Biology and Classification of

DWARF MISTLETOES (Arceuthobium)

by

Frank G. Hawksworth

Rocky Mountain Forest and Range Experiment Station

Fort Collins, Colorado

and

Delbert Wiens

Department of Biology

University of Utah

Salt Lake City, Utah

Agriculture Handbook No. 401

Library of Congress Catalog Card Number 72-609841

Forest Service

United States Department of Agriculture

Washington, D.C. February 1972

For sale by the Superintendent of Documents U.S. Government Printing Office Washington, D.C, 20402—Price $4.50

CONTENTS

DEDICATION v

ACKNOWLEDGMENTS vi

ABSTRACT viii

INTRODUCTION 1

Objectives and Scope 1

Taxonomic History 2

GENERALIZED LIFE CYCLE 3

BIOGEOGRAPHY 7

Paleobotany 7

Paleogeography 8

Extant Distribution 9

Ranges of the Genus and Species 9

Extension of Dwarf Mistletoe Ranges 11

Relation of Parasite to Host Distribution 12

Sympatry 13

HOST RELATIONSHIPS 16

Natural Hosts 16

Extra-Limital and Unnatural Hosts _ 25

Nonhosts 25

Host-Parasite Relationships 25

Host Reactions 26

Effects on Hosts 28

MECHANISMS AND TRENDS OF EVOLUTION 28

The Genetic System and Recombination

Potential 28

Progressive Evolution and Adaptive Radiation. _ 36

Absence of Hybridization and Polyploidy 36

Evolutionary Patterns and Taxonomic Structure 37

Evolutionary Trends 37

SYSTEMATIC^ OF ARCEUTHOBIUM 40

Taxonomic Considerations 40

Problems in Classification 40

Subspecific Classification 42

CRITERIA FOR CLASSIFICATION 42

Morphological Characters 42

Plant Size and Habit 42

Shoots 43

Inflorescence 45

Buds and Flowers 45

Anthers - 45

Fruit 45

Palynological Characters 45

Cytogenetical Characters 47

Physiological Characters 48

Phenology 48

Hosts 49

Witches' Brooms 49

Chemical Characters 49

Methods 51

Results 52

Discussion and Conclusions 55

NUMERICAL ANALYSES 56

Methods 56

Results _:_ 60

COLLECTING AND CURATING

TECHNIQUES 61

FORMAL TAXONOMY 62

Generic Description 62

Subgeneric Classification and Natural Keys to

the Species 26

Subgenus Arceuthohium 64

Subgenus Vaginata 64

Artificial Key to the New World Species of

Arceuthohium 67

Artificial Key to the Old World Species of

Arceuthohium 70

New World Taxa 90

1. Arceuthohium ahietinum 90

a. f. sp. concoloris 90

b. f. sp. magnificae 93

2. Arceuthohium ahietis-religiósae 93

3. Arceuthohium americanum 96

4. Arceuthohium. apachecum 101

5. Arceuthohium hicarinatum 104

6. Arceuthohium hlumeri 104

7. Arceuthohium californicum 106

8. Arceuthohium campylopodum 110

9. Arceuthohium cyanocarpum 114

10. Arceuthohium divaricatum 117

11. Arceuthohium douglasii 120

12. Arceuthohium gillii 124

a. Subspecies gillii 126

b. Subspecies nigrum 128

13. Arceuthohium glohosum 129

14. Arceuthohium guatemalense 132

15. Arceuthohium hondurense 135

16. Arceuthohium laricis 135

17. Arceuthohium microcarpum 138

18. Arceuthohium occidentale 141

19. Arceuthohium pusillum 145

20. Arceuthohium ruhrum 148

21. Arceuthohium strictum 149

22. Arceuthohium tsugense 153

23. Arceuthohium vaginatum 156

a. Subspecies vaginatum 157

b. Subspecies cryptopodum 159

c. Subspecies durangense 163
24. Arceuthohium verticilliflorum 163 LITERATURE CITED 172

Old World Taxa 166 APPENDIX 182

25. Arceuthohium chínense 166 Glossary 182

26. Arceuthohium minutissimum 167 Scientific and Common Tree Names 184

27. Arceuthohium oxycedri 167 Specimens Examined 186

28. Arceuthohium pini 169 Herbaria 186

Rejected Species 170 New World Taxa 186

MAJOR CONCLUSIONS 170 Old World Taxa 227

SUGGESTIONS FOR FURTHER RESEARCH _ 171 Exsiccatae. Index to Numbered Collections _ _ _ _ 228

IV
DEDICATION

This work is dedicated to the memory of Dr. Lake S. Gill (1900-

1969), Forest Pathologist with the U.S. Department of Agriculture

from 1923 to 1960. Dr. GilFs classic paper ^'Arceuthobium in the

United States'' (1935) was the first comprehensive treatment of the

North American dwarf mistletoes, and since its appearance has been

the authoritative publication on the genus.

Through his continued research and missionary zeal, Dr. Gill,

more than any other man, was responsible for the present awareness of

Arceuthobium as economically important forest pathogens. He demonstrated that dwarf mistletoes were not merely curiosities, but caused

significant mortality and reduction of growth rates in western conifers.

Dr. Gill followed our work with interest and encouraged our efforts.

We discussed many aspects of the study with him, but his death on

July 5, 1969 precluded his seeing the final manuscript. His enthusiasm

and interest was a constant source of encouragement for those with

whom he worked.

The use of trade, firm, or corporation names in this publication

is for the information and convenience of the reader. Such use

does not constitute an official endorsement of any product by the

U.S. Department of Agriculture to the exclusion of others which

may be suitable.
ACKNOWLEDGMENTS

We want to thank the curators of the many herbaria in North

America and Europe who allowed us to examine dwarf mistletoe

specimens; and the many colleagues who have collected the specimens,

reviewed the manuscript, suppHed photographs or drawings, assisted

with numerical analyses and Chromatographie work, and performed

innumerable other courtesies.

Canada

Joe A. Baranyay

Jan M. Bonga

Job Kuijt

Michael J. Larsen

John G. Laut

John A. Muir

Richard B. Smith

Honduras

Robert Armour

Antonio Molina R.

Mexico

Rudolfo Salinas Quinard

José de Jesus Valdivia Sanchez

Francisco Muños Sandoval

Netherlands

John Gremmen

United States

John W. Andresen

Southern lUinois University

Lyman Benson

Pomona College

Charles M. Drew

U. S. Navy, China Lake, Calif.

George F. Estabrook

University of Colorado

Ari Ferro

University of Utah

Henry S. Fleming

University of Colorado

David W. French

University of Minnesota

Robert L. Gilbertson

University of Arizona

Sharon Harris

University of Utah

John H. Hart

Michigan State University

Hugh H. Iltis

University of Wisconsin

Lee W. Lenz

Rancho Santa Ana Botanic Garden

Estella B. Leopold

U. S. Geological Survey, Denver

C. V. Morton

U. S. National Museum

J. R. Parmenter, Jr.

University of Cahfornia

David J. Rogers

University of Colorado

Lewis F. Roth

Oregon State University

Richard Snyder

University of Utah

Frank H. Tainter

University of Minnesota

R. K. Vickery, Jr.

University of Utah

Edward G. Voss

University of Michigan

William A. Weber

University of Colorado

VI
Personnel of The U.S. Department of Agriculture—Forest Service

Gerald W. Anderson David A. Graham Paul C. Lightle James L. Stewart

Ralph L. Anderson Donald P. Graham Elbert L. Little, Jr. John H. Thompson

Stuart R. Andrews Thomas E. Hinds Douglas R. Miller Harvey V. Toko

Wilmer F. Bailey Bohun B Kinloch ^ g Peterson Willis W. Wagener

Donald H. Brown Donald M. Knutson T^ u ^ 17 c^ u f TV/T.I .. T \XT..\..

T. W. Childs Richard G. Krebill I^^^ert F. Scharpf Melvyn J. Weiss

Oscar Dooling Thomas H. Laurent Keith R. Shea Ed F. Wicker

Vll
ABSTRACT

Arceuthobium (dwarf mistletoes), a distinctive genus of the family Viscaceae, is

parasitic on conifers of the Pinaceae in the Old and New Worlds and on Cupressaceae in the Old World. Although conifer forests in many parts of the Northern

Hemisphere are infected with dwarf mistletoes, those most seriously damaged are

in North America.

In this taxonomic revision, the 32 recognized taxa comprise 28 species, 5 subspecies, and 2 formae spéciales. Four taxa are known in the Old World; 28 in the

New. In North America, Arceuthohium ranges from central Canada and southeastern Alaska to Honduras and Hispaniola, but most species are found in western

United States and Mexico. Only A. pusillum, a parasite of Picea, occurs in eastern

North America.

Taxonomic characteristics presented for each of the 32 taxa include : morphology and color of shoots, fruits, and flowers; pollen features; phenology of flowering

and seed dispersal; hosts and host reactions; chromosomal features; and chromatography of shoot pigments.

The classification system is based on extensive field studies of all known New

World dwarf mistletoes in their natural state, examination of specimens at the

major herbaria in North America and Europe, and computer analyses of all taxonomic data collected.

The system, the first below-genus classification of Arceuthohium, divides the

genus into subgenera Arceuthohium and Vaginata, Subgenus Arceuthohium comprises all four Old World species {A. chinense, A. minutissimum, A. oxycedri, and

A. pini), and three New World species, but is not formally subdivided into sections.

Subgenus Vaginata, exclusively New World, contains three sections: Vaginata, 6

taxa; Campylopoda, 17 taxa; and Minuta, 2 taxa. Section Campylopoda contains

three series: Ruhra, 3 taxa; Campylopoda, 13 taxa; and Stricta, 1 taxon. Identification keys, artificial and natural, are provided.

The detailed botanical description of each dwarf mistletoe includes its distinctive

taxonomic features, hosts, a colored photograph, and a map of its geographic

distribution.

vni
Biology and Classification of Dwarf Mistletoes

{Arceuthohium)

by Frank G. Hawksworth^ and Delbert Wiens^

INTRODUCTION

The dwarf mistletoes {Arceuthohium) are^ a highly

specialized and clearly defined genus of aerial dicotyledonous parasites that occur on Pinaceae or

Cupressaceae. These mistletoes are small, leafless,

glabrous, and are characterized by a number of

unusual features: explosive, bicolored fruits; ringlike

archesporium; chlorophyllous endosperm; germinating radicle that contains stomata; and stems that

have no central vascular cylinder or sieve tube

elements.

Arceuthohium has been classically included in the

subfamily Viscoideae of the Loranthaceae. Tieghem

(1895) considered Arceuthohium so distinct from

related genera that he proposed its classification

as a separate family placed between the Viscaceae

and the Santalaceae; his proposal has never been

followed. Dixit (1962) and Barlow (1964) suggest

that the subfamilies Loranthoideae and Viscoideae

warrant family status. The two families differ in

many features of floral morphology, floral anatomy

and embryology. Previously supposed similarities

between the two groups are apparently the result of

convergence rather than common phyletic origin.

We accept full family status for the two groups as

formally proposed by Barlow^ (1964) and retained by

Thorne (1968). This treatment places Arceuthohium

in the Viscaceae.

Engler and Krause (1935) classified Arceuthohium

as a monogeneric tribe in the Viscoideae. They

placed Dendrophthora, Phoradendron, and KorthaU

sella in the tribe Phoradendreae; with Korthalsella in

subtribe Korthalsellinae, and Dendrophthora and

Phoradendron in subtribe Phoradendrinae. Regardless

of its relationship to other Viscaceae, Arceuthohium

1 Principal Plant Pathologist, Rocky Mountain Forest and

Range Experiment Station, Forest Service, U.S. Department

of Agriculture, with central headquarters maintained at Fort

Collins in cooperation with Colorado State University.

2 Associate Professor of Biology, Department of Biology,

University of Utah, Salt Lake City.

has clearly defined limits, and its generic status has

never been questioned. No formal subgeneric treatment of Arceuthohium has been published, although

Kuijt (1970) suggests that the genus is separable into

two natural groups on the basis of branching habit.

Arceuthohium is the only genus of the Viscaceae or

Loranthaceae that is clearly represented in both the

Old and New Worlds. Of four Old World species

discussed here, three have been described previously

—A. chinense in southwestern China, A. minutissimum in the Himalayas, and A, oxycedri, which

ranges from the Azores to the Himalayas and south

to Kenya; the fourth species is recorded in this

paper—A. pini of southwestern China and Tibet.

In the New^ World, 24 species (28 taxa) are distributed from central Canada and southeastern

Alaska to Honduras and Hispaniola.

The greatest species diversity is in northwestern

Mexico and the western United States, where 24 of

the 28 New World taxa occur. The four New World

species outside this area are A. hicarinatum of

Hispaniola in the Caribbean, A. hondurense and

A. guatemalense in Central America, and A. pusillum

in southeastern Canada, the Lake States, and the

northeastern United States. Fourteen taxa occur

in Mexico, six of which are found in the w^estern

United States. Of the 17 dwarf mistletoes found in

the United States, 5 are also common to Canada.

One species, A. douglasii, ranges from southern

British Columbia to southern Durango Mexico.

Objectives and Scope

With the increasing recognition of Arceuthohium as

destructive parasites on commercially important

forest trees, and the expanding interest in the biology

of the genus, the need for additional systematic work

on the group became apparent. For example, control of dwarf mistletoes in some areas is hampered by

inadequate knowledge of the identity of the parasite

involved.
Our initial concern in these studies was to clarify

the confusion surrounding the A. campylopodum

complex (see Hitchcock and Cronquist 1964), which

centered primarily around GiU's (1935) designation

of host forms in this group. When we discovered,

however, the relatively rich dwarf mistletoe flora in

IMexico (Hawksworth and Wiens 1965; Hawksworth,

Lightle, and Scharpf 1968), we expanded our

investigations to include comprehensive treatment

of the entire genus.

We have studied all known North American taxa,

plus the Himalayan species, A. minutissimum, in

their natural state. Since 1962 our field studies of

Arceuthobium have taken us by foot, hoof, wheel, and

wing over 100,000 miles throughout western North

America, Mexico, Central America, the Caribbean,

and the Himalayas. Over 1,500 dwarf mistletoe

specimens that we collected in our field work, plus

many early U.S. Department of Agriculture collections by J. R. Weir, G. G. Hedgcock, J. S. Boyce,

and L. S. Gill, are filed in the U.S. Forest Service

Mistletoe Herbarium at Fort Collins, Colorado.

Duplicates of our collections are deposited in many

North American herbaria, particularly the University of Colorado, Missouri Botanical Garden, U.S.

National Museum, University of Utah, and the

Instituto de Biología, Mexico City. In addition to

our own collections, we have studied dwarf mistletoes

at the major herbaria in North America and Europe

(see appendix, p. 186).

Our objectives in this study have been to develop

a useful, yet natural, system of classification for

Arceuthobium. Although the New World species

have been emphasized, the literature on the Old

World taxa has been reviewed, and results of our

field studies on one Old World species, A. minutissimum, are included.

This handbook is divided in 10 main sections:

(1) general life cycles, (2) biogeography, (3) host

relationships, (4) mechanisms and trends in evolution, (5) systematics of the genus, (6) classificatory

criteria, (7) numerical analyses, (8) formal taxonomic treatment, (9) major conclusions, and (10)

suggestions for further research. The appendix

contains the list of specimens that are cited for each

taxon in the taxonomic section, a glossary of some

of the less familiar terms used in the paper, and a

list of the trees mentioned in the text.

Taxonomic History

The first dwarf mistletoe described, Arceuthobium

oxycedri, was originally included in Viscum, but was

later segregated by Hoffman (1808) under the name

Razoumofskya. The name Arceuthobium, proposed

by Marschall von Bieberstein (1819), was generally

used until the early 1900^s. The Vienna Botanical

Congress in 1905 conserved Arceuthobium over

Razoumofskya, but because the American Code

emphasized strict priority, most botanists in the

United States continued to use Razoumofskya.

Arceuthobium finally displaced Razoumofskya as a

result of the 1930 Cambridge Botanical Congress

and GilFs publication (1935).

Humboldt and Bonpland's collection of a dwarf

mistletoe on Cofre de Perote, Veracruz, Mexico^ in

1804 was apparently the first in the New World. This

specimen was the type of Arceuthobium (as Viscum)

vaginatum that Willdenow described in 1806. David

Douglas, who explored the Pacific Northwest from

1823 to 1827, discovered two dwarf mistletoes in

1826. Although his journals were not published

until 1914, the plants he discussed seem certain to

be A. campylopodum (on Pinus ponderosaY and A.

americanum (on Pinus contorta). Hooker (1840), who

first discussed the taxonomy of Arceuthobium in

North America (north of :\léxico), compared

Douglas' specimens and another by Drummond

{A. americanum) with the European A. oxycedri,

but detected no specific differences except some inconsistent color variations. Engelmann was the

first to publish (Gray 1850) a formal description of a

dwarf mistletoe found in the United States, A.

americanum. As the taxonomic architect of the genus

in North America, Engelmann named most of the

American species of Arceuthobium during the latter

haff of the 19th century (Gray 1850, Watson 1880).

Between 1910 and 1920, George G. Hedgcock and

James R. Weir of the United States Department of

Agriculture each initiated studies to clarify the

taxonomy of Arceuthobium in the United States.

Although both scientists published several articles

on dwarf mistletoes, their main taxonomic works

were left unpublished, and are on file with the U.S.

Forest Service at Fort Collins, Colorado, and at

Moscow, Idaho.

