the Great Lakes area. Illustrations in Miller (1973). 

Bryum pseudotriquetrum (Hedw.) Gaertn., Meyer & Scherb.—Abun- 
dant, leafy plants and a few isolated leaves. Many plants were found 
matted together, and some individuals in such clumps carried a broken 
seta. Sporophyte bearing plants were searched for archegonia and/or 
antheridia, but out of approximately 25 specimens, only two could be 
shown to be synoicous and two dioicous (only archegonia seen). Plants 
of the two types were not intermixed in the same clump. One plant with 
a terminal antheridial cluster was also found. Dioicous collections of this 
species are sometimes referred to var. pseudotriquetrum and synoicous 
ones to var. binum (Schreb.) Lilj. Other species of Bryuwm may be present 
in the fossil materials, but in the absence of capsules and peristomes, 
identifications could not be made. Leaves of fossil B. pseudotriquetrum 
from New York State are illustrated in Miller (1973). 

*Mnium marginatum (With.) Brid. ex P. Beauy.—Rather abundant, 
leafy plants and isolated leaves, preservation good but the double row of 
teeth along the leaf margin difficult to demonstrate. A few isolated leaves 
were broadly ovate (cf. Fig. 18). Upper leaf cells of fossil material are + 
quadrate with thickened corners and vary in size from 20-28 pm. A wide- 
spread species in the Great Lakes region where it grows on soil and rock 
in open or more dense forests. Fig. 17-20. 

*Aulacomnium palustre (Hedw.) Schwaegr.—Rather abundant, leafy 
plants and isolated leaves, small marginal teeth sometimes apparent to- 
ward the leaf apex. A common moss in temperate and boreal America; 
occurring in a variety of usually moist to wet habitats including forests 
and open areas. 

*A. turgidum (Wahlenb.) Schwaegr.—Rare, one leaf-bearing stem tip 
and eight isolated leaves. Fossil materials identified as this species of 
Aulacomnium because the leaf apices are obtusely rounded and somewhat 
hooded. A predominantly arctic and subarctic moss in North America, 
and apparently not a member of the present flora of Wisconsin, the species 
occurs disjunctively near the north shore of Lake Superior. Fossil material 
of the species 12,100 yrs old is known from northwestern New York State 
(Miller, 1973, q.v. for illustrations); the species has been found in inter- 
glacial deposits on Banks Island, arctic Canada (Kuc, 1974). 

*Orthotrichum obtusifolium Brid.—Rare, two plants, preservation fair. 
Leaves of the fossils have bluntly rounded apices, unipapillose cells and 
plane margins. No brood bodies were seen. Identification probable but 
not positive because the fossil leaves in general have a shorter costa than 
that present in herbarium material. Orthotrichum obtusifolium occurs 
widely in north temperate and southern boreal North America, where it is 


34 NORTON G. MILLER 


a fairly common epiphyte, particularly on bark of species of Populus ( Vitt, 
1973), pollen of which occurs in the forest bed peat. The genus has ap- 
parently not been reported before from North American Quaternary 
deposits. 

*Myurella julacea (Schwaegr.) B.S.G.—Sparse, stem fragments with 
leaves, preservation fair to good. An uncommon species in the Great Lakes 
area where it grows on moist soil in forests, especially Thuja swamps, and 
sometimes on rock (e.g., crevices in cliffs) in + open woods. No other 
North American Quaternary fossil occurrences are known. Fig. 21 & 22. 

Thuidium abietinum (Hedw.) B.S.G.—Abundant, large plants and 
fragments, leaves sometimes decayed from stems and branches. Known 
from several Quaternary deposits elsewhere in North America. 

°T. recognitum (Hedw.) Lindb.—Abundant, complete plants and large 
fragments. Fossil material agrees in all characters with herbarium speci- 
mens. Not known with certainty from other North American Quaternary 
deposits. This moss and T. abietinum have been collected on the Door 
Peninsula and have a wide distribution in the Great Lakes area and else- 
where in North America. Thuidium recognitum occurs at open sites on 
soil and rock and less frequently in dry to moist, + open forests. Fig. 23-25. 

