A TAXONOMIC REVISION OF THE GENUS PHILONOTIS FOR NORTH AMERICA, NORTH OF MEXICO
by
WILLIAM MILTON ZALES
B. S. ed. Eastern Illinois University, 1966
M. S. Eastern Illinois University, 1967
A THESIS SUBMITTED IN PARTIAL FULFILMENT OF
THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
in the Department
of
Botany
We accept this thesis as conforming to the
required standard
THE UNIVERSITY OF BRITISH COLUMBIA
May, 19 73 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study.
I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission.
Department of BOTANY
The University of British Columbia Vancouver 8, Canada
Date 1 May 1973 i
ABSTRACT
The genus Philonotis (Musci) is revised for North /America, north of Mexico, and exclusive of Greenland,, following the last revision for this region by Seville Flowers in 1935.
Collections and field observations of all but two of the recognized taxa were made from a wide diversity of habitats throughout most of their geographic range in North America.
Numerous collections were cultured in a uniform environment to determine the stability of taxonomically diagnostic morphological characters. The position of papillae on the leaf cells, the structure of marginal cells, and to some degree the leaf shape and leaf cell shape are stable characters that are not modified by the environment. All other morphological
•characters examined are ecophenic and cannot be utilized to segregate taxa.
The present revision recognizes seven species and two varieties; Philonotis glaucescens (Hornsch.) Broth.,
P. longiseta (Rich, in Michx.) Britt., P. sphaerocarpa (Hedw.)
Brid., P. mavohioa (Hedw.) Brid., P. capillaris Lindb. ex
Hartm., P. fontana (Hedw.) Brid. var. fontana, P. fontana var. amevicana (Dism.) Flow, ex Crum, P. fontana var. pumila (Turn.)
Brid., and P. yezoana Besch. et Card, in Card, (new to North
America).
Chromosome counts of n = 6 for P. glaucescens and n = 12 for P. longiseta are new reports.
Keys, descriptions, illustrations, and distribution maps are provided for identification of the species. TABLE OF CONTENTS
Abstract, i
Table of Contents, ii-ila
List of Tables, Hi
List of Figures, iv-ivd
List of Maps, v
Acknowledgements, vi
I. Introduction, 1
II- Field Work and Herbarium Studies, 3
III. Nomenclatural History of North American Philonotis, 4
IV. Methods of Analyzing Morphological Diversity, 9
V. Morphological Analysis of Philonotis from Herbarium and
Field Specimens, 10
A. Sporophyte, 10
B. Gametophyte, 11
VI. Experimental Analysis of Morphological Diversity in
Philonotis, 21
A. History of Experimental Culture in Philonotis, 21
B. Methods and Materials, 22
C. Results, 2 3
1. Leaf size, 24
2. Leaf shape, 24
3. Leaf apex, 25
4. Leaf lamina, 25
5. Leaf margin, 29
•6. Costa, 31
7. Leaf arrangement, 32 8. Stem, 32
9. Rhizoids, 32
10. Propagula, 33
D. Conclusions, 33
VII. Species Concept, 39
VIII. Taxonomic Treatment, 40
A. Description of the Genus in North America, 40
B. Key to the North American Species of Philonotis,
C. Explanation of Species Figures, 45
Philonotis glaucescens, 46
Philonotis longiseta, 52
Philonotis sphaerocarpa, 60
Philonotis marchica, 63
Philonotis capillaris, 69
Philonotis fontana var. fontana, 73
Philonotis fontana var. americana, 82
Philonotis fontana var. pumila, 86
Philonotis yezoana, 90
IX. Phytogeography of Philonotis, 93
X. Distribution Maps, 95
XI. Excluded Taxa, 104
XII. Synonyms, 107
XIII. Summary, 111
XIV. Exsiccati and Selected Specimens Examined, 114
XV. Literature Cited, 162 • • •
LIST OF TABLES
±. North American Species of Philonotis, 6
II. Morphological Characters of Philonotis, 36
III. Descriptive Names Referring to Ecophenic Modifications, 38 LIST OF FIGURES
Figure
1-3 Philonotis fontana sporophyte and calyptra, 12
4, 5 Philonotis fontana stem and seta cross section, 13
6* 7 Philonotis fontana peristome teeth, 14
8-15 Scanning electron micrographs of the spores of
North American species of Philonotis , 15
16-25 Morphological variability in Philonotis fontana
induced by seasonal growth, 18
26-33 Comparison of various ecophenic expressions of
Philonotis fontana, 26
34, 35 Leaves of Philonotis glauoescens, 26
36-40 Types of lamina cells found in the North American species
of Philonotis, 30
41-43 Types of leaf margin cells found in the North American
species of Philonotis, 30
44, 45 Scanning electron micrographs of Philonotis fontana
rhizoids produced on the same stem, 34
46-48 Comparison of leaf shape and leaf margin in Philonotis
glauoescens 3 marchica, and sphaerooarpa, 49
49 Philonotis glauoescens, 51
50, 51 Mitotic chromosome configurations of Philonotis
longiseta and Philonotis glauoescens, 55
52, 53 Comparison of leaf shape and median leaf cell shape of
Philonotis marchica and Philonotis longiseta, 57
54 Philonotis longiseta, 59
55 Philonotis sphaerooarpa, 62 56, 57 Comparison of leaf shape, apex, and median leaf cells
of Philonotis marchica and Philonotis capillaris, 67
58 Philonotis marchica, 68
59 Philonotis capillaris, 72
'60 Philonotis fontana var. fontana, 81
61 Philonotis fontana var. americana, 85
62 Philonotis fontana var. pumila, 89
63 Philonotis yezoana, 92 V
LIST.OF DISTRIBUTION MAPS
Philonotis glauoescens, 95
Philonotis longiseta, 96
Philonotis sphaerooarpa, 97
Philonotis marchica, 98
Philonotis capillaris, 99
Philonotis fontana var. fontana, 100
Philonotis fontana var. americana, 101
Philonotis fontana var. pumila, 102
Philonotis yezoana, 103 vi ACKNOWLEDGEMENTS
I wish to express gratitude to my graduate advisor
Dr. W. B. Schofield for his influence and direction in demonstrating the value of thorough literature reviews, persistent field work, and for reading this manuscript throughout its production. I am also indebted to the members of the Botany Department of the University of
British Columbia who in numerous ways have assisted and encouraged my progress towards the completion of this research.
I wish to thank the curators of the herbaria listed on page 3 for their time and cooperation in sending specimens on loan for this study.
Acknowledgement is made of financial aid from the
National Research Council of Canada through grants to
Dr. W. B. Schofield for summer field expenses.
I am especially thankful for the patience and sympathetic understanding of my wife Dotty, and for the encouragement and assistance she has given during this project. 1 I. INTRODUCTION
The well defined moss genus Philonotis of the family
Bartramiaceae contains approximately 182 highly variable species that occur in moist or wet habitats throughout the world. In North America (exclusive of Greenland and Mexico) members of the genus are recognized by loose or dense tufted plants usually clothed in abundant tomentum, with short slender (1 cm high) stems varying to robust branched plants as large as 10 cm high that often possess whorled innovations and propaguliferous shoots or both. The highly variable leaves are ovate or triangular-lanceolate with an obtuse or acute apex and a long stout costa. The cells of the lamina are variously rectangular with prominent papillae and the
"margins are toothed with single or double teeth and usually rolled. The perigonia are discoid (rosette shaped) or gemmiform (bud shaped) and all but P. longiseta are dioicous.
The sporophyte is globose, inclined, and longitudinally furrowed on a long seta with a double series of sixteen peristome teeth.
Philonotis can tolerate a wide range of ecological conditions although it is unusual on living trees and rare on rotten wood. The wide ecological range is paralleled by a wide degree of morphological plasticity. This polymorphism has resulted in the description of many subspecific taxa. In the North American flora alone 99 names are involved. The descriptions accompanying some taxa match a number of specimens. Other diagnoses agree with the descriptions of 2
several different taxa, and the remainder fit no available
description. In the present treatment seven distinct species
and two varieties are recognized. Careful scrutiny of the
available literature exposes the fact that many errors and mis•
understandings have been perpetuated from one author's work
to another. Even more disheartening is the realization that
the identifications of previous monographers of portions of
the genus (Dismier, Flowers, Kindberg intev alia) do not
properly match their own keys and descriptions. The present
research, therefore, attempts to eliminate the ambiguity that
plagues the genus in North America and to clarify the species
concept.
The purpose of the present treatise is to define the
morphological variability of the genus Philonotis, to
circumscribe its taxa, and to provide keys, illustrations, and
maps to aid in their identification. To accomplish these
goals numerous collections cultured in a uniform environment were used to assess the natural variability of critical
morphological characters within the species. Such cultures
should reveal whether these characters are stable or whether
they are variable (ecophenic) depending on various ecological
parameters. An ecophene is defined as a plant or group of
plants that show a changed morphology induced by an altered
environment (ecologically modified phenotype). II. FIELD WORK AND HERBARIUM STUDIES
During the summers of 19 71 and 19 72 approximately 330 specimens of Philonotis were collected in southeastern, eastern, and western North America. At these times general field observations were recorded to obtain an appreciation of the ecological amplitude of each species collected.
All but three (P. yezoana, P. sphaevoaarpa, and
P. fontana var. pumila) of the nine taxa have been cultured in a uniform environment chamber, and the latter two have not been observed in the field.
Numerous specimens, including all available type specimens, have been examined from various herbaria in
North America, Europe, and Asia. Over 10,500 specimens were borrowed from the following herbaria: (standard abbreviations according to Lanjouw & Stafleu 1964) B, BM, BRNM, C, CAN, COLO,
CU, DPU, DUKE, F, FLAS, FSU, G, GL, GRI, GRO, H, IA, L, LAF,
LD, MANCH, MICH, MO, NFLD, NO, NY, OP, PC, SMS,,SMU, S-PA,
TENN, TEX, TRTC, UAC, UBC, UC, UPSV, US, UT, WIS, WJC, and
Tottori University, Japan (TOT). 4 III. N0F1ENCLATURAL HISTORY OF NORTH AMERICAN PHILONOTIS
Three species that are now included in the genus
Philonotis {fontanum, marchicum, and sphaericarpon) were described by Hedwig in 1801 as belonging to the large and inclusive genus Mnium. Hedwig also described two species of
Bartramia but failed to draw any relationships at the family level between these taxa and the three species of "Mnium", probably due to their difference in sexuality. This is understandable because of the acceptance of Linnaeus 1 dependence on sexual characters for the classification of flowering plants.
Bridel (1826) described the genus Philonotis separating it from Mnium and included in it P. fontana with varieties alpina and falcata, P. marchica, and P. sphaerocarpa. At this time no family relationships were established with the genus Bartramia.
In the important European work Bryologia Europaea,
(Bruch et al. 1842) many new families and genera were described. Such large and unnatural Hedwigian genera as
Mnium and Hypnum were divided into several more homogeneous genera. The family Bartramiaceae was also established and included Oreas, (now included in the Dicranaceae),
Bartramidula, and Bartramia. Bridel's species of Philonotis, however, were included in Hedwig's original, but now enlarged genus, Bartramia.
Schimper (1855) in the Corgllarium to Bryologia Europaea reinstated Bridel's genus Philonotis with the addition of two subgeneric categories, Philonotula, with gemmiform perigonia, and Philonotis, with discoid perigonia. Except for authors who have described new taxa within the genus, Schimper's treatment is where the generic concept has remained until the present. Table I lists the taxa included in catalogues of
North American mosses by various authors. These are arranged in chronological order. 6 TABLE I. NORTH AMERICAN SPECIES OF PHILONOTIS
Rail & Hervey 1880 Catalogue of North American Musci
Philonotis marchica Brid. {muehlenbergii Schwaegr.) fontana L. var. falcata (no authority cited) radicalis Beauv. var. porteri Aust. tenella Hedw. Lesquereux & James 1884 Manual of the Mosses of North America
Philonotis muehlenbergii Brid. var. tenella Brid. macounii Lesq. et James sp. nov. fontana Brid. var. alpina Brid. var. falcata Brid. calcarea Schimp. mohriana (C. Mull.) Jaeg.
Kindberg 1897 European and North American Bryineae
Philonotis marchica Willd. muehlenbergii Brid. macounii Lesq. et James arnellii Husn. vancouveriensis Kindb. sp. nov. glabriuscula Kindb. calcarea Br. Eur. var. caespitosa Wils. alpicola Jur. acutiflora Kindb. fontana L. var. seriata Mitt. mohrii C. Mull. adpressa Ferg. radicalis Beauv. tenella C. Mull. pumila Kindb. sp. nov.
Dismier 1910 Philonotis de I'Amerique
Philonotis radicalis (P. B.) Brid. gracillima Angstr. sphaerooarpa (Sw.) Brid. ssp. tenella (C. Mull.) Jaeg. var. terrestris Dism. var. nov. 7
marchica (Willd.) Brid. seriata (Mitt.) Lindb. ssp. americana Dism. ssp. nov. var. torquata (Ren. et Geh. ex Geh.) Dism. var. graciles cens Dism. var. nov. fontana Brid. var. heterophylla Card, et Ther. ssp. tomentella Mol. var. borealis (Hag.) Loeske var. compacta Dism. var. heterophylla Dism. var. nov. caespitosa Wils. var. compacta Dism. var. nov. var. laxa (Warnst.) Loeske et Warnst. var. adpressa Dism. var. heterophylla Dism. var. nov. ssp. fallax Dism. ssp. nov. oapillaris Lindb.
S. Flowers 1935 in Grout, Moss Flora of North America
Philonotis glauoescens (Hornsch.) Paris f. laxa Flow. /. nov. var. terrestris (Dism.) Flow. comb. nov. var. brevifolia (Dism.) Flow. comb. nov. gracillima Angstr. sphaericarpa (Sw.) Brid. var. terrestris Dism. uncinata (Schwaegr.) Brid. longiseta (Rich.) E. G. Britt. f. propagulicaulis Flow. f. polygama Flow. /. nov. var. porteri (Aust.) comb. nov. marchica (Willd.) Brid. muehlenbergii (Schwaegr.) Brid. oapillaris Lindb. caespitosa Wils. var. compacta Dism. var. adpressa Dism. var. laxa (Warnst.) Loeske
fontana Brid. f. dimorphophylla Flow. /. nov. var. heterophylla Card. & Ther. var. seriata Breidl. f. occidentalis Flow. /. nov. var. pumila Brid. f. longifolia Flow. nov. f. dimorphophylla Flow. /. nov. var. falcata Brid. var. adpressa (Ferg.) Limpr. var. borealis Hag. 8
var. laxa Flow. var. nov. f. tenuis Flow. f. nov.. amerioana Dism. f. taxa Flow. /. nov. f. dimorphophylla Flow. /. nov. var. torquata (Ren. et Geh.) Dism. var. graciles cens Dism. oalcarea f. occidentalis Flow. /. nov.
Crum, Steere, & Anderson 1965 A List of the Mosses of North America
Philonotis amerioana Dism. var. gracilescens Dism. var. torquata (Ren. et Geh. ex Geh.) Dism. caespitosa Jur. var. adpressa Dism. var. ampliretis (Dix.) Crum, Steere, et Anderson var. compacta Dism. oalcarea (B. S. G.) Schimp. capillaris Lindb. fontana (Hedw.) Brid. var. adpressa (Ferg. ex Hunt) Limpr. var. borealis I. Hag. var. falcata Brid. var. heterophylla Card, et Ther. var. laxa Flow. var. pumila Brid. var. seriata (Mitt.) Breidl. glaucescens (Hornsch.) Par. var. terrestris (Dism.) Flow. gracillima Angstr. longiseta (Michx.) Britt. var. porteri (Aust.) Flow. marchica (Hedw.) Brid. muehlenbergii (Schwaegr.) Brid. sphaerocarpa (Hedw.) Brid. uncinata (Schwaegr.) Brid. 9 IV. METHODS OF ANALYZING MORPHOLOGICAL DIVERSITY
Morphological diversity may be studied in several ways.
(i) Perennial plants like Philonotis may show variation due to seasonal changes in the environment. This variation is easily detected by examining one year's growth produced on a plant. Hi) Examination of several plants from a clonal population may show variation induced by competition for water, space, and minerals, or different degrees of exposure to sunlight. {Hi) Critical examination of large series of collections from throughout the entire range of a taxon may demonstrate that variation is either a continuum connecting several taxa with many intermediate forms or that there are definite groups of variants that may be recognized taxonomically. This is clearly the most important consideration in traditional or alpha taxonomy. Unfortunately many taxa of Philonotis have been described from a single specimen with no allowance given for variability. (iv) Mor• phological variability that is suspected to be environmentally induced can be tested by cultivating morphologically distinct specimens from numerous locations under uniform environmental conditions and comparing the new growth that develops. . All
four methods of analysis are considered herein, however the experimental results have proven to be the most reliable for revealing morphological diversity and its' causes in
Philonotis. 10 V. MORPHOLOGICAL ANALYSIS OF PHILONOTIS FROM HERBARIUM AND FIELD SPECIMENS
A. SPOROPHYTE
Although there is no a priori reason why the sporophytic characteristics of a moss should be less plastic or variable than the vegetative haploid gametophyte, the sporophyte is the most stable phenotypic expression of Philonotis. The initiation and production of gametangia, transfer of sperms, fertilization of the egg, and subsequent maturation of the sporophyte requires very specific environmental conditions in order to be successful. Unfortunately this narrow ecological tolerance drastically reduces the number of specimens collected with mature sporophytes. Not only are the sporophytes of little value as diagnostic characters due to their scarcity but the sporophyte is also morphologically uniform throughout the genus. This viewpoint is also held by
Dismier (1907a) .
The seta is green when young but becomes brown when mature and is twisted throughout when dry. Seta length is variable (1-6 cm) but generally longer in the larger specimens of P. fontana. The anatomy is similar in all North American species (fig. 5). The capsule is spherical and light green while developing (fig. 2). It becomes brown, deeply furrowed longitudinally, and constricted about the middle when mature and empty (fig. 3). The capsule is held obliquely and the peristome is positioned further off the vertical axis of the seta (fig. 2, 3). The exothecial cells are irregularly 11 rectangular, becoming irregularly cuboidal at the base where the stomata are located. Forty to 275 stomata have been counted on various capsules however the maximum to minimum ranges of all North American species overlap between 90 and
100 stomata per capsule. The cell walls are thicker on the ridges than in the furrows. The operculum is round-convex with a shallow apiculus in all species examined (fig. 2).
The peristome consists of two circles of sixteen teeth each that are similar for all species (fig. 6, 7). The chromosome number of Philonotis is constantly n = 6 in all species reported except for P. longiseta and P. socia which are n = 12.
(See discussion under P. longiseta.)
B. GAMETOPHYTE
Spore morphology in Philonotis does not provide any consistent diagnostic features. Spore size is variable depending on the size and maturity of the capsules they were produced in. Critical examination of spores with the scanning electron microscope (SEM) has revealed that the North American species are all similarly papillose and that the degree of papillosity is variable (fig. 8-15).
The small, smooth, and cucullate calyptra is shed very early in sporophyte development. The few calyptras examined seemed to be identical in several different species (fig. 1).
Their scarcity and similarity make them useless for taxonomic use within the genus Philonotis.
The perigonial and perichaetial leaves surrounding the gametangia have historically been employed as diagnostic Figures 1-3. Philonotis fontana sporophyte & calyptra
-1. Calyptra
-2. Fresh mature capsule with operculum
-3. Dry empty capsule 12
1 mm Figures 4, 5. Philonotis fontana stem & seta cross section -4. Stem cross section -5. Seta cross section
Figures 6,7. Philonotis fontana peristome teeth
-6. Portion of endostome
-7. Portion of exostome 14 Figures 8-15. Scanning electron micrographs of the
spores of North American species of
Philonotis
-8. P. glauoes oens
-9. P. longiseta
-10. P. sphaerocarpa
-11. P. marchica
-12. P. capillaris
-13. P. fontana var. fontana
-14. P. fontana var. amerioana
-15. P. fontana var. pumila
16
characters in Philonotis to an unusually great extent. The
two largest sections of Philonotis are organized according to
the arrangement of leaves forming the perigonia: section
Philonotis possesses discoid perigonia, and includes those
species with sguarrose-geniculate leaves forming a flat
rosette; whereas section Philonotula has gemmiform perigonia,
and includes those with erect leaves forming a bud. At the
sectional level this character (the arrangement of perigonial
leaves) is consistent, but at the specific level the perigonia are of slight value for two basic reasons. First,
specimens are not always collected in a sexual condition, and
second these leaves show an extreme polymorphism within each
general type. Janzen (1921), Loeske (1906), and Monkemeyer
(1927) have studied the variability of perigonial and
perichaetial leaves of Philonotis in great detail and also
agree that they are of little taxonomic value in distinguishing
species.
The flask shaped archegonia, oblong antheridia, and the
numerous elongate club shaped paraphases, which are similar to
most other bryophytes, are uniform throughout the North
American species.
Polymorphism as observed in the vegetative leaves of
Philonotis is also extreme and has resulted in the proposal
of many superfluous names. The size, color, contour, and to
some degree the shape of the leaves, and their position and
arrangement on the stem are all subject to modification by
the environment. Meusel (1935), and in greater detail Hagerup (1935), described and illustrated (fig. 16-25) the periodic or seasonal growth of P. fontana from field observations. They both remarked on the production of gametangia at the base of several innovations or shoots so characteristic of the genus.
Hagerup (19 35) continued to explain the presence of two kinds of annual shoots: spring shoots with short, blunt, closely appressed leaves (fig. 17, 23), and summer shoots with lanceolate, somewhat falcate leaves (fig. 20, 22). He made no taxonomic statements or suggestions even though P. fontana var. heterophylla Card, et Ther. corresponds to fig. 18, and
P. fontana f. dimorphophylla Flow, corresponds to fig. 20.
These two subspecific taxa are reciprocal dimorphic conditions produced in response to seasonal change, and should not be accorded separate taxonomic status.