Hedgcock completed a series of manuscripts in

1914 on the taxonomy of the dwarf mistletoes,

but for unknown reasons this work was never

published. This is unfortunate because Hedgcock

had a keen understanding of the genus. For example,

he first recognized that the dwarf mistletoe (our A.

californicum) on Pinus lambertiana was distinct

from A. cyanocarpum, and that A. campylopodum

was distinct from A. occidentale. Hedgcock (1915)

published a host list of the United States dwarf

mistletoes and later (Hedgcock and Hunt 1917)

reported on some new hosts of A. campylopodum and

A. occidentale based on artificial inoculations.

The only one of Weir's published papers that

3 A list of scientific names, authorities, and available

common names of the trees mentioned is given in the appendix

p. 184.
includes major taxonomic implications is a report

of a series of studies in which he artificially inoculated

several species of Northwestern dwarf mistletoes on

various hosts under field and greenhouse conditions

(Weir 1918a). A taxonomic treatise on Amreican

dwarf mistletoes that he began to write was never

completed. Weir did, however, publish observations

of hosts associated with several Northwestern dwarf

mistletoes."^

Nelson described two new dwarf mistletoes (Coulter

and Nelson 1909, Nelson 1913), and Tubeuf (1919)

published a review of the genus. Tubeuf's work

was based mainly on the literature, his extensive

knowledge of the European Arceuthohium oxycedri,

and results of a brief trip to the western United

States in 1913. His paper was a good summary of the

available literature, but it did little to clarify the

taxonomic relationships of the genus.

Following Nelson's and Tubeuf's work, taxonomic

understanding of the genus did not advance significantly until Gill's (1935) revision of the United

States species. Gill reevaluated Engelmann's works,

discovered flowering periods as an important taxonomic character, and reduced the rank of several

closely related species to host forms of A. campylopodum. Although Gill's paper clearly established

a taxonomic framework for the genus, he considered

his work to be provisionary 'Spending a complete

revision of the genus based on further field and

experimental evidence."

Kuijt^ rejected Gill's treatm.ent of A. campylopodum and A. vaginatum as being composed of host

forms. He adopted an even more conservative

philosophy than Gill for A. campylopoduin and A.

vaginatum, in which he preferred to consider each

taxon as only a single variable species. In his earhest

work, Kuijt (1955) even questioned the distinctness

of A. vaginatum from A. campylopodum, and stated

that they were separated ''only by a two- or- three

months difference in flowering time, no consistent

morphological differences having been discovered."

Furthermore he suggested, as did Gill (1935), that

A, campylopodum should include A. hicarinatum

from Hispaniola.

Hawksworth and Graham (1963) discussed the

difficulties of applying Gill's host-form concept to the

dwarf mistletoes that parasitize western spruces.

They also reported that Gill's forms in the A.

campylopodum complex can be distinguished morphologically. We (Hawksworth and Wiens 1964, 1965,

1970a, 1970b) have reported additional taxonomic

studies of the genus.

GENERALIZED LIFE CYCLE

Because we interpret discontinuities between fife

cycles as the basis for taxonomic distinction, we

place considerable emphasis on life cycles in this

paper (seep.40).

Shoots, flowers, and fruits of Arceuthohium are

illustrated in figures 1-3. Some important aspects of

the life cycle of one species (A. americanum) are

shown in figure 4. Taxa whose fife cycles have

been studied in some detail are: A. abietinum

(Scharpf and Parmeter 1967), A. americanum

(Hawksworth 1965b), A. vaginatum subsp. cryptopodum (Hawksworth 1961a, 1965b), A. campylopodum (Roth 1959, Wagener 1962), A. douglasii

(Wicker 1965, 1967a), A. laricis (Smith 1966a;

Wicker 1965, 1967a), A. tsugense (Smith 1966a)

and A. oxycedri (Heinricher 1915a, 1915b, 1924).

Although the dwarf mistletoes are typically

aerial parasites on branches or main stems of

coniferous trees, there are a few reports of their occurrence on roots. Scharpf (in Kuijt 1969) noted

parasitism of roots of Pinus sahiniana by Arceuthohium occidentale in California; also root parasitism

has been found on pines infected by A. glohosum

near jNIexico City,^ and on Volcán Tajumalco,

Guatemala (Steyermark specimen 36940 at the

University of Michigan). These are certainly

abnormal situations, however, and are presumably

due to vegetative spread into roots from infections on

the main stem near the groundline. Thus, the rare

root parasitism by Arceuthohium is not comparable

to that in the typical terrestrial members of the

Loranthaceae {Gaiadendron, Nuytsia) where initial

infection takes place through the roots.

Our discussion of dwarf mistletoe life cycles

begins with seeds. The normal Arceuthohium fruit

contains a single seed with one embryo, but rarely

fruits may contain two seeds or seeds with two

embryos (Hawksworth 1961b). Seed dispersal takes

place from midsummer to late fall depending on the

taxon involved. Seeds are explosively expelled from

the fruits at initial velocities of about 90 feet per

second for distances up to 50 feet, although the

average distance is considerably less (Hinds, Hawksworth, and AlcGinnies 1963; Hinds and Hawksworth

1965; Hawksworth 1961a). Birds may be involved in

4 Weir 1915a, 1915b, 1916c, 1917, 1918b.

' Kuijt 1955, 1960a, 1960b, 1963, 1964.

6 Personal communication, Rudolfo Salinas Q., Instituto

Nacional de Investigaciones Forestales, Mexico D. F., 1966.
oana

B

o

OQO

O o

Figure 1.—Shoots of dwarf mistletoe: A, Young shoot showing the decussate arrangement of the internodes; B,

Older shoot showing elongated internodes and typical decussate branching; C, Typical flabellate branching pattern

with cross section through dashed area; D, Typical verticillate or whorled type of branching with cross section

through the dashed area. All taxa show the type of primary branching (ß), but in most instances a secondary

type of branching also develops which may be either flabellate (C) or verticillate (D).

long-distance dispersal (see p. 9). The seeds are

viscous coated, and readily adhere to objects they

strike. Needles of coniferous trees are particularly

effective in intercepting the seeds in flight (Roth

1959, Hawksworth 1965b). Seeds that have been

intercepted usually remain on the needles until

the first rain lubricates the viscous coating, and

causes the seeds to slide down the needles. Although many fall from the needles to the ground,

some seeds become attached to the twigs by their

viscous coating.

Growth of dwarf mistletoe radicles is limited, so

only those seeds that germinate on or very near the

twigs can cause infection. The time of germination
:^ flowers

anther

nectary

B

Figure 2.—Flowers and fruits of Arceuthobium: A and

B, Staniinate shoots; C and D, Pistillate shoots.

A, Terminal part of staminate shoot showing flowers

prior to opening; ß, Terminal part ofstaminate shoot

showing open 3- and 4-nierous flowers; C, Terminal

portion of pistillate shoot showing flowers shortly

after pollination; D, Terminal portion of pistillate

shoot showing two crops of fruit, the terminal internodes with nearly mature fruits, and the lower internodes with fruit 1 year younger than the terminal

fruit.

is poorly known for most taxa, but most species

investigated germinate in late \\inter or spring. The

known exceptions are A. vaginatum subsp. crij-ptopodum and A. guatemalense which germinate in the

fall immediately after seed dispersal. Seeds of the

other Mexican and Central American species

probably also germinate soon after dispersal. The

germinating radicle forms a holdfast when it contacts an obstruction on the host branch, such as the

base of a needle (Bonga 1969b). The holdfast

develops a penetrating wedge of tissue into the host

(Scharpf and Parmeter 1967), and thus initiates the

infection process.

For most host-parasite combinations that have

been investigated infection can take place only

through the most recent tissues, usually segments'

less than 5 years old. An exception is A. americanum,

which can become established on segments at

' We refer to the single year's growth length of a conifer

branch or stem, sometimes incorrectly called an "internode,"

as a segment. The area of branchlet emergence or bud scales,

sometimes incorrectly called a "node," is termed a girdle

(see Thoday and Johnson 1930, Kuijt 1960b).

9

endosperm

embryo

viscin cells

Figure 3.—Mature fruit and seed dispersal in Arceuthobium: A, Diagrammatic cross-section of mature fruit

and seed (left) and fruit just discharging its seed (right); B, Photograph of seed in flight immediately after

discharge from fruit (lower right). Photograph taken at 5 millionths of a second (Hinds, Hawksworth, and

McGinnies 1963).
megasporogenes

fertilization

first slnoots

Figure 4.—Generalized life cycle of a dwarf mistletoe as exemplified by Arceuthobium americanum on lodgepole

pine.

least 60 years old (Hawksworth 1954). Once the

penetrating wedge has entered the cortex, an endophytic (or rootlike) system ramifies throughout

the bark. Some parts of the endophytic system

(the ^^sinkers'O become embedded in successive

layers of xylem.^ Once infection is established, an

incubation period of usually 2 to 5 years elapses

before aerial shoot formation begins. A swelling at the

point of infection often precedes shoot production

^ The endophytic systems of various species of Arceuthobium

have been intensively studied by Heinricher 1924; Heil 1923;

Thoday and Johnson 1930; Cohen 1954, 1963; Kuijt 1960b;

Srivastava and Esau 1961.

by a year or more, but this varies with the mistletoe,

the host, and other factors. Usually 1 to 2 years pass

after initial shoot appearance before flowering

begins.

Meiosis may immediately precede flow^er production or it may occur approximately 5 to 8 months

previously (Wiens 1968). Flowering may occur

as early as February and March (A. glohosum) or as

late as November and December (A. occidentale).

For a given taxon and locality, flowering usually

lasts 4 to 6 weeks, but most pollen is dispersed

within a 2- to 3-week period (Hawksworth, unpublished data).
The dwarf mistletoes have features characteristic

of insect-rather than wind-pollinated plants (Whitehead 1969): they bear a relatively limited amount

of spined pollen that is shed in clusters, produce

nectar, and emit odors from both staminate and

pistillate flowers. Experimental work on dwarf

mistletoe pollination is in progress (Gregor and

Wiens, unpublished data). Thrips seem to be involved in the pollination of A. vaginatum subsp.

cryptopodum (Hawksworth 1961a), but different

types of insects are apparently more important in

other species (Stevens and Hawksworth 1970).

Some wind pollination may be effected, at least at

very close distances. The staminate flowers and the

terminal parts of the shoots are usually shed a few

weeks after anthesis. In species such as A. verticilliflorum, the entire flowering spikes are deciduous

after anthesis. In species other than A, pusillum,

individual shoots may produce crops of flowers over

several successive years.

The time required for fruits to mature varies

considerably; it is only about 5 months for A.

pusillum, a year or more in other taxa, and as long as

18 to 19 months in A. gillii subsp. gillii. The minimum time from infection to initial seed production

averages 6 years in A. americanum and 7 to 8 years

in A. vaginatum subsp. cryptopodum. There have

been reports (Peirce 1905, Korstian and Long 1922)

that the shoots die after the fruits mature, but this

is certainly not the normal situation. We have

observed individual shoots of A. americanum and

A. vaginatum subsp. cryptopodum that have produced

successive crops of fruits for 5 years and are still

living. Also, Kuijt (1970) reports that several taxa

have relatively long-lived shoots. With seed production, our discussion of the life cycle is completed.

All dwarf mistletoes are dioecious; the pistillate

and staminate plants occur in approximately equal

numbers in the taxa that have been investigated:

A. vaginatum subsp. cryptopodum (Hawksworth

1961), A, campylopodum (Wicker 1967b), and A.

americanum (Hawksworth, unpublished data).

Several biotic and abiotic factors affect the shoots,

fruits, and seeds of the dwarf mistletoes. A number

of fungi attack these plants and, although they are

damaging and abundant in some areas in certain

years, they seem to exert relatively little biological

control (Wicker and Shaw 1968, Wilson 1969).

Little information is available on the effects of the

many insects associated with the dwarf mistletoes

(Gill and Hawksworth 1961, Stevens and Hawksworth 1970). Squirrels, porcupines, birds, and other

animals eat dwarf mistletoe shoots, but their effects

seem to be rather minor. Various climatic factors

such as temperature extremes and winter drying

affect dwarf mistletoe shoots, but no data on the

importance of these factors on populations are

available.

BIOGEOGRAPHY

Paleobotany

The fossil record of Arceuthohium is meager. The

oldest known fossil pollen is from the middle Eocene^

in East Germany. Identified as Spinulaepollis

arceuthohiodes (Krutzsch 1962), it was found at more

than 60 East Germany stations. Most grains were in

upper Eocene to Miocene strata, with extremes from

middle Eocene to the Pliocene. Stuchlik (1964),

working in Polish Miocene deposits, considered the

»The Cenozoic era, or approximately the last 65 million

years of geologic history, is divided by the U.S. Geological

Survey as follows: (data provided by Dr. Estella B. Leopold,

1970).

Period

Quaternary

Series or epoch

^Holocene

j Pleistocene

Years before present

10,000

3 million

[Pliocene 3-12 million

Miocene 12-26 million

Tertiary ' Oligocène

Eocene

Paleocene

26-38 million

38-53 million

53-65 million

pollen similar to, if not identical with, the living

Arceuthohium oxycedri. He also found a rarer, larger

type of pollen described as a new subspecies: Spinulaepollis arceuthohioides subsp. major.

The fossil subspecies had pollen grains 24 to SO/x

in diameter compared with 20 to 24jLt for the species.

Possibly swelling due to fossiUzation or to laboratory

preparation procedures could have affected pollen

size. The A. oxycedri pollen that we have examined

has a diameter of 18 to 20^. From our pollen studies a

range from 20 to 30^ is unusual within a taxon;

therefore the larger grained type may represent an

extinct species. The northern limits of A. oxycedri

are now about 500 miles south of the East German

and Polish fossil pollen sites.

The fossil genus Patzea (Caspary 1872), which was

considered by Engler and Krause (1935) to be

synonymous with Arceuthohium, was described from

OUgocene amber formations on the Baltic Coast.

However, judging by the drawings of Patzea published by Conwentz (1886), we believe that Patzea

is not congeneric with Arceuthohium.
Leopold^^ noted two Miocene Arceuthobium pollen

records from North America: (1) early or middle

Miocene pollen from the north flank of the Alaska

range, associated with Pinus and Picea pollen

(Wahrhaftig, Wolfe, Leopold, and others 1969), and

(2) pollen in the middle Miocene Troublesome

formation in north-central Colorado; with associated

coniferous pollen, predominantly Picea and Pinus,

and some Abies (Weber 1965).

Leopold^^ also found Arceuthobium pollen in sediment dated from 5,000 to 10,000 years before present

(B.P.) in cores from near the center of Searles Lake

in the Mojave Desert in California. The species of

Arceuthobium has not been determined, but it is

probably A. divaricatum which occurs today in the

same area on Pinus monophylla.

Adam (1967) reported the occurrence of Arceuthobium pollen over the last 10,000 years at four sites

in the central Sierra Nevada of California. He noted

irregular distribution of Arceuthobium pollen during

this period and found that the peaks occurred when

pines were declining. Arceuthobium was perhaps

more abundant during drier periods because dwarf

mistletoe, at least on Pinus ponderosa, now seems to

be more abundant in the drier regions of California.

Data published by Janssen (1968) show a similar

trend for A. pusillum on spruce in Minnesota; thatis,

peaks in Arceuthobium pollen occurred when spruce

was declining.

Baker (1969) found Arceuthobium pollen in

sediments near Lake Yellowstone, Wyoming, that

dated to about 11,000 years B.P. The species was

presumed to be ^. americanum because this is the

only species now common in the region and the pollen

grains were associated with fossilized needles of

Pinus contorta, the principal host of this dwarf

mistletoe.

Pleistocene Arceuthobium pollen (presumably A.

pusillum), has been found in the southeastern

United States about 300 miles south of the present

southern hmits of A. pusillum. Fossil pollen (radiocarbon dated at 38,000 years B.P.) has been found in

North Carolina and South Carolina (Whitehead

1963, 1964; Whitehead and Barghoorn 1962; Whitehead and Doyle 1969) and in northwestern Georgia

(radiocarbon dated at about 15,000 to 23,000 years

B.P.) (Watts 1970). Whitehead and Barghoorn

(1962) consider the possibility that the Arceuthobium

pollen represents a species from the western United

States or an extinct species. However, since the

dwarf mistletoe pollen is associated with spruce

pollen and needles, and A. pusillum is the only

eastern species, it seems almost certain to be this

taxon. Also, A. pusillum is the only species now

Impersonal communication, Dr. Estella B. Leopold, U.S.

Geological Survey, 1969.

occurring within about 1,200 miles of the fossil sites.

Fossil pollen of Arceuthobium pusillum has also

been found within the present range of the species

in Minnesota. AIcAndrews (1966) reported it at

Lake Itasca, Clearwater County (radiocarbon dated

at about 5,000 B.P.) and Janssen (1968) found it at

Myrtle Lake, Koochiching County (radiocarbon age

from present to 9,000 years B.P.).

The only reported macrofossils of Arceuthobium

are those from Pleistocene formations of Coastal

California (Chaney and Mason 1930, 1933; Mason

1934). Axelrod (1966) dated the sites as late Pleistocene, and obtained radiocarbon dates of 30,000

to 40,000 years B.P. These fossils, found at Carpenteria and Santa Cruz Island in Santa Barbara

County, and at Tomales Bay, ]Marin County,

were abundant at the latter two localities. The

dwarf mistletoe was associated with Pinus radiata

and P. muricata, and was reported to be indistinguishable from modern A. occidentale (as "A.

campylopodum''), which presently parasitizes these

pines. This mistletoe does not now occur at the

fossil sites in Santa Barbara County, although P.

muricata is still present on Santa Cruz Island. The

closest modern mistletoe population on these

pines is about 100 miles north near Cambria on

P. radiata. In ]Marin County, A. occidentale is

still present on P. muricata on Inverness Ridge

adjacent to Tomales Bay. Chaney and Mason (1933)

note that the fossil plant had primarily 4-merous

staminate flowers. This character is quite variable,

however, and the 4-merous condition is predominant

in many modern A. occidentale populations.