*Campylium polygamum (B.S.G.) C. Jens.—Very abundant, leafy 
plants. Widely distributed in North America; in the Great Lakes region 
occurring on soil in bog forests, swamps, and wet open areas. Fig. 26 & 27. 

C. stellatum (Hedw.) C. Jens.—Abundant, fossil materials sometimes 
recognizable at a glance by their rich, coppery brown color, but otherwise 
determinable by the leaves which have a short double costa (vs. costa 
strong and single in C. polygamum). Extending northward from the 
temperate zone in North America, C. stellatum often grows in + open, 
calcareous habitats, especially rich fens, but also sometimes at forested 
sites. Fig. 28 & 29. 

*Amblystegium serpens (Hedw.) B.S.G.—Abundant, leafy stems mixed 
with other mosses or solitary. A common, widespread species of wet places, 
often on soil or rotting wood, in both forest and nonforest habitats. Not 
apparently recorded before as a subfossil in North America. Fig. 30. 

Drepanocladus aduncus var. polycarpus (Bland. ex Voit) Roth—Rather 
abundant, leafy plants. An aquatic or semiaquatic moss, usually of open 
or wooded calcareous places, found throughout the Great Lakes region 
and northward. The variety is not known from other deposits of the 
forest bed, although vars. aduncus and pseudofluitans Sanio occurred in 
Wilson’s samples (Cheney, 1930, 1931). Numerous reports of D. aduncus 
and some of its varieties are reported from Quaternary sediments else- 
where in glaciated portions of North America. 

D. revolvens (Sw.) Warnst.—Rare, only two leafy fragments. Leaves of 
fossil material are nonstriolate and lack inflated alar cells. Recent collec- 


QUATERNARY BRYOPHYTE SUBFOSSILS 35 


tions of this and the following species have been made in Door County, 
Wisconsin (Cheney & Evans, 1944). 

D. uncinatus (Hedw.) Warnst.—Very abundant, large fragments or 
intact plants, sometimes with one or several setae that lack capsules. 
Dense mats occurred in some sediment blocks. A common moss of open 
or dense boreal conifer forests; in the Great Lakes area occurring par- 
ticularly in Thuja swamps. Fig. 31. 

Scorpidium turgescens (T. Jens.) Loeske—Rare, two leafy plants. Un- 
known in the present flora of Wisconsin but found at several places in 
Michigan and adjacent areas, there reaching its present southern limit of 
distribution in central North America. Throughout its range, this moss 
occurs most frequently at wet, nonforest, calcareous sites, particularly rich 
fens. Illustrations of fossil material from New York State are in Miller 


(1973). 


27 


o.5mm o.5mm 


Fics. 26-30. Subfossil mosses (cont.).—26. Campylium polygamum (B.S.G.) C. Jens., plant. 
—27. same, three leaves from same plant.—28. C. stellatum (Hedw.) C. Jens., plant.—29. same, two 
leaves from same plant.—30. Amblystegium serpens (Hedw.) B.S.G., leaf. 


36 NORTON G. MILLER 


Tomenthypnum nitens (Hedw.) Loeske—Abundant, entire plants and 
fragments, one individual with two setae. Another calciphile, occurring in 
open or forested, wet habitats; found in North America from the Great 
Lakes region northward into the Arctic. Fossil material illustrated in 
Miller (1973). 

*Brachythecium turgidum (C. J]. Hartm.) Kindb.—Abundant, leafy 
plants; large fragments (3-4 cm) have a few short branches. Stem and 
branch leaves are entire and in them the costa extends to just beyond the 
center of the leaf. Apparently unknown in the present Wisconsin flora, 
although the species has been collected in Michigan, Culberson (1955) 


Fics. 31-35. Subfossil mosses (cont.).—31. Drepanocladus uncinatus (Hedw.) Warnst., plant 
with two setae.—32. Brachythecium turgidum (C. J. Hartm.) Kindb., fragment of plant.—33. same, 
three leaves from same plant.—34, Eurhynchium pulchellum (Hedw.) Jenn., fragment of plant.—35. 
same, two leaves. 


QUATERNARY BRYOPHYTE SUBFOSSILS 37 


has reported B. salebrosum (Hoffm.) B.S.G. from samples of the forest 
bed. Fig. 32 & 33. 