If an entire tuft or sod is examined rather than a single plant, a better appreciation is gained of the polymorphism within a taxon. It may be assumed that the plants of a tuft have the same genotype as they are often the product of vegetative reproduction. If the tuft is fertile and reproducing sexually we may usually assume that the parents are of the same gene pool and that all of the individuals belong to the same taxon. Plants in the center of a tuft will generally be more robust, more tomentose, and in some species posses larger, falcate, or plicate leaves with rolled margins. Plants at the edge of a population are depauperate, lax, lighter in color, less erect, often Figures 16-25. Morphological variability in Philonotis
fontana induced by seasonal growth
-17&23. Spring shoot and leaf
-20&22. Summer growth on the same shoot and leaf
24. One innovation (branch) of a male plant
becoming fertile in the second year
25. Mature sporophyte produced on a female
plant during the third year, and a
young sporophyte developing at the end
of the same year's growth
After Hagerup (19 35 p. 8&10) 18 19 etiolated, and with flat appressed leaves. These extremes in a population correspond to described varieties, forms, and even other species, such as adpressa, falcata, taxa,
graailescens3 eaespitosa, and oompaota. It is noteworthy that these taxa are rarely collected with sporophytes.
Compared to the narrow ecological requirements necessary to produce mature sporophytes, the gametophyte can tolerate a wide range of environmental conditions. Those conditions that favor sporophyte production will also produce the healthiest gametophytes whereas other environmental conditions will produce entire populations that correspond to these sterile varieties, forms, and "species".
Polymorphism is even more confusing when plants from widely separated populations are compared. Then the assumption that these taxa actually belong to a single species is even more difficult to support unless numerous series of collections are examined, and the described taxa are shown to be connected by a complete line of intermediate and intergrading forms.
After examining all available North American and many extra-North American specimens, it seems plausible to agree with Flowers (s. a.).
"It is time to give up the pretense that
certain segments of this array of plants
represents discrete genetical groups and face up
to the fact that we are dealing with a highly
involved situation where the interaction of 20 heredity and environment is producing a maze
of forms stemming from essentially the same
stock." VI. EXPERIMENTAL ANALYSIS OF MORPHOLOGICAL DIVERSITY IN PHILONOTIS A. HISTORY OF EXPERIMENTAL CULTURE OF PHILONOTIS
Experimental studies using bryophytes in culture have been concerned almost exclusively with the physiology of a
very few common plants like Funaria hygrometvica or
Marohantia polymorpha.
H. Buch (1922) conducted extensive studies on the hepatic family Scapaniaceae in uniform environment chambers
and found that water and light influenced the plant's morphology considerably.
More recently Lodge (1960) cultured Drepanooladus
fluitans and D. exannulatus and found that the shape and
arrangement of the leaves and the length of the costa are highly variable depending mostly on the amount of moisture
and light.
Davy de Virville (1927) studied the action of the
environment on several mosses and found that Philonotis
fontana in a saturated atmosphere produces many rhizoids, has
shorter leaves with thinner cell walls, smaller teeth, and
fewer chloroplasts. When the plant is grown submerged it has light green stems 2-4 cm long with few rhizoids, variable
leaves exhibiting an attenuated apex, thin cell walls, and
large chloroplasts.
Zastrow (19 34) grew P. fontana in liquid culture and
found that it tolerated a wide pH range, developed long stems with widely spaced erect leaves that had flat margins, a thin 22 percurrent costa without a central strand, and lax cells, but
retained their original shape.
Extensive research has been conducted by many workers
on spore viability, mode of germination, and the physiological effects that control or influence spore
germination. Only Matteri (19 70) has studied members of the
Bartramiaceae and she found that three species of Philonotis
develop the typical heterotrichous type of protonema and no
intrageneric differences were present.
No one has cultured Philonotis for the purpose of
studying the validity of its taxonomic characters, although
several authors have suggested that this type of experimental
research might be fruitful. (Nyholm 1960, Florschiitz 1964,
Bowering 1966).
The present research shows that some morphological
features of the genus Philonotis are stable, while others
that have traditionally been thought to be important are too
variable to be of taxonomic value.
B. METHODS AND MATERIALS
A uniform environment chamber was constructed for the
purpose of culturing large collections of Philonotis from
diverse populations. The chamber measured 48 X 96 inches
and had a capacity of approximately 270 petri dishes. The
cultures were illuminated by eight 36 inch cool white
fluorescent and eighteen 7 watt incandescent white bulbs
suspended 20 inches above the surface of the dishes. A total
illumination of approximately 200 foot candles was produced.
The chamber was automatically adjusted to maintain an 23 18/6 hour day/night cycle. The temperature varied with that of the ambient air and increased approximately 10°F during each day cycle. High humidity was maintained by frequent watering in the dishes with 50% Hoagland's solution (Hoagland
& Arnon 19 38).
Small populations of plants, approximately 5X5 cm, from various habitats and various locations throughout North
America were placed on wet paper toweling in standard plastic petri dishes. After several weeks the new growth was transferred to sterile Hoagland's solution solidified with
2% agar. This process was continued until obvious fungal and algal contamination was eliminated.
All observations, measurements, and illustrations were made from slides prepared by first relaxing the plants in boiling water. The plants were then placed in lacto-phenol and the leaves were removed from the stem. Lacto-phenol is composed of equal parts of phenol, lactic acid, glycerin, and distilled water (Johansen 1940). The lacto-phenol was drained and the specimens were made permanent in Hoyer's mounting medium (Anderson 1954) . C. RESULTS
In culture all of the taxa show a high degree of morphological stability, although their appearance may be drastically different from the original collection in nature.
This demonstrates that a specific set of environmental conditions will produce a specific set of morphological responses. The comparison of specific characters as they 24 appear in nature to their appearance in culture can be used to assess their diagnostic value.
Comparison of several morphologically distinct plants from various habitats in nature with their new growth produced in a uniform environment has shown that their distinctive differences are no longer present, and that they have become morphologically identical. These "distinctive" differences can then be assumed to be ecophenic or environmentally induced, and the plants can be assumed to belong to the same taxon.
Diagnostic characters of Philonotis are analyzed and evaluated individually based on the results obtained from culture experiments. 1. LEAF SIZE
Leaf size is one of the most variable characteristics of Philonotis. In general the leaves found in nature are much larger than the ones that develop in culture, although plants with small leaves are often collected. Unless fertile plants are collected, leaf size is misleading and other characters must be sought for analysis. 2. LEAF SHAPE
The general leaf shapes are consistent in some respects and variable in others. Polymorphism, produced by differential seasonal growth, observed in some species in nature, is lost in culture; dimorphic and heteromorphic forms
(fig. 16-25) produce only one kind of leaf in a uniform environment. Plants with falcate leaves in nature produce only straight leaves in culture (fig. 28, 30). Very broadly ovate-lanceolate leaves become lanceolate in culture
(fig. 27, 29, 31). Species with triangular-lanceolate leaves in general are stable and can be distinguished from species with ovate-lanceolate leaves. From the analysis of culture studies, leaf shape is a suitable character only when used in combination with other characters.
3, LEAF APEX
The leaf apex remains the same in culture as in the field in all taxa except P. glaucescens. Plants of
P. glaucescens with an acuminate apex and percurrent costa in the field will develop a rounded apex with a shorter costa in culture (fig. 34). Plants with a rounded apex in nature maintain this condition in culture but when transplanted to a suitable environment in the field develop an acuminate apex (fig. 35) . When these plants from the field were returned to the culture chamber they again produced the rounded leaf apex. This latter condition appears to be produced as a response to high humidity and low light intensity.
4. LEAF LAMINA
There are many characters suitable for analysis in the broad blade of vegetative leaves, excluding the margin and costa. The areolation is directly responsible for the size and shape of the leaf. Cell size is extremely variable, being proportionately larger in larger leaves. Plants of the same taxon with various size leaf cells from different Figures 26-33. Comparison of various ecophenic expressions
of Philonotis fontana. The first three
leaves in each figure were drawn from
plants as they appeared in the field;
(Scale A) the adjacent two leaves were
drawn from the same collection after
being cultured at the same time in a
uniform environment (Scale B). 26
Scale A Scale B Figure 34. Leaves of Philonotis glauoescens having
an acute apex in the field (Florida,
Zalesj 206 8) developed an obtuse apex
in culture.
Figure 35. Leaves of Philonotis glauoescens having
an obtuse apex in the field (Florida,
Zales} 2041) continue to have an obtuse
apex in culture. These same cultured
plants developed leaves with an acute
apex when transplanted to the field, but
developed an obtuse apex when returned
to the culture chamber. 27 28 environments in the field develop the same size cells in
culture. Cell shape is consistent and a more useful diagnostic
character than cell size. Some species have narrow very elongate cells (those with triangular-lanceolate leaves),
and others have rectangular, quadrate, to slightly irregular
cells (those with ovate-lanceolate leaves). These features
remain constant in culture.
Cell wall thickness varies directly in response to environmental changes. All species of Philonotis growing in wet or densely shaded habitats or both will produce thin walled lax cells. This lax condition can be produced by
growing plants in water culture. Philonotis laxa and variety laxa (taxa based on the lax condition of several
species) are ecophenic and should not be recognized taxonomically.
The leaf cell papillosity is an important family
characteristic and also is the most stable specific
character in the genus. The papillae may be found at the upper, middle, or lower cell ends of certain species
(fig. 36-43), never at both ends on the same surface as
implied by some authors and illustrators. The leaves must be cleared with lacto-phenol if an accurate assessment of papillae position is to be made. Regardless of all other variation in Philonotis, the papillae position on leaf cells
remains consistent when cultured plants are compared to the
same plants in nature.
The size of individual papillae is generally smaller in 29 cultured material due to thinner cell walls. Occasionally papillae will occur at opposite ends of a cell on opposite surfaces of a leaf. In longitudinal section the two cells appear to be overriding each other (fig. 39).
Longifttudinal folds or plicae are evident in some species. This feature has been employed as an important specific character but, as demonstrated by culture experiments (fig. 26-30), is not always present throughout the same plant. Plication in the leaf lamina is a valid character only in fertile or very robust plants.
Cell contents have rarely been used for taxonomic purposes. The size and number of chloroplasts are variable.
Plants in culture will produce fewer but larger chloroplasts per cell than those in the field. Oil bodies are occasionally produced but exhibit no correlation to the environment, age, or stature of the plants in culture or in the field. Furthermore, the chloroplasts and oil bodies are lost during lacto-phenol treatment. Cell chemistry may be of diagnostic importance if proper analytical methods are employed.
A distinctive alar region is not differentiated in
Philonotis. 5. LEAF MARGIN
Two features of the leaf margin are easily detectable; they are papillosity and inrolling. Papillosity is an important and stable taxonomic character. The papillae may be at the upper cell end only in some species (fig. 42, 43), Figures 36-40. Types of lamina cells found in the
North American species of Philonotis
in surface view and longitudinal
section.
-36. P. longiseta
-37. P. marchica
-38. P. capillaris
-39. P. fontana
-40. P. yezoana
Figures 41-43. Types of leaf margin cells found in the
North American species of Philonotis .
-41. P. fontana
-42. P. marchica
-43. P. (j-Zauceseens
31 or at both ends forming "zwillingsmamillen" of Janzen (1921)
(fig. 41). These "zwillingsmamillen" or double teeth should not be confused with margins that appear to be doubly toothed due to a rolled margin (fig. 43). The presence or absence of double teeth is apparent in large fertile plants but less obvious in smaller or cultured plants. This size difference is not important, but their occurrence is consistent in the field as well as in culture and of basic taxonomic value.
Many small sterile plants collected in nature will show flat leaf margins. These same plants would have produced rolled margins if they had continued to develop into robust plants. This ecophenic variation is demonstrated by the fact that rolled leaf margins are absent from most species in culture but present in the material collected in situ.
(fig. 26-32). The presence or absence or degree of inrolling of the margin is too variable to be of much, taxonomic value. 6. COSTA
Costa anatomy in Philonotis is relatively consistent.
All species have a strong costa that reaches nearly to the apex, is percurrent, or excurrent. In cross section the costa is composed of one, two, or three guide cells surrounded by several layers of incrassate stereids. Cultured plants in general will have fewer stereids with thinner cell walls than the same plants from the field. The intraspecific variation observed in costa anatomy is too great to be employed as an interspecific character. 7. LEAF ARRANGEMENT
Phyllotaxis and spacing is variable throughout Philonotis
Plants growing in a uniform environment show a regular spiral
phyllotaxy. Regular geometric phyllotaxis is rarely observed
in nature, instead the leaves may appear homomallous, somewhat
five-ranked, irregular to scattered, or sometimes arranged in
vertical rows. The spacing is consistent in culture but highly variable in nature. Very wet or shaded conditions will produce stems with greater distances between the leaves
(etiolated plants). Considering this evidence, these
modifications are all ecophenic and invalid as taxonomic
characters. 8. STEM
Stem structure is relatively uniform in this genus.
There is an outer layer of inflated cortical cells
surrounding an area several cells in thickness composed of
very thick walled cells. The center of the stem is composed
of large thin walled cells and a central strand of small
thin walled cells (fig. 4). The stems may be larger with
a more prominent central strand in robust plants when
compared to the same plants in culture. Pigmentation is
more evident in older and healthier stems than in young,
shaded, aquatic, or cultured plants. 9. RHIZOIDS
Rhizoids are too variable to be of taxonomic importance.
Their presence is influenced by moisture. Plants growing in
a dry atmosphere or in extremely wet to aquatic habitats will 33 have few stem rhizoids. Plants growing in a humid
atmosphere, as in the cultures or in some habitats in nature, will produce an abundant growth of tomentum. The
SEM has shown that the fine structure of Philonotis
rhizoids is also variable. Some parts of the rhizoids may be smooth while others from the same stem may be quite scabrous (fig. 44, 45). 10. PROPAGULA
Propagula of some form occur on all species of
Philonotis. They are not consistently present but develop in response to changing environmental conditions. Abundant propagule production can be induced by allowing plants in
culture to slowly desiccate. In nature as in the cultures, propagula are important asexual reproductive organs. They occur most frequently on members of the section Philonotula.
Those found in the section Philonotis may fall off and
function as propagules or remain attached and develop into the typical whorl of small branches or innovations. Taylor
(1958) has described small residual leaves that are present
at the point of propagule attachment. These tiny structures
are not unusual in any plants developing propagules, but
are ecophenic and occur too sporadically to be of diagnostic value.
D. CONCLUSIONS
Philonotis, a common and widespread genus in North
America, can tolerate a great diversity of soil and rock types, moisture regimes, and light intensities. As is true of many Figures 44, 45. Scanning electron micrographs of
Philonotis fontana rhizoids
produced on the same stem.
other waterloving plants, the broad ecological amplitude
of Philonotis is paralleled by its extreme morphological
plasticity.
Table II lists those characters that are typical of
all species in the genus, those that are specific for
certain species or groups of species, and those characters
that are variable depending on environmental conditions.
The nature of stem anatomy, presence of rhizoids, and
lanceolate leaves with toothed margins and papillose
rectangular cells are important gametophytic characteristics
that, in combination, distinguish Philonotis from other
genera. The shape of these lanceolate leaves and rectangular
cells in various combinations with the position and number
'of papillae are stable characters that are always present
and can be used to describe and identify the North American
taxa. The size of leaves, cells, and stems; the abundance
of rhizoids, propagula, plicae, and rolled margins; or leaf
arrangements are all variable ecophenic modifications.
These ecophenic modifications have, in the past, been
used to recognize taxa with formal taxonomic status. This
has caused much confusion and ambiguity in the understanding
of the genus.
Table III lists the descriptive names and their
meanings, without author citations, that have been applied
to one or more of the recognized taxa of Philonotis.
Monkemeyer (19 27) stated;
"Solche Wuch-, Farben-oder Anpassungsformen TABLE II, MORPHOLOGICAL CHARACTERS OF PHILONOTIS
ECOPHENIC CONSISTENT THROUGHOUT GENUS CONSISTENT WITHIN SOME SPECIES
lax cells plicate leaves leaf arrangement rolled or flat margin presence of propagulae leaf size size & color of stems....stem cross section amount of rhizoids presence of rhizoids cell size .. ..± rectangular cells elongate rectangular vrs. quadrate rectangular presence of papillae position of papillae toothed margin single teeth vrs.double teeth lanceolate leaves... ovate-lanceolate vrs. triangular-lanceolate sollte man der Einfachheit wegen,
um den nebensachlichen Formelfram
der Benennung und Prioritatsfragen
umgehen zu konnen, je nachdem . . .
und . . . ohne Autorzitierung
bezeichnen, sie sind sofort ohne
langatmige Beschreibungen verstandlich
und das ist ja die Hauptsache."
Buch (1928) also has proposed a similar system for
designating ecophenic variation, the modificatio, in the
leafy hepaticae. These proposals are practical and the
use of descriptive adjectives to point out various ecophenic expressions within a taxon should be based on simple
ecological and morphological observations but should avoid
the rigidity of taxonomic nomenclature.
Crundwell (19 70) suggests that modificatio be used for
variants with no genetic basis because taxa at this level
are often not mutually exclusive. TABLE III. DESCRIPTIVE NAMES REFERRING TO ECOPHENIC MODIFICATIONS
MODIFICATIO APPEARANCE (descriptive name) Referring to the size of the plant tenera delicate, etiolated mollis soft, flagella form terrestris small, widely spaced compacta dense tomentose tufts ampliretis splendid, large erect stems gracilescens slender tenuis thin, narrow laxa etiolated
Referring to the leaves falcata bent, usually to one side torquata twisted adpressa close to the stem laxa inflated cells with thin walls laxifolia inflated cells with thin walls pumila small heterophy11a dimorphic, blunt apex grading into acute dimorphophylla dimorphic, acute apex grading into blunt brevifolia small leaves longifolia long leaves
Referring to asexual reproductive structures gemmiclada gemmae present propagulicaulis gemmae present
Referring to sexuality polygama synoicous 39 VII. SPECIES CONCEPT
Now that ample experimental data are available to demonstrate the phenotypic plasticity within the North
American species of Philonotis, it is possible to exclude
formal taxonomic recognition of ecophenic modifications.
For the purpose of the present study a species is considered to be a series of biotypes that can be recognized by a combination of stable morphologically distinct characters. A variety is a closely related,
although morphologically and geographically segregated series of biotypes within a species. Modificatio adjectival names may be added to describe variable conditions within a species or variety, but they should be used only for convenience or to clarify the appearance of various ecophenes or growth forms as such are names without formal taxonomic status. VIII, TAXONOMIC TREATMENT ' A. DESCRIPTION OF THE GENUS IN NORTH AMERICA Genus Philonotis Brid. Bryol. Univ. 2:15. 1827.
Derivation of the name Philonotis is from the Greek philo - to love and notis = moisture, alluding to the habitat of most species.
The North American plants are small and slender (about
1 cm long), or robust (to 10 cm), bright green to yellowish, almost prostrate or usually ascending, irregularly branched or more frequently with whorled fastigiate innovations below the gametangial heads, in loose or usually dense tufts forming small patches or extensive sods, the stems clothed in ± dense reddish tomentum with a prominent central strand and a single layer of vesicular turgid cortical cells. Small reproductive branches are occasionally present and subtended by minute, ecostate, ovate residual leaves.
Leaves arranged in five to many ± spiral rows, widely spaced to usually crowded, appressed-imbricate to erect spreading, narrowly triangular-lanceolate to broadly ovate- lanceolate, 0.5-3.5 mm long, + plicate, sometimes falcate, acuminate and acute or occasionally bluntly ovate. Margins variously serrulate with single or double teeth nearly throughout, inrolled, or flat. Costa subpercurrent to excurrent. Lamina unistratose, except for perigonial leaves, composed of cells variously elongate rectangular to quadrate rectangular, somewhat rounded in lax plants. Papillae at upper or lower cell ends or middle, on one or both surfaces, 41 rarely smooth. Some species' have dimorphic stem leaves with typical leaves on sterile and female plants and appressed, smaller, and more distant leaves with blunt tips on male plants.
Dioicous, (monoicous in P. longiseta), sporangia
arising terminally becoming lateral as the stems develop, perigonia gemmiform (bud shaped) or discoid (rosette shaped), bracts broadly ovate, sheathing at the base, abruptly narrowed with weakly developed costae, paraphyses abundant
and filiform. Seta single, elongate 1-7 cm long, straight, or flexuose. Calyptra small, glabrous, cucullate, fugacious.
Capsules ovoid to subglobose, bright green when developing becoming dark rust brown, strongly inclined to horizontal,
asymmetric, ribbed, or strongly furrowed longitudinally and constricted when empty and dry, short necked with numerous stomata, operculum convex-conic or bluntly apiculate, annulus
absent. Peristome double, sixteen teeth in each circle,
exostomial teeth lanceolate, dark reddish or orange brown,
appearing trabeculate with prominent horizontal lamellae
occasionally with round tori in the upper segments, segments
of the endostome pale yellowish, striate papillose, or
smooth, with a well developed basal membrane, cilia 1-3, well
developed, irregular, and occasionally cohering. Spores
globose or reniform, densely papillose, brown to orange brown, 20-35y.
Hygrophytic mosses growing on various substrates
rarely containing much organic material, some species frequenting shaded ditches, cliffs, or ravines and others most often in exposed meadows, lake and river banks, or springs.
Type species: Mnium fontanum (= Philonotis fontana
(Hedw.j Brid.) Hedw., Sp. Muse. 195. 1801. 43
B. KEY TO THE NORTH AMERICAN SPECIES OF PHILONOTIS
(Leaves must be cleared in lacto-phenol)
1. Leaf cells papillose at upper or lower ends 2
1. Leaf cells papillose in the center throughout
P. yezoana p. 90
2. Leaf cells papillose at. the upper ends throughout.... 3
2. Leaf cells papillose at the lower ends at least in
lower portion of leaf.. 7
3. Leaf cells elongate (6-20:1) in + vertical rows..4
3. Leaf cells quadrate (1:1) to rectangular (4:1) in
less obvious vertical rows 5
4. Leaves long triangular-lanceolate, straight or
falcate and flat, cells long and narrow (9-15:1),
papillae pointed at the extreme upper cell ends,
monoicous P. longiseta p. 52
4. Leaves triangular to slightly ovate-lanceolate,
carinate, cells rectangular (less than 9:1), papillae
round near the upper cell ends, dioicous .