Paleogeography

The occurrence of Arceuthobium pusillum on

eastern spruces is perplexing, because dwarf mistletoes have not colonized the eastern and southern

pines. Arceuthobium pusillum may have evolved

somewhere outside the present range of the parasite,

perhaps in northwestern North America. According

to Mirov (1967), the pines in eastern and western

North America have been essentially isolated since

the Phocene; hence, the dwarf mistletoes may not

have had an opportunity to colonize the eastern

pines.

Presumably the Pleistocene glaciations eliminated

A. pusillum from the northern and western parts of

the range of spruce. The dwarf mistletoe was pushed

southward into spruce refugia in the Appalachians

(as witnessed by Pleistocene Arceuthobium pollen

in the Carolinas), and possibly on the Gaspe Peninsula in Ontario. Since the glacial period, A. pusillum

has again moved northward, but its spread has been

slower than that of its hosts. Both Picea mariana

and P. glauca now occur as far northwestward as

8
Alaska, but A. pusillum just enters eastern Saskatchewan. Possibly the mistletoe is still migrating

northward and westward in Canada.

Picea apparently arrived in North America in two

or more migration waves (Gordon 1968, Wright

1955). The available information on dwarf mistletoe

parasitism of Picea tends to confirm this. The species

that arrived in the Cretaceous or earlier (relatives of

Picea hreweriana, P. chihuahuana, and possibly P,

sitchensis) do not have principal dwarf mistletoes

associated with them. However, spruces that

evolved later (early Tertiary?) such as forms related

to P. pungens, P. engelmannii, P. mariana, and P.

glauca, are now severely parasitized. These^ data

indicate that Arceuthohium may have arrived in the

New World in the early Tertiary.

Three species of Arceuthohium {A. americanum,

A. pusillum, and A. tsugense) now have far northern

distributions. Their northward migration since the

last Wisconsin glaciation is difficult to explain.

Current migration rates of dwarf mistletoes through

host stands are reported to average only about 1 to

2 feet per year (Hawksworth 1958, 1961a). If, during

Wisconsin glaciation, the parasites were pushed to

the presumed ni^rthern forest Hmits as mapped by

Dillon (1956), it indicates that all three dwarf mistletoes have migrated 1,000 miles or more in the last

10,000 to 12,000 years. This equals a rate of spread

(on the order of 0.1 mile per year) many times greater

than that recorded in present stands, and suggests

different means of dispersal. Spread data in present

stands are based only on spread due to the explosive

fruits, and do not account for possible long distance

vectors such as birds. Perhaps birds were primarily

responsible for this northward migration, but the

explosive mechanism enabled the plants to intensify

locally once they became established in new areas.

No definitive studies have been made on the importance of birds as dispersal agents of the dwarf

mistletoes,!! but these characteristics should be kept

in mind: (1) dwarf mistletoes are dioecious, so at

least two seeds must germinate, cause infection,

flower, and fruit before a new population can be

established, and (2) these plants mature their seeds

in the fall, so bird migration patterns would presumably tend to disseminate seeds southward.

The occurrence of A. hicarinatum on the Island of

Hispaniola in the Caribbean is of unusual biogeographic interest. This species occurs only on the

relatively high-altitude species, Pinus occidentalis.

Probably this parasite reached this area during

Miocene time when land connections may have

existed between islands of the Caribbean and

Central America (Mirov 1967). The recent discovery

11 Gill 1935; Hawksworth 1961a, 1967; Kuijt 1955; Zavarin,

Critchfield and Snajberk 1969.

of A. hondurense in the uplands of Honduras is

possibly of paleogeographic significance in this

regard. Arceuthohium hicarinatum and A. hondurense,

which are now separated by about 1,000 miles, may

represent remnants of what was once a common

stock in ]Miocene time.

The lack of dwarf mistletoe on the low-elevation

Caribbean pines is difficult to explain because these

trees were undoubtedly exposed to infection during

the migration of A. hicarinatum to Hispaniola.

Pinus carihaea is a host of A. glohosum in the uplands

of British Honduras but the dwarf mistletoe apparently does not occur on the more extensive lowelevation stands of this species elsewhere in the

Caribbean. Perhaps the Arceuthohium type of seed

dispersal is not adaptable to low, moist, tropical

chmates, although A. tsugense is abundant in the

cool, very wet climate of the northwestern coast of

North America.

Extant Distribution

Ranges of the Genus and Species

In the Old World Arceuthohium is represented by

four known species: A. oxycedri on Juniperus, A.

minutissimum on Pinus grißthii, A. chinense on

Ahies and Keteleeria, and A. pini on P. tahulaeformis.

Arceuthohium oxycedri, the only member of the

genus exclusively parasitic on Cupressaceae, ranges

from the Himalayas, throughout the Mediterranean

region, to the Azores, and south to Kenya. This

extreme range is perhaps explained because its

main distribution is east-west and in regions relatively unaffected by Pleistocene glaciation.

Arceuthohium minutissimum is a markedly reduced

species that is known only on Pinus griffithii in the

western Himalayas. In size, systemic habit, and high

specificity it resembles A. pusillum and A. douglasii

of the New World. However, the similarity is

probably due to convergence rather than close

phyletic relationship.

The dwarf mistletoe known as A. chinense consists

of two taxa: A. chinense sensu stricto on Ahies and

Keteleeria and the newly described A. pini on Pinus

tahulaeformis (Hawksworth and Wiens 1970b).

Arceuthohium chinense has some verticillate branching (Kuijt 1970). Although the branching type has

not been determined for A. pini, specimens we have

seen showed no secondary branching. Both species

are poorly known, but they deserve intensive study

because they occur near the presumed area of origin

of the genus.

In the New World, Arceuthohium is widely distributed and is represented by 28 taxa. The known

range of Arceuthohium in North America (fig. 5) indi-
Figure 5.—Distribution of Arceuthobium in North America. The number of taxa in various areas is indicated. Two areas of highest concentration of taxa are in Durango, Mexico, and in the vicinity of Mount

Shasta in northern California. Outline map copyrighted by Denoyer-Geppert Co., Chicago (used by

permission).

10
cates that the genus is concentrated mainly in

Mexico and the western United States. Also indicated

in figure 5 is the number of taxa within various parts

of the range of the genus. For example, although

five taxa occur in Colorado, no more than three

are found within a local area. Two North American

regions have high numbers of taxa. In western

Durango, Mexico, nine taxa are found: A. hlumeri^ A.

douglasii, A. gillii subsp. nigrum, A. glohosum, A.

vaginatum subsp. vaginatum and durangense, A.

veriicilliflorum, A. rubrum and A. stridum] the latter

four are endemic to this region. The second area

where Arceuthohium is highly concentrated is in

northern California where these nine taxa occur:

A. ahietinum (both formae spéciales), A. cyanocarpum, A. californicum, A. douglasii, A. americanum, A. campylopodum, A. occidentale, and A.

tsugense.

Of the 17 taxa in the United States, all but one are

western, and six are endemic to the western United

States (table 1); 10 are found in California, 9 in

Arizona, and 8 in Oregon. The only eastern species,

A. pusillum, is found in Maine, Vermont, New

Hampshire, Massachusetts, Rhode Island, Connecticut, New York, New Jersey, Pennsylvania, Michigan, Wisconsin, and Minnesota.

Five of the species found in the United States also

occur in Canada :

A. americanum—Ontario, Manitoba, Saskatchewan, Alberta, British Columbia

A. douglasii—British Columbia

A. laricis—British Columbia

A. pusillum—Newfoundland, Prince Edward Island, Nova Scotia, New Brunswick, Quebec,

Ontario, Manitoba, Saskatchewan

A. tsugense—British Columbia.

Of the 15 taxa found in Mexico (table 2), 7 range

into the western United States. Dwarf mistletoes are

reported from 20 of the 32 States or territories in

Mexico, but information is lacking on the distribution of many Mexican taxa. Two species are found in

Guatemala, and one each in Honduras, British

Honduras, and Hispaniola.

Extension of Dwarf Mistletoe Ranges

No intercontinental transfer of dwarf mistletoes,

such as the accidental introduction of the European

Viscum album into California (Howell 1966), has

been reported. Occasionally, dwarf mistletoes have

become established outside their natural ranges,

when infected trees have been transplanted. Most

such introductions have been for relatively short

distances, but some from 100 to 300 miles have

TABLE 1.—Distribution o/Arceuthobium in the western United States

Taxon c3

1

<

tí

1

Ö

O

2

*tíu

s

6

o3

>

t—1

c3

Ö

tí

o

bßtí

'ao 43

o

03

O

o

tí

o

<

8

CO

03

X

A. ahietinum f. sp. concoloris

ahietinum f. sp. magnificae

americanum -_-.

apachecum

X

"x"

X

X

X

X

X

X

X X X

4

4

1 X X X X X

A X

X

X

A hlumeri

A calijornicum

, campylopodum

, cyanocarpum

, divaricatum

, douglasii

, gillii suhsp. giïlii _ _

"x"

X

X

1

"x "

X

X

X

X

X

A X

X

X

X

X

A X X X X

X

X

X

X

X

X

X

X

X

A X

X

2

X

A,

A

X X X

A , laricis

microcarpum

X X X X

A . X X

A . occidentale

, tsugense

, vaginatum subsp. cryptopodum

""x"______ "x"" X

A X

A. . X X X X X

Niimbfir of taxa -_ 1 6 8 10 5 5 4 3 6 5 9 5 2

Note: Data marked by leaders (__) not known in State.

^ Reported in State but locality unknown.

2 Probably occurs within State.

11
TABLE 2.—Known distribution of Arceuthobium in Mexico (dwarf mistletoe records are available for onlij 20 of

the 32 States)

•^

§

Estado

535

OH

n

X2

e S

^ ^ -r^ Ö

OH

^

^ 1«^

X

Baja California N

Chiapis

Chihuahua

Coahuila

D. F

Durango

Hidalgo X

Jalisco X

Mexico X

Michoacán

Nayarit

X X

X

X

X

X

X

X X X

X

X

X X X

X

X

X

_x_

X

X

X

X

X

X

X

X

X X

Nuevo León X

Oaxaca

Puebla_

X X

X X

X

X

X

X

X

Siiialoa X _x_

Sonora

Tamaulipas x

Tlaxcala X

Veracruz X X X

Zacatecas X

X

X

Note: Data marked by leaders (__) not known in State.

been reported. Examples of such transfer include

Arceuthobium occidentale on Pinus radiata at Stanford University and North Berkeley, California

(Peirce 1905, Offord 1946b), and our observations of

A. divaricatum on P. edulis in Albuquerque, New

Mexico; A. vaginatum subsp. cnjptopodum on P.

ponderosa in Denver, Colorado ; and A. microcarpum

on Picea pungens in Alpine, Arizona. Thomas (1954)

found A. pusillum in planted windbreak and ornamental Picea glauca in an area in southern Manitoba

where the closest natural infection is about 6 miles

away.^^ In all these instances, the dwarf mistletoes

matured fruit in the new areas, and in most cases had

spread to nearby planted trees of the same species.

Boyce (1961) emphasized that dwarf mistletoes

should be eradicated from the vicinity of conifer

nurseries to eliminate the possibility of introducing

these parasites into new areas on planting stock.

Because the holding period is longer, the danger

seems greater in nurseries where trees are grown as

ornamentals than in those for forest plantings. Weir

(1916b) found a dwarf mistletoe-infected Pinus

ponderosa in a denuded planting site near Wallace,

Idaho, and assumed that the tree was infected at the

nursery near Boulder, Alontana. The only dwarf

mistletoe that occurs in this area of Montana is the

lodgepole pine dwarf mistletoe, A. americanum,

which sometimes occurs on Pinus ponderosa. The

longest range extension reported to date^^ is a small

dwarf mistletoe-infected Picea pungens found in a

nursery near Riverside, Cahfornia. The parasite,

not yet definitely identified, appears to be Arceuthobium tsugense. The tree was presumably infected

in a nursery near Portland, Oregon.

Relation of Parasite to Host Distribution

The distribution of a dwarf mistletoe is generally

centered within the range of its principal host or

hosts. Probably climatic history is primarily involved in the four exceptions to this ''central

distribution'^ rule:

ARCEUTHOBIUM PUSILLUM on PICEA MARIANA and

P. GLAUCA.—Possibly this spruce dwarf mistletoe

^2 Personal communication, J. G. Laut, Canada Dep.

Fisheries and Forestry, 1969.

13 Personal communication, Dr. L. J. Farmer, California

Department of Agriculture, 1969.

12
was centrally distributed within the range of its

hosts prior to the Wisconsin glaciations, but now

only the southern ranges of spruce range are infected.

Picea glauca and P. mariana migrated northward

after the glacial period faster than the dwarf mistletoe, and vast areas in the northern parts of the ranges

of these spruces are now free of A. pusillum. Whether

the mistletoe has reached its northern climatic limit

or is still migrating northward is not known.

ARCEUTHOBIUM MICROCARPUM on PICEA ENGELMANNii and P. PUNGENS.—This western spruce

dwarf mistletoe occurs in western montane coniferous

forests near the southern limits of its hosts' ranges.

A. microcarpum is confined to four general regions in

Southwestern United States: (1) the north rim of the

Grand Canyon, (2) the San Francisco peaks, (3) the

Graham mountains, and (4) the eastern IMogollon

Rim. The distribution of this dwarf mistletoe

appears relictual on spruces in the Southwest.

Perhaps A. microcarpum once had a more northern

distribution, but for unknown reasons did not survive

in the higher latitudes where the two spruces are now

most abundant. The phyletic affinities of A. microcarpum are anomalous, but since it does not seem

to be closely related to the other dwarf mistletoes

now in the Southwest, the spruce dwarf mistletoe

probably did not evolve from extant species presently in the region. Arceuthobium microcarpum

probably evolved in another area, migrated southward, and eventually became isolated in the southern

parts of its hosts' ranges.

ARCEUTHOBIUM AMERICANUM on Pmus BANKSIANA.—The range of A. americanum coincides closely

with that of Pinus contorta which is its major host.

However, the parasite is also common on Pinus

banksiana, but only in the western parts of this tree's

range. Jack pine extends for about 1,500 miles east

from the known limits of the mistletoe to western

Ontario. Presumably, A. americanum evolved as a

principal parasite of Pinus contorta and spread

relatively recently to P. banksiana through central

Alberta where the two trees occur together and

frequently hybridize. Possibly A. americanum is still

spreading eastward on P. banksiana. This situation

is discussed in more detail in the Formal Taxonomy

section.

ARCEUTHOBIUM ABIETINUM f. sp. CONCOLORIS on

ABIES CONCOLOR.—The main distribution of A.

abietinum f. sp. concoloris is in California where

Abies concolor obtains its maximum development.

However, the host extends for about 500 miles

beyond the known eastern limits of the parasite in

Grand Canyon, Arizona, and in southern Utah.

Possibly the scattered distribution of Abies concolor

in the Great Basin and Southern Rockies has

prevented the spread oí A. abietinum into these areas.

Sympatry

We consider sympatry (the occurrence of two

taxa together) to be important in the taxonomic

understanding of Arceuthobium. If two taxa are sympatric and if their flowering periods overlap, hybridization of the two taxa should occur if they are

genetically compatible and have mutual pollinators.

However, as will be discussed later, no evidence of

hybridization exists in Arceuthobium.

Dwarf mistletoes observed within 100 feet of each

other are arbitrarily considered to be sympatric

(fig. 6) because interspecific pollinations should be

possible within such limits. However, observations of

other taxa within }i mile are also shown in the

figure. Little is known of the effective distance of

pollen transfer in Arceuthobium, but the limited data

available suggest that the 100-foot distance may be

conservative. Potter and Rowley (1960) found a

single Arceuthobium pollen grain at a station in the

treeless San Augustin Plains of New Mexico.

Judging by the locality and time of year when the

grain was caught, we estimate that the closest dwarf

mistletoe populations are at least 3 miles away.

Also Leopold^^ discovered Arceuthobium pollen in

8 of 11 pollen traps in a transect across Searles Lake

in the Alojave Desert in California. The closest

Arceuthobium has not been determined but the

closest probable host in this region, Pinus monophylla, occurs on Argus Peak, about 10 miles north

of Searles Lake.

Most North American taxa are known to be

sympatric with at least one other member of the

genus except A. hondurense, A. bicarinatum, and

A. guatemalenscy although the latter may occur with

A. globosum in Guatemala. Nothing is known of

sympatry in the Old World species but A. minutissimum and A. oxycedri possibly occur together in the

Indian Himalayas where their elevational ranges

overlap (Brandis 1907). Also, A. chinense and A. pini

may be sympatric in southwestern China.

We observed several instances where three species

of Arceuthobium occur sympatrically, and one case

where four species were involved (table 3).

An interesting aspect of dwarf mistletoe parasitism

is the occurrence of two species on an individual host

tree (table 4). This phenomenon is rare, perhaps

because of an exclusion principle in which infection

of a tree by a secondary parasite is rare if the principal parasite is present (see p. 25).

^^ Personal communication, Dr. Estella B. Leopold, U.S.

Geological Survey, 1970.