*Eurhynchium pulchellum (Wedw.) Jenn.—Rather abundant, branched 
leafy plants and isolated branches. A species widespread in North America 
that usually grows on soil and rotting logs in forests; also extending north- 
ward to the tundra. A common moss of Thuja swamps and bog forests in 
the upper Great Lakes region. Fig. 34 & 35. 

*Hypnum bambergeri Schimp.—Abundant, fragments with and without 
short branches. A short double costa (occasionally single) and an elevated 
cluster of reddish-brown, thick-walled alar cells characterize leaves of 
the fossils, which match recent herbarium materials in all important 
characters. The present North American range of this moss is limited 
mostly to arctic and subarctic regions. The species is an apparent calci- 
phile. Ilustrations of fossil material from the Wisconsin locality are in 
Miller (1976). 

*Hypnum pallescens (Hedw.) P. Beauv.—Rare, one plant with branches 
and several unbranched leafy fragments; identified on the basis of ser- 
rulate leaves with a double costa and quadrate alar cells. At present found 
widely in north temperate and boreal North America (and extending 
southward in the mountains), this moss occurs in forests on the bases of 
trees and sometimes on humus or other substrata. 

*Hylocomium splendens (Hedw.) B.S.G.—Abundant, some nearly com- 
plete plants and fragments. A common moss of moist forest soil throughout 
boreal North America but also found northward into the tundra and 
southward in mountain forests. Recent collections from places on the Door 
Peninsula are cited by Cheney and Evans (1944). 


DISCUSSION 


The vegetation cover of northeastern Wisconsin during Two Creeks 
time has been interpreted as closed canopy spruce forest (West, 1961) or 
open boreal woodland (Schweger, 1969), two more or less floristically 
similar vegetation types occurring today at places in the North American 
boreal forest. Local variation in Twocreekan plant communities existed 
because fossil assemblages from different deposits vary in composition. 
Spruce, however, is uniformly the major arboreal component. Pollen or 
cones of white spruce predominate in some instances (West, 1961, this 
report ); black spruce is more abundantly represented in others (Schweger, 
1969; Wilson, 1936). In present-day boreal America white and black 
spruce usually occupy dry and wet soil sites respectively, and the same 
situation no doubt prevailed during Two Crecks time. Nonarboreal pollen 
(NAP) totals, indicative of the amount of open, nonforest vegetation, also 
vary. Spectra from some samples have as little as 5% NAP, others contain 
20-25% or more. The existence of communities with herbs and shrubs is 


38 NORTON G. MILLER 


compatible with white and black spruce forests because similar mixtures 
of nonforest and forest vegetation now occur in parts of boreal America. 

At the Peters’ borrow pit megafossil material of white spruce is most 
abundant and lesser quantities of tamarack and black spruce occur. Forest 
communities on drier soil types must therefore have been prominent in the 
area during accumulation of forest bed sediments. Nonarboreal pollen is 
also sufficiently abundant to indicate the presence of plant communities 
containing or perhaps dominated by herbs, especially members of the 
Cyperaceae. The large assemblage of mosses found in the peat (Table 3) 
provides a way to define the communities more precisely, since all of the 
mosses represent extant species whose habitat preferences are known. 
The species are about evenly divided between those characteristic of 
forests and those occurring in nonforest habitats, several of which are 
indicated. Species listed more than once in the table are those that exhibit 
broad habitat tolerances. 

The forest mosses support the view that dry to moist but not wet soil 
conditions prevailed. Species of dense to open forest stands are present. 
Some of them grow on litter or rotting logs in existing spruce forests (e.g., 
Drepanocladus uncinatus, Eurhynchium pulchellum, Hylocomium splen- 
dens); others occur on mineral or humus-rich soil (Dicranella heteromalla, 
Bryoerythrophyllum recurvirostrum, Mnium marginatum). It can be in- 
ferred that similar habitats existed in forests of Two Creeks time. Two 
epiphytic species, Orthotrichum obtusifolium and Hypnum pallescens, 
were found, although the latter occasionally grows on soil or rocks also. 
Some of the forest mosses are calciphiles. These perhaps grew on or close 
to calcareous substrata beneath the forest bed. Other species typical of 
acid substrata indicate that humus accumulation was fairly advanced. 
The variety of habitats indicates well established forest and not pioneer 
communities. 