P. marchica p. 63
5. Papillae obvious and numerous, margins rolled and
strongly scabrous, costa long excurrent, rare, Florida
only P. sphaerocarpa p. 60
5. Papillae faint, margins + rolled and toothed, costa
variable 6
6. Costa percurrent or ending short of the apex, margins
composed of rounded papillose cells appearing doubly
serrate, southeastern U.S.A P. glaucescens p. 46 44
6. Costa long excurrent, margins singly serrate,
Pacific Coastal P. oapillaris p. 69
7. Margins doubly serrate, (sometimes very faint)
P. fontana 8
7. Margins never doubly serrate.P. oapillaris p. 69
8. Plants usually large (4-10 cm), forming loose tufts
or sods (very slender in aquatic or shaded plants),
± tomentose , costa variable, holarctic 9
8. Plants small (less than 5 cm), forming compact turfs
held together by abundant tomentum, costa excurrent,
northern or alpine P. fontana var. pumila p. 86
9. Leaves broadly ovate and shortly lanceolate,
widely spaced with several plicae on each side
of the costa, stem tips twisted, western
P. fontana var. amerioana P« 82
9. Leaves variable, ovate-lanceolate + falcate,
rarely with shallow plicae, stem tips not
twisted, cosmopolitan
P. fontana var. fontana p. 73 C. EXPLANATION OF SPECIES FIGURES
To facilitate reference and comparison of the illustrations, all figures have been arranged and lettered uniformly. The details of the leaf cross section, lamina, margin, basal angle, and apex were all drawn at the same magnification and reduced in proportion to the indicated measurement on each plate.
A. Leaf
B. Median leaf cross section
C. Cells in upper portion of lamina
D. Cells in median portion of lamina
E. Cells of leaf base
F. Median marginal leaf cells
G. Apical leaf cells 46
Philonotis glauoescens (Hornsch.) Broth., Bih. K. Svensk.
Vet. Ak. Handl. 21 Afd. 3(3):27. 1895. Fig. 49.
Bartramia glauoescens Hornsch. in Mart., Fl. Brasil.
1(2):40. 1840. TYPE: "in trucis vetustis silvarum
prope Sebastianopolin. Brazil. Beyrioh,"
(Holotype BM seen).
Philonotis glauoescens var. brevifolia Flow, in Grout,
Moss. Fl. No. Amer. 2(3):168. 1935. nom. nud.
P. glauoescens f. laxa Flow, in Grout, Moss Fl. No.
Amer. 2 (3) -.168. 1935. TYPE: "Florida, Smith,"
(type reported by Flowers to be at US but not
locatable). syn. nov.
Bartramia tenella C. Mull., Syn. Muse. Frond. 1:481.
1849. TYPE: "Insula Trinitatis Antillarum, Cruger,"
(not seen).
Philonotis tenella (C. Mull.) Jaeg., Adumb. 1:541.
1873-4.
P. muehlenbergii var. tenella Brid., Bryol. Univ. 2:23.
1827.
P. sphaerooarpa (Hedw.) Brid. ssp. tenella (C. Mull.)
Jaeg. var. terrestris Dism., Bull. Soc. Bot. France
10. Mem. 17:14. 1910. TYPE: "Florida, Beresford,
Hulst 6a & 6b," (not seen).
P. glauoescens var. terrestris (Dism.) Flow, in Grout,
Moss Fl. No. Amer. 2(3):168. 1935. syn. nov.
P. uncinata var. glauoescens (Hornsch.) Florsch., Fl.
Suriname 6(1):205. 1964. syn. nov. 47
P. graoillima Angstr. , Oefv. Kgl. Sv. Vet.-TAkad. Forh.
33(4):17. 1876. TYPE: "Brazil, Caldas. A. F.
Regnell 38," (Holotype S-PA seen). syn. nov.
P. unoinata var. graoillima (Angstr.) Florsch., Fl.
Suriname 6(1):205. 1964. syn. nov.
Plants 0.5-2.0 cm, mostly small pale glaucous green to yellowish green forming thin and scattered patches on soil, rarely very tomentose.
Leaves 0.4-1.2 mm long (mostly 0.7-8.0) by 0.1-0.3 mm wide, crowded at the top to widely spaced below, erect becoming erect spreading when moist, ovate-oblong or triangular-lanceolate occasionally subfalcate; apex bluntly rounded-obtuse (in young leaves and in plants of shaded sites) to slenderly acute; costa percurrent or ending below the apex, papillose near the apex on the abaxial surface; margins of one or two rows of inflated cells, bluntly papillose at the upper ends only and appearing doubly serrate, rarely flat; lamina of pellucid, usually thin walled oblong-rectangular to rhomboidal cells (2-5:1) becoming quadrate below, rounded papillae at upper cell ends restricted to the adaxial surface, usually throughout but occasionally faint or absent.
Dioicous, rarely with sporophytes in North America, perigonial leaves long acuminate forming a bud enclosing the antheridia (gemmiform).
Seta 1-2 cm long; capsule 1.5-2.0 mm, spherical; stomata
50-100 at base of capsule; outer peristome teeth as for the 48 genus, occasionally producing spherical interlamellar thickenings, inner peristome evenly papillose; spores
22-26y bluntly rounded papillose.
CHROMOSOME NUMBER: n = 6. J. Snider has generously permitted inclusion of the report of this count he made at
Duke University from a culture of P. glaucescens (Florida,
Hernando Co., Brooksville, Zales 2041 UBC) (fig. 51).
HABITAT: On basic mineral substrata (limestone, sandstone, sandy soil, mud, or rarely charred wood), in ditches or sinkholes, along river or creek banks, and at the margins of springs, waterfalls, or dams, predominantly in shaded locations.
DISTRIBUTION: Common in the northern two thirds of
Florida and rare in Texas, Louisiana, and Georgia. Reported from the West Indies, Mexico, Central and South America
(Map fig. 64).
Philonotis glaucescens and P. gracillima were distinguished solely on the basis of leaf apex shape and costa length. Most of the collections examined, including the type specimens of both species, show both rounded-obtuse leaves with the costa ending just below the apex and acuminate leaves with a percurrent costa. The separation of these two taxa was usually a subjective judgement dependent on which kind of leaf was more abundant. Culture experiments demonstrate that these two characters are ecophenic (fig. 34,
35) .
Occasionally P. glaucescens has been confused with Figures 4 6-48. Comparison of leaf shape and leaf margin
-46. Philonotis glauoescens
-47. Philonotis marchica
-48. Philonotis sphaerooarpa
50
P. marchica. Very young and lax P. marchica may have a blunt apex with faint papillae on the lamina but the triangular-lanceolate shape and flat leaf margin of oblong pointed teeth will distinguish it from the lanceolate shape and usually rolled leaf margin composed of short, blunt, and somewhat inflated teeth of P. glaucescens (fig. 46, 47).
Florschutz (1964) has treated P. glaucescens and
P. graoillima as varieties of P. uncinata with the reservation that even varietal status may be too high.
Because P. sphaerocarpa does not occur in the area of
Florschutz' monograph he missed the opportunity of synonymizing it with P. uncinata. Philonotis sphaerocarpa however has priority. He is correct in combining P. graoillima and P. glauces cens but overlooked several distinctive features separating them from P. sphaerocarpa (uncinata). Philonotis glaucescens is a small pale plant rarely exceeding 2.0 cm with ovate-oblong or triangular-lanceolate leaves of usually thin walled cells with faint papillae and bluntly toothed margins. Philonotis sphaerocarpa is generally robust, yellowish green, and 2.0-5.0 cm high with concave oblong- lanceolate leaves of thick walled cells with prominent papillae and a scabrous appearing margin due to the large papillae on the revolute laminar margin (fig. 46, 48). Figure 49. Philonotis glaucescens 52
Philonotis longiseta (Rich, 'in Michx.) Britt., The
Bryologist 14:44. 1911. Fig. 54.
Bartramia longiseta Rich, in Michx., Fl. Bor. Amer.
2:301. 1803. TYPE: "in Carolina ex D. Bosc et in
Virginia ex D. Beauvois," (Isotype NY seen).
Philonotis longiseta var. propagulaecaulis Flow., The
Bryologist 38:8. 19 35. TYPE: "wet soil, Louisiana,
Lafayette, Leon 1353," (Holotype US seen). syn. nov.
P. longiseta f. propagulioaulis Flow, in Grout, Moss Fl.
No. Amer. 2(3):172. 1935. nom. nud.
P. longiseta var. polygama Flow, in Grout, Moss Fl.
No. Amer. 2(3):171. 1935. TYPE: "near Careyville,
Campbell Co., Tennessee, on moist roadside bank,
Sharp 34154," (Holotype UT seen). syn. nov.
Bartramia radicalis P. Beauv., Prodr. Aetheog. 44. 1805.
Philonotis radicalis (P. Beauv.) Brid., Bryol. Univ.
2:17. 1827.
Bartramia radicalis var. porteri Aust., Bull. Torrey Bot.
Club 6 (36) :190. 1877. TYPE: "on wet rocks and banks,
western Pennsylvania, Garter', Rockdale (Lehigh Valley)
Wolle 1874; Oneida, New York, Warne 1873; Ohio,
Beardslee & Biddlecome," (Lectotype Wolle 1874 F seen).
Philonotis longiseta var. porteri (Aust.) Flow, in Grout,
Moss Fl. No. Amer. 2(3):171. 1935. syn. nov.
Plants 0.5-3.0 cm high (mostly 2.0), pale yellowish green to green, forming loose to dense, usually small rounded tufts, branching especially below the sex organs, 53 usually rust brown tomentose, often producing deciduous propaguliferous shoots.
Leaves 1.0-2.0 mm long, by 0.1-0.3 mm wide, erect and occasionally secund when dry becoming spreading when moist, linear triangular-lanceolate to lanceolate, slenderly acuminate with an excurrent costa; cells long linear (9-15:1) in. vertical rows becoming gradually enlarged below, not inflated, papillose at the extreme upper ends of all cells on the adaxial surface only; margins plane or narrowly revolute + the entire length usually on both sides,, strongly serrate with blunt teeth at upper cell ends only.
Autoicous or synoicous, perigonia gemmiform (composed of erect leaves enclosing the gametangia and forming a "bud").
Seta 1.5-3.5 cm long, capsule rounded, stomata 67-146
(average 100) per capsule, restricted to the lower one quarter, peristome as for the genus, spores 22-33y, reniform, papillose.
CHROMOSOME NUMBER: n = 12. Heitz (1942) and Lowry
(194 8) reported for the genus Mnium, that those species which have the basic chromosome number are generally dioicous and those species which are polypoids are monoicous. This also seems to be the case in Philonotis. Ochi (1963) has shown that the dioicous and monoicous races of P. socia in
Japan are the same and Yano (1955) has described n = 6 and n = 12 populations of P. socia, but it is not clear from the literature if the dioicous populations are n = 6 and the monoicous populations are n = 12. This generalization is supported by the monoicous P. longiseta. J. Snider, using the facilities at Duke University, has established a chromosome number of n = 12 for a polygamous specimen of
P. longiseta (Florida, Leon Co., Tallahassee, Zales 1941
UBC) (fig. 50). All of the species of Philonotis that have been counted except for P. socia and P. longiseta are dioicous and n = 6. Ironically there is no n = 6 dioicous taxon that corresponds morphologically to P. longiseta, at least not in North America.
HABITAT: On sandstone, limestone, or phosphate rock, sandy or clay soil low in organic material, in ditches, ravines, or on river banks and bluffs, around springs, or along roadsides. Usually at low elevations in shaded places that are wet for some part of the year.
DISTRIBUTION: Predominantly southeastern, common in the Ozarks and southern Appalachian mountains extending north along the rivers into Iowa, west into central Texas, south to central Florida, and northeast to Pennsylvania.
Reported from Guatemala, Martinique, Mexico, and Puerto
Rico (Map fig. 65).
Philonotis longiseta is a clearly demarcated species, geographically, sexually, cytologically, and morphologically.
In North America P. longiseta occurs east of the 100th meridian and mostly south of the 40th parallel. It is the only North American species of Philonotis that is monoicous or has a haploid chromosome number of 12, and it has long lanceolate-triangular leaves with revolute margins and Figures 50, 51. Mitotic chromosome configurations
of Philonotis
-50. P. longiseta
-51. P. glauoes cens 55 56
papillae at the extreme upper ends of most cells.
The perigonial leaves are variable and of little
taxonomic value. They consist of broad, lax, almost
ecostate inner leaves that give the perigonium an almost
discoid appearance to leaves that are similar to enlarged
comal leaves.
Philonotis longiseta has occasionally been confused with P. mavchioa. If the leaves are cleared with
lacto-phenol it is obvious that the former has longer
narrower cells with papillae at the extreme upper cell ends,
and the latter has papillae on or near the upper cell ends.
Philonotis mavehiaa also has a shorter broader leaf.
Extremely young and lax or etiolated plants may have shorter
and broader leaves with slightly rounded cells, plane
margins, and a percurrent costa. These can be recognized
as P. longiseta by the position of the papillae on the
leaf cells (fig. 36, 37, 52, 53).
Philonotis longiseta var. porteri was described from
several collections possessing erect, slightly tapering
leaves, shorter stems and seta, and a more erect capsule. All
but one of the specimens cited in Austin's description (1877)
of Bartramia radicalis var. porteri have been seen: western
Pennsylvania, Garber (not seen); Rockdale, Lehigh Valley,
Vlolle 1874 (Lectotype F) is a lax expression of the species;
Oneida, New York, Warne 1873 (NY) is P. longiseta, 1874 (NY)
is a mixture of P. longiseta and P. marchica', Ohio, Beardslee
(NY) is P. longiseta, Biddlecome 1877 (US) is P. marchica. Figures 52, 53. Comparison of leaf shape and median
leaf cell shape
-52. Philonotis marchica
-53. Philonotis longiseta
Besides the original description, several pages of Austin's notes are housed in the New York Botanical Garden Herbarium which compare B. radicalis var. porteri with B. striata and
P. marchica but give no indication of its difference from
B. radicalis . The minor size differences seen in the syntypes do not support the recognition of this variety as being distinct from the species. 59
Figure 54. Philonotis longiseta 60
Philonotis sphaerocarpa (Hedw.) Brid., Bryol. Univ. 2:25.
1827. Fig. 55.
Mnium sphaericarpon Hedw., Sp. Muse. 197. 1801.
TYPE: "In Jamaica, Swartz," (Isotype NY seen).
Philonotis sphaericarpa var. terrestris Flow, in
Grout, Moss Fl. No. Amer. 2(3):168. 1935. nom. nud.
Bartramia uncinata Schwaegr., Suppl. Sp. Muse. 1(2):60.
1816. TYPE: "In monte sulphurifero Guadalupae et
in Martinica lectum et curiose observatum dedit,
Richard," (Holotype Guadalupae PC seen).
Philonotis uncinata (Schwaegr.) Brid., Bryol. Univ..
2:22. 1827. syn. nov.
Plants 2.0-5.0 cm, relatively robust, yellowish green erect tufts or dense mats, abundantly rust brown tomentose below, often branched below the sex organs.
Leaves 1.0-2.0 mm long, usually crowded, erect or erect-spreading, straight or subfalcate especially at the stem tips, lanceolate to oblong-lanceolate often concave at the base, acuminate with the costa short excurrent to strongly spinulose-excurrent, margins usually strongly revolute appearing doubly or more serrate nearly to the base, upper cells linear or oblong-linear (3-7:1) becoming short rectangular below, densely scabrous with papillae at upper cell ends on the adaxial surface.
Dioicous, perigonia small and bud shaped (gemmiform).
Seta 1.5-3.0 cm long, capsule spherical, 1.5-2.5 cm, stomata 90-150 restricted to the base, peristome as for the 61
genus, spores 20-26u papillose.
CHROMOSOME NUMBER: Not reported.
HABITAT: On damp soil and rock banks and cliffs at various elevations in the American tropics.
DISTRIBUTION: One station in North America (Sanford,
Florida), West Indies, Mexico, Central and South America
(Map fig. 66).
Examination of the type material of P. sphaerooarpa
and P. uncinata and many authentic specimens from the West
Indies, Mexico, Central and South America indicate that the
range of variability of these taxa overlap to such a great
extent that they are impossible to separate. The only major
difference between the type specimens is the amount of
curvature in the leaves which in the genus Philonotis is a
notoriously variable character.
The single waif specimen from North America was
collected by S. Rapp in 1905 in a hummock near Sanford,
Florida. All other specimens identified as P. sphaerooarpa
have been assigned to P. longiseta^ glauoescenssor marchica.
For the characters separating P. sphaerooarpa from
P. glauoescens, see the discussion under P. glauoescens.
63
Philonotis marchica (Hedw.) Brid., Bryol. Univ. 2:23 & 735.
1827. Fig. 58.
Mnium marchicum Hedw., Sp. Muse. 196. 1801.
Philonotis glabriusoula Kindb. in Mac. et Kindb., Cat.
Can. PI. 6:107. 1892. pro. parte. TYPE: "by
springs in a gully, Canaan Forks, Queen's Co., N. B.,
January 2nd, 1890. J. Moser," (Syntype CAN seen,
S-PA seen).
P. subcapillaris Kindb., Eur. No. Amer. Bryin. 326.
189 7. pro. parte. TYPE: "Switzerl. near Lugano,"
(not seen).
Bartramia muehlenbergii Schwaegr., Suppl. Sp. Muse.
1(2):58. 1816. TYPE: "in Pennsylvania,
Muhlenberg," (not seen).
Philonotis muehlenbergii (Schwaegr.) Brid., Bryol. Univ.
2:22. 1827.
Plants 0.5-6.0 cm high, mostly small and yellowish tufts in dry habitats, becoming robust and brighter green forming erect mats in moist areas and high elevations. Stems brown tomentose below and occasionally clothed with abundant propagula.
Leaves 0.9-2.2 mm long by 0.3-0.5 mm wide, erect spreading, more divergent when moist, occasionally arranged in five spiral rows in robust plants, triangular-lanceolate mostly straight, carinate; margins plane or only narrowly recurved, serrate nearly to the base with papillae at the upper ends only; costa narrow, shortly excurrent to clearly 64
excurrent; upper cells linear-oblong (5:1), 4-8u wide,
lower cells oblong (3-4:1), 6-15u wide, papillae obvious at
upper ends of cells throughout, on the adaxial or dorsal
surface only.
Dioicous, perigonia discoid (forming a rosette).
Seta 2-4 cm long; capsule spherical, 1.0-2.5 mm, stomata
two or rarely four celled, 90-200 mostly 130 or more per
capsule, restricted to the base; outer peristome as for the
genus, inner peristome papillose in lines, cilia variable;
spores 20-30y, papillose.
CHROMOSOME NUMBER: Not reported.
HABITAT: On rocks and sandy soil in wet places, usually
in slowly running water and forming small to large mats
depending on the moisture, at various elevations. Reported
on calcareous schist, limestone, sandstone, travertine, bogs (?), and roadside ditches.
DISTRIBUTION: Rare in the Rocky Mountains from
southern British Columbia and Alberta south to Mexico, absent
from the interior plains but reappearing in the central
highlands of Missouri and adjacent states, the mountains of
eastern North America from southern Newfoundland to Georgia,
and in scattered localities around the Great Lakes. Reported
from central and southern Europe and the western Mediterranean
(Map fig. 67).
The existing literature implies confusion between
P. marchica and P. capillaris. However, the habit of this
species is usually much more erect and rigid than P. capillaris. 65
The stem leaves are closer and more erect and have a triangular base compared to the ovate base of P. oapillaris.
The leaf cells of P. marchica are longer, narrower, and distinctly mammillose throughout (fig. 56, 57). Besides obvious morphological differences the range of P. marchica and P. oapillaris are exclusive except possibly in central
Idaho and southern British Columbia.
Philonotis muehlenbergii (Schwaegr.) Brid. was originally described (Schwaegrichen 1816) from a single specimen collected from Pennsylvania by Muhlenberg. This specimen appears to no longer be in existence. The species has persisted in the literature to the present time in spite of Dismier's review (1910), and the subsequent translation and publication of the same paper in The Bryologist by
Britton (1911), which demonstrated the lack of distinct characters separating it from P. marchica. Flowers (e. a.) has synonymized P. muehlenbergii and P. marchica for the state of Utah; "I have found that those with triangular to oblong-lanceolate leaves, broadest at the insertion, ultimately have shown some of the lower cells to be papillose at the upper ends and referrable to P. marchica."
Recent taxonomic works have relied on the absence or presence of only rudimentary cilia to separate
P. muehlenbergii from P. marchica. Considering the variability observed in a large series of specimens throughout their geographic range and the ease with which the fragile cilia are broken and lost, there is no reason to maintain 66
P. muehlenbevgii as a separate taxon.
For a comparison of P. marchica and P. glaucescens see the discussion under P. glaucescens. Figures 56, 57. Comparison of leaf shape, apex, and
median leaf cells
-56 . Philonotis marchica
-57. Philonotis capillaris
69
Philonotis capillaris Lindb. ex Hartm., Skand. Fl. ed. II
10:40. 1871. TYPE: "R. Spruce, Lassaron, Pyreneerna,"
(Isotype M7ANCH seen, S-PA seen). Fig. 59.
Philonotis capillaris Lindb., Hedwigia 6:40. 1867.
nom. nud.
P. fontana (Hedw.) Brid. var. capillaris (Lindb.)
Lindb., cf. Dism. Soc. Nat. Sci. Nat. Math.
Cherbourg Mem. 36:392. 1907.
P. macounii Lesq. et James, Man. Mosses No. Amer. 208.
1884. TYPE: "Vancouver Island, Macoun, Can. Musci
152," (Lectotype US seen, Isotype H seen).
P. vancouveriensis Kindb., Eur. No. Amer. Bryin. 2:326.
1897. pro. parte. TYPE: "Vancouver Island and
Alaska, J. M. Macoun," (Syntype S seen, Isotype
CAN seen).