13
SYMPATRY

• Within 100 feet

A Within 1/4 mile

1

1

1

1

1

Í

1

{Q.

(A

n

3

f

1—

1

3

1

1 I

1 1 t

1

1

o 1'il

î

i

I

6.

w

.o

3

w

1

&

1 t

.o1

1

1

1

1 1

t 1 i

i

1

1

1

1

1

t1

1

1

1

1

io

6. M

1

1

Â. abietinum f. sp, concoloris • A • A • •

A.abietinum f. sp. magnificae A • 8•

A. a bieti s - re/ígiosae • • M

A.americanum • • • • • M •

A. apachecum • • A Ä

A. blumeri •• • •

«i •

A.californicum m A •

A.campylopodum • • A ííiíií• • •

A.cyanocarpum • WH •

A. divaricatum • W: A A

A. douglasii • •• H •• A •• •• •

A. gill il %Mb%p,gi/lii • • W: •

A. gillii subsp. nigrum •Ü

A. globosum • II• •

A.laricis ¡i • •

A.microcarpum • miííííí • %

A. occidentale A M A

A. pusi Hum •

A. rubrum •^

A.strictum • !»

A. tsugense M A • A

A. vaginatum subsp. vaginatum • • • Á • •

A.vaginatum subsp. cryptopodum ^1 • • ••• • ••• •

A. vaginatum subsp. durangense Ä •

A. verticilliflorum •

Figure 6.—Sympatry of the North American dwarf mistletoes.

14
TABLE 3.—Instances of observed sympatry of three or four species 0/Arceuthobium {occurrence within 100 feet)

Dwarf mistletoes Hosts Locality

THREE SPECIES:

A. abietinum f. sp. concoloris Abies concolor Grand Canyon, Arizona

A. douglasii Pseudotsuga menziesii

A. vaginatum subsp. cryptopodum Pinus ponderosa var. scopulorum

A. cyanocarpum Pinus flexilis Bryce Canj^on, Utah

A. douglasii Pseudotsuga menziesii

A. vaginatum subsp. cryptopodum Pinus ponderosa var. scopulorum

A. apachecum Pinus strobiformis Mogollón, New Mexico

A. douglasii Pseudotsuga menziesii

A. vaginatum subsp. cryptopodum Pinus ponderosa var. scopulorum

A. apachecum Pinus strobiformis Alpine, Arizona

A. douglasii Pseudotsuga menziesii

A. microcarpum Picea pungens

A. americanum Pinus contorta var. latifolia Tieton Lake, Washington^

A. douglasii Pseudotsuga menziesii

A. laricis Larix occidentalis

A. abietis-religiosae Abies vejarii Cerro Potosí, Nuevo León, Mexico

A. blumeri Pinus strobiformis

A. vaginatum subsp. vaginatum Pinus ponderosa var. arizonica

A. blumeri Pinus strobiformis El Salto, Durango, Mexico

A. globosum Pinus cooperi

A. vaginatum subsp. vaginatum Pinus cooperi

A. blumeri Pinus strobiformis La Junta, Chihuahua, Mexico

A. gillii subsp. gillii Pinus leiophylla

A. vaginatum subsp. vaginatum Pinus ponderosa var. arizonica

FOUR SPECIES:

A. apachecum Pinus strobiformis Big Lake, Arizona

A. douglasii Pseudotsuga menziesii

A. microcarpum Picea pungens

A. vaginatum subsp. cryptopodum Pinus ponderosa var. scopulorum

^ Also observed near Growden, Washington by Iild F. Wicker, personal communication, 1969.

TABLE 4.—Known instances of dual parasitism in Arceuthobium and Arceuthobium with other mistletoes

Mistletoes Host Locality Reference

ARCEUTHOBIUM:

A. americanum & A. campylopodum Pinus ponderosa Idaho Weir 1916a

Do. Pinus contorta do. Do.

A. americanum & A. laricis do. British Columbia Kuijt 1954

Do. do. Washington Wicker^

Do. do. Montana Hawksworth and Wicker^

Do. Pinus sylvestris Washington Graham and Leaphart 1961

A. americanum & A. vaginatum subsp. Pinus ponderosa Colorado Hawksworth and Peterson 1959

cryptopoduTn

Do. Pinus contorta do. ^

A. cyanocarpum & A. vaginatum subsp. Pinus ponderosa do. 2

cryptopodum

A. globosum & A. vaginatum subsp. Pinus spp. Mexico Hawksworth and Wiens 1965

vaginatum

A. rubrum& A. vaginatum aubsp. Pinus herrerai Durango, Mexico ^

durangense

^ Personal communication, Ed F. Wicker, 1969.

2 First reported in this paper.

15
TABLE 4.—Knonn instances of dual parasitism in Arceuthobium and Arceuthobium with other mistletoes

Mistletoes Host Locality Reference

ARCEUTHOBIUM AND OTHER MISTLETOES:

A. abietinum f. sp. concoloris & Phoraden- Abies concolor

dron holleanum subsp. pauciflorum

A. bicarinatum & Dendropemon Pinus occidentalis

pycnophyllus

California Gill 1935 and our observations

Dominican Republic ^

^ Personal communication, Ed F. Wicker, 1969.

2 First reported in this paper.

HOST RELATIONSHIPS

Natural Hosts

A summary of the principal host groups of the 32

dwarf mistletoes (fig. 7) includes the four Old World

species. Each species is restricted to a certain host

group, with the exceptions of ^. occidentale (on pines

of sections Insignes, Oocarpae, and Sahinianae),

A. glohosum (on pines of sections Australes, Oocarpae,

and Ponderosae) and the poorly known A. chinense

(on Abies and Keteleeria).

Although previous dwarf mistletoe literature has

mentioned the susceptibility of trees in terms of

common, principal, uncommon, and rare hosts, these

classifications have not been specifically defined.

The susceptibility classes that we devised are based

on the determination of an '^infection factor " or the

percentage of trees of the species in question that are

infected wdthin 20 feet of heavily infected^^ principal,

or main, hosts of a dwarf mistletoe. Infection-factor

determinations should be made only in stands older

than 20 to 30 years.

This classification, based entirely on natural

susceptibility, may not necessarily reflect the abundance of a dw^arf mistletoe on a particular host. For

example, we have classed Pinus cembroides as a

principal host of A. divaricatum (table 5). This tree

^^ Heavily infected trees are class 5 or 6 of the 6-class system

described by Hawksworth (1961a). For this system the live

crown is divided into thirds, and each third is rated as: 0, no

mistletoe; 1, light mistletoe (less than one-half of branches

infected); and 2, heavy mistletoe (more than one-half of

branches infected). The ratings of each third are added to

obtain a total for the tree. For example, a tree heavily infected

in the lower one-third of the crown, lightly infected in the

middle one-third, and not infected in the upper third would

be class 3. A tree heavily infected in each third would be class

6.

rarely occurs within the range of the dwarf mistletoe,

but it is as susceptible as the other pinyons in the

few areas where this host-parasite combination is

found.

The five susceptibility classes used here

Host class

I. Principal

II. Secondary

III. Occasional (or

tertiary)

IV. Rare

V. Immune

Infectionfactor

At least 90 percent;

usually nearly

100 percent

50 to 90 percent

5 to 50 percent

More than 0 but less

than 5 percent

0

are:

Remarks

Uninfected trees are

seldom found

within the 20-foot

zone unless they

are small and

stunted

Potential host trees

not infected even

where the dwarf

mistletoe in question is common

Dwarf mistletoe parasitism is a dynamic process,

and some variation is to be expected. A tree species

that is heavily infected in one area may sometimes be

less severely parasitized in another part of the

dwarf mistletoe's range. This could be due to ecotypic

variation in the host or parasite or to environmental

factors. Also, a tree species may be more frequently

infected when it occurs in certain habitats. For

example, Daubenmire (1961) found that ponderosa

pine w^as commonly parasitized by Arceuthobium

campylopodum on habitat types where the tree was

associated with xerophytic grasses but not on the

more mesic habitat types associated with the

shrubs Physocarpus and Symphoricarpos. In spite of

such occasional differences, however, the relatively

consistent host relationships of each dw^arf mistletoe

make overall susceptibility ranking meaningful and

useful.

16
ARCEUTHOBIUM

PINUS

fHAPLOXYLONK

PINUS

(DIPLOXYLON)

OTHER GENERA

m

•

c

'S.

•

£

1

^ 1 1

j 1 i

1 1 1

{

1 1 }

A.abietínum f. sp. concoloris •

A.labietinum f. sp. magnificat •

A. abistis - religiosas •

A.americanum •

A.apachecum •

A. bicarinafum •

A. blumeri •

A.californicum •

A.campylopodum •

A. chínense • •

1 A.cyanocarpum •

A, divaricafum •

A.douglasiJ •

A.gillil tubsp.gillii •

A.gillh subsp.nigrum •

A.globosum • • •

A.guatemalense •

A. hondurense •

A.laricis •

A.microcarpum •

A,minufissimum •

A. occidentale • •

A. oxycedri •

A, pini •

A. pusi Hum •

A.rubrum •

A.strictum •

A Isugense •

A. vaginaturn su bsp. vaginatum •

A. vaginatums\xbsp.cryptopodum •

A. yaginatumsúbsp. durangense •

A. verticilliflorum • 1 1

Figure 7.—Principal hosts of the New and Old World dwarf mistletoes. (Divisions of Diploxylon according to Critchfield and Little 1966.)

17
TABLE b,-—Susceptibility of hosts of North American dwarf mistletoes based on natural infection of native trees

(a question mark indicates that we have not determined the appropriate susceptibility class or that the reported

host-parasite combination has not been confirmed)

Arceuthohium Principal host Secondary host Occasional host Rare host Immune^

A. abietis-religiosae Abies religiosa

Abies vejarii

Pinus hariwegii

Pinus leiophylla var.

leiophylla

Pinus montezumae

Pinus ponderosa var.

arizonica

Pseudotsuga menziesii

A. abietinum f.

concoloris

sp. Abies concolor

Abies grandis

Abies amabilis?

Picea breweriana

Abies lasiocarpa

var. lasiocarpa

Pinus contorta subsp.

murrayana

Pinus lambertiana

Pinus moniicola

Abies magnifica

Picea engelmannii

Picea pungens

Pinus flexilis

Pinus monophylla

Pinus muricata

Pinus ponderosa var.

ponderosa

Pinus ponderosa var.

scopulorum

Pinus washoensis

Pseudotsuga menziesii

Tsuga heterophylla

A. abietinum Î. sp.

magnificae

Abies magnifica

Abies procera?

Abies concolor

Abies grandis

Tsuga meriensiana

A. americanum Pinus banksiana

Pinus contorta

subsp. latifolia

Pinus contorta

subsp. murrayana

Pinus contorta

subsp. contorta?

Pinus ponderosa

var. scopulorum

Pinus albicaulis Picea engelma,nnii Abies grandis

Pinus aristata Picea glauca Abies lasiocarpa var.

Pinus flexilis Picea pungens lasiocarpa

Pinus ponderosa Pinus attenuata Abies magnifica

var. ponderosa Pseudotsuga menziesii Larix occidentalis

Tsuga mertensiana

A. apachecum Pinus strobiformis Abies concolor

Abies lasiocarpa var.

arizonica

Picea pungens

Pinus edulis

Pinus engelmannii

Pinus ponderosa var.

arizonica

Pinus ponderosa var.

scopulorum

Pseudotsuga menziesii

^ Includes only members of the Pinaceae in a particular stand, but associated Cuppressaceae or Taxodiaceae are likewise not

infected.

18
TABLE 5.—Susceptibility of hosts of North American dwarf mistletoes based on natural infection of native trees

(a question mark indicates that we have not determined the appropriate susceptibility class or that the reported

host-parasite combination has not been confirmed)—Continued

Arceuthohium Principal host Secondary host Occasional host Rare host Immune^

A. hicarinatum Pinus occidentalis

A, hlumeri Pinus strobiformis Abies vejarii

Pinus cooperi

Pinus engelmannii

Pinus leiophylla var.

leiophylla

Pinus ponderosa var.

arizonica

Pinus ponderosa var.

scopulorum

Pseudotsuga menziesii

A. californicum Pinus lambertiana Pinus montícola Picea breweriana Abies concolor

Pinus attenuata

Pinus coulteri

Pinus ponderosa var.

ponderosa

A. campylopodum Pinus ponderosa

var. ponderosa

Pinus Jeffreyi

Pinus attenuata

Pinus coulteri

Pinus ponderosa

var. scopulorum

Pinus contorta

subsp. contorta

Pinus contorta

subsp. latifolia

Pinus contorta

subsp. murrayana

Pinus lambertiana Abies concolor

Abies grandis

Pinus monophylla

Pinus montícola

Pinus quadrifolia

Pinus sabiniana

Pseudotsuga menziesii

A. cyanocarpum Pinus flexilis

Pinus aristata

Pinus albicaulis Pinus monticola Picea engelmannii?

Pinus balfouriana

Pinus contorta

subsp. latifolia

Pinus ponderosa

var. scopulorum

Abies lasiocarpa var.

lasiocarpa

Pinus lambertiana

Pinus strobiformis

Pseudotsuga menziesii

A. divaricatum Pinus edulis

Pinus monophylla

Pinus quadrifolia

Pinus cembroides

Pinus Jeffreyi

Pinus ponderosa var.

scopulorum

Pseudotsuga menziesii

A. douglasii Pseudotsuga Abies lasiocarpa Abies grandis

var. arizonica

Abies concolor

Abies lasiocarpa

var. lasiocarpa

Picea engelmannii

Picea pungens

Pinus flexilis

Abies magnifica

Abies vejarii

Larix occidentalis

Pinus edulis

Pinus ponderosa var.

ponderosa

Pinus ponderosa var.

scopulorum

Pinus strobiformis

1 Includes only members of the Pinaceae in a particular stand, but associated Cuppressaceae or Taxodiaceae are likewise not

infected.

19
TABLE 5.—Susceptibility of hosts of North American dwarf mistletoes based on natural infection of native trees

(a question mark indicates that we have not determined the appropriate susceptibility class or that the reported

host-parasite combination has not been confirmed)—Continued

Arceuthobium Principal host Secondary host Occasional host Rare host Immune^

A. gillii subsp. Pinus leiophylla Pinus leiophylla

gillii var, chihuahuana var. leiophylla

Pinus lumholtzii

Pinus ponderosa

var. arizonica

Pinus ponderosa

var. scopulorum

Pinus cembroides

Pinus engelmannii

Pinus strohiformis

A. gillii subsp.

nigrum

Pinus leiophylla

var. leiophylla

Pinus leiophylla

var. chihuahuana

Pinus lumholtzii

Pinus teocote

Pinus montezumae Pinus cooperi

Pinus engelmannii

A. globosum Pinus cooperi

Pinus douglasiana

Pinus durangensis

Pinus engelmannii

Pinus hartwegii

Pinus lawsonii

Pinus michoacana

Pinus montezumae

Pinus pringlei

Pinus pseudostrohus

Pinus rudis

Pinus tenuifolia

Pinus ponderosa

var. arizonica

Cupressus sp.?

Pinus teocote

Abies guatemalensis

Abies religiosa

Pinus leiophylla var.

leiophylla

A. guatemalense Pinus ayacahuite Abies guatemalensis

Pinus pseudostrobus

A. hondurense Pinus oocarpa Pinus pseudostrobus?

A. laricis Larix occidentalis Abies lasiocarpa

var. lasiocarpa

Larix lyallii?

Pinus contorta

subsp. latifolia

Tsuga mertensiana

Picea engelmannii

Pinus albicaulis

Pinus ponderosa

var. ponderosa

Abies grandis

Pinus monticola

Abies concolor

Pinus contorta subsp.

murrayana

Pseudotsuga menziesii

Tsuga heterophylla

A. microcarpum Picea engelmannii

Picea pungens

Abies lasiocarpa

var. arizonica

Abies concolor

Abies lasiocarpa var.

lasiocarpa

Pinus ponderosa var.

scopulorum

Pinus strobiformis

Pseudotsuga menziesii

A. occidentale Pinus sabiniana

Pinus radiata

Pinus muricata

Pinus attenuata

Pinus coulteri

Pinus contorta

subsp. bolanderi

Abies grandis

Pinus ponderosa var.

ponderosa

Pseudotsuga menziesii

Tsuga heterophylla

^ Includes only members of the Pinaceae in a particular stand, but associated Cuppressaceae or Taxodiaceae are likewise not

infected.