Of the nonforest mosses recovered from the peat, species of rich fen 
communities, which develop in association with shallow, calcium-rich 
water, are well represented. The distribution and character of rich fens 
are less fully known in North America than in Europe, although descrip- 
tions of some to the west of James Bay, Canada, are available (Sjérs, 1961, 
1963; and Persson & Sjérs, 1960, for ecological data on mosses in this 
region). Species characteristic of wetter areas of rich fens are separated 
in the table from those often occurring in drier parts and emphasize the 
relative scarcity of aquatic mosses in the peat samples. The predominantly 
aquatic moss genera Calliergon, Drepanocladus and Scorpidium are better 
represented in forest bed exposures along the shore of Lake Michigan 
(Wilson, 1932, 1936) than at the borrow pit, perhaps indicating a greater 
abundance of poorly drained sites in the former area during Two Creeks 
time. A few mosses listed in the fen margin category (e.g., Aulacomnium 
palustre, A. turgidum, Sphagnum napillosum) prefer more acid substrata, 


QUATERNARY BRYOPHYTE SUBFOSSILS 


TABLE 3. SUBFOSSIL MOSSES, FROM TWO CREEKS FOREST BED EXPOSED AT 
PETERS BORROW PIT, BROWN COUNTY, WISCONSIN, BY HABITAT TYPE 


Forest: mostly humus or mineral soil 


Dicranella heteromalla Campylium polygamum 

Bryoerythrophyllum recurvirostrum Amblystegium serpens 

Bryum pseudotriquetrum Drepanocladus uncinatus 

Mnium marginatum Eurhynchium pulchellum 

Orthotrichum obtusifolium (tree bark ) Hypnum pallescens (tree bark ) 

Thuidium abietinum Tomenthypnum nitens 

T. recognitum Hylocomium splendens 
Nonforest: rich fen, aquatic or semiaquatic 

Campylium stellatum Scorpidium turgescens 

Drepanocladus aduncus var. polycarpus Tomenthypnum nitens 


D. revolvens 


Nonforest: fen margin, + moist, mostly calcareous 


Sphagnum papillosum Aulacomnium turgidum 
Dicranella heteromalla Myurella julacea 
Distichium capillaceum Campylium polygamum 
Ditrichum flexicaule C. stellatum 

Tortella fragilis Amblystegium serpens 
Bryoerythrophyllum recurvirostrum Tomenthypnum nitens 
Bryum pseudotriquetrum Brachythecium turgidum 
Mnium marginatum Hypnum bambergeri 


Aulacomnium palustre 


Nonforest: moist rocks or mineral soil 


Distichium capillaceum Trichostomum tenuirostre 

Encalypta procera Mnium marginatum 

Tortella fragilis Aulacomnium turgidum 

Bryoerythrophyllum recurvirostrum Myurella julacea 
Nonforest: + xeric, well-drained soil, e.g., sand 

Tortella tortuosa Thuidium abietinum 

T. inclinata T. recognitum 


Tortula ruralis 


39 


40 NORTON G. MILLER 


and these perhaps grew on + dry peat hummocks or other noncalcareous 
sites within fens. Mosses limited to rock or mineral soil in open or semi- 
forested situations were few (e.g., Encalypta procera, Trichostomum 
tenuirostre ), although certain other species present in the peat sometimes 
grow in these habitats. The small group of mosses generally characteristic 
of xeric, open areas may have grown on well-drained dune or beach sand, 
which probably accumulated during fluctuations in the level of the lake 
then occupying the Lake Michigan basin. 