P. arnellii Husn., Muse. Gall. 268. 1890.
Plants slender, 1-3 cm high, in loose tufts or scattered, often procumbent, tomentose below.
Leaves 0.7-1.5 mm long by 0.3-0.5 mm wide, erect when dry, erect spreading when moist, lanceolate from an ovate base, acuminate, without plicae; margins plane or narrowly recurved above the ovate base, distinctly serrate nearly to base with single teeth at upper ends of cells only, basal border of five to ten or more quadrate cells; costa excurrent to long excurrent, upper cells (2-3:1) 6-lly wide, lower cells (2-1) 8-14y wide; papillae obscure in lower half of leaf on the adaxial surface, rarely at lower cell ends or 70 overlapping on opposite surfaces, cell walls thick, the lumen rounded.
Dioicous, often collected with sporophytes, perigonia discoid (forming a rosette).
Seta 2-3 cm long, capsule rounded, furrowed when dry; stomata 40-115 per capsule, restricted to the base, mostly less than 100; outer peristome as for the genus, inner peristome papillose in lines, cilia variable; spores 20-26u, papillose.
CHROMOSOME NUMBER: Not reported.
HABITAT: On wet sandy soil, humus, or various thin soils over rock on slopes, cliff ledges, or ditches.
Preferring wet shaded places mixed with small vascular plants and other bryophytes or both.
DISTRIBUTION: The Pacific Coast from Alaska to central California and east to northern Idaho and southeastern
British Columbia. Reported from Europe and Asia Minor
(Map fig. 68) .
The type collection is sterile and immature material collected by R. Spruce from the Pyrenees. However, it does demonstrate the specific characters. The margins of
P. oapillaris are composed of single teeth, all projecting at the upper cell ends as compared to the P. fontana complex where the margins are doubly toothed by cells projecting at both ends. Philonotis oapillaris has been reported from several stations in central and eastern North America, however, these have all ultimately shown some cells with two projections, and thus cannot be included in this species. The papillae of the broadest part of the lamina are obscure and must be cleared with lacto-phenol to be observed at all. Papillae are at the upper ends of the cells but occasionally may be in the lower ends as in the type material.
The small size of fertile plants, long excurrent costa flat almost smooth lamina, ovate leaf base with single toothed margins, and Pacific Coast distribution in North
America make this species easily recognizable.
The European P. arnelli may be conspecific with
P. oapillaris, as stated by Dismier (1907b, 1910),
Brotherus (1923), and Monkemeyer (1927). Certainly all
North American specimens labeled P. arnellii are
P. oapillaris.
Philonotis oapillaris occasionally possesses propaguliferous shoots that would normally develop into branches (innovations typical of the genus) if the plant continued to grow. Propaguliferous shoots in. P. oapillaris have also been reported by Field (1965) and Dismier (190 7b) 72
Figure 59. Philonotis capillaris 73
Philonotis fontana (Hedw.) Brid. var. fontana, Bryol. Univ.
2:18. 1827. Fig. 60.
Mnium fontanum Hedw., Sp. Muse. 195. 1801.
Bartramia fontana (Hedw.) Turn., Muse. Hib. 107. 1804.
Philonotis adpressa Ferg. in Hunt, Mem. Lit. Phil. Soc.
Manchester ser. 3(5):102. 1872.
P. fontana var. adpressa (Ferg.) Limpr., Krypt. Siles.
1:116. 1876. syn. not).
P. fontana ssp. adpressa (Ferg.) Dix., Stud. Handb.
Brit. Moss. 295. 1896.
P. aoutiflora Kindb. in Roll, Hedwigia 35:67. 1896.
TYPE: "Victoria, Vancouver Island, J. Roll 52,"
(Lectotype S-PA seen, Isolectotype MANCH seen).
P. acutifolia Kindb. ex Dism., Bull. Soc. Bot. France
10, Mem. 17:27. 1910. nom. inval. err. pro.
P. aoutiflora Kindb.
P. alpioola var. tenuis Kindb. in Moll., The Bryologist
27:75. 1924. nom. nud.
P. braohychypus Kindb. in Moll., The Bryologist 27:75.
1924. nom. nud.
P. tomentella Mol. f. brachycarpa (Kindb.) Moll., The
Bryologist 27:75. 1924. nom. nud.
P. fontana var. brachyphylla Kindb. in Mac. et Kindb.,
Cat. Can. PI. 6:107. 1892. TYPE: "P. E. I.,
Barkley Pt. in bogs, 28 June 1888, Macoun," (Isotype
CAN seen). 74
P. glabriuscula Kindb. in Mac. et Kindb., Cat. Can.
PI. 6:107. 1892. pro. parte. TYPE: "by springs
in a gully, Queen's Co., N. B., Jan. 2, 1890,
J. Moser," (Syntype C/AN seen, MANCH seen, WIS seen).
P. microcarpa Kindb., Rev. Bryol. 32:37. 1905. TYPE:
"eastern slope of Rocky Mts., bogs along Pipestone
Cr. alta 6,000 ft., 8 July 1904, 75," (Lectotype
S-PA seen).
P. fontana var. mi or obi asta C. M. et Kindb. in Mac. et
Kindb., Cat. Can. PI. 6:107. 1892. TYPE: "Selkirk
Mt. Roger's Pass, along R. R., 30 July 1890, Macoun,"
(Lectotype S-PA seen, Isolectotype CAN seen).
P. fontana var. microthamnia Kindb., Ottawa Nat. 23:183.
1910. TYPE: "Yukon, Hunker Creek, peat bog, July
1902, 152," (Isotype CAN seen).
P. fontana var. columbiae Kindb. in Mac. et Kindb., Cat.
Can. Pi. 6:107. 1892. (published as Columbia)
TYPE: "on wet rocks below the Illicillewaet Canyon,
Revelstoke, B. C., 18 May 1890, Macoun," (Holotype
S-PA seen).
P. fontana var. serrata Kindb. in Mac. et Kindb., Cat.
Can. PI. 6:107. 1892. TYPE: "Selkirk Mountains,
swamps in Roger's Pass, 24 August 1885, Macoun',
Ounalaska & Atkah Islands, Behring Sea, 1891, Macoun,"
(Lectotype Roger's Pass CAN seen, Isosyntype Ounalaska
& Atkah Islands CAN seen). 75
P. fontana var. heterophylla Card, et Ther., Univ.
Calif. Publ. Bot. 2:300. 1906. TYPE: "Unalaska,
Setchell 2 & 30," (Lectotype 2 UC seen, Syntype 30
UC seen), syn. nov.
P. vanoouveriensis Kindb., Eur. No. Amer. Bryin. 2:326.
1897. pro. parte. TYPE: "Vancouver Island &
Alaska, Macoun," (Syntype S-PA seen, Isosyhtype CAN
seen).
P. tenella C. Mull. var. ooloradensis Ren. et Card.,
Bot. Gaz. 22:51. 1896. orth. err. P. venella.
TYPE: "Colorado, Boulder Co., Springdale, M.
Eolzinger 1892," (Lectotype NY seen, Isolectotype
CAN seen, WIS seen).
P. fallax Dism., Bull. Soc. Bot. France 10. Mem. 17:35.
1910. TYPE: "Canada, B. C., Sicamous, Finlayson
1583 . . . Idaho, Lake Pend d'Oreille, Leiberg 49,"
(Lectotype Finlayson NY seen, Syntype Leiberg CAN
seen, NY seen).
P. fontana f. dimorphophylla Flow, in Grout, Moss Fl.
No. Amer. 2(3):175. 1935. TYPE: "Alaska," (not
seen).
P. fontana var. seriata Breidl. f. oooidentalis Flow.
in Grout, Moss. Fl. No. Amer. 2(3):176. 1935. nom. nud.
P. calcarea (B. S. G.) Schimp. f. oooidentalis Flow, in
Grout, Moss Fl. No. Amer. 2(3):179. 1935. TYPE:
"Texas, Harvard.s. n.," Not in US as reported by
Flowers, (Isotype SMU seen). 76
P. fontana var. flaoata Brid., Bryol. Univ. 2:21. 1827.
P. caespitosa Jur., Verh. Zool.-Bot. Ges. Wien 11:235.
1862. syn. nov.
P. fontana var. caespitosa (Jur.) Limpr., Krypt. Fl.
Schlesien 1:116. 1876.
P. caespitosa var. compacta Dism., Bull. Soc. Bot.
France 10. Mem. 17:28. 1910. TYPE: "Conn., Ledyard
& Hamdem, Nichols," (not seen). syn. nov.
P. caespitosa var. compacta Jaeg., Ber. S. Gall. Naturw.
Ges. 1873-74:86. 1875. (Ad. 1:548). nom. nud.
P. caespitosa var. heterophylla Dism., Bull. Soc. Bot.
France 10. Mem. 17:28. 1910. No specimen cited.
P. caespitosa var. adpressa Dism., Rev. Bryol. 34:68.
1907. syn. nov.
P. caespitosa var. ampliretis (Dix.) Crum et al., The
Bryologist 68:434. 1965. syn. nov.
P. fontana var. ampliretis Dix., Jor. Bot. 40:73. 1902.
P. caespitosa var. laxa Loeske et Warnst. in Loeske,
Hedwigia 45:104. 1906.
P. laxa Warnst., Krypt. Fl. Mark Brendenb. 2:619. 1895.
horn, illeg. (P. laxa Limpr., Laubm. Deutschl. 2:563.
1893. Name not North American = P. marchica).
P. fontana var. laxa Flow, in Grout, Moss Fl. No. Amer.
2(3):178. 1935. horn, illeg. (P. fontana var. laxa
Vent. Rev. Bryol. 9:45. 1882). P. fontana var. laxa f. tenuis Flow, in Grout, Moss
Fl. No. Amer. 2(3):178. 1935. TYPE: "collected in
New York, Pennsylvania and Colorado," (Lectotype
Holz. Musci Aero. Bor.-Amer. 661 NY seen, Isolectotype
CAN seen).
P. fontana f. flowersii Crum, The Bryologist 72:244.
1969 . syn. nov.
P. fontana var. aristinervis (Monk.) Diener, Cas. Nar.
Mus. Praha 104:122. 1930.
P. fontana f. aristinervis Monk., Laubm. Eur. 584. 121c.
1927.
P. amerioana var. graoilesoens Dism., Bull. Bot. Soc.
France 10. Mem. 17:23. 1910. TYPE: "Canada, Gaspe
Co., Mt. Albert, P. Q., J. F. Collins 4172,"
(Holotype NY seen). syn. nov.
Plants extremely variable, 2.0-10.0 cm high, (rarely
larger) usually robust erect ± irregularly branched, often
producing whorls of short branches below the sex organs, bright yellow green to green, forming small scattered tufts
to extensive dense sods matted with rust brown tomentum.
Leaves erect, appressed to erect-spreading ± falcate and
secund, 0.7-2.5 mm long, by 0.5-1.5 mm wide, mostly ovate-
lanceolate with a broad base gradually or abruptly tapering
to a ± slender acuminate apex (occasionally blunt), costa
stout, percurrent to excurrent, margins revolute or plane,
serrate with some cells, projecting at both ends (doubly
serrate) becoming plane at the base and singly serrate near 78 the apex, lamina usually concave with one or several shallow plicae on either side of the costa, usually plane, upper leaf cells linear or oblong-linear (4-10:1) papillose at the upper cell ends on the abaxial surface, lower cells becoming shorter and broader and papillose at the lower cell ends on the abaxial surface. Male stems usually rust brown with distant, broad, and appressed leaves with blunt apexes and lax cells with faint papillae.
Dioicous, perigonia discoid (rosette shaped) bracts large, lax, and highly variable.
Seta 2.0-6.0 cm long, capsule rounded, stomata
110-275 restricted to lower quarter, peristome as for the genus, spores 20-30u, papillose.
CHROMOSOME NUMBER: n = 6 Anderson & Crum (1958),
Vaarama (1953, 1956), Smith & Newton (1966).
HABITAT: Philonotis fontana occurs in almost any wet habitat, but is most abundant and luxuriant in seepage water in exposed locations such as ditches, sloping banks of rivers or highways, and around springs or small waterfalls, also wet meadows and lake shores, and growing over mineral substrates such as mud, sand, gravel, or rocks of various composition. Plants growing in shaded, dry, or completely aquatic sites show ecophenic variation.
DISTRIBUTION: Widespread throughout most of North
America especially in regions of variable topography, generally absent from the northern tundras, central prairies, and southern portions of the states bordering on the Gulf of Mexico. 79
Widespread in Europe, Asia, northern Africa, and extending
into Mexico (Holarctic) (Map fig. 69).
Philonotis fontana has a phenotype that is easily
modified by various environmental conditions. So many of
these modified plants have been described and named that
positive identification has become difficult and confusing.
Experimental cultures have shown that leaf shape, size,
color, arrangement on the stem, thickness of cell walls, and
presence or absence of tomentum are highly variable ecophenic
characteristics, and thus useless as diagnostic characters.
The names P. adpressa and var. adpressa of several species
have been used for plants with broad and short leaves with
blunt apexes, closely appressed to the stem. This is a
common condition of male or semiaquatic plants growing in full
exposure to the sun. The names tomentella and borealis used
at the specific or varietal level (sensu European workers)
have been applied to plants that have narrow elongate leaves with slender excurrent costae. According to the habitat data
on the specimens this seems to be a response to low
temperature and high elevation. These plants often have
leaves swept to one side which have been included under
descriptions of var. falcata. Philonotis caespitosa and
several lesser varieties of P. caespitosa have been described
from small plants that usually have flat leaves with plane
margins. These plants are almost always sterile and represent
small plants of P. fontana. The epithet laxa has been
applied by several authors to several species and varieties of Philonotis. The nomenclature is confused because of different interpretations. They all refer to plants growing in aquatic or semiaquatic habitats that are heavily shaded causing the plants to become etiolated, pale in color, have inflated cells with thin cell walls, and lack tomentum.
N. C. Kindberg (1905, 1910) has described several specie and varieties, mostly from western Canada, that are only minor variations of P. fontana. These names are represented by relatively few specimens throughout national herbaria.
Philonotis fontana is readily separated from other species of Philonotis by the large size of the plants, the presence of obvious papillae at the lower ends of the leaf cells, and the doubly serrate margins. The distinctive features of varieties americana and pumila are discussed under the respective taxa.
82
Philonotis fontana var. amerioana (Dism.) Flow, ex Crum,
The Bryologist 72:244. 1969. Fig. 61.
P. amerioana Dism., Bull. Soc. Bot. France 10. Mem.
17:35. 1910. TYPE: "high up on the Selkirk Mts.,
near Armstrong, B. C., Wilson 19 July 1904 as
P. fontana," Holzinger, Musci Aero. Bor.-Amer. 70b,
(Lectotype NY seen, Isolectotype CAN seen).
P. seriata ssp. amerioana Dism., Bull. Soc. Bot. France
10. Mem. 17:22. 1910.
P. torquata Geh., Rev. Bryol. 23:61. 1896.
P. amerioana var. torquata (Ren. et Geh.) Dism., Bull.
Soc. Bot. France 10. Mem. 17:23. 1910.
P. macounii var. torquata Ren. et Geh., Rev. Bryol.
23:61. 1896. TYPE: "Washington, Olympia, Blacker
40," (not seen).
P. amerioana f. laxa Flow, in Grout, Moss Fl. No. Amer.
2(3):178. 1935. TYPE: "Selkirk, Canada," (not seen).
P. amerioana f. dimorphophylla Flow, in Grout, Moss Fl.
No. Amer. 2(3):178. 1935. TYPE: "Alaska, under
snow banks in cold water," (not seen).
Largest of the North American species of Philonotis, stem 5-10 cm or more long, erect, simple, or with whorled branches, interwoven with brown tomentum, forming deep yellowish green tufts or mats.
Leaves usually distant, characteristically diverging from the stem at a wide angle with the upper portion curving upward, often twisted forming a spiralling stem tip, 83 broadly ovate, concave, straight, or falcate, shortly acuminate with a percurrent or excurrent costa; lamina plicate with rounded rectangular cells papillose at the lower end becoming longer near the leaf apex; margins usually recurved and strongly doubly serrate.
Dioicous, perigonia large, discoid (rosette shaped).
Seta 3.0-6.0 cm long; capsule rounded, furrowed when dry; stomata 130-2 30 restricted to the base of capsule; peristome and spores as for the species.
CHROMOSOME NUMBER: Not reported.
HABITAT: Occurring in similar habitats as the species and reaching its maximum development on exposed slopes or mountain meadows where water slowly seeps through the extensive Philonotis mats.
DISTRIBUTION: Endemic to North America and restricted to the Coastal and Rocky Mountains from California to the
Aleutian Islands of Alaska (Map fig. 70).
Philonotis fontana var. americana was originally described by Dismier (1910) in his revision of the North
American species of Philonotis. His diagnosis and description in French on page 22 clearly established the name as a subspecies of P. seriata Mitt. To comply with the
1906 Vienna Code of botanical nomenclature, Dismier provided a Latin description on page 35, however, this description was for P. americana sp. nov. Although the original publication is ambiguous, all of Dismier's annotation labels in NY read "P. americana Dism." which indicates that he 84 considered this taxon a species rather than a subspecies.
After the French description, Dismier cited 41 specimens
(syntypes) of which 27 are at NY. Of these 27, only 11 are beyond the range of variation included in the species
concept of P. fontana. Considering the extreme variation of the species, only robust "typical" plants of var. americana can be identified with certainty.
Persson (1954) has clarified the exclusion of
P. seriata from North America, however, his opinion that ssp. americana is closely related to this extra-North
American species is erroneous.
Philonotis fontana var. americana can be recognized by its large size (5-10 cm) which is the largest of the genus in North America, the widely spaced erect spreading
leaves with several plications on each side of the costa, and the very broadly ovate-lanceolate leaves that form a spiralling stem tip. 85
Figure 61. Philonotis fontana var. amerioana 86
Philonotis fontana var. pumila (Turn.) Brid. Bryol. Univ.
2:20. 1827. Fig. 62.
Bartramia fontana var. pumilum Turn., Muse. Hib. 107.
1804. TYPE: "in paludivus," (not seen).
Philonotis fontana var. pumila f. longifolia Flow, in
Grout, Moss Fl. No. Amer. 2(3):176. 1935. TYPE:
"Nevada, Baker 15 86," (Not in US as reported by
Flowers).
P. fontana var. pumila f. dimorphophylla Flow, in
Grout, Moss Fl. No. Amer. 2(3):176. 1935. nom. nud.
P. tomentella Mol. in Lor., Moosstud. 170. 1864.
syn. nov.
P. fontana var. tomentella (Mol.) Jaeg., Ber. S. Gall.
Naturw. Ges. 1874-75:86. 1875.
P. fontana ssp. tomentella (Mol.) Dism., Bull. Soc.
Bot. France 10. Mem. 17:26. 1910.
P. fontana ssp. tomentella var. heterophylla Dism.,
Bull. Soc. Bot. France 10. Mem. 17:26. 1910.
TYPE: "B. C, Selkirk Mts. , Macoun 37 8," (not seen).
P. fontana var. pumila f. heterophylla (Dism.) Flow, in
Grout, Moss Fl. No. Amer. 2(3):177. 1935. syn. nov.
P. borealis (I. Hag.) Limpr., Laubm. Deutschl. 2:565.
1893.
P. fontana var. borealis I. Hag., Kongl. Norske. Vidensk.
Selsk. Skrift. 1888-90:13. 1889. TYPE: "In alpe
Galdh altit. ca. 1800 M. ad terram humidam 11/8,
1887," (not seen). 87
P. tomentella var. borealis (I. Hag.) Loeske., Verh.
Bot. Ver. Brandenburg 47:338. 1906. (IV).
P. fontana var. oompaota Schimp., Syn. Muse. ed.
2:520. 1876.
P. tomentella var. compacta (Schimp.) Dism., Mem. Soc.
Sc. Nat. Cherbourg 36:413. 1908.
P. fontana var. alpina (Brid.) Brid., Bryol. Univ.
2:20. 1827.
P. alpicola var. borealis (I. Hag.) Podp., Veroeff.
Gepbot. Inst. Rubel Zurich 1:254. 1924.
P. alpicola var. tomentella (Mol.) Torre et Sarnth.,
Fl. Tirol 5:412. 1904.
Plants 1.0-6.0 cm high (mostly less than 5.0 cm), erect, yellowish to dark green, forming small mats containing
5.0-15.0 mm of green leafy stems over an accumulation of densely tomentose and closely arranged straight stems with dead brown leaves.
Leaves 0.6-1.5 mm long by 0.3-0.7 mm wide, erect not crowded; + long lanceolate from a somewhat ovate concave base, straight or falcate; apex blunt in very short leaves but usually acuminate with a percurrent or shortly excurrent costa occasionally becoming hyaline; margins revolute or rarely plane, doubly serrate with cells projecting at both ends, less serrate at the base and singly serrate at the apex; lamina usually concave lacking plicae, upper leaf cells oblong-rectangular (2-8:1), papillose at the lower cell ends on the abaxial surface, lower cells similar, 88 becoming proportionately larger but shorter (2-4:1).
Dioicous, perigonia discoid (rosette shaped) bracts highly variable.
Sporophyte small, seta 10.0-20.0 mm long; capsule spherical 1.0-2.0 mm; stomata 50-120 at base of capsule; peristome as for the genus; spores 18-24u, papillose.
CHROMOSOME NUMBER: Not reported.
HABITAT: Common in arctic or alpine regions on clay, silt, or rock, near creeks or seepage slopes growing with other bryophytes and vascular plants, or as pure sod.
DISTRIBUTION: Circumpolar, extending south in the alpine regions of the Rocky Mountains. Reported from arctic Europe, Asia, Iceland and Greenland (Map fig. 71).