20
TABLE 5.—Susceptibility of hosts of North American dwarf mistletoes based on natural infection of native trees

{a question mark indicates that we have not determined the appropriate susceptibility class or that the reported

host-parasite combination has not been confirmed)—Continued

Arceuthohium Principal host Secondary host Occasional host Rare host Immune^

A. pusillum Picea mariana

Picea glauca

Picea rubens Larix laricina Pinus strobus

Pinus resinosa

Pinus banksiana

Abies balsamea

A. ruhrum . Pinus cooperi

Pinus durangensis

Pinus engelmannii

Pinus teocote

Pinus herrerai

Pinus strobiformis

A. strictum Pinus leiophylla

var. chihuahuana

Pinus teocote Pinus engelmannii

A. tsugense Tsuga heterophylla Abies amabilis Abies grandis Picea engelmannii Abies magnifica

Tsuga mertensiana Abies lasiocarpa

var. lasiocarpa

Abies procera

Pinus albicaulis

Pinus contorta

subsp. contorta

Pinus monticola

Picea breweriana Picea sitchensis Larix occidentalis

Pinus contorta subsp.

murrayana

Pinus Jeff reyi

Pseudotsuga menziesii

A. vaginatum Pinus durangensis Pinus culminicola Pinus leiophylla var.

subsp. z; aginatum Pinus cooperi

Pinus engelmannii

Pinus hartwegii

Pinus herrerai

Pinus montezumae

Pinus ponderosa

var. arizonica

Pinus ponderosa

var. scopulorum

Pinus rudis

Pinus lawsonii

chihuahuana

Pinus leiophylla var.

leiophylla

Pinus lumholtzii

Pinus strobiformis

Pinus teocote

Pseudotsuga menziesii

A. vaginatum Pinus ponderosa Pinus contorta Pinus flexilis Abies concolor

subsp. crypto- var. scopulorum subsp. latifolia Pinus strobiformis Abies lasiocarpa var.

podum Pinus ponderosa

var. arizonica

Pinus engelmannii

Pinus arizonica

var. stormiae

Pinus aristata arizonica

Picea pungens

Pinus cembroides

Pinus edulis

Pinus leiophylla var.

chihuahuana

Pseudotsuga menziesii

A. vaginatum Pinus montezumae Pinus herrerai? Pinus leiophylla var.

subsp. durangense Pinus durangensis leiophylla

Pinus lumholtzii

1 Includes only members of the Pinaceae in a particular stand, but associated Cuppressaceae or Taxodiaceae are likewise not

infected.

21
TABLE 5.—Susceptibility of hosts of North American dwarf mistletoes based on natural infection of native trees

(a question mark indicates that we have not determined the appropriate susceptibility class or that the reported

host-parasite combination has not been confirmed)—Continued

Arceuthohium Principal host Secondary host Occasional host Rare host Immune^

A. verticilliflorum Pinus cooperi

Pinus engelmannii

Pinus leiophylla var.

chihuahuana

Pinus leiophylla var.

leiophylla

Pinus teocote

1 Includes only members of the Pinaceae in a particular stand, but associated Cuppressaceae or Taxodiaceae are likewise not

infected.

Classification of host susceptibility for each North

American dwarf mistletoe (table 5) is based primarily

on actual infection-factor determinations. In some

cases, however, no data are available, and we have

placed trees in the susceptibility class which seems

most appropriate on the basis of our field experience.

No such susceptibility classification is currently

possible for the Old World species, because we have

not studied all of them in the field.

Also, when we rated the trees by susceptibility

classes, we based our decision on the most common

condition for a particular tree throughout the range

of the dwarf mistletoe in question. This, of course,

may not necessarily apply in individual instances.

For example, near Snow Valley in the San Bernardino Mountains, California, Pinus coulteri is heavily

infected by Arceuthobium campylopodum, but at

lower elevations (where the mistletoe is severe on

associated P. attenuata), P. coulteri is only occasionally attacked. We have placed P. coulteri in the

"secondary host^' category because it is typical of

most situations where this host-parasite combination

occurs. Variation as extreme as this example is

exceptional and most dwarf mistletoe populations

readily fall into the susceptibility classes indicated

in table 5.

As shown in table 5, some trees are parasitized by

several dwarf mistletoes: eight are known on Pinus

ponderosa (table 6), eight on Pinus contorta (table 7),

and six on Pinus engelmannii in Mexico (Hawksworth and Wiens 1965). Some dwarf mistletoes are

quite specific, for example, A. blumeri and A.

apachecum, which are restricted to Pinus strobiformis, but occur in different parts of the host range.

Others have broad host ranges, such as A. laricis,

which not only parasitizes Larix but also Abies,

Pinus, Picea, and Tsuga. The principal host of A.

douglasii is Pseudotsuga menziesii, and it is the only

dwarf mistletoe that parasitizes Pseudotsuga. Arceuthobium douglasii, however also grows on Abies and

Picea,

The five dwarf mistletoes that occur principally on

the North American white pines (table 8) are: A.

cyanocarpum on Pinus flexilis and P. aristata; A.

californicum on P. lambertiana; A. guatemalense on

P. ayacahuite; and A. blumeri and A. apachecum on

P. strobiformis. The Central American Pinus

chiapensis is the only member of this group not

known to be parasitized.

Pinus monticola is the only widespread western

white pine that is not a principal host for any dwarf

mistletoe. Although six taxa parasitize this tree, these

are mainly crossover situations (table 8). It may be

of evolutionary significance that none of the members of the Pinus chiapensis-strobus-monticola phylad

(Andresen 1966) has been colonized as a principal

host by Arceuthobium. This may suggest that the

phylad originated in an area outside the range of

Arceuthobium. If this is true, P. monticola may not

have occupied its present range until after the

principal radiation of Arceuthobium onto the various

conifers of the area had already occurred. This

could explain why various dwarf mistletoes sometimes utilize P. monticola as a lesser host, and why it

is not the principal host of any species.

Dwarf mistletoes parasitize nearly all western

firs {Abies spp.) (table 9). Only Arceuthobium

abietinum in the western United States and A.

abietis-religiosae of Mexico, however, are principal

parasites of Abies. Arceuthobium abietis-religiosae

is known only from Abies religiosa and A. vejarii,

but is to be expected on other Mexican true firs.

In addition to the firs listed in table 9, we have

examined one collection on the rare bristlecone fir

Abies bracteata, from the Santa Lucia Mountains of

California. We believe that the specimen is a crossover infection of a pine dwarf mistletoe, probably A.

occidentale. The specimen is in the Weir collection

at the University of Illinois, and was obtained by

Raymond in 1915. Positive identification is impossible because the specimen is fragmentary and

poorly documented.

22
AU North American spruces, except the recently

described Mexican species Picea chihuahuana and

P. mexicana (Martínez 1963), are known hosts for

Arceuthohium (table 10). Picea sitchensis, however,

is a very rare host. We have a recent collection on this

tree from Oregon and this host-parasite combination

has been reported from Alaska (Laurent 1966)

and British Columbia (Molnar, Harris, Ross, and

Ginns 1968).

TABLE 6.—Susceptibility of members of the Pinus ponderosa complex to

dwarf mistletoes^' ^

Arceuthohium

Pinus ponderosa variety

ponderosa scopulorum arizonica "stormiae'^^

A. americanum III

A. cyanocarpum

A. campylopodum I

A. gillii subsp. gillii

A. glohosum

A. laricis III

A. vaginatum subsp.

cryptopodum

A. vaginatum suhsp. vaginatum

II

IV

II

IV IV

II

I

I

11 = Principal host, II = secondary host, III = occasional host, and IV = rare

host; "-J' dwarf mistletoe not observed within range of the tree.

2 In addition, Smith and Craig (1968) have artificially infected Pinus ponderosa

(presumably var. ponderosa) with Arceuthohium tsugense.

3 The tree known in northern Mexico as Pinus arizonica Engelm. var. stormiae

Martinez (1948) has not been formally described as a variety of Pinus ponderosa, but

our field studies in Coahuila indicate that it is quite distinct from the tree known in the

United States as Pinus ponderosa var. arizonica.

TABLE 7.—Susceptibility of members of the Pinus contorta complex to dwarf

mistletoes^

Pinus contorta subspecies

Arceuthohium

contorta latifolia murrayana holanderi

A. ahietinum f. sp. concoloris.

A. americanum

A. campylopodum

A. cyanocarpum

A. laricis

A. occidentale

A. tsugenseA. vaginatum subsp.

cryptopodum

III

II

__ IV

I I

III III

IV __

II V

III

III

^ I = Principal host, II = secondary host. III = occasional host, IV = rare host,

and V = immune; ''__" dwarf mistletoe not observed within range of the tree.

2 Presumably on this subspecies in the Oregon Mountains, southern Josephine

County, Oregon (Weir 1917), but this has not been confirmed.

23
TABLE 8.—Susceptibility of North American white pines to dwarf mistletoes^

Pinus

Arceuthobium

albicaulis aristata* ayacahuite halfouriana flexilis lambertiana monticola strohiformis strohus

A. ahietinu7n f. sp. concoloris

A. americanum III III

A. apachecum

A. blumcri

A. californicum

A. campylopodum

A. cyanocarpum II I

A. douglasii

A. guatemalense

A. laricis III

A. pusillum

A. tsugense II

A. vaginatum subsp.

cryptopodum __ III

IV

V

FTT

IV IV --

-- -- I

I

I II

__ IV V

I V III V

V -- -- V

-- -- IV

II ;;

IV

IV IV

^ I = Principal host, II = secondary host, III = occasional host, IV = rare host, and V = immune; "__" dwarf mistletoe

not observed within range of the tree.

2 Susceptible in greenhouse inoculations at Fort Collins, Colorado.

* Recent studies (Bailey, D. K., Phytogeography and taxonomy of Pinus subsection Balfourianac, Ann. Mo. Bot. Gard. 57:

210-249, 1970) have shown that ^'Pinus aristata^' as formerly understood comprises two species: Pinus longacva D. K. Bailey in

the Great Basin (California, Nevada, and Utah) and Pinus aristata sensu stricto in the Rocky Mountains (Colorado, New IVlexico,

and Arizona). Pinus longaeva is frequently parasitized by Arceuthobium cyanocarpum in southern Utah and Nevada. Pinus

aristata is an occasional host for A. americanum and A. vaginatum subsp. cryptopodum in Colorado. Arceuthobium cyanocarpum

is not known on Pinus aristata in Colorado or New Mexico but it may be the taxon on this tree in Arizona (see p. 117).

TABLE 9.—Susceptibility of western North American and Mexican species of true firs (Abies) to dwarf mistletoes^

Abies

Arceuthobium

aînabilis concolor

lasiocarpa lasiocarpa

grandis var. var. ynagnifica procera

lasiocarpa arizonica

religiosa vejarii

A. abietinum f. sp. concoloris II?

A. abietinum f. sp.

magnificae

A. abietis-religiosae

A. laricis

A. microcarpum

A. tsugense II

A. douglasii

I I III -- V

V V -- I

V IV II

V __ V IV __

III II __ V

V III IV II V

I?

II

^ I = Principal host, II = secondary host. III = occasional host, IV = rare host, and V = immune;

not observed within range of the tree.

dwarf mistletoe

24
TABLE 10.—Susceptibility of North American spruces to dwarf mistletoes^

Arceuthohium

breweriana engelmannii

A. abietinum f. sp. concoloris.- - II

A. americanum

A. californicum III

A. cyanocarpum

A. douglasii

A. laricis

A. microcarpum

A. pusillum

A. tsugense III

V

IV

IV?

IV

III

I

IV

Picea

mañana pungens rubens sitchensis

IV

V

IV

IV

I

II

IV

1 j ^ Principal host, II = secondary host, III = occasional host, IV = rare host and V = immune; ''__" dwarf mistletoe

not observed within range of the tree.

2 On planted trees in Maine (U. S. Dep. Agr. 1960).

Extra-Limital and Unnatural Hosts

Several workers have inoculated dwarf mistletoes

on trees that do not occur naturally within the

range of a particular species of Arceuthohium

(extra-hmital hosts). The results of their studies,

summarized in table 11, are discussed further under

the dwarf mistletoes involved in the Formal Taxonomy section.

Perhaps conditions for infection and development of dwarf mistletoes are more favorable

under artificial than natural situations, and plants

grown under such conditions have occasionally

become estabhshed on hosts that are not naturally

infected. Weir (1918a) artificially grew A. campylopodum on Ahies concolor, and we grew A. cyanocarpum on Pinus strohiformis. In nature, these trees

are exposed to the two dwarf mistletoes, but we have

not found naturalinfection of them.

Inoculation experiments are most useful for determining susceptibility of extra-Jimital hosts. Such

studies do not reveal, however, how the parasite will

act in the natural environment of a new host. We

question whether hfe-cycle data could be based on

artificial inoculations, because of the accelerated

growth rates we noted in the greenhouse. For

example, when we inoculated Pinus strohiformis

with A. cyanocarpum in the greenhouse we obtained

shoots within 5 months, and flowering within 7

months, of the date of inoculation. In nature, A.

cyanocarpum shoots do not appear until 2 to 3 years

after infection, and flowering does not begin until

after at least one more year.

Observations of natural susceptibility of trees from

outside the range of a mistletoe are sometimes

economically important (table 12). For example.

Graham and Leaphart (1961) reported that plantation-grown Pinus sylvestris was so severely parasitized by the larch dwarf mistletoe {A. laricis) in

Washington, that this tree should not be planted in

areas where A. laricis occurs.

Nonhosts

Since the dwarf mistletoes parasitize so many

western conifers we have recorded those in the western United States and Mexican Pinaceae not yet

known to be naturally infected by Arceuthohium'}^

Ahies: (True firs)

durangensis

guatemalensis

hickeli

mexicana

oaxacana

Picea: (Spruces)

chihuahuana

mexicana

Pinus: (Pines)

chiapensis

greggii

maximartinezii

nelsonii

patula

pinceana

rzedowskii

torreyana

washoensis

Pseudotsuga: (Douglas-fir)

macrocarpa

Host-Parasite Relationships

A pecuHar feature of dwarf mistletoe parasitism,

at least in the central Rocky IMountains, is the

existence of a type of host exclusion between taxa

(Hawksworth 1968). If the principal parasite of a

certain host is present in an area, other dwarf16 Kuijt (1960a) .successfully inoculated Piniis torreyana

with A. occidentale; A. abietis-relig-iosae possibly parasitizes

Abies guatemalensis.

25
TABLE 11.—Extension of host ranges 0/Arceuthobium

as determined by artificial inoculations on unnatural

hosts

Host Dwarf mistletoe Reference

Abies

concolor A. campylopodum Weir 1918a

grandis do. Do.

Larix

europaca A. larids Do.

leptolepis do. Do.

occidentalis A. campylopodum Do.

Do. A. tsugense Smith 1970a

Picea

abies A. campylopodum Weir 1918a

Do. A. tsugense Smith 1965

glauca do. Do.

engelmannii do. Smith 1970b

pungens do. ?

1

Pinus

banksiana A. occidentale Hedgcock and

Hunt 1917

bungeana do. Do.

mugo (as P. A. campylopodum Weir 1918a

montana)

Do. A. americanum Do.

palustris A. occidentale Hedgcock and

Hunt 1917

Pinus

pinea do. Do.

ponderosa A. tsugense Smith and Craig

1968

radiaia do. Do.

resinosa A. campylopodum Weir 1918a

strobiformis A. cyanocarpum 1

strobus do. 1

sylvestris A. campylopodum Weir 1918a

Do. A. tsugense Smith and Craig

1968

torreyana A. occidentale Kuijt 1960a

virginiana do. Hedgcock and

Hunt 1917

1 suga

canadensis A. tsugense Weir 1918a

^ From studies reported in this paper.

mistletoes rarely parasitize that tree. Conversely,

if the principal parasite is absent, the chances of

parasitism by other dwarf mistletoes is greatly increased. This is exemplified by A. americanum

(principal host, in this instance, Pinus contorta

subsp. latifolia) and A. cyanocarpum (principal host,

Pinusfiexilis). In mixed stands where A. americanum

is present, parasitism of P. contorta by A. cyanocarpum has not been found. Conversely, in stands

where there is no A. americanum, A. cyanocarpum

frequently occurs on this host. A comparable, and

reciprocal, situation exists for P. flexilis. Another

example is the relative frequency of A. americanum

on Pinus ponderosa var. scopulorum and A. vaginatujn

subsp. cryptopodum on Pinus contorta, which shows a

similar relationship to that above (Hawksworth

1968).

Wiens (1962) has recorded a comparable situation

in Phoradendron parasitizing junipers. The explanation for this host exclusion principle is unknown and

presents a fascinating field for research.

The dwarf mistletoes seem to show an ''all or

nothing'' parasitism of trees that are rarely infected.

Such trees are usually immune, but when infection

does occur such individuals are often moderately

to highly parasitized. An example of this is the

parasitism of Picea engelmannii by A. americanum.

Usually less than 1 percent of the spruces are infected in stands where the mistletoe is common on

the associated Pinus contorta. However, the individual P. engelmannii trees that are parasitized

may bear 100 infections or more (Hawksworth and

Graham 1963).

Our field studies suggest that some very rare trees

have little resistance to dwarf mistletoes. For

example, Pinus culminicola is readily parasitized by

Arceuthobium vagin atum subsp. vaginatum, and

Picea breweriana is infected by A. abietinum f. sp.

concoloris^ (fig. 8), A. californicum, and A. tsugense

in every instance where we have seen them exposed

to these dwarf mistletoes. The situation for Abies

bracteata may be similar (because Arceuthobium has

been collected on it), but we have not observed this

tree associated with dwarf mistletoe. No dwarf

mistletoe occurs within the range of Pinus torreyana,

but Kuijt (1960a) inoculated this tree with A.

occidentale and found it susceptible.

Features that often indicate host-parasite incompatibility are poor (sometimes lack of) shoot

development and unusually large swelHngs at the

point of infection (table 13).

Host Reactions

The first external symptom that a dwarf mistletoe

has become established in a branch is usually a

swelling of the host tissues at the point of infection.

As the infection becomes older, the swelUngs enlarge

and eventually become fusiform. Typically, dwarf

mistletoe infection leads to the production of profuse,

dense masses of distorted host branches called

'Vitches' brooms'' (figs. 8-14).