Peat samples from the Peters’ borrow pit have yielded a moss flora con- 
sisting of 32 extant species. The following seven additional mosses have 
been reported by previous workers from other exposures of the forest bed 
(Cheney, 1931, 1932; Culberson, 1955): Brachythecium salebrosum (Web. 
& Mohr) B.S.G., Bryum tortifolium Funck ex Brid., Calliergon cordifolium 
(Hedw.) Kindb., C. stramineum (Brid.) Kindb., Drepanocladus sendtneri 
(Schimp.) Warnst., D. vernicosus (Lindb. ex C. Hartm.) Warnst. and 
Scorpidium scorpioides (Hedw.) Limpr. No one ecological grouping of 
species dominates the flora, although aquatic or semiaquatic mosses may 
be more abundant at some localities. All species except seven are present 
members of the Wisconsin flora. Of these, however, five are known at 
stations in northern Michigan (Crum, 1973), and it is probable that they 
also occur in Wisconsin. Based on records cited in Cheney and Evans 
(1944) and Forman (1967), 16 of the 39 species (41%) are represented by 
recent collections from Brown, Door, Kewaunee and Manitowoc counties, 
which comprise the Door Peninsula. 

Two mosses that occur in the borrow pit peat, Aulacomnium turgidum 
and Hypnum bambergeri, are far out of place based on their present 
ranges. While several disjunct stations for the former are known in the 
Thunder Bay District of Ontario, about 500 km north of Green Bay, 
the southern limit of its continuous distribution in North America roughly 
coincides with the northern edge of the boreal forest. The species also 
occurs southward in alpine regions both in the East and West. Found as 
far south as the Gaspé Peninsula, Quebec, in the East and near Banff, 
Alberta, in the West, Hypnum bambergeri also has a predominantly arctic 
and subarctic distribution in North America. Based on material in five 
herbaria (CANM, FH, MICH, NY, US), this moss presently is found no farther 
south in the midcontinent region than northern Manitoba, some 1200 km 
north of the fossil occurrence. 

The presence of arctic and subarctic mosses in Wisconsin during Two 
Creeks time with others still found in the upper Great Lakes region and to 
the south is phytogeographically important. Such a mixed assemblage 
establishes that the Two Creeks flora was composed of species of different 
current geographical affinities. Existing disjunct stations for certain arctic 
species in the upper Great Lakes area may thus relate to Two Creeks 
time when such northern species occurred beyond their present ranges. 


QUATERNARY BRYOPHYTE SUBFOSSILS 4] 


Other arctic and subarctic mosses present as fossils in deposits about 
12,100 and 13,300 years old in northern Michigan and northwestern New 
York State are discussed in Miller (1973) and Miller and Benninghoff 
(1969). 

While 95% of the Twocreekan moss flora of northeastern Wisconsin is 
still represented in this area or nearby, some of the calcicolous species 
currently are rare in the upper Great Lakes region and attain their greatest 
abundance northward. Certain of them and other species of similar distri- 
bution also occur to the east in the two deposits mentioned above. The 
species include Catoscopium nigritum, Distichium capillaceum, Ditrichum 
flexicaule, Meesia uliginosa, Myurella julacea, Tortella fragilis, Scorpidium 
turgescens and others. Their presence in the assemblages indicates that 
much of the Great Lakes region 13,300 to 11,850 years ago was edaphically 
(and climatically) suited to the occurrence of such calcicolous species. 
Since the mosses now occur in the same general region as they did during 
the waning phases of glacial activity in late Wisconsinan time, their present 
distribution in the Great Lakes area can be viewed as an example of per- 
sistence at suitable habitats. That the species may be recent immigrants is 
also possible but perhaps less likely because of evidence provided by the 
fossil record. Similar assemblages of subfossil mosses from postglacial 
deposits, i.e., those less than about 10,000 years old, will help establish the 
more or less continuous presence of the species. This topic is more fully 
discussed in Miller (1976). 

The moss flora of northeastern Wisconsin during Two Creeks time was 
surprisingly diverse considering that it developed at a time of active 
glaciation. However, based on the richness of the flora as it is now known, 
study of material from other localities will continue to add species and 
improve the vegetation interpretation presented here and by other workers. 