Philonotis fontana var. pumila is a distinct arctic- alpine variety recognized by its small size, strict, erect, and closely packed stems clothed with abundant tomentum and small sporophytes. This taxon, treated in the broad sense, has been variously recognized as P. fontana var. alpina, P. alpicola var., P. borealis, P. tomentella and all manner of combinations of these. Philonotis tomentella, sensu European workers, includes many of the small expressions of var. fontana. Fertile plants are easily placed in either var. fontana or var. pumila depending on their size and
± erect habit, while sterile specimens beyond the distributional limits of var. pumila should be included in the species.
90
Philonotis yezoana Besch. et Card, in Card., Bull. Soc.
Bot. Geneve ser. 2, 1:123. 1909. TYPE: "Japon: Mori
(n. 3505, ser. I); Kominato (n. 42, ser. I); Aomori (n. 406, ser. I; Kinashi, n. 3, 8, 46, 52); Riishiri (n. 665);
Hokodate (n. 1901, 2738); Osorezan (n. 2123); Ikaregaseki
(n. 2669). Coree: Ouen-San (n. 591)," (not seen). Fig. 63.
Plants 1.0-3.0 cm high, forming loose dark green tufts, strongly brown tomentose below, occasionally producing elongate deciduous propaguliferous shoots.
Leaves 0.8-1.2 mm long by 0.4-0.7 mm wide, appearing lax, concave, and somewhat appressed, broadly ovate- lanceolate, slightly acuminate; margins narrowly recurved near the base, serrate with single teeth or with low double teeth; costa stout, percurrent; upper cells elongate
(2-4:1) 6-9u wide, lamina cells larger, cuboidal, rhomboidal, or irregularly many sided (1-2:1) 8-14u wide, some leaves possessing borders 2-6 cells wide of oblong or linear cells, all basal cells coarsely papillose over the middle of the lumen on both surfaces, less strongly on the abaxial surface, becoming progressively weaker toward the apex.
Dioicous, perigonia discoid (forming a rosette).
Seta 2.5-4.0 cm long; capsule spherical, 2.0-3.0 mm, stomata approximately 10 0, restricted to the capsule base; peristome as for the genus; spores 19-24y, papillose. All of the North American material examined was sterile, therefore this description was made from Japanese plants.
CHROMOSOME NUMBER: Not reported. 91
HABITAT: Over rock in shaded stream canyons, cliffs, or steep slopes that are wet by seepage water.
DISTRIBUTION: Apparently rare, Vancouver Island,
British Columbia mainland, California, Montana, Vermont,
Washington, and Japan (Map fig. 72).
These citations represent the first report of
P. yezoana from North America. Due to the dependence on u leaf shape and other variable characters for identification these few collections have been previously misinterpreted.
I have examined Japanese specimens and they all compare well to the North American material, not only in the characters mentioned in the key but also in the plants stature, dark green color, widely spaced stems forming loose turfs, and abundant tomentum.
This rare species is distinguished by having papillae exactly in the middle on almost all of the shortly rectangular or irregularly cuboidal cells, .and by having a border of two to six cells that are two to three times as long as the lamina cells.
Ochi (1962) has combined this taxon from Japan (along with several misidentified specimens of P. fontana from
North America) with P. fontana var. seriata (Mitt.) Brid.
I have examined the type specimen of P. seriata Mitt, in NY, and when considering the stability of papilla position, marginal cell shape, and the presence of a border of elongate cells, it is clear that Ochi's concept of P. fontana var. seriata is too broad and that P. yezoana deserves specific status. 92
Figure 63. Philonotis yezoana 93
IX, PHYTOGEOGRAPHY OF PHILONOTIS
The nine North American taxa of Philonotis display several noteworthy phytogeographic patterns. Those taxa with gemmiform perigonia (section Philonotula) are endemic to the Americas and reach their northern limit of distribution in the southeastern United States. There is a great diversity of species, including a considerable number of endemics (44), in Central and South America, and in the
West Indies. Philonotis longiseta, glauoescens, and sphaerooarpa represent northern extensions from this center of species diversification.
Those taxa with discoid perigonia (section Philonotis) are Holarctic. Philonotis fontana is the most widespread of all the North American species, and is distributed almost throughout the Holarctic. Philonotis fontana var. pumila occurs in the arctic and extends south only in alpine regions.
Philonotis fontana var. americana is the only endemic taxon in North America, and is distributed throughout the Rocky and Coastal Mountains. Philonotis oapillaris and P. marchica occur sporadically throughout Europe and adjacent regions while in North America each shows a distinct distribution pattern; P. oapillaris is restricted to the Pacific Coast and
P. marchica is widespread in eastern North America and the southern Rocky Mountains. Philonotis yezoana is apparently rare in North America and is disjunct from Japan.
It is difficult to say exactly where the North American taxa of the section Philonotis originated. However, the largest number of taxa occur in Africa and southern Asia
(approximately 40 species) which may indicate another center of species diversification and the subsequent migration to
North America.
The affinities of the North American species of
Philonotis are with those of Eurasia (P. fontana, marchica, and capillaris), the West Indies, Central and South America
(P. longiseta, glaucescens, and sphaerocarpa) , and Japan
(P. yezoana) . 95 96 97 98
' >fO i-j IM cO ao 30
Map fig. 67 Philonotis marchioa (Hedw.) Brid. 99 100 101 102
Map fig. 71 Philonotis fontana var. pumila (Turn.) Brid. 103 104
XI. EXCLUDED TAXA
Philonotis caloarea(B. S. G.) Schimp., Coroll. 86. 1856.
Grout (19 35) , Lesquereux & James (1884), and Jennings
(1951) have reported this species from North America with reservations about its positive identification. Holzinger issued an exsiccati collection, reported by Bailey (1926), as No. 414 of his Musoi Acrocarpi. This collection consists of robust plants of P. fontana var. fontana. All other collections identified as P. calcarea are referable to other
North American taxa and do not compare at all to authentic
European material.
Philonotis leiophylla Kindb. in Mac, Bull. Torrey Bot. Club
15:185. 1888. nom. nud.
Philonotis mohriana (C. Mull.) Jaeg., Ber. S. Gall. Naturw.
Ges. 1873-74:90. 1875.
P. mohvii C. Mull, ex Kindb., Enum. Bryin. Exot. 70.
1888.
This plant, which belongs in the genus Bveutelia, was collected in Mexico and mistakenly credited to Louisiana as thoroughly explained by Steere (19 49).
Philonotis pumila Kindb., Eur. No. Amer. Bryin. 2:328. 1897.
TYPE: "Can. Ontario near Belleville, Macoun," (Lectotype S seen).
The probable type was seen from Kindberg's herbarium.
It is apparently a species of Oncophorus, and clearly not a species of Philonotis. 105
Philonotis seriata Mitt., Jour. Linn. Soc. Bot. Suppl. 1:63.
1859. TYPE: "In Helvetia, Schleicher. In Britannia in monte Ben-na-Bourd, W. Gardiner," (Holotype Gardiner NY seen).
P. fontana ssp. seriata (Mitt.) Dix., Stud. Handb. Brit.
Moss. 294. 1896.
P. fontana var. seriata (Mitt.) Kindb., Bih. K. Svensk.
Vet. Ak. Handl. 7(9):255. 1883.
There are no North American specimens of Philonotis that compare with the type of P. seriata. Many of the European and extra-European specimens also do not compare well with the type, which indicates that this problematic taxon needs further study on a world wide basis.
Philonotis wilsonii (B. S. G.) Mitt., Jour. Sinn. Soc. Bot.
7:153. 1863.
This little moss does occur in eastern North America but belongs in the genus Bartramidula.
Philonotis falcata (Hook.) Mitt., Jour. Linn. Soc. Bot. Suppl.
1:62. 1859.
Bartramia falcata Hook., Trans. Linn. Soc. London 9:317.
1808.
I have seen P. falcata from Japan, Taiwan, the
Philippines, and India. It is very similar to P. marchica as also suggested by Mitten (1859), Kabiersch (1937), and Ochi
(1962). The only characters separating the two species are the somewhat larger size of both the gametophyte and sporophyte, and the arrangement of leaves in vertical rows in
P. falcata. It is not always possible to distinguish small and sterile specimens except on the basis of geographic distribution. Philonotis falcata and P. marchica are probably vicarious if not conspecific. 107
XII. SYNONYMS
Bartramia
B. fontana = Philonotis fontana
B. fontana var. pumilum - P. fontana var. pumila
B. glauces cens = P. glauces cens
B. longiseta = P. longiseta
B. muehlenbergii - P. marchica
B. radicalis = P. longiseta
B. radicalis var. porteri = P. longiseta
B. tenella = P. glaucescens
B. uncinata = P. sphaerocarpa
Mnium
M. fontanum = P. fontana
M. marchicum = P. marchica
M. sphaericarpon = P. sphaerocarpa
Philonotis
P. acutiflora = P. fontana
P. acuti folia = P. fontana
P. adpressa = P. fontana
P. alpicola var. borealis = P. fontana var. pumila
P. alpicola var. tenuis = P. fontana
P. alpicola var. tomentella = P. fontana var. pumila
P. amerioana = P. fontana var. amerioana
P. amerioana var. gracilescens = P. fontana
P. amerioana var. torquata = P. fontana var. amerioana
P. amerioana f. dimorphophylla = P. fontana var. amerioana
P. amerioana f. Zaxa = P. fontana var. amerioana 10 8
P. arnellii = P. oapillaris
P. borealis = P. fontana var. pumila
P. brachy chypus = P. fontana
P. caespitosa = P. fontana
P. caespitosa var. adpressa = P. fontana
P. caespitosa var. ampliretis = P. fontana
P. caespitosa var. compacta Dism. = P. fontana
P. caespitosa var. compacta Jaeg. = P. fontana
P. caespitosa var. heterophylla = P. fontana
P. caespitosa var. Zaxa = P. fontana
P. calcarea f. Occidentalis = P. fontana
P. fallax = P. fontana
P. fontana var. adpressa =• P. fontana
P. fontana ssp. adpressa = P. fontana
P. fontana var. alpina = P. fontana var. pumila
P. fontana var. ampliretis = P. fontana
P. fontana var. aristinervis = P. fontana
P. fontana f. aristineruis = P. fontana
P. fontana var. borealis = P. fontana var. pumila
P. fontana var. brachyphylla = P. fontana
P. fontana var. caespitosa = P. fontana
P. fontana var. oapillaris = P. oapillaris
P. fontana var. columbiae = P. fontana
P. fontana var. compacta = P. fontana var. pumila
P. fontana f. dimorphophylla = P. fontana
P. fontana var. falcata = P. fontana
P. fontana f. flowersii = P. fontana 109
P. fontana var. heterophylla = P. fontana p. fontana var. laxa = P. fontana p. fontana var. laxa f. tenuis = P. fontana p. fontana var. miaroblasta = P. fontana p. fontana var. microthamnia = P. fontana p. fontana var. pumila f. dimorphophylla = P. fontana var.
pumila p. fontana var. pumila f. heterophylla = P. fontana var.
pumila p. fontana var. pumila f. longifolia = P. fontana var.
pumila p. fontana var. seriata f. oocidentalis = P. fontana p. fontana var. serrata = P. fontana p. fontana var. tomentella = P. fontana var. pumila p. fontana ssp. tomentella = P. fontana var. pumila p. fontana ssp. tomentella var. heterophylla = P. fontana
var. pumila
P. glabriuscula = P. marchica p. p. P. fontana
P. glaucescens var. brevifolia = P. glaucescens
P. glaucescens f. Zaxa = P. glauces cens
P. glaucescens var. terrestris = P. tyZ-auceseens
P. graoillima = P. glauces cens
P. longiseta var. polygama = P. longiseta
P. longiseta var. porteri = P. longiseta
P. longiseta var. propagulaecaulis = P. longiseta
P. longiseta f. propagulicaulis = P. longiseta
P. macounii = P. capillaris 110
P. macounii var. torquata = P. fontana
P. microcarpa = P. fontana
P. muehlenbergii = P. marchica
P. muehlenbergii var. tenella = P. glaucescens
P. radicalis = P. longiseta
P. seriata ssp. amerioana = P. fontana var. amerioana
P. sphaerocarpa ssp. tenella var. terrestr-is = P. glaucescens
P. sp ft a ere car-pa var. terrestris = P. sphaerocarpa
P. subcapillaris = P. marchica
P. tenella = P. glauces cens
P. tenella var. coloradensis = P. fontana
P. tomentella = P. fontana var. pumila
P. tomentella var. borealis = P. fontana var. pumila
P. tomentella f. brachycarpa = P. fontana
P. tomentella var. compacta = P. fontana var. pumila
P. torquata = P. fontana var. amerioana
P. uncinata = P. sphaerocarpa
P. uncinata var. glauces cens = P. glauces cens
P. uncinata var. graoillima = P. gZaueeseens
P. vancouveriensis = P. capillaris p. p. P. fontana Ill XIII, SUMMARY
As with many other hygrophytic plants, Philonotis exhibits a great degree of environmentally induced phenotypic polymorphism. This phenotypic plasticity has resulted in the description of many ecophenic modifications, which have been given formal taxonomic status. The present research was initiated to evaluate the taxonomic value of morphological characters of Philonotis.
The experimental procedure adopted was to compare plants from the field with each other before and after culturing in a uniform environment. Numerous collections from various types of habitats produced plants that were morphologically different in some characters. These plants were then cultured at the same time in the same environment. Careful comparisons of these plants revealed that some characters are easily modified by the environment while other characters are stable within a taxon regardless of the environment. Also, some plants appearing quite different in nature (from different habitats) appear identical after culturing in a uniform environment.
This research shows, for the. North American species of
Philonotis, that the position of papillae oh the leaf cells
(top, middle, or bottom), nature of the marginal cells
(singly or doubly serrate), shape of the leaf (triangular or ovate-lanceolate), and the shape of the leaf cells (quadrate to elongate-rectangular), are stable characters not modified by the environment and therefore can be vised to support a 112 valid classification. Leaf size, arrangement, and amount of falcation; the amount of curvature on the margin, leaf apex shape, costa length, and plication; presence or absence of rhizoids, propagula, or peristome cilia; and the shape and size of the perigonal bracts are all ecophenic characters too variable to have diagnostic value.
Using the available experimental data, 29 names recognized by Flowers in Grout (19 35) are shown to be based on ecophenic modification and may be reduced to synonymy.
After careful examination of type specimens and many extra-
North American specimens, seven taxa may be excluded from the
North American flora. One taxon (P. yezoana) not previously recognized outside Japan is newly reported from North America.
Most species of Philonotis are dioicous with a chromosome number of n = 6. Philonotis longiseta is monoicous, and the chromosome number of n = 12 is a new report.
After plotting the distribution of the. North American species it can be shown that P. glauoescens, longiseta, and sphaerooarpa are predominantly in the southeastern United
States, and have their closest affinities with taxa of the
West Indies, and Central and South America. In North America
P. oapillaris is restricted to the Pacific Coastal region while P. marchica is much more widespread; both species occur in Europe. Philonotis yezoana is rare in North America and also occurs in Japan. Philonotis fontana is very widespread in North America and throughout the Holarctic. Philonotis fontana var. americana is endemic to the Pacific Coastal and Rocky Mountains, while P. fontana var. pumila is arctic and alpine. 114 XIV. EXSICCATI AND SELECTED SPECIMENS EXAMINED
Philonotis glauoescens Exsiccati Examined
Grout, No. Amer. Musci Perf. 69 as P. uncinata (CAN, DPU,
MICH, NO, SMU, TENN, TEX, UC, US), some specimens contain
P. longiseta also, and some labels have inconsistent
corrections.
Renauld & Cardot, Musci Amer. Sept. 299 as P. tenella (CAN,
MICH).
Philonotis glauoes cens Selected Specimens Examined
U. S. A.
FLORIDA:
Alachua Co., Devils Millhopper, Zales 2067, 2070, 2082
(UBC), Griffin 1750 (FLAS, GRO, LAF), Breen 1267 (FLAS,
MICH, UT), McFarlin A36 58 as P. gracillima (CAN, FLAS,
FSU), Anderson & Crum 13288, 13292 (CAN), Gainesville,
Schornherst 2622 as P. gracillima (FSU).
Citrus Co., Crystal River, Schallert 1933 as P. gracillima
(CU) .
Gadsden Co., Chattahoochee, Schornherst 1931 as P.
gracillima (FSU).
Hardee Co., Wauchula, McFarlin 22 Feb. 1938 (UT).
Hernando Co., Brooksville, Zales 2041 (UBC).
Leon Co., Tallahassee, Pursell 300 MF48 (DUKE), Breen 440
(UT), Schornherst 1607 as P. graoillima (FSU).
Levy Co., Suwannee R. , Reese 734 (LAF), Manatee Springs St.
Pk., Breen 3041 (FSU, NY), 1267 (UT), 3040 (FSU), 115
Fannin Springs, Schornherst 1713 as P. gracillima (FSU).
Liberty Co., Rock Bluff, McFarlin 1713 (UT), Rock Springs,
Schallert M123 as P. sphaerooarpa (IA).
Madison Co., Withlacoochee R., Schornherst 1354 as
P. gracillima (FLAS).
Manatee Co., Manatee R., Grout 22 Jan. 1941 (IA), March
1926 (UT), Ellenton, McFarlin 573 as P. tenella (FSU,
MICH), Bradenton, Robinson 15 July 1953 as P. gracillima
(US) .
Marion Co., Juniper Springs, Conard 18 Feb. 1956 (IA),
Lake Weir, Underwood 211 (IA, NY), Rainbow Springs,
Breen 2874, 2879, 2898 (FSU), Silver Springs, Breen
3363A (FSU).
Orange Co., Apopka, Breen 3256 (FSU).
Pasco Co., Elfers, Breen 2950 (FSU).
Polk Co., Highlands City, Redfearn 431-50-55 (SMS)., Lake
Wales, McFarlin 96 as P. tenella (UT), Winterhaven,
McFarlin 205 (FSU).
Seminole Co., Sanford, Rapp 125 (TENN), 22 Feb. 1921 (FSU).
Suwannee Co., Ichatucknee Springs, Griffin 11574 as P.
gracillima (FLAS), Redfearn 176-55 (SMS).
GEORGIA:
Monroe Co., High Falls St. Pk., Lampton 2420, 2423 (LAF),
Stewart Co., Lumpkin, Grice 298-55 (SMS).
LOUISIANA:
Caldwell Parish, Columbia, Reese 9239 as P. gracillima
(LAF). 116
Iberia Parish, New Iberia, Langlois 1893 as P. tenella
(MICH).
MISSISSIPPI:
Wilkinson Co. , Woodville, Reese 11172 as P. graoillima (LAF).
TEXAS:
Anderson Co., Palestine, Whitehouse 22373 (SMU, UT).
Burnet Co., Granite Mt., McVaugh 7651 (MICH, TENN, UBC, US).
Comal Co., New Brumfels, Clover 24 Nov. 1933 as P. tenella
(MICH), Whitehouse 18103a, 26213 as P. graoillima (SMU).
Sabine Co., Milen, Reese 3790 (LAF). 117
Philonotis longiseta Exsiccati Examined
Grout, No. Amer. Musci Perf. 69 as P. uncinata, some
specimens contain P. glauoescens (CAN, DPU, IA, MICH, US).
Grout, No. Amer. Musci Perf. 264 as P. radicalis (DPU, IA,
LAF, MICH, SMU, TENN, TEX, UBC, UC, US, UT).
Hall, Amer. Plains Flora 1870 as B. radicalis (F).
Holzinger, Musci Aero. Bor.-Amer. 94 as P. radicalis (IA, L,
MICH, OP, UC, US, WIS).
Jermy, Fl. Texensis 379 as B. radicalis (MO, US).
Langlois, Fl. Ludoviciana 442 as B. radicalis (US).
Rapp, Florida Mosses 11 (OP).
Renauld & Cardot, Musci Amer. Sept. 54 as B, radicalis (NY).
Sullivant & Lesquereux, Musci Bor. Amer. 169 (ed. I 1856) as
B. radicalis (MICH, MO).
Sullivant & Lesquereux, Musci Bor. Amer. 254 (ed. II 1865) as
B. radicalis (CAN, MICH).
Philonotis longiseta Selected Specimens Examined
U. S. A.
ALABAMA:
Baldwin Co., Spanish Fort, Zales & Jamieson 1904, 1909 (UBC)
Conecuh Co., Evergreen, Webster & Wilbur 769 as P.
glauoescens (MICH, TENN, UBC).
Dale Co., Clayhatchee, Reese 2791 (LAF).
Geneva Co., Hartford, Breil 1564 as P. sphaerooarpa (FLAS).
Lee Co., Auburn, Earl & Baker 17 April 1897 as P. radicalis
(NY), Letterman 133 as B. radicalis (MO). 1X8
Marion Co., Winfield, Harvill 7138 as var. porteri (UT).
Mobile Co., Mobile, Mohr 5403 as P. radicalis (UT),
Bellingrath Gardens, Zales & Jamieson 1896, 1897 (UBC).
Tuscaloosa Co., Warrior R., Harper 3298 as P. radicalis
(DUKE, MO, NY, TENN, US).
ARKANSAS:
Faulkner Co., Cadron R., Hedfearn 25918 (SMS).
Garland Co., Crystal Springs, Demaree 21992 (TENN), Hot
Springs Nat. Pk., Demaree 22734 (US), Sharp 22734 (TENN).
Merian Co., Buffalo R., Redfearn 15246 (SMS).
Montgomery Co., Cedar Glade, Anderson 11761 (CAN, UBC).
Newton Co., Ponca, Redfearn 12569 (MO, SMS, US).
Saline Co., Owensville, Redfearn 24311 as P. graoillima
(MICH, SMS).
Scott Co., Waldron, Whitehouse 23357 (DUKE, SMU, US, UT).
Stone Co., Allison, Anderson 11673, 11675, 11693, 11698 (UT).
FLORIDA:
Alachua Co., Gainesville, Murill 3 March 1938 (DUKE, UT),
Zales, Jamieson S Griffin 2061-2065 (UBC).
Calhoun Co., Mill Creek, Wilson 3973 as P. graoillima
(FSU, NY).
Gadsden Co., Ochlocklonee R., Latina 52 (UBC).
Hernando Co., Brooksville, Zales 2047, 2053, 2055 (UBC).