Two basic types of witches' brooms are formed

(Kuijt 1960b, Hawksworth 1961a): (1) Systemic

(isophasic) types in which growth of the endophytic

system keeps pace with growth of the infected

26
TABLE 12.—Reports of extra-limital hosts naturally infected by dwarf

mistletoes

Host Arceuthobium Locality Reference

Cupressus

niacrocarpa A. oxycedri U.S.S.R. Zefirov 1955

Larix

europaea A. tsugense British Columbia Kuijt 1964

Picea

abies A. laricis Idaho U. S. Dep. Agr. 1963

pungens A. pusillum Maine U. S. Dep. Agr. 1960

Pinus

banksiana A. laricis Idaho Graham 1959a

pinaster A. campylopodum California Kuijt 1960b

resinosa A. laricis Idaho U. S. Dep. Agr. 1962

sylvestris do. Washington Graham and Leaphart

1961

Do. A. americanmn do. Do.

Do. do. Alberta Powell 1968

Do. A. campylopodum Idaho J. R. Weir, unpubl.^

^ Specimen at the University of Illinois.

branch, dwarf mistletoe shoots are scattered along

the host branch, often concentrated at the branch

girdles and, (2) nonsystemic (anisophasic) types in

which the shoots remain concentrated near the

original site of infection. These two types of broom

formation, however, are species constant and

therefore of taxonomic value. Arceuthobium americanum, A. douglasii, A. guatemalense, A. minutissimum, and A. pusillum consistently form systemic

brooms. Herbarium specimens revealthat A. chinense

TABLE 13.—Reported incompatible host-parasite

relationships

Arceuthobium Host Reference

A.. abietinum f. sp.

concoloris Pinus lambertiana R. F. Scharpfi

Do. Pinus contorta D. A. Grahami

A.. americanum

subsp. ynurrayana

Pinus albicaulis Weir 1918b

Do.

Do.

Picea pungens

Pseudotsuga

menziesii

2

R. S. Peterson!

A.. pusillum Larix laricina Tainter and French

1967

A,. tsugense

Do.

Larix europaea

Larix occidentalis

Kuijt 1964

R. B. Smithi

Do.

(inoculations)

Picea engelmannii 2

Do. Picea sitchensis Laurent 1966

! Personal communications, 1965-69.

2 First reported in this paper.

and A. pini also cause systemic brooms, but whether

this host reaction is consistent is not known. The nonsystemic type is much more common, although most

dwarf mistletoes will sometimes cause systemic

brooms. In a few host-parasite combinations, little or

no witches' brooms of either type are formed.

In Pinus contorta branches older than about 5

years, A. americanum shoots on systemic brooms are

formed only at the girdles and not on the segments

(Kuijt 1960b). Similarly, our limited observations of

systemic infections of A. abietis-religiosae on Abies

religiosa suggest that here, also, the dwarf mistletoe

shoots are produced only at the girdles.

Within a host genus, the parasite and not the host

determines the type of brooms formed (Hawksworth

1956a, Kuijt 1960b, Weir 1916). For example, the

witches' brooms formed by A. aynericanum on any

pine are basically similar, but are distinctly different

on spruce (Hawksw^orth and Graham 1963, Kuijt

1960b).

Although systemically infected branches are

usually immune to secondary infection, ]Muir (1968)

reported several secondary pistillate and staminate

infections on a pistillate systemic broom of A.

americanum on Pinus banksiana in northeastern

Alberta. The secondary infections were detectable

because of the location of their shoots, differences in

shoot color, and formation of swellings. We have

observed similar situations (one instance each) in the

following: A. americanum on Pinus contorta in

Colorado, A. vaginatum subsp. cryptopodum on P.

ponderosa in Arizona, and A. guatemalense on P.

ayacahuite in Guatemala.

27
Kuijt (1960b) reported that cones usually are not

produced on witches' brooms, particularly systemic

ones. Bonga (1964), however, noted fertile cones on

an aberrant broom in Picea mariana caused by

Arceuthohiwm pusillum. We have observed cones on

several systemic witches' brooms on Pinus contorta

(caused by A. americanum) but all were sterile.

Effects on Hosts

The dwarf mistletoes are serious pathogens of

North American forest trees, and in many parts of

the West they are the most damaging disease agents.

They not only cause extensive losses through direct

mortality, but also adversely affect growth rates

of infected trees. In addition, the dwarf mistletoes

reduce seed production and wood quality, and in

some hosts, particularly Abies and Tsuga, provide

entrance points for decay fungi. The extent of the

damage caused by these parasites is estimated to

total 3.2 bilUon board feet annually in the western

United States (Shea and Howard 1969). At 1970

prices, this annual loss totals about $75 million. The

commercially important trees that are most seriously

damaged are Pinus ponderosa, P. contorta, P.

banksiana, P. lambertiana, Pseudotsuga menziesii,

Abies magnifica, A. concolor, Larix occidentalis, and

Tsuga heterophylla. Arceuthobium pusillum, considered the most serious cause of timber loss in

Picea mariana in the Lake States, also causes serious

losses in Picea glauca along the coast of Maine.

Dwarf mistletoe infection reduces the tree's

growth rate in both height and diameter, but there

is no significant reduction until the upper half of the

tree's crown is parasitized. Then the growth rate

declines rapidly as the degree of infection in the

upper crown increases (Hawksworth 1961a).

Heavy dwarf mistletoe infection eventually kills

the tree. However, the length of time required for the

parasite to kill a tree varies considerably, and

depends on many factors: the host and dwarf

mistletoe species involved; the amount of infection

Hosts

Abies magnífica

Pinus contorta, P. banksiana

P. flexilis, P. alhicaulis

Pseudotsuga menziesii

Larix occidentalis

Picea engebnannii, P. pungens

Picea mariana, P. glauca

Pinus ponderosa

in a tree; the vigor of the tree; the ecological and

climatic situation under which the tree is growing;

and activity of secondary pests, particularly bark

beetles, which often attack and help kill heavily

infected trees. Some host-parasite combinations in

which tree mortality is particularly high are :

Dwarf mistletoe

A.abietinum f. sp. magnificae

A. americanum

A. cyanocarpum

A. douglasii

A. laricis

A. microcarpum

A. pusillum

A. vaginatum subsp.

cryptopodum

How dwarf mistletoes affect the growth of their

host trees is not fully understood. Important factors,

however, might be the appropriation of water,

minerals, and other nutrients by the parasite and

infected parts of the lower crown at the expense of

upper parts of the crown. Once a branch becomes

infected, it utilizes more than its share of nutrients.

Thus, the diameter growth of infected branches is

greatly enhanced. Infected branches may become

several times as large as uninfected branches in the

same whorl, and they tend to live long after the

adjacent uninfected branches are shaded out.

As progressively more and more nutrients are

appropriated by infected branches, the vigor of the

crown declines. Eventually, the effective photosynthetic surface area of noninfected branches is

reduced below that necessary to sustain the tree, and

death results.

Jjconard and Hull (1965) found that several dwarf

mistletoes in California obtained large quantities of

photosynthate from their hosts at all seasons of the

year. Although the dwarf mistletoes do contain

chlorophyll and can manufacture a small part of their

required carbohydrates, neither material is translocated to the host. Physiological work on the

water balance of dwarf mistletoe-infected trees is

needed.

MECHANISMS AND TRENDS OF EVOLUTION

The Genetic System and

Recombination Potential

Arceuthobium has a base chromosome number of

X = 14 (that is, the lowest extant haploid number).

Polyploidy is unknown. A base chromosome number

oi X = 14 is relatively high in the angiosperms, and

one might suspect that x = 14 is a tetraploid on a

base of seven. This is unlikely, however, because in

the Viscaceae as a whole the lowest known number

is x = 10 (in Viscum). Viscum shows an aneuploid

series ranging from x = 10-13 and may be the

primitive genus in the Viscaceae. The putatively

derived genera, PJioradendron, Korthalsella, and

Arceutliobium, are all characterized by chromosome

numbers of x = 14. Chromosome evolution in the

Viscaceae as a whole is discussed in detail by Wiens

and Barlow (1971). In view of the patterns of

28
Figure 8.—Effects of dwarf mistletoes on various hosts: A, Arceuthobium abietinum f. sp. concoloris on Picea

breweriana, severely broomed tree killed by the parasite, Siskiyou National Forest, Oregon; B, A. douglasii on

Pseudotsuga menziesii, most trees have been killed by the parasite, Mescalero Apache Reservation, New Mexico;

C, A. occidentale on Pinus sabiniana, no brooms formed even though trees are heavily infected, both trees died

within 2 years after the photograph was taken. North Fork, California; D, A. microcarpum on Picea pungens,

note the small brooms and the trees killed by the mistletoe. Grand Canyon National Park, Arizona.

29
Figure 9.—Witches' brooms caused by Arceuthobium

americanum: A, on Pinus ponderosa (left) and P.

contorta (right) note similarity of the brooms; B, on

Picea engelmannii, brooms are distinctly different

from those in pines caused by same mistletoe; C, on

Pinus contorta; (A, B, C, in Roosevelt National

Forest, Colorado.) D, on Pinus banksiana. Golden

Ridge, Saskatchewan; photograph by J. G. Laut,

Canadian Department of Fisheries and Forestry.

f|»i^^^^'P"^

30
Figure 10.—Witches' brooms on pines caused by five

dwarf mistletoes: A, Arceuthobium strictum on Pinus

leiophylla var. chihuahuana (left), and A. verticilliflorum on P. engelmannii (right) show difference in

host reaction to the two mistletoes; B, A. rubrum

on Pinus teocote (right), not in P. engelmannii (left)

but elsewhere this tree is parasitized by A. rubrum;

C, A. gillii subsp. nigrum severely damaging Pinus

leiophylla; (A, B, C, near El Salto, Durango, Mexico.)

D, A. californicum on Pinus lambertiana. Sierra

National Forest, California.

31
Figure 11.—Effects of dwarf mistletoe on various pines:

A, B, C—Arceuthobium vaginatum subsp. cryptopodum. A, on Pinus engelmannii. El Largo, Chihuahua, Mexico; B, massive witches' broom in Pinus

ponderosa. Grand Canyon National Park, Arizona;

C, group of Pinus ponderosa killed by dwarf mistletoe,

Grand Canyon National Park, Arizona; D, A. guatentalense on Pinus ayacahuite. Sierra Cuchumatanes,

Guatemala.

32
Figure 12.—Witches' brooms caused by two southern dwarf mistletoes. A, B, Arceuthobium bicarinatum on Pinus

occidentalis, Constanza, Dominican Republic; C, D, A. hondurense on P. oocarpa, near Tegucigalpa, Honduras.

In D, the dense clumps in the lower crown are epiphytic bromeliads.

33
Figure 13.—Witches' brooms caused by larch and hemlock dwarf mistletoes: A and B, Arceuthobium laricis

on Larix occidentalis. Blue Mountains, Oregon. Photograph B by J. R. Weir. C, A. laricis on Tsuga mertensiana. Coeur d' Alene National Forest, Idaho;

D, A. tsugense on Tsuga mertensiana, Stanislaus

National Forest, California.

34
^v

Figure 14.—Dwarf mistletoes on various conifers: ^, Arceuthobium

pusillum on Picea mariana, Minnesota; B, A. pusillum on P.

glauca, Minnesota. Photographs A and B by D. W. French,

University of Minnesota. C, A. campylopoduni on Pinus ponderosa, near Spokane Washington; photograph by J. R. Weir; D, A.

minutissimum on Pinus griffithii, Kashmir:

35
chromosome numbers and change in the family, the

X = 14 in Arceuthohium appears to be the product of

an increasing aneuploid series, rather than a tetraploid based on x = 7.

The apparent chromosome system in Arceuthohium

is diploid with X = 14. The genetic system is presumably sexual and obHgately out-crossing (the

genus being dioecious). No evidence of asexual

reproduction exists. These characteristics, coupled

with the relatively high diploid chromosome number

{x = 14) and high chiasma frequency (averaging

perhaps 2 to 3 chiasma per bivalent), suggest a

relatively high recombination index (Darlington

1958).

speciaHzed (reduced) morphologically, and also

occupy narrow ecological niches, or host trees, to

w^hich they are almost exclusively confined. Furthermore, these hosts (Pinus contorta and Pseudotsuga

menziesii) usually are not climax species, as are the

hosts for the more primitive species, but are intermediates in the successional sequence of the ecosystems in which they occur. This suggests that the

climax host species may have been colonized first

by the primitive species of dwarf mistletoes. As

these ecological niches were filled, further evolution

required the more advanced species to occupy

niches (host trees) characteristic of pioneering or

intermediate stages of the ecological sere.

Progressive Evolution and

Adaptive Radiation

The significance of a high recombination index

depends on the evolutionary opportunities of the

group. For the North American dwarf mistletoes,

numerous habitats (host trees) became available

for colonization because of the large development of

Pinus during the iMiocene (Mirov 1967). The genetic

system of Arceuthohium—diploid with high recombination potential—theoretically should be well

suited to progressive evolution.

If a large number of ecological niches (host

species) are available for colonization, evolution

should eventually produce an adaptive radiation

into the available niches. The high proportion of

North American Pinaceae presently parasitized by

Arceuthohium (see table 5) indicates that such an

adaptive radiation has occurred. With the exception

of the eastern pines of North America, most of the

major species of Pinaceae are parasitized by

Arceuthohium.

A factor conceivably important in this adaptive

radiation was the apparent lack of competition with

other plants for the available niches. With the

possible exception of Phoradendron holleanum subsp.

pauciflorum, which sometimes occurs on Ahies

concolor with A. ahietinum, and a few tropical species

of Dendropemony Psittacanthus, and Struthanthus,

no extant groups compete with Arceuthohium.

Competition with epiphytes might be expected in

tropical cloud forest situations, but this does not

occur, presumably because only young shoots are

susceptible to infection by dwarf mistletoes. Generally, emerging shoots are infected by dwarf

mistletoe before epiphytes can become established.

On the basis of our criteria, a number of dwarf

mistletoes have reached a high degree of specialization. This suggests that the basic adaptive radiation

has already taken place, and that the adaption is

now into increasingly more specialized niches. For

example, A. americanum and A. douglasii are highly

Absence of Hybridization and

Polyploidy

One of the most significant factors in the evolution

of vascular plants is the occurrence of natural

hybridization. So pervasive is this phenomenon, that

most groups of vascular plants studied by contemporary workers appear to show some evidence of

natural hybridization.

In Arceuthohium, however, no evidence of natural

hybridization has been discovered. In our cumulative years of field experience (and those of several

knowledgeable colleagues) we have never encountered a single instance of natural hybridization,

although we have consciously searched for such

evidence, especially in our studies of sympatry

(see table 3). Overlap occurs in a number of characters, especially quantitative traits which might

be interpreted as evidence for natural hybridization.

We believe, however, that the overlap is the result of

evolutionary convergence. While a single character

of one taxon might overlap with a trait of another,

the majority of character combinations defining

each taxon do not occur discordantly (Anderson

1951). Discordant variation occurs through hybridization when character combinations typical of more

than one taxon occur together. Traits are concordant

in Arceuthohium; they remain associated with their

regular constellation of characters.

The absence of hybridization is apparently typical

of both familes of mistletoes (Barlow and Wiens

1971, Wiens and Barlow 1971), and is not unique to

Arceuthohium. In Phoradendron, a single clear

instance of natural hybridization is known (Wiens

1962, Vasek 1966), and the hybrid is apparently

sterile. The absence of natural hybridization in

Arceuthohium as well as other mistletoes can be

explained by occurrence of strong interspecific

isolating mechanisms, particularly seasonal isolation.

A factor which might explain the apparent

absence of hybrids in Arceuthohium is the lack of

suitable habitats for their establishment. Most

36
dwarf mistletoes have a principal host (see table 5)

to which the species must be adapted. Because

hybrids combine the genetic characteristics of two

species, theoretically they should be adapted to

intermediate habitats. However, intermediate habitats in parasitic groups theoretically could be

produced only through hybridization of the hosts.

In other words, ''hybridization" of the habitat, as

Anderson (1948) expressed it, would literally be

necessary. Various pinaceous hosts of Arceuthohium

do hybridize, but we are unfamihar with any situations where two species of dwarf mistletoes parasitize

hybridizing host populations. Where host pines do

hybridize, for example, P. jeffreyi X P. ponderosa

and P. contorta X P. hanksiana, the hybridizing pair

of species are so closely related that they are parasitized by the same dwarf mistletoe.

Another factor that may inhibit hybrid establishment also involves habitat. If intermediate hosts are

not available, then hybrids of the dwarf mistletoes,

even if they occurred, would be without a suitable

habitat and might not survive. Hence, hybrids in

Arceuthohium might not be detected. Whether or

not hybridization in dwarf mistletoes is precluded

by strong isolating mechanisms, or because hybrids

fail to become established, the result is the same. In

either case, there should be strong selective pressures

against gene combinations that allow hybridization

(Dobzhanskyl951).

The presumed absence of natural hybridization

(or hybrid estabhshment) is an important basis for

further evolutionary considerations in Arceuthohium.

Not only have we failed to discover hybrids in

dwarf mistletoes, but they are not reported in the

extensive literature of the group. But if hybrids do

not occur, we could hardly expect to find literature

on the subject.

A theoretical explanation proposed for the absence

of polyploidy in Arceuthohium (Wiens 1968) is

based on the assumption that most polyploids are

alloploids (Stebbins 1950), and depend on natural

hybridization for their origin. If hybridization does

not occur, alloploidy is precluded. Autoploidy might

still occur, but its presence is probably of little

evolutionary significance; probably the few cases of

known polyploidy in the Loranthaceae and Viscaceae

are of autoploid origin (Barlow and Wiens 1971,

Wiens and Barlow 1971).