ACKNOWLEDGMENTS 


Samples used in this study were collected by Hugh H. Iltis (University of Wisconsin, 
Madison) who wishes to acknowledge with appreciation support from that university’s 
Research Committee of the Graduate School and Dean of Letters and Sciences. IItis 
had earlier discovered the organic deposit in the company of Walter Biichmann. 
Samples of the organic bed were sent to me through the courtesy of W. C. Steere. I am 
grateful to Prof. R. F. Black of the University of Connecticut and Prof. Iltis for 
supplying descriptions of the borrow pit and of sediments exposed there. Howard 
Crum and L. E. Anderson examined some of the specimens on which the moss identi- 
fications are based, and I thank them for their comments. The habit drawings of mosses 
are the work of Marion Seiler whose care in their preparation is appreciated. 


LITERATURE CITED 


BENNINGHOFF, W. S. 1947. Use of trisodium phosphate with herbarium material and 
microfossils in peat. Science 106: 325, 326. 

Biack, R. F. 1970. Glacial geology of Two Creeks Forest Bed, Valderan type locality, 
and Northern Kettle Moraine State Forest. Wisconsin Geol. Nat. Hist. Sur. Inf. 
Cire. 13. 40 pp. 


42 NORTON G. MILLER 


——————— . 1974. Late Pleistocene shorelines and stratigraphic relations in the Lake 
Michigan basin: Discussion. Geol. Soc. Amer. Bull. 85: 659, 660. 

——————— and M. Rusin. 1968. Radiocarbon dates of Wisconsin. Trans. Wisconsin 
Acad. Sci. Arts Letters 56: 99-115. 

Broecker, W. S. and W. R. Farranp. 1963. Radiocarbon age of the Two Creeks 
Forest Bed, Wisconsin. Geol. Soc. Amer. Bull. 74: 795-802. 

CHENEY, L. S. 1930. Wisconsin fossil mosses. Bryologist 33: 66-68. 

——————— . 1931. More fossil mosses from Wisconsin. Bryologist 34: 93, 94. 

——————~ and R. Evans. 1944. The mosses of Wisconsin. Trans. Wisconsin Acad. 
Sci. Arts Letters 36: 171-224. 

Crum, H. 1973. Mosses of the Great Lakes forest. Contrib. Univ. Michigan Herb. 10. 
iii + 404 pp. 

Cu.Berson, W. L. 1955. The fossil mosses of the Two Creeks Forest Bed of Wisconsin. 
Amer. Midl. Nat. 54: 452-459. 

Evenson, E. B. 1973. Late Pleistocene shorelines and stratigraphic relations in the 
Lake Michigan basin. Geol. Soc. Amer. Bull. 84: 2281-2297. 

———— , W. R. Farranp and D. F. EscuMan. 1974. Late Pleistocene shorelines 
and stratigraphic relations in the Lake Michigan basin: Reply. Geol. Soc. Amer. 
Bull. 85: 661-664. 

Farcri, K. and J. Iversen. 1964. Textbook of pollen analysis. 2nd Rev. Ed. 237 pp. 
Munksgaard. Copenhagen. 

Forman, R. T. T. 1967. New and uncommon Wisconsin mosses. Bryologist 70: 1L15—- 
117. 

Kuc, M. 1974. The interglacial flora of Worth Point, western Banks Island. Geol. Sur. 
Can. Paper 74-1B: 227-231. 

Miuier, N. G. 1973. Lateglacial plants and plant communities in northwestern New 
York State. Jour. Arnold Arb. 54: 123-159. 

——————— . 1976. Quaternary fossil bryophytes in North America: A synopsis of the 
record and some phytogeographic implications. Jour. Hattori Bot. Lab. in press. 

a and W. S. BEeNNINGHOFF. 1969. Plant fossils from a Cary—Port Huron 
Interstade deposit and their paleoecological interpretation. Geol. Soc. Amer. 
Special Paper 123: 225-248 + pl. 3. 

Persson, H. and H. Syérs. 1960. Some bryophytes from the Hudson Bay Lowland of 
Ontario. Sv. Bot. Tidskr. 54: 247-268. 

Preston, R. S., E. Person and E. §. DeEvey. 1955. Yale natural radiocarbon measure- 
ments II. Science 122: 954-960. 

Rusin, M. and C, ALExANDER. 1960. U.S. Geological Survey radiocarbon dates V. 
Amer. Jour. Sci. Radiocarbon Suppl. 2: 129-185. 