Highlands Co., Highlands Hammock St. Pk., Griffin 11573
as P. glaucescens (FLAS).
Homes Co., Ponce de Leon, Zales & Jamieson 1918, 1920 (UBC).
Leon Co., Tallahassee, Sohornherst 1550 (FLAS, FSU, MICH), u 119
1016 (FSU, MICH), Zales & Jamieson 1931, 1939, 1940,
1941, 1948 (UBC).
Levy Co., Suwannee R., Schornherst 1291 as P. glauoescens
(MICH, US).
Liberty Co., Torreya St. Pk., Zales 1930 (UBC), Redfearn
192-55 as P. glauoescens (SMS).
Manatee Co., Manatee, McFarlin 433 as P. glauoescens (FSU).
Marion Co., Eureka Springs, Schornherst 2026 as P.
sphaerooarpa (FSU, MICH).
Polk Co., Bartons, McFarlin 25 (FSU, MICH, UT).
Seminole Co., Sanford, Rapp 125 (NY), Oviedo, Zales &
Jamieson 2020, 2024, 2027, 2028, 2030 (UBC).
Sumter Co., Wildwood, Robinson 13 July 1953 as P.
glauoescens (US).
Suwannee Co., Suwannee R., Schornherst 1390 (FLAS, FSU, MICH).
Wakulla Co., Newport, Schornherst 1852 as P. glauoescens
(NY) , 1853 (FSU) .
GEORGIA:
Carroll Co., Yellow Dirt Creek, Zales
Whooping Creek, Zales & Lampton 2133 (UBC).
Charlton Co., Folkston, Crosby 1868 (DUKE, MO).
Fulton Co., Palmetto, Zales 2109, 2110, 2111, 2114 (UBC).
Lowndes Co., Quarterman 3 (IA).
Monroe Co., High Falls St. Pk., Zales 2086 (UBC).
Rabun Co., Tallulah Falls, Small 222 (DUKE, UBC).
IOWA:
Allamakee Co., Union City, Savage 1 June 1899 (IA). '-. 120
Marion Co., Red Rock, Coriard 14 May 1949 (IA).
Muscatine Co., Wildcat Den, Savage 13 May 1899 (BRNM, IA,
MICH, UT), Sweetland Creek, Shimek 8 May 1903 (DUKE, IA,
MICH, TENN, US).
KANSAS:
Montgomery Co., Elk'City, Smith 303 (NY).
KENTUCKY:
McCreary Co., Cumberland Falls St. Pk., Norris 129A (TENN).
LOUISIANA:
Avoyelles Parish, Mansura, Pennebaker 22 March 1940 (DUKE,
UT) .
Calcasieu Parish, Lake Charles, Shimek 29 Feb. 1936 (BRNM,
DPU, DUKE, IA, MICH).
Evangeline Parish, Easton, Reese 2662, 2665 (LAF).
Iberia Parish, Weeks Island, Zales, Jamieson & Reese 1876,
1879, 1890, 1893, 1894 (UBC) .
Lafayette Parish, Lafayette, Leon 30 March 19 31 (UT),
Jan. 1931 (UBC), March 1933 (L, UBC), St. Martinsville,
Langlois March 1891 (US), Vermilion R., Zales 1861, 1863,
1866 (UBC).
Ouachita Parish, Luna, Reese 8377 as P. glauoescens (LAF).
Plaquemines Parish, Point a La Hache, Langlois 442 as
B. radicalis (US).
Rapides Parish, Zimmerman, MoWilliams <5 Davis 419 (LAF) .
Sabine Parish, Many, Reese 67 71 (LAF).
St. Landry Parish, Grand Coteau, Reese 7 867 (LAF).
St. Martin Parish, Duchamp, Langlois 15 March 1892 as 1.21
B. radicalis (US).
St. Mary Parish, Cote Blanche Is., Reese 7200 (LAF).
West Feliciana Parish, Grant's Bayou, Reese 10802 as
P. glauces cens (LAF).
Winn Parish, Atlanta, Reese 6904 (LAF).
MISSISSIPPI:
Adams Co., Natchez, Shimek June 1898 (IA).
Harrison Co., Wolf R. & Miss. Hwy. 53, Reese 3101 (LAF).
Oktibbeha Co., Noxubee Wildlife Refuge, Zales 1836 (UBC).
Perry Co., Ragland Hills, Rogers 3676 as P. uncinata (FLAS).
Wilkinson Co., Woodville, Reese 2557 (LAF).
MISSOURI:
Butler Co., Poplar Bluff, Russell 78 (MO).
Cedar Co., Stockton, Redfearn 12831 (SMS).
Dade Co., Sylvania, Redfearn 8206 (SMS).
Jefferson Co., Pacific, Redfearn 20728 (SMS).
St. Louis Co., Allenton, Nelson 2158, 1627 (DUKE, S-PA),
7646 (NY).
Vernon Co., Nevada, Hall 1870 as B. radicalis (F).
NORTH CAROLINA:
Alleghany Co., Roaring Gap, Schallert 9872 as P. rigida
(DUKE).
Haywood Co., Lake Junaluska, Anderson 294 (DUKE).
Jackson Co., E of Cherokee, Zales & Jamieson 2374 (UBC),
Whitewater Falls, Sharp 24 Oct. 1959 (TENN).
Jones Co., Trenton, Anderson 10792, 10794 as P. graoillima
(DUKE). 122
Lee Co., Sanford, Anderson 20,554 (DUKE).
McDowell Co., Woodlawn, Anderson & Jones 9671 (DUKE).
Sampson Co., Newton Grove, Conard 17 March 1953 as
P. sphaerooarpa (IA).
Swain Co., Cherokee, Whitehouse 26821 (UT), Bryson City
Anderson 819 (DUKE).
OHIO:
Hocking Co., Laurel Twp., Bartley & Pontius 412 (NY).
Jackson Co., McCune, Bartley 609 (US).
OKLAHOMA:
Greer Co., Granite, Koch 5192 (MICH).
LeFlore Co., Big Cedar, Redfearn 19786 (SMS), Ponteau,
Redfearn 19592 (SMS), Short Mt., Redfearn 24723 (SMS).
McCurtain Co., McCurtain Co. Game Preserve, R.edfearn
18720 (SMS).
PENNSYLVANIA:
LeHigh Co., Rockdale, Wolle Dec. 1874 (F) .
Wyoming Co., Tunkhunnock, Glowenke 856 (IA).
SOUTH CAROLINA:
Charleston Co., Charleston, Smith April 1877 (US).
Marion Co., Rains, Wonderly 867 (IA).
Oconee Co., Sumter Nat. Forest, Wagner 9 June 1949 (TENN).
Richland Co., Columbia, McCorkle B-98 (IA), 17 April 1950
(UT) .
TENNESSEE:
Blount Co., Abrams Falls, Sharp 341040 as f. polygama
(TENN, UT) , Kinzel Springs, Sharp 34598 (F, MICH, TENN), 123
Rich Mt., 24 June 1934 (FSU).
DeKalb Co., -Caney Fork Creek, Sharp 51T14 (UT).
Monroe Co., Tellico R., Sharp 1 May 1932 (DPU).
Polk Co., Hiwassee R., Sharp 4911 (DPU, DUKE, MICH, TENN,
UBC) .
Sevier Co., Tremont, Grout & Sharp 2643 264a (LAF, UBC,
US) .
TEXAS:
Bastrop Co., Bastrop, McAllister 19 April 19 36 (DUKE, SMU,
TEX) .
Brazos Co., White Creek, Parks 15 March 1947 (F, IA).
Cherokee Co., Rusk, Whitehouse 22919a (SMU).
Denton Co., Grapevine, Whitehouse 17918, 17943 (SMU, US).
Gillespie Co., Grape Creek, Jermy 379 as B. radicalis
(MO, US).
Harrison Co., Caddo Lake St. Pk. , Whitehouse 27988 (SMU, US).
Llano Co., Fredricksburg, McAllister 4 June 1919 (SMU, TEX),
Austin, McAllister April 1938 (SMU, UT).
Rusk Co., Mt. Enterprise, Whitehouse 20821a (SMU).
San Jacinto Co., Pointblank, Reese 3597 (LAF, US).
Tarrant Co., Hurst, Whitehouse 26292 (DUKE, SMU, US),
26531, 26538 (SMU).
VIRGINIA:
Carroll Co., Fancy Gap, Gleason 47195 (MICH).
Prince Edwards Co., Smith April 1878 (US).
WEST VIRGINIA:
Raleigh Co., Whitesville, Reed 74998 (US). 124
Philonotis sphaerocarpa Exsiccati Examined
Holzinger, Musci Aero. Bor.-Amer. 415 as P. uncinata (L, MICH,
MO, UBC, UC, US, WIS).
Husnot, PI. Antilles 147 as P. uncinata (L, WIS).
Philonotis sphaerocarpa Selected Specimens Examined
U. S. A.
FLORIDA:
Seminole Co., Sanford, Rapp. 128 as P. uncinata (OP). 125
Philonotis mavchiaa Exsiccati Examined
Austin, Musci Appal. 223 p. p. as B. fontana (CAN).
Austin, Musci Appal. 225 as Bartramia, some specimens
contain P. fontana (CAN, MICH, UBC, US, WIS).
Holzinger, Musci Aero. Bor.-Amer. 471 as P. graoillima (CAN,
L, MICH, MO, NY, OP, UBC, UC, US, WIS).
Holzinger, Musci Aero. Bor.-Amer. et Eur. 543 as P. capillaris
'(CAN, L, MICH, MO, NY, OP, UBC, UC, US, WIS).
Macoun, Can. Mosses 173 (CAN) .
Sullivant & Lesquereux, Musci Bor.-Amer. 168 (ed. I 1856)
as B. muehlenbergii (CAN, MICH, MO, WIS).
Sullivant & Lesquereux, Musci Bor.-Amer. 253 (ed. II 1865)
as B. muehlenbergii (CAN, MICH).
Philonotis marchica Selected Specimens Examined
CANADA
ALBERTA:
Banff Nat. Pk., E Entrance, Crum & Schofield 5331 as
P. fontana (CAN).
Ghost Ranger Station, Bird & Glenn 11299a as P. fontana
var. falcata (UAC).
BRITISH COLUMBIA:
Halcyon Hot Springs, MacFadden 4056, 19994 (CAN), 392 (UBC).
NEW BRUNSWICK:
Bass River, Fowler 1873 (L).
NEWFOUNDLAND:
Cow Head, St. Barbe, Tuomikoski 5021 (H). 126
NOVA SCOTIA:
Anapolis Co., Kejimkujik Nat. Pk., Ireland 12511 (UBC).
Inverness Co., Margaree Forks, Lamb 7012 as P. caespitosa
(DPU, UBC) .
ONTARIO:
Bruce Co., Sauble Beach, Cain 1325 (UBC, UT).
Carleton Co., Macoun 7 July 1900 as P. fontana (CAN).
Elgin Co., Sparta, James 35 as P. oapillaris (CAN).
Essex Co., Leamington, Macoun 173 (CAN, MICH, US), 351 (CAN).
Grey Co., Owen Sound, Crum 11614 as P. muehlenbergii (CAN).
Hastings Co., Belleville, Macoun 5 June 1865 (CAN).
Norfolk Co., Turkey Point, Cain 1330 (DUKE, IA, UT).
Northumberland Co., Macoun 22 June 1864 as P. calcarea (CAN).
Ontario Co., Lake Couchiching, Hand 818 (CAN, DUKE).
Parry Sound Dist., Sundridge, White 249 as P. fontana (CAN).
Wellington Co., Elora, Moxley 7 June 1933 (DUKE).
QUEBEC:
Gaspe, Fabius 4490 as P. oapillaris (DUKE, NY).
U. S. A.
ALABAMA:
DeKalb Co., DeSoto St. Pk., Conard 5-8 June 1957 as
P. sphaericarpa (IA).
ARIZONA:
Apache Co., Three Forks, Phillips & Mason 1572 (CAN, UT),
as P. muehlenbergii (CAN).
Cochise Co., Huachuca, Gooding 10329t 10345, 10030 (UBC,
UT) . 127
Coconino Co., Oak Creek Canyon, Johnson 673 (UT), Marble
Canyon, Clover & Jotter 2249 (CAN, MICH).
Pima Co., Sycamore Canyon, Haskell 11 March 1945 as
P. muehlenbergii (UT).
Santa Cruz Co., White House Canyon, Bartram 101 as
P. capillaris (CAN, DUKE, OP, US), Sycamore Canyon,
Haring 3325 as P. muehlenbergii (UT).
ARKANSAS:
Boone Co., Harmony, Redfearn 25678 as P. muehlenbergii (SMS).
Crawford Co., Natural Bridge, Redfearn 169 81 as
P. muehlenbergii (SMS).
Fulton Co., Mammoth Springs, Redfearn 15621 (SMS).
Garland Co., Hot Springs, Scully 1101 (DUKE, IA, UT).
Newton Co., Jasper, Anderson 11986 as P. muehlenbergii
(DUKE), Ponca, Redfearn 12918 (SMS).
Poinsette Co., Marked Tree, Demaree 17545 (US).
Stone Co., Allison, Anderson 11616 as P.• muehlenbergii
(DUKE, SMS).
COLORADO:
Gunnison Co., Gothic, Bunoe 92 as P. muehlenbergii (US).
LaPlata Co., Hesperus, Pursell 3211 (CAN, DUKE, US).
Summit Co., Tenmile Creek, Zales 3023 (UBC).
FLORIDA:
Jefferson Co., Monticello, Lightipe 117 (US).
GEORGIA:
Crisp Co., Cordele, Lamp ton 2278a as P. graoillima (LAF).
Walker Co., Cloudland Canyon St. Pk., Anderson 12,973 as 128
P. uncinata (DUKE).
IDAHO:
Elmore Co., Atlanta, MaoFadden 19317 (CAN, DUKE, NY, UT).
ILLINOIS:
Lake Co., Lake Bluff, Hill 22 Nov. 1904, 22 June 1905 (MICH).
Menard Co., Wolf (US).
INDIANA:
Jasper Co., Remington, Welch 10116 (DPU).
Jefferson Co., Hanover, Wolle 56 (DPU).
Monroe Co., Bloomington, Welch 17420 (DPU).
Putnam Co., Limedale, Baylor Spring 1904 as P. fontana (UC),
Greencastle, Welch 10115 (DPU, MICH).
Warren Co., Little Pine Creek, Ireland 5401 as P. gracillima
(DPU, US) .
IOWA:
Allamakee Co., Quandahl, Welch 17270 as P. longiseta (DPU).
Delaware Co., Delhi, Kurth Aug. 1946 (IA, SMU).
Dubuque Co., Pine Hollow, Cavanagh 7290 (WJC).
Emmet Co., Wellingford, Conard 7 Aug. 1946 (IA).
VanBuren Co., Lucey-Keosauqua St. Pk., Conard 15 Oct. 1932
as P. longiseta (IA).
Winneshisk Co., Bluffton, Conard 11 June 1949 (IA).
KANSAS:
Crawford Co., Farlington Lake, Gier 2376 (IA as P. longiseta,
SMU) .
Montgomery Co., Elk City, McGregar 2388 (IA). 129
KENTUCKY:
Edmundsen Co., Brownsville, Price 28 as P. fontana (NY).
Powell Co., Natural Bridge St. Pk., Conard 3 Sept. 1941 (IA) .
LOUISIANA:
West Feliciana Parish, Plettenberg, Reese 10486, 10491 as
P. uncinata (LAF).
MAINE:
Cumberland Co., Peak's Is., Taylor 2875 (MICH).
MARYLAND:
Calvert Co., Scientist's Cliff, Chase 8412 as P. fontana
(CAN).
Garrett Co., Swallow Falls St. Pk., Worthley 27 Sept. 1964
(US) .
MICHIGAN:
Algar Co., Munsing, Conard 29 Aug. 1939 (IA).
Barry Co., Lawrence Lake, Redfearn 22213 (MICH, SMS).
Cheboygan Co., E of Burt Lake, Nichols 1920 (CAN, MICH).
Chippewa Co., Scotty Bay Creek, no coll. cited, 30 July
.1920 (MICH) , Upper Tahquamenon Falls, Shetler 28 July
1958 (US).
MINNESOTA:
Blue Earth Co., Leiberg 393 as P. fontana (US).
Hubbard Co., LaSalle Springs, Stein 58163 as P. muehlenbergii
(UBC) .
MISSISSIPPI:
Clarke Co., Dunn Falls, Rogers 5762 as P. glaucescens (LAF).
Clay Co., Hwy. 50 & Tombigbee R., Zales 1846 (UBC). 130
MISSOURI:
Barry Co., Seligman, Redfearn 10269 (SMS).
Benton Co., Avery, Redfearn 6922 as P. muehlenbergii (SMS).
Boone Co., Ashland, Drew 3734 as P. oapillaris (MICH).
Cedar Co., Bear Creek, Redfearn 12861 as P.
muehlenbergii (SMS).
Dade Co., Greenfield, Redfearn 6422 as P. muehlenbergii
(SMS), Bona, Redfearn 7484, 6835 as P. muehlenbergii (SMS).
Dallus Co., Celt, Redfearn 12788 (SMS).
Douglas Co., Jackson's Mill, Redfearn 8571 as
P. muehlenbergii (SMS).
Laclede Co., Sleeper, Redfearn 8262 (SMS).
Madison Co., Minela Motte, Redfearn 159 58 as P. longiseta
(CAN, DUKE, SMS).
Newton Co., Boulder City, Redfearn 6676 (SMS).
Phelps Co., James, Redfearn 7165 as P. muehlenbergii (SMS).
Reynolds Co., Sec. 25, Gier 4609 as P. muehlenbergii (NY).
Ripley Co., Buffalo Creek, Redfearn 6058 as
P. muehlenbergii (SMS).
Stoddard Co., Wapparello, Anderson 12567 as P. muehlenbergii
(CAN, DUKE).
Taney Co., Ozark Dam, Gier 462 as P. muehlenbergii (NY).
Texas Co., Arroll, Redfearn 9 517 as P. muehlenbergii (SMS),
Cabool, Redfearn 5182 as P. muehlenbergii (SMS).
NEBRASKA:
Cherry Co., Bog Valley, Walker 6 as P. fontana (DUKE).
Thomas Co., Seneca, Kiener 20692 (CAN, MICH). 131
NEW MEXICO:
San Miguel Co., Las Vegas, Arsene 18714 (US).
NEW YORK:
Fulton Co., Canada Lake, Ketchledge 555 (MICH, UBC, UT),
Stoner Lake, Ketohledge 561 (MICH, UBC).
St. Lawrence Co., Massena, Andrews 15 Sept. 1948 as
P. capillaris (CU).
Sullivan Co., Willowemoc, Smith 37214 as P. fontana (MICH).
Wayne Co., Wolcott, Cook July 1887 (NY).
NORTH CAROLINA:
Forsyth Co., Winston Salem, Chapman 2911 as P. graoillima
(DUKE).
Franklin Co., Riley, Beaven 677 as P. graoillima (DUKE),
Weldon's Mill, Beaven 563 as P. graoillima (DUKE).
Haywood Co., Lake Junaluska, Watson 16 June 1928 as
P. fontana (DUKE).
Jackson Co., Whitewater R., Anderson 8669 as P. graoillima
(DUKE).
Macon Co., Coweeta, Webster 25 Nov. 1969 as
P. muehlenbergii (TENN), Cullasaja Gorge, Schofield
9898 as P. graoillima (CAN, DUKE, UBC).
Watauga Co., Blowing Rock, Zander 419 as P. graoillima
(DUKE), Boon, Zales 2454 (UBC).
OHIO:
Athens Co., Athens, Robinson & Hall 25 May 1953 as
P. longiseta (IA, US).
Clark Co., Springfield, Biddlecome 26 Sept. 1877 as 132
B. radicalis var. porteri (US).
Lake Co., Kirtland, Shipman 24 Oct. 1920 (NY).
OKLAHOMA:
Canadian Co., Devils Canyon, Little 13 as P. fontana (TENN).
Cherokee Co., McSpadden Falls, Bird 3227 (UT).
Murray Co., Turner Falls, Ikenberry 2080 as
P. muehlenbergii (IA), Davis, Emig Aug. 1917 as
P. calcarea (NY).
Washington Co., Springdale, Anderson 12287 (CAN).
PENNSYLVANIA:
Allegheny Co., Douthitt, Jennings 26 April 1908 as
P. fontana (MO, NY).
Northampton Co. , Bethlehem, -Rau 17 May 19 47 as
P. muehlenbergii (TENN), as P. fontana (NY).
SOUTH CAROLINA:
Lancaster Co., Taxahaw, Huntley 156 (DUKE).
TENNESSEE:
Davidson Co., Donelson, Clebsch 14595 as P. longiseta f.
propagulicalis (TENN).
Greene Co., Tusculum, Byers 10 Ap ril 1949 as P. fontana (NY).
Montgomery Co., Clarksville, Clebsch 808 (IA).
Sevier Co., Laurel Falls, Zales 210 (UBC).
TEXAS:
Burnet Co., Burnet, Whitehouse 18502 as P. longiseta (SMU).
Dallas Co., Dallas, Whitehouse 26302 (SMU, US).
Gillespie Co., Palo Alto, Jermy 847 (MO).
Mason Co., Mason, Harvill 2902 as P. longiseta (UT).
Nacogdoches Co., Nacogdoches, Arnwere 24- (IA). 133
San Seba Co., Gorman Falls, Harvill 2915 as P. longiseta
(UT) .
UTAH:
Kane Co., Escalante Ft. , Flowers 5624 (UT) , Kanab Canyon,
Flowers 3248 (UT).
Salt Lake Co., Brighton, Flowers 572 (UT).
San Juan Co., Temple of Music, Flowers 4822 (UT).
VERMONT:
Winsor Co., Norwich, Hutchinson 18 Oct. 1949 (DUKE, IA, SMU).
VIRGINIA:
Albemarle Co., Jones Run, Hermann 15173 as P. graoillima
(CAN, DUKE, US).