Evolutionary Patterns and

Taxonomic Structure

Arceuthohium is apparently a sexual, outcrossing

group which could be expected to produce a relatively

wide pattern of variation. A high recombination

potential and open niches on the Pinaceae could

provide the fundamental conditions to produce an

adaptive radiation onto most North American

Pinaceae. In the absence of hybridization and

polyploidy, the species have apparently maintained

distinct evolutionary lines. This feature has important evolutionary and taxonomic consequences,

because distinct phyletic lines should produce

dendritic evolutionary patterns.

Such evolutionary patterns are typical of most

animals. In vascular plants, however, reticulate

evolution is the rule, and is almost certainly produced through natural hybridization. Alloploidy

may stabihze hybrid genomes into true-breeding

species. Hybridization and polyploidy, therefore,

have the result of combining the characteristics

of different evolutionary fines and thereby produce

reticulate evolutionary patterns characteristic of

vascular plants.

In contrast, a dendritic evolutionary pattern

appears to characterize Arceuthohium, but the situation is obscured by the extreme morphological

reduction attendant with the parasitic habit. The

species of Arceuthohium are relatively distinct, but

the characters separating them are often cryptic or

discernible for only short periods of the life cycle.

Even though individual characters are not always

clearly evident, when the characters are analyzed

in their totality, the taxa become well differentiated.

This is evident from our numerical analyses which

show that all taxa have a relatively high degree of

integrity and tend to be more distinct than in

other groups where hybridization has played an

important evolutionary role (Sokal and Sneath 1963).

The two subgenera or phyletic lines (based on

branching patterns) which Kuijt (1970) proposed seem

valid, but the reduction of shoots in species such as

A. pusillum sometimes obscures the relationship. In

Arceuthohium the evolutionary pattern is further

characterized by three stages of evolutionary divergence. These three stages appear to be present in both

the New and Old Worlds, but are better developed in

the former.

Because of the dendritic evolutionary patterns,

these evolutionary stages can be defined taxonomically and basicafiy correspond to the primitive,

intermediate, and advanced species groups we

propose in the New World subgenera (see discussion

under the subgenera and fig. 15). We have constructed a subgeneric taxonomy based on what

appear to be definable evolutionary stages in

distinct phyletic fines. We befieve that the taxonomic

treatment given here generally represents the

phyletic history of Arceuthohium.

Evolutionary Trends

Evolutionary trends in Arceuthohium are inferred

(Gin 1935), but the course of evolution is far from

37
%IV

^Of^LD

,^v^' ,.ceut

^ârf. * v9rticilliflorum

Sec. Campylopoda

Ser. I Stricta

Ser. 2 Campylopoda

Ser. 3 A'í/^AíJ

ARCEUTHOBOIDKORTHALSELLOID

ANCESTOR

Figure 15.—Presumed phyletic evolution of Arceuthobium showing two primary lines of diversification—subgenera

Arceuthobium and Vaginata. Subgenus Arceuthobium is divided into Old World and New World groups with three

and four species, respectively. Subgenus Vaginata contains a primitive group (Sec. Vaginata), an intermediate

group (Sec. Campylopoda), and an advanced group (Sec. Minuta).

understood. Some evolutionary tendencies that we

recognize are summarized below:

Characteristic

Host specificity

Shoot size

Fruit maturation period

Shoot longevity

Polymorphism

Flowering group

Branching type

Witches' brooms

Primitive

Low

Large

Long

Long

High

Direct

Verticillate

Nonsystemic

Advanced

High

Small

Short

Short

Low

Indirect

Flabellate

Systemic

By most of Gill's criteria, A. pusillum and A.

douglasii are the most advanced species, and A.

vaginatum and A. campylopodum the most primitive.

We use the concepts of Davis and Heywood (1963)

that primitive groups are not necessarily ancient:

'Trimitive or advanced groups may be ancient or

recent according to whether they became separate

entities early or late in evolution/' ''Primitive''

merely connotes a character that occurs in both the

modern species and its ancestors, while ''advanced"

implies derived characters not present in ancestral

stock.

Wiens (1968) studied the flowering characteristics of the New World dwarf mistletoes. He

hypothesized that the different flowering groups

arose as the genus moved northward from Mexico

(the present area of greatest species diversity) into

latitudes of greater annual variation in photoperiod.

He concluded that taxa with indirect, spring flowering (A. americanum, A. douglasii, A. pusillum) were

derived from those with direct, summer flowering (the

campylopodum group, A. rubrum, and A. stridum),

which in turn are derived from those with direct,

spring flowering {A. abietis-religiósae, A. globosum,

A. verticilliflorum, and A. vaginatum).

Kuijt (1970) presents evidence that flabellate

branching pattern in Arceuthobium, common in

most North American taxa, is a derived character.

The more primitive, verticillate type of branching is

38
present in the two Old World species in which

this trait was studied (A, oxycedri and A. chínense).

The diminutive A. minutissimum, and the New

World A. pusillum, are so reduced that their basic

branching type is not determinable. In the New

World, A. americanum and A. abietis-religiósae

exhibit verticillate branching. Arceuihohium verticilliflorum is anomalous because the flowers are

verticillate, but the stems show no secondary

branching. However, Kuijt (1970) states the branching pattern in A. verticilliflorum is basically

verticillate.

The formation of systemic (or isophasic) witches'

brooms (see p. 26) is an additional feature that

probably indicates an advanced degree of evolution.

Although systemic types of witches' brooms are

occasionally induced by several dwarf mistletoes,

they are consistently formed by only five species:

A, pusillum, A. douglasii, A. guatemalense, and A.

americanum in the New World, and A. minutissimum in the Himalayas. Some host-parasite combinations, where witches' brooms are not consistently

formed, may represent a primitive state that preceded nonsystemic broom formation, which, in turn,

may have preceded systemic broom formation.

Systemic witches' brooms seem to be of adaptive

advantage because they greatly increase the reproductive potential of the dwarf mistletoe. By

formation of systemic witches' brooms, the endophytic system from a single seed can ramify

through hundreds of linear feet of host branches

and produce a profusion of shoots, flowers, and fruit.

Most dwarf mistletoes eventually kill their hosts,

but mortality occurs much sooner in some hostparasite combinations than in others. Parasites

that are less lethal to their hosts are presumably

better adapted. This factor is apparently not related, however, to the relative stage of advancement

as measured by other factors. For example, A.

pusillum, A. americanum, and A. douglasii are

presumably advanced species, but are ultimately

very damaging to their hosts. However, this could

be explained if these species are relatively newly

evolved and neither the host nor the dwarf mistletoe

are yet co-adapted—the dwarf mistletoe for less

pathogenicity and the host for greater resistance.

Within the campylopodum group there are some

taxa that cause severe host mortality (for example,

A. laricis on Larix occidentalis), and some in which

host mortality is relatively low {A. divaricaium

onpinyons).

We consider A, pusillum and A, douglasii to be

the most advanced New World species, because they

exhibit the following features: (1) very small shoots,

(2) very specific host associations, (3) derived

flowering type, (4) flabellate branching, in A.

douglasii, and probably in A. pusillum, and (5)

consistent formation of systemic witches' brooms.

Conversely, the Mexican species, A. verticilliflorum,

probably represents the most primitive species

because of its (1) large shoots and extremely large

fruits, (2) less specific association, (3) primitive

flowering type, (4) verticillate branching, and (5)

formation of nonsystemic witches' brooms.

Arceuihohium is primarily a parasite of Pinus,

since 22 of the 32 taxa occur principally on this

genus (see fig. 7). Of the 22 taxa, 15 parasitize

Diploxylon and 7 Haploxylon pines. Mirov (1967)

recognizes 73 Diploxylon and 32 Haploxylon pines,

so about one-fifth of each group is parasitized. The

oldest known pine fossils are from the Jurassic period,

and even then the pines had differentiated into the

subgenera Haploxylon and Diploxylon, the former

possibly preceding the latter (Mirov 1967).

In both the Old and New Worlds, Pinus (Diploxylon and Haploxylon) and Abies are hosts of

Arceuihohium. However, four other genera (Larix,

Picea, Pseudotsuga, and Tsuga) that occur in both

hemispheres are parasitized only in the New World.

Juniperus, which is common throughout the Northern Hemisphere, is a host only in the Old World.

Keteleeria, a genus not found in the New^ World, is

parasitized in China.

We surmise that the primitive Arceuihohium

stock was a relatively large, verticillately branched

parasite of pines in northeastern Asia. Chromosomal

and morphological studies suggest that Korthalsella

is the closest related living genus, and that the two

genera arose from common ancestral stocks (Wiens

and Barlow 1971). One species, Korthalsella dacrydii,

occurs on conifers (Podocarpus) and even has mildly

explosive fruit (Danser 1937). Korthalsella is now

distributed from Hawaii through the South Pacific

to Africa (Danser 1937). Arceuihohium, conversely,

spread westward to the Mediterranean region, south

into Africa, and northeastward through the Bering

Straits region into the New World. In the New

World, the genus apparently found ideal conditions

for rapid expansion, and many new species evolved.

Generally, in the Tertiary period, the angiosperms

have evolved more rapidly than the gymnosperms

(Leopold 1967). Consistent with this trend, the

dwarf mistletoes probably evolved much more

rapidly than their coniferous hosts.

39
SYSTEMATICS OF ARCEUTHOBIUM

Taxonomic Considerations

Arceuthohium has long been considered to be a

taxonomically difficult genus. This is primarily

because the genus has undergone extreme morphological reduction due to its parasitic habit. Also,

the general morphological aspects of all members

of the group are similar and there are few gross

discontinuous variables. Many of the morphological

features such as leaves and trichomes commonly

used in the classification of other mistletoe groups

are not present in the genus. In all species, the

flowers are small (2-4 mm. in diameter) and generally

similar in form.

Gill (1935) segregated the members of the Arceuthobium campylopodum and A. vaginatum groups

into host forms, that is, ^'taxa delimited exclusively

on the basis of the host relationships, without regard

to biological parity.^^ Although Gill's system provided an effective and facile method for naming the

various dwarf mistletoes, it obscured the systematic

difficulties in the group. Gill realized his classification

system was unnatural, but available data were too

limited for a more natural treatment. Gill noted

that the '^forms are not all of equal rank in the

sense of being biologically distinct. Some are welldeveloped strains showing a marked affinity for a

limited group of host species, while others are admittedly artificial categories, into which infrequent

or even accidental host relationships of the better

defined forms have been cast.'' Gill further stated

that a system of forms based exclusively on hosts

^ ^though without good taxonomic precedent, should

be tolerable as a temporary device, pending a complete revision of the genus based on further field and

experimental evidence." As will be shown, we consider Gill's host forms in Arceuthohium campylopodum

and A. vaginatum to be morphologically distinct;

hence, they warrant higher taxonomic rank.

Danser (1950) stated that the function of taxonomy is to classify life cycles. Danser was a student

of the Old World mistletoes, and how much his

experience with mistletoes influenced his thinking

is unknown. However, such a concept is particularly

well suited to the classification of Arceuthohium^

where general reduction and convergence of gross

features of the sporophyte have obscured the

relationships of the species. The idea that all aspects

of an organism are potentially useful taxonomically

is not new, but we have not seen the idea expressed

in quite the same context as Danser. We believe that

Danser's idea of classifying life cycles provides a

useful theoretical model on which to base systematic

studies. As a corollary, discontinuities between life

cycles should determine taxonomic units.

The term ''life cycle" is used in this paper to

encompass all aspects of the biology of a dwarf

mistletoe, and includes interactions with host and

environment. All phases of the life cycle are considered here as potentially vahd taxonomic criteria,

and the first priority is to determine the basic

features of the life cycle in the group and to isolate

discontinuities between them. The test of valid

discontinuities rests on genetic control and consistency of occurrence. Other vahd taxonomic

criteria we have considered are physiological characters such as time of meiosis, anthesis, period of

seed dispersal, time of seed germination, host

specificity, and brooming response of the host

because they are measurable and consistent. We

therefore disagree with Davis and Heywood (1963)

who question the use of physiological characters in

taxonomy. Quantitative and qualitative morphological characters may also be utilized in classification. The quantitative difference, while often small,

may be statistically and taxonomically significant.

Previous difficulties in the genus are therefore the

result of the extreme reduction and lack of precise

data on morphology and life cycles. There appear

to be few other plant groups in which a knowledge

of living populations is so essential for their

classification.

At certain stages of their life cycle, some dwarf

mistletoes may be difficult to identify. The herbarium

taxonomist, confronted by an individual specimen,

might well experience difficulty in identification

because of the particular stage of the life cycle

represented in the specimen before him. For example,

in a nonflowering specimen the time of anthesis

might not be obvious. Color, generally consistent

in living plants, may be altered in drying; A.

ruhrum is a dark, shiny red in nature but becomes

dull brown when dried. Characteristics of habit,

obvious in living plants, such as A. glohosum which

forms open spherical masses, may be completely

obliterated by careless pressing or fragmentation.

Because Arceuthohium is difficult to preserve and

fragments easily, general botanists seldom collect

these plants. Some suggestions for collecting and

curating dwarf mistletoes are given on page 61.

The monographer's duty is to extensively study

the life cycles of various taxa to determine their

significant differences and similarities. Whether

these differences are easily discernible in a particular

specimen in no way affects their intrinsic taxonomic

value. The distinctions between classification and

ease of identification should not be confused.

40
Fortunately, characters such as host specificity may

provide an easy method of identification, but this

does not infer that our classification is based on hosts

alone, because no characters in our system are

weighted.

The key we have prepared reflects natural relationships, and includes characters manifest at

various stages of the life cycle. In addition, an

artificial key to facilitate identification is included.

Natural hybrids apparently do not exist in

Arceuthohium, so the taxa should be easily delimited

unless they represent intermediate stages of gradual

evolutionary divergence. In Arceuthohium the taxa

appear to be reasonably well defined, and significant

discontinuous variation occurs in a number of

characters. The amount of infra-taxon variation is

about what one would expect in a sexual, outcrossing

group. Even the subspecific categories appear to^ be

demarcated by relatively sharp discontinuities,

although of less magnitude than those separating

species.

Problems in Classification

Based on our exhaustive field observations and

experimental studies, we believe the taxa Gill (1935)

classified as host forms in the A. campylopodum

complex are distinct species. The most convincing

evidence to support our position is that when these

dwarf mistletoes occur on trees other than the principal

host, they maintain their morphological integrity and

are identifiable. Natural dwarf mistletoe crossovers

from principal hosts to secondary, occasional, or

rare hosts have been observed in all but 4 of the 13

members of the main A. campylopodum group (A.

apachecum, A. blumeri, A. guatemalense, and A.

divaricatum),

Various examples illustrate difficulties with GilFs

host-form concept. At McKenzie Pass, Oregon,

where Arceuthohium tsugense occurs on Pinus

alhicaulis, Ahies lasiocarpa, and the principal host,

Tsuga mertensiana, the parasities on all hosts are

identifiable as A. tsugense. Under GilFs system,

however, three different names (f. cyanocarpum, f.

ahietinum, and f. tsugensis, respectively) were

assigned according to the host plant. At Priest

River Experimental Forest, Idaho, A. laricis,

common on its principal host (Larix occidentalis),

is also found on Pinus contorta and on three introduced conifers, P. hanksiana, P. resinosa, and

Picea ahies. Here, too, the morphological integrity

of the dwarf mistletoe is retained. The host-form

concept is difficult to apply when dwarf mistletoes

occur on introduced hosts since, by definition, each

form is restricted to a specific host. Strict adherence

to such an unnatural system would require that a

new name be coined for each new host-parasite

combination.

Another disadvantage to GilFs treatment is that

whenever more than one member of the A. campylopodum group parasitized the same host, all had the

same name. For example, where A. campylopodum,

A. laricis, and A. cyanocarpum parasitized Pinus

ponderosa, all three were classified as A. campylopodum f. camplyopodum. Each dwarf mistletoe,

however, is morphologically distinct on P. ponderosa

as well as on any other of its hosts. This evidence also

supports our contention that these are vaHd taxa

rather than host forms, because of discontinuous

variations among them.

In addition to morphological integrity, dwarf

mistletoes show definite host preferences, even

though they occasionally occur on trees other than

their principal hosts. The reasoning employed

when we assigned taxonomic ranks, particularly in

the troublesome A. campylopodum group, can be

illustrated by A. campylopodum and A. occidentale.

Pinus ponderosa and P. sahiniana, respectively, are

their principal hosts, and these dwarf mistletoes are

restricted to their principal hosts even where the

hosts of the other mistletoe occur sympatrically. This

could be interpreted as evidence that these dwarf

mistletoes simply represent ecotypic races; however,

morphological and physiological differences (other

than host) also characterize the two plants. Where

the two populations occasionally occur together,

the morphological type of each dwarf mistletoe

occurs on its respective host. Arceuthohium occidentale

also occurs on Pinus radiata and P. muricata, but not

P. ponderosa. In these cases, it is easy to identify as

that which predominately occurs on P. sahiniana.

Arceuthohium campylopodum, whose principal host

is the typical variety of P. ponderosa, is also found on

some other pines such as P. jeffreyi, P. attenuata,

P. coulteri, but not P. sahiniana. Again, the morphological and physiological characteristics of the dwarf

mistletoe remain relatively stable and identifiable.

The peak flowering periods of A. campylopodum

and A. occidentale differ, but there is some overlap.

If mutual pollinating agents are available, gene

exchange should occur if the plants are crossable.

The problem of sympatry is complicated in parasitic

groups (Dobzhansky 1951). However, we consider

these two dwarf mistletoes to be sympatric because,

in terms of temporal and spatial characteristics,

gene exchange should be possible unless they are

genetically isolated. Although one is tempted to

weight sympatry in classification, we have not done

so since the present data show sufficient diversification between the two taxa to warrant specific status.