ScuweceEr, C. E. 1969. Pollen analysis of Iola bog and paleoecology of the Two Creeks 
forest bed, Wisconsin. Ecology 50: 859-868 + 1 pl. 

SyOrs, H. 1961. Forest and peatland at Hawley Lake, northern Ontario. Natl. Mus. 
Canada Bull. 171: 1-31. 

——————— . 1963. Bogs and fens on Attawapiskat River, northern Ontario. Natl. Mus. 
Canada Bull. 186: 45-133 + pls. 3-6. 

Virr, D. H. 1973. A revision of the genus Orthotrichum in North America, north of 
Mexico. Bryophytorum Bibliotheca 1. 208 pp. + 60 pls. 

West, R. G. 1961. Late- and postglacial vegetational history in Wisconsin, particu- 
larly changes associated with the Valders readvance. Amer. Jour. Sci. 259: 766— 
783 + 2 pls. 

Witson, L. R. 1932. The Two Creeks Forest Bed, Manitowoc County, Wisconsin. 
Trans. Wisconsin Acad. Sci. Arts Letters 27: 31-46. 

a . 1936. Further fossil studies of the Two Creeks Forest Bed, Manitowoc 
County, Wisconsin. Bull. Torrey Bot. Club 63; 317-325. 


A NEW COMBINATION IN THE GENUS GYMNOMYCES 
Donatp H. PFISTER 


While studying some members of the Hynangiales I examined speci- 
mens of Octaviania redolens Cunn. Based on the reexamination of the 
holotype, I propose that it be referred to the genus Gymnomyces Massee 
and Rodway. 

Gymnomyces redolens,! known only from New Zealand, represents a 
distinct, well-delimited species of the genus. It differs from G. pallidus 
Massee and Rodway, described from Tasmanian material, in the size and 
shape of the basidiospores and in the number of them produced on each 
basidium. The basidia of G. pallidus produce four spores each, as opposed 
to G. redolens in which the basidia consistently produce but a single spore. 
In this respect G. redolens resembles G. monospora Stewart and Trappe 
(1975). Both species are unlike other members of the genus since they 
produce single-spored basidia. However, they differ from each other in 
several features. The basidiospores of G. redolens have amyloid material 
covering many of the ornamentations and have low amyloid connecting 
lines between the spines. On the other hand, the basidiospores of G. mono- 
spora were described as having heavy amyloid deposits in the form of dots 
or collars at the base of the spores. Basidiospore size also distinguishes 
the two species. The basidiospores of G. redolens are globose, rarely sub- 
globose, 12-15 wm in diam, excluding ornamentations. Those of G. mono- 
spora are subglobose to broadly ellipsoid, 15-21 12-18 pm, excluding 
ornamentations. Additionally, the basidiospore ornamentations in G. 
monospora are in the form of unconnected spines rather than as those in 
G. redolens which join toward their bases to form a low, incomplete 
reticulum. 

Cunningham’s (1942, 1944) descriptions of Octaviania redolens fail to 
mention the presence of sphacrocystis in the tramal plates. He mentioned 
only the presence of interwoven hyphae. There are, however, numerous 
sphaerocystis in this layer. 

I wish to thank Joan M. Dingley, Auckland Plant Disease Division, for 
providing the specimens for study. 

SPECIMENS EXAMINED. Holotype: on ground, Te Aroha, 250 ft., Auckland, May, 


1940, G. H. Cunningham, 10141 (ax). Three specimens collected by J. M. Dingley 
in Auckland (Ak 6355, 12334, 28585 ). 


LITERATURE CITED 


CunnincuaM, G. H. 1942. Two additional New Zealand Gasteromycetes. New Zealand 
J. Sci. Technol. 23: 172B-173B. 
——-—--— . 1944. Gasteromycetes of Australia and New Zealand. Published by the 
author. 226 pp. 
Stewart, E. L. and J. M. TRAPPE. 1975. Gymnomyces monospora sp. Nov. Mycotaxon 
2: 209-213. 
1Gymnomyces redolens (Cunn.) Pfist. comb. nov. = Octaviania redolens Cunn., New Zealand 


J. Sci. Technol. 23: 172B. 1942. 
43