Botetourt Co., Hollins, Patterson 3 July 1943 as
P. muehlenbergii (UT).
Greene Co., South R. Falls, Hermann 15132 as P. graoillima
(US) , Ireland 3855 as P. glauces cens (US) .
Isle of Wight Co., Burwell's Bay, Correll 11487 (DUKE).
James City Co., Williamsburg, Mukula 4647 (NY).
Madison Co., Shenandoah Nat. Pk., Flowers 6426 (UT).
Smyth Co., East Marion, Small 18 May 1892 as P. fontana (NY).
WEST VIRGINIA:
Mineral Co., Elk Garden, Ammons 1460 as P. muehlenbergii (UT).
Mingo Co., Myrtle, Reed 70533 as P. muehlenbergii (US).
Summers Co., Hinton, Morris 966 (NY, US, UT).
WISCONSIN:
Ashland Co., Copper Falls St. Pk., Hulbary 853 as
P. longiseta (IA). Grant Co., Potosi, Cheney 11336 as P. fontana (WIS)
WYOMING:
Park Co., Clark Fork, Whitehouse 27497 (SMU). 135
Philonotis capillaris Exsiccati Examined
Holzinger, Musci Aero. Bor.-Amer. 440 (CAN, H, L, MICH, MO,
NY, UBC, UC, US, WIS).
Macoun, Can. Musci 152 as P. macounii (CAN, MICH, MO, NY,
UAC, UBC, UC, US, WIS).
Macoun, Can. Musci 416 as P. longiseta (MANCH).
Renauld & Cardot, Musci Amer. Sept. 29 8 as P. macounii
(CAN, MICH, NY).
Philonotis opaillaris Selected Specimens Examined
CANADA
BRITISH COLUMBIA:
Brittania Beach, Schofield 12839 (UBC).
Chilliwack, base of Mt. Cheam Zales & Schofield 1235 (UBC).
Harrison Lake, Zales 2696 (UBC).
Hope, Schofield 1669 8 (DUKE, UBC).
Lund, Schofield & Boas 18157 (CAN, DUKE, UBC).
Prince Rupert, Schofield & Boas 21490 (CAN, UBC).
Skeena River, Schofield & Boas 20976 (UBC).
Slocan City, MacFadden 9 June 192 7 (MO, UBC, UC).
Taft, Schofield 29564 (UBC).
Vancouver Island, Nootka Sound, Schofield & Halbert 40207
(UBC) .
Queen Charlotte Islands, Graham Is., Skidegate Channel,
Schofield 14001 (UBC), Moresby Is., Schofield 25266 (UBC).
U. S. A.
ALASKA:
Juneau, Canby 487 (MO, NY, US). 136
Gold Creek Canyon, Cooley 31 July 1891 (NY).
Kodiak, Trelease 1841 (MO, NY).
Unalaska, Setchell 10 (UC).
CALIFORNIA:
Humboldt Co., Eel Rock, Koch 80 (MICH), Mad R., Schofield
28652 (UBC).
Mendocino Co., Comptche, Zales 2899 (UBC), Fort Bragg,
Zales 2893 (UBC), Leggett, Zales 2880, 2887 (UBC),
Rockport, Schofield & Thomas 28 883 (CAN, DUKE, UBC).
San Mateo Co., Lake San Andreas, Howe 136 (NY, UC, US).
Santa Cruz Co., Big Basin Redwoods, Koch 2055 (MICH).
Sonoma Co., Cazadero, Howe 14 March 1896 as P. fontana
(NY) .
Trinity Co., Salyer, Tracy 16839 (UBC, UC).
IDAHO:
Kootenai Co., Little North Fork, Leiberg 172 (US),
Tourlle R., Leiberg 96 (NY).
OREGON:
Lane Co., Leaburg, Leiberg 1697 (DUKE, US), Mapleton,
Leiberg 1892 (US).
Tillamook Co., Siuslaw Nat. Forest, Zales 2821 (UBC).
WASHINGTON:
King Co., North Bend, Ireland 5843 (CAN).
Snohomish Co., Darrington, Schofield & Hermann 21828 (CAN,
UBC), Gold Basin, Eyerdam 3007 (MICH, UBC), Mt. Pilchuck,
Ireland 5590 (CAN), Stillaguamish R., Schofield, Ireland
& Boas 18458 (CAN, UBC). 137
Philonotis fontana var. fontana Exsiccati Examined
Austin, Musci Appal. 223 as Bartramia p. p. P. marchica
(CAN, MICH, NY, UBC, US, WIS), 224 as Bartramia (CAN, MICH,
NY, UBC, US, WIS), 226 as Bartramia marchica (MICH, UBC,
US, WIS).
Backer, Pacific Slope Bryo. 1586 (DUKE, MO, NY, UC, US, WIS),
766 (DUKE, MO, NY, UBC, UC, US, WIS).
Bryhn, Mosses W. Minn. 22-26 June 1901 as P. alpicola (DUKE,
MO, NY, US).
Flowers, Mosses of Utah 8 (SMU).
Frye, Moss Exs. 88 (CAN, MICH, NY, SMU, UBC).
Grout, No. Amer. Musci Perf. 13 (CAN, MICH, NY, TEX, UC, US),
13c (CAN, DPU, LAF, MICH, NY, SMS, SMU, UBC).
Grout, Hand-lens Mosses 46 (NY).
Holzinger, Mosses N,-E. Minn. 15 Aug. 1902 as P. fontana var.
caespitosa (BRNM, DUKE, NY, OP).
Holzinger, Musci Aero. Bor-Amer. 70 (NY, OP, US), 316 as var.
caespitosa (BRNM, CAN, L, MICH, MO, NY, OP, UC, US, WIS),
369 as var. seriata (CAN, L, MICH, MO, NY, UBC, UC, US, WIS),
414 as P. oalcarea (CU, L, MICH, MO, UBC, UC, US, WIS),
439 as P. amerioana (CAN, CU, L, MICH, MO, NY, UBC, UC, US,
WIS), 441 as P. tomentella (CAN, MO, US), 441 as
P. alpicola (CU, L, MICH, NY, UC, WIS), 517 as P. fallax
(CAN, L, MO, NY, OP, UC, US, WIS).
Holzinger, Musci Aero. Bor.-Amer. et Eur. 661 (CAN, L, MO, NY,
UBC, UC, US, WIS).
Leiberg, Musci Leibergiana 49 (CAN, DUKE, MICH, NY, UC, US,
WIS) . 138
Macoun, Can. Mosses 170a as 'P. alpicola (CAN), 153 (UC, US).
Nelson & Macbride, PI. Idaho 609 (MO, NY, US).
Nelson & Holzinger, Pi. Yellowstone Nat. Pk. 6062 (GRO, MO,
NY, UC, US), 6110 as var. alpina (GRO, MO, NY, UC, US).
Nelson & Holzinger, Pi. Wyo. 1955 (MO, NY, UC, US), 6381
(MO, NY, US), 7718 (NY), 8770 (CU, MO, NY, UC, US).
Sandberg, PI. Idaho 1115 (NY).
Shaw, Selkirk Fl. 983 as P. marchica (IA, L, MO).
Small, Mosses So. U. S. 37 (CAN, DUKE, MO, NY, UBC, UC, US,
WIS) .
Sullivant, Musci Allegh. 123 as Bartramia (L, MICH, NY, US).
Sullivant & Lesquereux, Musci Bor.-Amer. (ed. I 1856) 165 as
Bartramia calcarea (MICH, MO, NY, WIS), 166 as Bartramia
(MICH, MO, NY, WIS), 167 as Bartramia (MICH, MO, NY, WIS).
Sullivant & Lesquereux, Musci Bor.-Amer. (ed. II 1865) 250 as
Bartramia calcarea (CU, MICH, NY, WIS), 251 as Bartramia
(CAN, MICH, NY, WIS), 252 as Bartramia (CAN, MICH, NY, WIS).
Philonotis fontana var. fontana Selected Specimens Examined
CANADA
ALBERTA:
Banff Nat. Pk., Kicking Horse Lake, Macoun 20 July 1885
(CAN), Lake Louise, Porsild & Breitung 17822 (CAN, DUKE).
Cypress Hills Prov. Pk., Elkwater Bird 4454 (CAN, DUKE, NY,
UAC) .
Jasper Nat. Pk., Jasper, Crum 3657 (CAN, MICH).
Waterton Lakes Nat. Pk., Goat Lake, Bird & Lakusta 16561
(CAN, UAC). BRITISH COLUMBIA:
Barkerville, Schofield 39097 (CAN, DUKE).
Brackendale, Macoun 193 (CAN).
Cheam View, Fraser R., Schofield & Boas 17667 (CAN, TENN,
UAC, US).
Eagle Pass, Macoun 121 (CAN).
Encampment, Svihla 42271 as var. pumila (CAN).
Galiano Island, Salamanco Point, Schofield & Boas 17470
(CAN).
Haines Hwy., Crum & Schofield 9194, 9213, 9615 (CAN).
Hope, Williams 300 (CAN).
Kootenay Lake, Lockheart Beach, Band 57-124 (CAN, DUKE).
Liard Hot Springs, Porsild 22147, 22189 (CAN).
Lillooet, Noaxe Lake, Dill 1 Aug. 1961 (CAN, DUKE).
Manning Prov. Pk., Allison Pass, Schofield 24764 (CAN).
Macleod's Lake, Macoun 243 (CAN).
New Denver, MacFadden 40 4 3 (CAN).
New Westminster, Macoun 6 89 (CAN).
Nicola R., Cougar Lake, Brinkman 10 July 1910 (CAN).
Quesnel, Macoun 29 May 1875 (CAN).
Rogers Pass, Selkirk Mts., Macoun 174 (CAN).
Selkirk Mts., Armstrong, Wilson 70b (CAN).
Slocan City, MacFadden 4054 (CAN).
Vancouver Island, Elk Lake, Macoun 90 (CAN), Mt. Benson,
Macoun 184 (CAN), Shawnigan Lake, Macoun 170a as
P. alpicola (CAN), Victoria, Macoun 140 (CAN).
Wells Gray Prov. Pk., Battle Mt., Ahti 13305, 14254 (CAN). 140
Whistler Mt., Schofield 37897 (CAN), Zales 1199, 2614 (UBC).
Yoho Prov. Pk., Hector, Crum & Schofield 4499 as var.
falcata (CAN, MICH, UBC).
MANITOBA:
Gillam, Kettle Rapids, Crum 7543 (CAN, UBC).
Herb Lake, Wekusko Lake, Scoggan 104 (CAN, OP, SMU).
Spruce Woods Prov. Pk., Melbourne, Bird 585 8 (CAN, MICH,
NY, UAC).
York Factory, Lower Hayes R., Ritchie 4229 (CAN).
NEW BRUNSWICK:
Albert Co., Fundy Nat. Pk., Ireland 11599 (DUKE, MICH,
NFLD, UAC).
Kent Co., Bass River, Fowler 18 June 1872 (L).
Victoria Co., Grand Falls, Habeeb 939 (MICH).
NEWFOUNDLAND:
Avalon Peninsula, Colinet, Smith 427 (CAN).
Battle Harbour, Waghorne 30 Aug. 1891 (CAN).
Bonne Bay, Elkington Bill (NY).
Cape Caribou, Grand Lake, Kallio 18 July 1963 (CAN).
Churchill Falls, Brassard 5032, 5238, 5487 as P. tomentella
(NFLD) .
Hampden, Tuomikoski 3089 (H).
LaPoile, Tuomikoski 2066 (H).
• Lejeune Lake & Abel Lake (between), Harper 3388 (CAN, US).
Main Brook, Damman TD6027-4 (CAN).
NOVA SCOTIA:
Colchester Co., Falleigh Lake, Prince 6098 (MICH, OP). 141
Inverness Co., Cape Breton Highlands Nat. Pk., Ireland
11868 (DUKE, MICH, NFLD, UAC, UBC).
Kings Co., Black Hole, Schofield 10954 (NY).
Victoria Co., Indian Brook, Schofield 6009 (NY, US).
ONTARIO:
Algoma Dist., Spanish Mills, White 242 (CAN), Montreal
R., Lerstern 16 July 1960 (FLAS).
Bruce Co., Hay Bay, Crum 11395 (CAN, MICH), Lions Head,
Hand & Moxley 137 (CAN).
Cockburn Island, Grassl 69 70 (MICH).
Gray Co., Markdale, Crum 11254 (CAN, MICH).
Kenora Dist., Nextor Falls, Hand 1159 (CAN).
Muskoka Dist., Coopers Falls, Cain 1326 as P. caespitosa
(IA, MICH, UAC).
Niagra Falls, Drummonds Island, 15 May 1901 (CAN).
Owen Sound, West Hill, Moxley July 1926 (CAN).
Parry Sound Dist., South R., Hand 843 (CAN).
Peel Co., Orangeville, Cain 4956 (CAN, NO).
Renfrew Co., Deep River, Crum 1 June 1955 (CAN).
Simeoe Co., Sparrow Lake, Cain 1327 as P. amerioana (DUKE,
IA, MICH, NO, NY, TENN, UAC, UBC).
Thunder Day Dist., Black Fox Lake, Brodo 13806 (CAN, MICH),
Ouimet Canyon, Garton 4477 (CAN), Nipigon, Macoun
26 June 1884 (CAN).
PRINCE EDWARD ISLAND:
Prince Co., Richmond, Ireland 10320 as P. caespitosa
(MICH, UBC) . 142
QUEBEC:
Gaspe Peninsula, Grosses-Roches, Jones 31953 (CAN).
Gaspesian Prov. Pk., Mt. Albert, Crum & Williams 10582
(CAN, LAF, MICH, UBC), 10723 (CAN).
Hudson Bay, Great Whale R., Brisson & Forest 20633, 21051,
21310 (CAN, NFLD, UBC).
Levis, St. Henri, Masson & Gagnon 8597 (CAN).
Luskville, Luskville Falls, Ireland & Ley 10025 (FLAS,
MICH, UAC, UBC).
Montmagny, Notre Dame du Rosaire, Masson & Gagnon 10090
(CAN).
Richmond Gulf, Wiachewan Bay, Marr M371 (MICH).
Riviere-du-Loup, Blouin 6550-1 (CAN).
Terrebonne Co., Saint Jovite, LacMercier, Crum 9960 (CAN,
LAF) .
LaVerendrye Prov. Pk. (S of), Crum & Williams 10184 (CAN).
SASKATCHEWAN :
Qu'Appelle Valley, Elbow Forest Preserve, DeVries 913 (CAN).
YUKON TERRITORY:
Dawson, Calder & Billard 3870 (NY).
Dezadeash River, Alaska Hwy., Crum & Schofield 8658, 9508
(CAN).
Mayo, Keno Hill, Gillett & Calder 4378, 4266 (NY).
U. S. A.
ALABAMA:
Randolph Co., Wedowee, Hermann 10034 (MO, US).
Shelby Co., Montevallo, Blaokiston 6 April 1926 (NY). 143
ALASKA:
Adak Isalnd, Shagak Bay, Jordal 2761 (CAN, MICH).
Alaska Range, Mt. McKinley Nat. Pk., Smith 2202A (CAN, UC).
Attu Island, Chicagof Harbor, Sohaack 691 (CAN, MICH,
UBC, US).
East Oumalik, Steere 15502 (CAN, MICH).
Glacier Bay Nat. Monument, Muir Inlet, Argus & Chunys
6545, 6547 (CAN, MICH).
Juneau, Strong 55 as var. pumila (NY).
Kenai Peninsula, Kenai R., Lutz 50-5, 50-6 (NY).
Kodiak Island., Olga Bay, Looff E24S, E281 (CAN).
Mt. McKinley Nat. Pk., Mt. Eielson, Weber & Viereck 10132
as P. tomentella (CAN, DUKE, SMS, SMU).
Prince William Sound, Cooper 5 Sept. 1935 (NY).
Skagway Quadrangle, Haines, Hermann 21799 (CAN).
Unalaska Island, Maooun 303 (CAN).
ARKANSAS:
Conway Co., Petit Jean Mt., Anderson 11884 (DUKE).
Crawford Co., Winfrey, Redfearn 21093 (SMS).
Faulkner Co., Martinville, Redfearn 19399 (SMS).
Garland Co., Hot Springs, Russell 4 (NY).
Newton Co., Ben Hur, Anderson 11933 as P. caespitosa (MICH).
Polk Co., Mena, Anderson 11512 as P. muehlenbergii (DUKE,
MICH).
Stone Co., Ozark Nat. Forest, Redfearn 23164 (SMS).
VanBuren Co., Shirley, Redfearn 16568 (SMS).
Washington Co., Odell, Redfearn 16920 (MO, SMS). 144
White Co., Steprock, Redfearn 19427 (SMS).
Yell Co., Dardanelle, Redfearn 23917 (SMS).
ARIZONA:
Cochise Co., Rucker Canyon, Pultz, Barrow, Gould & Phillips
2765 (NY).
Coconino Co., Grand Canyon Nat. Pk., Raring & Bryant 3721
(NY), S of Jacob Lake, Zales 2982 (UBC).
Gila Co., Payson, Smith 9 May 1945 (NY).
Graham Co., Mt. Graham, Pultz, Barrow & Phillips 3 Sept.
1944 (NY).
Pima Co., Romero Canyon, Baring 3260, 3262 (NY).
Santa Cruz Co., Baldy Trail, Bartram 172 as P. tomentella
(CAN, DUKE, OP, US).
CALIFORNIA:
Butte Co., West Branch Campground, Kowalski 10 Oct. 1967
(CAN).
Calaveras Co., Stanislaus R., Koch 1529 as var. pumila (NY).
Eldorado Co., Upper Echo Lake, Beetle 4054 (NY).
Fresno Co., Shaver Lake, Ikenberry 1602 (CAN).
Humboldt Co., Grouse Mt., Tracy 14088% (UC).
Inyo Co., Mt. Whitney, Baring 4523 (NY).
Marin Co., Mt. Tamalpais, Koch 2291 (MICH).
Mariposa Co., Yosemite Falls, Koch 1706 (MICH).
Mono Co., Leevining Grade, Koch 17 79 (NY, UC).
Napa Co., Napa-Monticello Rd., Koch 1366 (NY, UC, US).
Placer Co., Donner Lake, MacFadden 16958 as var. pumila
(CAN), Canyon Creek, MacFadden 16954 as var. pumila
(CAN). 145 Plumas Co., Feather River Canyon , 27 April 1970 (CAN),
LaPorte, Traverse 505 (CAN).
Riverside Co., San Jacinto Mt., Hall 2388 (UC).
San Bernardino Co., Snow Valley, Haririg 4515 (NY).
Shasta Co., Stillwater, Backer & Nutting 18 May 1894 (NY).
Siskiyou Co., Mt. Shasta, Frye 10 July 1933 (SMU).
Stanislaus Co., Oakdale, Zales 2922 (UBC).
Tualumne Co., Sonora Pass, Howell 5 June 1954 (CAN, US).
Tulare Co., E Fork Kaweah R., Koch 2168 (NY, US).
COLORADO:
Boulder Co., Boulder, Conard 41-214 (CAN).
Chaffee Co., Monarch Pass, Zales 3014 (UBC).
El Paso Co., Manitou, Jewett 883 (NY).
Grand Co., Rabbit Ears Pass, Schofield 7177 (CAN).
Larimer Co., Fort Collins, Hermann 16968 (CAN, DUKE, US),
Bear Lake, Zales 3058 (UBC).
Mineral Co., Pagosa Peak, Baker 121 (NY)-.
Ouray Co., Red Mt., Summit, Griffin 121 (FLAS).
Park Co., Jefferson, Weber B-4391 (CAN, NY).
Pitkin Co., Aspen, Weber B-10912 (CAN, LAF), B-10898,
10899 (LAF).
San Juan Co., E Lime Creek, Zales 3000s 3002 (UBC).
Summit Co., Blue Lake, Weber & Anderson B-34256 as
P. tomentella (CAN), Tenmile Creek, Zales 3027 (UBC).
CONNECTICUT:
New Haven Co., Carmel Mt., Eaton 5 May 1880 as Bartramia
(NY) . 146
GEORGIA:
Carroll Co., Yellow Dirt Creek, Zales 2119-2122 (UBC).
Dekalb Co., Little Stone Mt., Small 95773 9589 (NY, UBC).
Rabun Co., Tallulah Falls, Small 9523 (NY).
Walker Co., Cloudland Canyon St. Pk., Zales 2147 (UBC).
Whitefield Co., Rock Face Mt., Harper 279a (NY, US).
IDAHO:
Bonner Co., Lake Pend d'Oreille, Leiberg 49 (CAN, NY, US).
Custer Co., Fourth of July Creek, MaoFadden 18737 (CAN,
DUKE, MICH, NY, SMS).
Elmore Co., Queens River, MaoFadden 18739 (CAN, MICH, NY, US).
Idaho Co., Burgorf, Hermann 20291 as P. amerioana (NY).
Kootenai Co., Leiberg 377. as. var. gracilescens (CAN, NY).
Latah Co., Beal's Butte, Mueggler 8M-37 (CAN, DUKE).
Shoshone Co., Marks Butte, Mahler 2313 (SMU).
Teton Co., Victor, Payson 2181 (NY).
ILLINOIS:
Johnson Co., Goreville, Redfearn 18739 (SMS).
LaSalle Co., Starved Rock St. Pk., Voth & Richards 958
(DPU, NY).
Pope Co., Burden Falls, Chase 11676 (CAN).
INDIANA:
Elkhart Co., Simonton Lake, Deam 54644 (DPU).
Lagrange Co., Mongo, Welch 7635 (DPU, MICH).
Tippecanoe Co., Lafayette, Welch 10132 as var. falcata (DPU).
KENTUCKY-:
Powell Co., Clay City, Wharton 124 (CAN, NY). 147
MAINE:
Franklin Co., Rangeley, Oakes s. n. as P. caespitosa
(IA, SMU) .
Hancock Co., Mount Desert Island, Faxon & Band June 1890
as P. muehlenbergii (NY).
Kennebec Co., Monmouth, Merrill 46 (NY).
Oxford Co., Sumner, Parlin 12382 (NY).
Penobscot Co., Lake Matagamon, Allard 6941 (NY, US).
Piscataquis Co., Mt. Katahdin, Crane 562 (CAN).
Sagadahoc Co., Georgetown, Friend 10505 (NY).
Somerset Co., Skowhegan, Allen 6 July 1877 (NY).
York Co., Alfred, Crane 10 (CAN).
MARYLAND:
D. C. Rock Creek Park, Maxon 6290 (NY).
MASSACHUSETTS:
Dukes Co., Nashawena Island, Northrop 5 July 1901 (NY).
Essex Co., Amesbury, Huntington 10 March 1901 (NY).
Suffolk Co., Blue Hill, Faxon 197 (NY).
MICHIGAN:
Alger Co., Munising, Nichols 38 (MICH, NY).
Barry Co., Yankee Springs, Mazzer 15 April 1962 (CAN, DUKE).
Cheboygan Co., Iron Bridge, Ireland 4674 (CAN, US).
Chippewa Co., Sugar Island, Crum July 1969 as P. capillaris
(MICH).
Delta Co., Big Bay de Noc, Gleason 2423 (DUKE, MICH, NY).
Emmet Co., Cecil Bay, Wynne 2674 (NY).
Gratiot Co., Alma, Davis 22 Oct. 1892 (LAF). 148
Keweenaw Co., Copper Harbor, Hermann 16146 (CAN, US).
Mackinac Co., Bois Blanc Island, Wynne 2581 (NY).
Marquette Co., Marquette, Britton 83 (NY).
Presque Isle Co., Ocqueoc Falls, Crum 6 July 1947 (CAN).
MINNESOTA:
Big Stone Co., Ortonville, Holzinger 22-26, 1901 as
P. alpicola (NY).
Cook Co., Grand Marais, Holzinger 316 as var. caespitosa
(CAN).
Lake Co., Two Harbors, Gleason 1940 as P. caespitosa
(NY, UBC) .
MISSOURI:
Cedar Co., Cedar Springs, Ireland 3492 (CAN, US).
Dade Co., Jordan Creek, Redfearn 6995 (SMS, US).
Iron Co., Annapolis, Russell 63 (NY).
Jasper Co., Neck, Palmer 3052 (CAN).
Jefferson Co., Pacific, Redfearn 25514 (SMS).
Lincoln Co., Poley, Steyermark 104 (DUKE, NY).
Madison Co., French Mills, Redfearn 20880 (SMS).
Ripley Co., Pleasant Grove, Bush 10 Aug. 1899 (NY).
Shannon Co., Eminence, Redfearn 8412 (FSU, LAF, SMS, TENN,
US) .
St. Charles Co., Schluersburg, Steyermark 156 (DUKE, NY).
St. Clair Co., Blackjack, Redfearn 6853 (FSU, US).
St. Genevieve Co., Pickle Springs, Redfearn 18386 as
P. capillaris (MICH, SMS, TENN). 149
MONTANA:
Flathead Co., McDonald Falls, Schofield 12192 (CAN, DUKE).
Gallatin Co., Bozeman, Chase 13858 (CAN).
Glacier Co., Logan Pass, Hermann 18062 (CAN).
Lake Co., Poison, Schofield 11740 (CAN, DUKE, TENN, US),
Crane Creek Waterfall, ZaZ.es 3122 (UBC).
Lincoln Co., Cabinet Mts., Schofield 11977 (CAN, DUKE).
Madison Co., Pony, Rydberg <£ Bessey 1897 (NY).
Missoula Co., Lolo Nat. Forest, Hermann 20183 (US).
NEBRASKA:
Garden Co., Oshkosh, Kiener 16227 as P. americana (MICH,
TENN).
NEVADA:
Elko Co., Schoonover Creek, Lawton 2716 (CAN, DUKE, US).
Ormsby Co., Kings Canyon, Backer 915 as P. aoutiflora
(DUKE, NY).
Washoe Co., Mt. Rose, Lawton 3081 (CAN, -DUKE, TENN, US).
White Pine Co., Backer Creek, Lawton 2818 (CAN, DUKE,TENN,
US) .
NEW HAMPSHIRE:
Coos Co., Gorhara, Ogden 2367 (CAN, DUKE, MICH, UAC).
Crafton Co., Holderness, Faxton 192, 193 (NY).
NEW JERSEY:
Hunterdon Co., Lanibertville, Best 108 (CAN).
Sussex Co., Montague Township, Nash 4 July 1909 (NY).
NEW MEXICO:
Cafron Co., Gila Nat. Forest, Baad 1391 (MICH). 150
Dona Ana Co., Organ Mts.,•Wooton 1500 as P. amerioana (NY).
Lincoln Co., Ruidosa Creek, Wooton 1516 as P. amerioana (CAN).
Santa Fe Co., Santa Fe Canyon, Bartram 56 (CAN).
NEW YORK:
Essex Co., St. Huberts, Hermann 14804 (CAN, DUKE, NY).
Green Co., Stony Clove, Smith 32687 (CAN, NY).
Hamilton Co., Blue Mountain Lake, Hermann 15692 (CAN, US).
Nassau Co., Massapequa, Cain 34 (NY).
Rensselaer Co., Schaghticoke, Backer 2915 (NY).
Rockland Co., Suffern, Austin June 1866 (NY).
Steuben Co., Hornell, Smith & Ogden 13836 (NY).
Tioga Co., Waverly, McNeilus 24 (FLAS).
Ulster Co., Moose Lodge, Beats 661 (CAN).
" Warren Co., Thurman, Smith 31049 (CAN, NY, US).
NORTH CAROLINA:
Alexander Co., Heddenite, Keever 490 (DUKE).
Alleghany Co., Roaring Gap, Schallert 1195 (DUKE).
Avery Co., Grandfather Mt., Anderson 12746 (CAN, DUKE, MICH).
Davidson Co., Lexington, Anderson 9205 (DUKE).
Durham Co., Durham, Blomquist 4535 (DUKE).
Forsyth Co., Schallert 5-10-1930 (NY).
Franklin Co., Bunn, Beaven 714 (DUKE).
Jackson.Co., Oakland, Hermann 15340 (NY).
Macon Co., Highlands, Ireland 2674, 2774, 2945 (CAN, US),
Bridal Veil Falls, Zales 2331, 2334, 2335 (UBC),
Whiteside Mt., Zales 2350-2352 (UBC).
McDowell Co., N Fork Catawba R., Zales 2452, 2453 (UBC). 151
Transylvania Co., Cashiers, Ireland 2580 (CAN, US).
OHIO:
Fairfield Co., Sugar Grove, Werner 3 Sept. 1892 (NY).
Hocking Co., Cantwell Cliff St. Pk., Redfearn 25278 (SMS).
Jackson Co., Liberty Township, Bartley & Pontius 23 (NY).
Lake Co., Painesville, Werner 20 July 1892 (NY).
OKLAHOMA:
LeFlore Co., Cedar Creek Lake, Redfearn 24751 (SMS).
Sequoyah Co., Black Fox Creek, Redfearn 24592 (SMS).
OREGON:
Clackamas Co., Timberline Trail, Thomas 317A (CAN).
Benton Co., Mary's Peak, Lawrence 1381 (NY).
Clamath Co., Buck Lake, Copeland 3648 as P. tomentella (NY).
Mount Hood, Cloud Cap Inn, Miller 8428 (CAN).
Multnomah Co., Multnomah Falls, Zales 2767 (UBC).
Union Co., Elgin, Sheldon 8731 (NY, US).
PENNSYLVANIA:
Bucks Co., Upper Black Eddy, Taylor 3614 (MICH).
Clearfield Co., Penfield, Moul 6029 as var. falcata (DUKE).
Dauphin Co., Gratz, Altland 11 March 1938 (NY).
McKean Co., Balivar River, Burnett 2556 (NY).
Mercer Co., Grove City, Barbour 14 (BRNM, MICH, OP).
Pike Co., Dingman's Falls, Wiegmann 10 Sept. 1944 (NY).
Wayne Co., Five Mile Creek, Moul, s. n. (NY).
RHODE ISLAND:
Cumberland, Olney, s. n. (NY).
SOUTH DAKOTA:
Pine Ridge Reservation, Visher 2235 (NY). 152
TENNESSEE:
Sevier Co., Charlie's Bunion, Schofield 8845 (CAN, DUKE),
Clingmans Dome, Zales 2245 (UBC), Elkmont Campground,
Zales 2161, 2167 (UBC), Laurel Falls, Zales 209, 2209
(UBC), Newfound Gap,.Zales 2313 (UBC).
Unicoi Co., Unaka Springs, Sharp UM-691 (DUKE, MICH, TENN,
US) .
TEXAS:
Gillespie Co., Coal Creek, Jermy 1890 as P. muehlenbergii
(NY, SMU, US).
Jeff Davis Co., Mt. Livermore, Hinckley as var. pumila
1136 (MICH, NY, TEX), 1137, 1147 (NY, TEX), 1148 (TEX).
UTAH:
Salt Lake Co., Brighton, Darker 5763 (MICH).
Washington Co., Zion Nat. Pk., Zales 2972 (UBC).
VERMONT:
Lamoille Co., Lake of Clouds, Grout 7-3-1896 (DUKE, NY).
Rutland Co., Rutland, Tollison 10619 (NY).
Windham Co., Bellows Falls, Githens 2735 (CAN, UAC).
VIRGINIA:
Albemarle Co., Jones Run Falls, Ireland 2198 (CAN, LAF, US),
2201, 2213 (CAN).
Craig Co., Patterson R-673 (NY).
Giles Co., Mountain Lake Biological Station, Ireland 1128
(CAN, US).
Lee Co., Cumberland Mt., Patterson SW-11 (CAN).
Madison Co., Shenandoah Nat. Pk., Ireland 1903, 1918, 1928, 153
(CAN, US), 1927 (CAN, FSU, US).
Page Co., Lewis Falls, Ireland 2147 (CAN, DUKE, US).
Rockbridge Co., Cypress Falls, Carroll Nov. 1948 as
P. caespitosa (NY).
Shenandoah Co., Devils Hole Mt., Allard 4585, 4605, 6479
(CAN, DPU, MICH).
Smyth Co., Bear Creek, Vail & Britton 7 June 1892 (NY).
St. Paul Island, Macoun 1 Aug. 1892 (CAN).
WASHINGTON:
Chelan Co., Stevens Pass, Lawton 2399 (CAN), Zales 2627
(UBC) .
King Co., W of Stevens Pass, Zales 2628 (UBC).
Kittitas Co., Lake Keechelus, Frye 88 (CAN).
Lewis Co., Nisqually River, Schofield & Boas 22228 (CAN,
DUKE).
Olympic Nat. Pk. , Soldac Hot Springs, Schofield 19451
(CAN, DUKE).
Pierce Co., Nisqually Glacier, Cowles 795 (CAN).
Skamania Co., Skamania, Ireland & Simpson 9611 as var.
falcata (CAN).
Snohomish Co., Glacier Peak, Schofield & Hermann 21838
(CAN, DUKE).
Whatcom Co., Mt. Baker, Schofield 16061 (CAN, DUKE), Zales
2567 (UBC).
WEST VIRGINIA:
Adams- Co., Dells, Taylor 37 (NY).
Pocahontas Co., Marlinton, Gray M-509 (OP). 154
Preston Co., Erwin, Gray 7512 (NO, NY).
Tucker Co., Canaan Valley, Allard 10092 (CAN, SMU, US).
WISCONSIN:
Barron Co., Barron, Cheney 9981 (WIS).
Bayfield Co., Drummond, Cheney 4055 (WIS).
Douglas Co., Amnicon Falls St. Pk., Schroeder M158 (LAF).
Jackson Co., Black River Falls, 1859 (CAN, NY).
Lincoln Co., Grandfather, Cheney 2612 (WIS).
Milwaukee Co., Henderson 17 May 1884 (NY).
Vilas Co., Wisconsin River, Cheney 608 (WIS).
Waukesha Co., Mukwonago, 10229 (WIS).
Wood Co., Wisconsin Rapids, Chase 6470 (CAN).
WYOMING:
Albany Co., Brooklyn Lake, Porter 1427 (NY).
Carbon Co., Ryan Park, Hermann 17179 as var. pumila (CAN,
DUKE, NY, US).
Park Co., Beartooth Butte, Conard 23 Aug. 1953 (CAN).
Teton Co., Grand Teton Nat. Pk., Zales 3091 (UBC). 155
Philonotis fontana var. americana Exsiccati Examined
Baker, Pacific Slope Bryo. 592 as P. alpicola (DUKE, MO, NY,
UBC, UC, US, WIS).
Allen, Mosses Cascade Mts. Wash. 52 as P. fontana (CAN, DUKE,
DPU, MICH, NY, SMU, UBC, US).
Holzinger, Musci Aero. Bor.-Amer. 70b (BRNM, CAN, L, MICH, NY,
OP, UC, WIS).
Philonotis fontana var. americana Selected Specimens Examined
CANADA
ALBERTA:
Jasper Nat. Pk., Pyramid Mt., Ostafichuk 22 July 1970 as
P. fontana (CAN).
BRITISH COLUMBIA:
Hastings, Macoun 4 May 1889 (CAN).
Mt. Revelstoke Nat. Pk., Hamilton Cr. Brodo 7703 (UBC).
Queen Charlotte Islands, Graham Island, Schofield 14 231
(DUKE).
Vancouver Island, Port Hardy, Fort Rupert, Schofield &
Williams 26778 (CAN, DUKE).
SASKATCHEWAN:
Cypress Hills Prov. Pk., Lone Pine Creek, Bird 4679 (NY,
SMS, UAC).
U. S. A.
ALASKA:
Adak Island, Shagak Bay, Jordal 2760 (DUKE, TENN, UBC, US).
Kodiak Island, Rigg June 1913 as P. fontana var. seriata
(NY). 156
Juneau, Meh.au May 1904 as P. seriata (NY) .
Popof Island, Kinoaid 8-19 July 1899 (US).
Unalaska, Coville & Kearney 1744 as P. fontana (NY).
Yakobi Island, Shacklette 3937 (MICH).
CALIFORNIA:
Butte Co., Jonesville, Copeland 1159 as P. tomentella (UBC).
Del Norte Co., Bear Basin, Norris 9005 (DUKE, MICH).
Lassen Co., Pine Creek, Baker & Nutting 12 July 1894 as
P. fontana (NY).
Madera Co., Yosemite Nat. Pk., Ikenberry 1088 (SMU, UAC).
Modoc Co., Jess Valley, Grant 8258 (MICH).
San Bernardino Co., San Bernardino Mts., Blesdale 66 as
P. fontana (NY).
Santa Cruz Co., Camp Elvirs, Wagner & Ranzoni 2343 (MICH).
Shasta Co., Lassen Nat. Pk., Koch 1933 (CAN, NO, NY, UC).
Tehama Co., Lassen Nat. Pk., Koch 1945 (MICH, NY, UC).
Tuolumne Co., Sonora Pass, Wiggins C-8 as P. fontana var.
pumila (UBC) .
IDAHO:
Benewah Co., Potlatch, Alexander 7 July 1961 (SMU, UAC).
Custer Co., Stanley, Hermann 19938 (US).
Kootenai Co., mountain stream Leiberg 49 (NY).
Latah Co., St. Joe Nat. Forest, Mahler 1860 (SMU, UBC).
Valley Co., Lakefork Creek, Smith 5 July 1942 as
P. fontana (CAN, DUKE, MICH, TENN, UBC, US).
MONTANA:
Glacier Nat. Pk., Ole Creek, Maquire & Piranin 5334 (UC). 157
Missoula Co., Lolo Hot Spring, Nixon 8 July 1951 (UBC).
Yellowstone Nat. Pk., NW corner, Cain 4952 (NO, UAC, UBC).
NEVADA:
Washoe Co., Mt. Rose, Billings 1265 (DUKE, TENN).
OREGON :
Clackamas Co., Mt. Hood, Hermann 18738 (US).
Douglas Co., Crystal Springs, Redfearn 11659 (SMS).
Hood River Co., Timberline Trail, Ireland 7114 as
P. fontana var. -pumila (UBC).
Lane Co., Blue R., Becking 24 July 1963 (UBC).
Linn Co., E of Cascadia, Gilkey 6-43-A (IA).
Multnomah Co., Larch Mt., Miller 8500 (OP).
WASHINGTON:
Chelan Co., Stevens Pass, Lawton W5479 as P. fontana (SMU).
Grays Harbor Co., Quinalt R., Meyer 91 as P. fontana (DPU,
SMU) .
Pierce Co., Fairfax, Frye 20 May 1934 (CAN).
Snohomish Co., Mt. Pilchuck, Eyerdam 3046 (UBC).
Widbey Island, Gardner 1898 as P. fontana (US).
WYOMING:
Albany Co., Centennial, Koepper 5-10 July 1937 (DPU).
Bighorn Mts., Paintrock Creek, Jack 4 Aug. 1900 as
P. fontana (NY).
Teton Co., Grand Teton Nat. Pk., Zales 3104, 3112 (UBC).
Yellowstone Nat. Pk., Obsidian Creek, Castle 14 July 1921
(CU), NW corner on Hwy. 191, Zales 3121 (UBC). 158
Philonotis fontana var. -pumila Selected Specimens Examined
CANADA
ALBERTA:
Banff Nat. Pk., Moraine Lake, Crum & Schofield 3946 as
P. fontana (CAN, MICH, UAC, UBC), Lake Louise, Crum &
Schofield 3833 as P. fontana (CAN, DUKE, MICH, UAC, UBC).
Jasper Nat. Pk., Tonquin Valley, MaoFadden 20068 as
P. fontana var. tomentella (CAN).
BRITISH COLUMBIA:
Sikanni R., N of Fort St. John, Correll 11930 (DUKE).
LABRADOR:
Port Burwell, Polunin 1082a-10 as P. tomentella (MICH).
W Turnavik Island, Nutt 13 (US).
MANITOBA:
Fort Churchill, Twin Lake Hill, Crum & Schofield 6686
(CAN, NY, UBC).
NORTHWEST TERRITORIES:
Baffin Island, Arctic Bay, Polunin 2590b as P. tomentella
(MICH), Pangnirtung, Polunin 2643c as P. tomentella (MICH).
Bathurst Island, Bracebridge Inlet, Tener & Harington 37 5
(CAN).
Belcher Island, Tukarak Island, Doutty 30 July 19 38 as
P. caespitosa (NY).
Bylot Island, Eclipse Sound, Coombs 9 as P. tomentella (CAN).
Devon Island, Basecamp Lowlands, Barrett 179, 199, 394, 399,
(UBC), Truelove Lowland, Vitt 4388 (NFLD).
Ellesmere Island, Tanquary Fiord, Brassard 1505, 1717, 1807a, 159
1884 (CAN, NY, UBC), 2531 (CAN, NFLD), Lake Hazen,
Powell 8 (CAN, DUKE), Ella Bay, Mohl 31 July 1965 (UBC).
Great Bear Lake, Port Radium, Steeve 10032, 10124, 10230
as P. tomentella (CAN, MICH), 10230, 10503 (DUKE, MICH,
• UBC).
MacKenzie R. Delta, Eskimo Lakes, Scotter 9 583 as
P. tomentella (UAC).
Meighen Island, Kuo 23 Aug. 1968 as P. tomentella (CAN).
Southampton Island, Coral Harbor, Ritchie 27 July 1954 as
P. tomentella (CAN).
QUEBEC:
Hudson Strait, Wakeham Bay, Polunin 1479a as P. fontana
(MICH).
U. S. A.
ALASKA:
Bering Strait Dist., Ogotoruk Creek, Viereck & Bucknell
4618 as P. tomentella (CAN, UBC, US), Cape Dyer, Viereck
& Bucknell 4185 as P. tomentella (CAN).
Delta Junction, Donnelly Dome, Hermann 21142 (MICH).
East Oumalik, Steere 15330, 15437 (CAN, MICH).
Gubic, Chandler & Colville R., Steere 16121 (NY).
Hubbard Glacier, Disenchantment Bay, Coville & Kearney 1073
(NY) .
Jago Lake, Jago River, Cantlon & Gills 1463C (CAN).
Okpilak Valley, Okpilak Lake, Cantlon & Malcolm 0216 (CAN).
Meade.River Camp, Steere 15671, 15685, 15703 (CAN, MICH). 160
COLORADO:
Park Co., Mt. Lincoln, Weber 8829, 8824 as var. fontana
(DUKE) .
Pitkin Co., Roaring Fork R., Weber B-10989 as P. fontana
(CAN).
MONTANA:
Flathead Co., McDonald Creek, Hermann 18044 (CAN, NY, UBC).
WYOMING:
Park Co., Beartooth Lake, Welch 16794 (DPU). 161
Philonotis yezoana Selected Specimens Examined
CANADA
BRITISH COLUMBIA:
Vancouver, Indian Arm, Zales & Schofield 1376 (UBC).
Vancouver Island, 16 mi. W of Jordan River, Halbert &
Price 2671, 2693 (UBC), Strathcona Prov. Pk., Zales
2505 (UBC).
U. S. A.
ALASKA:
Sitka, Sharp, Sharp & Iwatsuki 5468a, 5492a (TENN).
CALIFORNIA:
Mariposa Co., Yosemite Nat. Pk., Vernal Falls, MaoFadden
17418 as P. fontana var. laxa f. tenuis (CAN), Tioga
Rd., Flowers 5461 as P. caespitosa (UT).
MONTANA:
Flathead Co., Lake McDonald, Williams 446 as P. seriata
(NY), Mt. Lottie Stanton, Holzinger & Blake 21 July
1898 (NY).
Glacier Co., St. Marys Lake, Whitehouse 25497 as
P. americana (DUKE, NY), Flowers 6127 as P. caespitosa
(UT) .
VERMONT:
Mt. Mansfield, in the Notch, 19 Aug. 1882, no coll. cited
(NY). The label cites New Hampshire however Mt.
Mansfield is in Vermont.
WASHINGTON:
Snohomish Co., Glacier Big Four, Schofield 20024 as
P. fontana (UBC) . 162 XV. LITERATURE CITED
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