All members of the A. campylopodum group are

sympatric with at least one other member of the

group (see fig. 6). In all cases the morphological

41
integrity of each dwarf mistletoe is maintained.

Our research indicates that the number of species

in Arceuthohium is greater than that recognized by

previous workers. We believe two criteria are

fundamental in delimiting these taxa as species.

First, and perhaps most critical, is that dwarf

mistletoes maintain their morphological integrity

when growing on trees other than their principal host.

Secondly, taxa, recognized as species, are almost

always sympatric, yet show no evidence of

hybridization.

Some taxonomists may question why many taxa

we recognize as species should not be considered

ecotypes. Ecotypes are usually defined as genetically

distinct races physiologically adapted to locaHzed

habitats and sometimes differentiated morphologically, particularly by quantitative characters.

The taxa we recognize as species should not be considered ecotypes because of their morphological

integrity on nonprincipal hosts, and the lack of

intermediates between sympatric taxa. AJso, ecotypes

are not reproductively isolated and usually are not

distributed as widely as the taxa we recognize as

species. A. microcarpum has the most restricted

range of any dwarf mistletoe outside of Latin

America; it is found only within a 300-mile area in

Arizona and New Mexico. Most dwarf mistletoes

have ranges at least twice this large. Conceivably,

some of the dwarf mistletoes we recognize as species

might correspond to regional ecotypes, but most

taxonomists consider regional ecotypes to be comparable to subspecies so the taxonomic treatment

would not be greatly different.

We beheve that ecotypes occur in Arceuthohium,

but they do not exhibit the level of differentiation

of taxa we recognize as species or subspecies. Species

of dwarf mistletoe with more than one principal

host probably are ecotypically differentiated in

various parts of their range. For example, Pinus

ponderosa is one of the principal hosts of A. campylopodum, and this tree is parasitized from Washington

to southern Cahfornia. In parts of northern California and southern Oregon, however, where the

principal host of A. campylopodum is Pinus attenuata,

associated P. ponderosa is seldom infected. Within a

dwarf mistletoe, however, cross infection between the

principal hosts occurs. Therefore, we do not believe

a useful purpose is served by taxonomic recognition

of populations we consider as ecotypes.

Subspecific Classification

Subspecies in Arceuthohium are interesting because

they do not conform to commonly accepted definitions. Geographically associated morphological variation occurs in two species of North American

mistletoes, A. gillii and A. vaginatum. Usually no

gradation of characteristics or ''shading-off'' occurs

between the Arceuthohium population systems that

constitute subspecies, as is common in many plant

groups. The presence of discontinuous variation

between such population systems might suggest

that they should be species, even though the differences are small. This situation might be comparable to that in Carex where the species are often

based on small, yet apparently consistent, differences. We believe, however, that geographically

restricted populations, delimited by relatively few

but consistent variations, are best classified as

subspecific units.

In A. ahietinum, we have used the category of

forma specialis, to avoid confusion with the usuaJ

concept of forma which denotes morphological

distinction. Special forms designate physiological

races without morphological differences. Scharpf

and Parmeter (1967) have shown experimentally

that the dwarf mistletoe on Ahies magnifica will not

infect Ahies concolor, and vice versa. However, the

form on A. concolor will also infect Ahies grandis,

and the form on A. magnifica will parasitize A.

procera (see discussion under A. ahietinum). This

high specificity is confirmed from field studies in

mixed fir stands. If the races of A. ahietinum could

parasitize either species of fir, their presence would

be expected in mixed stands where millions of dwarf

mistletoe seeds from each type of fir are deposited

yearly on the other species. Yet, cross infection does

not occur.

CRITERIA FOR CLASSIFICATION

In our treatment of Arceuthohium, we considered

many morphological, palynological, cytological,

physiological, and chemical features of the genus.

Some traits, for example, branching and flowering

season, are classical characters used by Engelmann

and Gill. Other characters such as pollen, flowering

group, and shoot pigments, have not been previously

used in the classification of the genus.

Morphological Characters

Plant Size and Habit

Shoot size varies considerably within the genus

from a mean of about 0.5 cm. for A. minutissimum

to over 70 cm. for A. glohosum. Thus the size of

42
mature plants may be over 100 times larger in some

species than in others. Hooker (1886) reported A.

minutissimum as the smallest dicotyledonous plant,

but Pilostyles thurberi (Rafflesiaceae), a parasite of

leguminous trees in Southwestern deserts, probably

is smaller.

Within a taxon, shoots on systemic witches^

brooms are somewhat smaller than nonsystemic

infections on the same host. In our descriptions,

(see Formal Taxonomy section) the shoot dimensions

for the most common situation for each species are

given; that is, for systemic brooms for A. americanum, A. douglasii, A. giiatemalense, A. minutissimum, and A. pusillum, but for nonsystemic

infections for all other taxa. Also, shoot dimensions

as given are for both pistillate and staminate plants.

Plants of one sex are occasionally larger than those

of the other, but the differences usually are not

significant.

Plant size varies somewhat because the vigor of the

dwarf mistletoe depends directly on the vigor of

the host plant. Mean dimensions shown for each

species are based on the most frequently observed

situations; in addition, we have indicated the tallest

shoots measured for each taxon.

Growth habit is a distinguishing character for

some species. Shoots may form dense spherical

masses (A. globosum and A. occidentale), be scattered

along the stem (as in most taxa), or so dense that the

host twigs are obscured {A. cyanocarpum and A.

apachecum).

Shoots

Gill (1935) used branching as a major taxonomic

character in classifying Arceuthobium. All species

have decussate primary branching (Kuijt 1970),

and in some taxa branching proceeds no further

(A. pusillum, A. minutissimum, and A, verticilliflorum). In most taxa, however, secondary branching is apparent and is of two basic types: verticillate

or flabellate (see fig. 1). Kuijt (1970) considers the

verticillate habit to be a primitive feature present

in the Old World taxa, A. oxycedri, A. chinense, and

presumably in A. minutissimum, and in three New

World taxa, A. americanum, A. abietis-religiósae,

and A. verticilliflorum. The latter species does not

actually exhibit secondary branching, although

the flowers are verticillately arranged. The remaining New World taxa show flabellate branching.

A. pusillum, however, is so reduced that the basic

branching type cannot be determined, but it probably belongs in the flabellate group. Some taxa

show little secondary branching, for example,

A. rubrum (flabellate) or A, abietis-religiósae

(verticillate).

Internode dimensions are useful criteria for

distinguishing several species. We have consistently

used the third internode from the base for this

statistic. Length and standard deviation of the third

internode are given for each taxon in the Formal

Taxonomy section. Frequently the basal and second

internodes are not normally elongated. Because

dwarf mistletoe shoots have a basal meristem, individual shoot internodes elongate for several years.

For this reason, Kuijt (1970) has questioned the

validity of our use of internode dimensions as a

taxonomic character. However, the overall mature

internode dimensions among taxa differ so much

that internodal elongation does not negate the usefulness of the characters in certain cases (fig. 16). When

we analyzed the length/width ratio of the third

internode (table 14), we found less variation than

in length alone.

Some species show sexual dimorphism in shoot

characteristics. For example, the staminate shoots of

the two subspecies of A. gillii are more divaricate

than the pistillate ones. In A. strictum, the staminate

shoots show no branches in annual growth segments,

TABLE 14.—Length/width ratio of third internode of

the North American taxa of Arceuthobium

Taxon Length/width Number of

ratio collections

A. americanum 10.1

A. abietinum f. sp. magnificae. _ _ 8.8

A. occidentale 7.1

A. calijornicum 7.0

A. abietinum f. sp. concoloris 6.7

A. guatemalense 6.7

A. microcarpum 6.2

A. divaricatum 6.1

A. laricis 6.1

A. tsugense 6.1

A. hondurense 6.1

A. campylopodum 5.6

A. abietis-religiosae 5.5

A. blumeri 5.5

A. bicarinatum 5.2

A. apachecum 4.8

A. cyanocarpum 4.7

A. gillii subsp. gillii 3.8

A. globosu7n 3.7

A. douglasii 3.6

A. rubrum 3.4

A. vaginatum subsp. durangense. 3.3

A. gillii subsp. nigrum 2.9

A. vaginatum subsp. vaginatum _ _ 2.9

A. vaginatum subsp.

cryptopodum 2.5

A. pusilluîn 1.9

A. strictum 1.6

A. verticilliflorum 0.9

20

5

11

8

15

3

9

19

12

15

3

27

4

8

5

12

15

9

6

29

5

5

6

11

25

13

5

5

43
A.globosum

A. a biefi s - rellgiosae

A. abief, f. sp. magnificae

A, vag. subsp. durangense

A, vag. subsp. vaginatum

A, abief. f. sp. concoloris

A.occidentale

A.americanum

A.campylopodum

A.guafemalense

A, gillii subsp. nigrum

A. gil I a subsp. giiiii

A,bicarinafum

A.californicum

A. divaricafum

A.microcarpum

A. fsugense

A, blumeri

A.tiondurense

A. larieis

A. vag, %\xbsp,crypfopodum

A,apactiecum

A, rubrum

A.cyanocarpum

A. douglasi i

A, sfriefum

A. verficilliflorum

A, pusi Hum

A, minufissimum

MEAN

+

J_

PLUS AND MINUS ONE

STANDARD DEVIATION

-L

10 15 20 25

LENGTH OF THIRD INTERNODE (MM)

30

Figure 16.—Length of the third internode of the New World taxa of Arceuthobium and the Old World A, rninutissimum.

but the pistillate shoots are abundantly branched.

Several species can be readily distinguished by

shoot color (see color photographs). Shoot color is a

useful taxonomic character even though it may

vary slightly within a taxon, and the shade of the

staminate shoots may be different from the pistillate

shoots (for example, in A. campylopodum). Colors

range from black, purple, brown, red, orange, green,

to light gray, and carefully preserved dried specimens

usually retain the original color. An exception is A.

ruhrum; the bright red living shoots become dull

brown when they are dried.

Shoots are typically erect, but they may be

somewhat pendant in the larger individuals of the

A. vaginatum group. Shoots of most species live for

many years, but only for 1 or 2 years in some of the

smaller species such as ^. pusillum.

A single shoot axis that originates from a basal

cup characterizes all species except A. minutissimum;

in this species numerous shoots arise from the basal

cup.

44
Inflorescence

Characters of the inflorescence are of taxonomic

value. For example, the size of staminate spikes

prior to flowering is useful in separating some

members of the A. camyylopodmn group; the

branching pattern of the staminate spikes helps

distinguish the subspecies of A. vaginatuin.

Buds and Flowers

The lateral staminate buds of A. doucjlasii and

A. americanwn are spherical, but those of the other

taxa are lenticular.

The flowers are pedicillate in some taxa and sessile

in others. Gill (1935) used the ''pedicillate joint'' of

the staminate flowers as taxonomic features of A.

americanum and, to a lesser extent, A. douglasii.

Kuijt (1970) has shown that this ''pedicillate joint''

is an annual dichasial unit. Kuijt (1970) first reported pedicillate pistillate flowers on the main axis

in some taxa. We consider this feature to be of taxonomic value because it is present in some groups

(the A. campylopodu7n complex) and absent in

others {A. vaginatuîn, A. gillii, and A. glohosurn).

Both the staminate and pistillate flowers of

Arceuthobium are structurally simple and remarkably

uniform throughout the genus (Cohen 1968, 1970).

The color, size, and number of perianth lobes of the

staminate flowers are useful taxonomic characters.

The inner surface of the staminate flowers is the

same color as the shoots in all species except A.

abietis-religiosae, A. douglasii, and A. hondurense.

The staminate flowers are about 2-3 mm. in diameter for most taxa but average 4 mm. in A.

verticilliflorum. The lateral staminate flowers are

predominantly 3-merous except for A. verticilliflorum

(4-merous), A. guatemalense (2- to 3-merous), A.

hlumeri (4- to 6-merous) and A. stridum (3- to 7-

merous). The terminal flowers are frequently 4-

merous in most taxa. Length and width of staminate

perianth segments, closely related to overall flower

size, are sometimes useful taxonomic characters.

Staminate flowers that do not fully expand is a

unique feature oí A. rubrum.

A nectary or "central cushion" is present in the

center of each staminate flower (Cohen 1968).

We have not found any valid taxonomic features of

this nectary, although it is more prominent in some

specimens than in others.

Anthers

Anther diameter is discontinuously variable among

some species, and may be as large as 1 mm. in A.

verticilliflorum, and less than half this size in others.

Engelmann (in Watson 1880) used the location of the

anther on the perianth segment to help distinguish

some taxa, but our analyses indicate that this

character has little taxonomic value.

Fruit

The mature fruit in most species ranges from 3 to

5 mm., but is 15 mm. long in A. verticilliflcrum.

All species have bicolored fruits, and the relative

size of the proximal and distal portions is a useful

taxonomic character. The distal portion of the fruit

may account for 45 percent of the total fruit length

in A. gillii, but only 25 percent in A. abietis-religiosae.

Dimensions of dried fruits are usually 15 to 25 percent less than that of fresh specimens. Shrinkage is

more pronounced in the distal portion (25 to 35

percent) than the proximal portion (10 to 20 percent). Thus the proximal/distal ratio tends to be

somewhat higher in dried plants.

The fruit surface, also useful in taxonomy, may

be either glaucous, nonglaucous and dull, or nonglaucous and shiny. Seed size is directly correlated

with fruit size so it was not analyzed separately,

but it does vary among taxa.

Kuijt (1970) noted that some species, for example,

A. bicarinatum, t3i3Ícally have one or two sterile

nodes just below the terminal fruit on the main

axis of each pistillate shoot. This condition was

evident in all populations of A. bicarinatum we have

studied.

In most taxa, the base of the mature fruit fuses

smoothly into the receptacle of the pedicel. In A.

guatemaleuse and A. vaginatum subsp. diirangense,

however, the receptacle, slightly wider than the base,

forms a characteristic ring at the base of the fruit.

Palynological Characters

No comprehensive pollen studies have previously

been made in Arceuthobium, although pollen grains

of a few taxa have been measured or illustrated:

A. chinense (Lecomte 1915), A. minutissinium

(Bhandari and Xanda 1968), A. oxycedri (Erdtman

1952, Heinricher 1915a), and A. pusillum (Gill 1935,

Pomerleau 1942, Whitehead 1963, Whitehead and

Barghoorn 1962).

Our preliminary fíndings are reported here, but

we plan to pubUsh details later. In collaboration

with Dr. Estella B. Leopold of the U.S. Geological

Survey, we plan to use standard palynological

techniques, including electron scanning photomicrography, to analyze all members of the genus.

We have examined all Arceuthobium taxa, although

some of our study samples were meager. We analyzed

45
one to eight collections of each taxon and measured

at least 10 pollen grains in each collection. Our

analyses here are based primarily on the appearance

of dried pollen grains mounted in glycerin; however,

we have used the electron scanning system to

study two taxa, A. pusillum and A. verticilliflorum.

The pollen grains of Arceuthobium can be easily

distinguished from other genera (Erdtman 1952).

Approximately 20 to 30M in diameter, the grains are

roughly spherical in shape, but the equatorial diameter is 5 to 15 percent larger than the polar diameter

(fig. 17). The grains are divided into six alternating

spined and smooth sections that converge at the

poles. The grains are 3-colpate (grooved), and as the

pollen dries, these colpae become deeply grooved.

Lying parallel to the colpae on the intervening

walls are three pseudocolpae, or short grooves, which

do not reach the poles, but become more prominent

during drying.

We have confirmed Gill's (1935) suggestion that

pollen characteristics might be of taxonomic value,

although we did not find, as he reported, that A.

pusillum has larger pollen grains than the other

species. In our studies, pollen diameters ranged

from a mean of 18/i in A. minutissimum to 28iu in

A. verticilliflorum. Intra-taxon variation is usually

limited to within 1 or 2ß of the mean. Another readily

measured pollen feature of taxonomic value is the

height of spines in relation to wall (exine and intine)

thickness. In some species the spine height may be

three times the wall thickness (A. tsugense), while

in others the wall thickness greatly exceeds the spine

height {A. verticilliflorum). These taxa and a few

others are so distinct that they may be identified

solely on the basis of pollen characteristics.

Our electron scanning photomicrographs of two

species reveal detail in Arceuthobium pollen grain

morphology (fig. 18) that has not been apparent in

light microscope studies. For example, the wall of the

grain in A. pusillum is scabrate (with low, widely

spaced warts), while the wall in A. verticilliflorum

is more uniformly roughened. The spines of A.

verticilliflorum are smaller and less abundant than

those of A. pusillum. Spines are absent in the grooves

20 I B

20 fi

Figure 17.—Arceuthobium pollen grains: A, Diagrammatic drawing of pollen. Upper, polar aspect showing crosssectional and external views. Lower, lateral aspect showing cross-sectional and external views. B, Arceuthobium

americanutn pollen showing polar (upper) and lateral views.

46
in A. verticilliflorum but present in A. pusillum. Also,

the bases of the spines seem to be more distinct with

vertical sides in A. pusillum but are more spreading

in A. verticilliflorum.

Cytogenetical Characters

The chromosome number of all dwarf mistletoes

studied is n = 14 (fig. 19), and the base generic

chromosome number is considered to be x = 14.

In the New World, Dowding (1931) first reported

n = 14 for Arceuthobium americanum which Wiens

(1964) confirmed. Wiens also obtained the same

number for five additional taxa, and in 1968, reported counts for 10 additional New World taxa.

Chromosome counts for 12 more taxa are included

in this paper, so 23 of the 28 New World taxa have

now been studied. The five taxa not yet examined

Figure 18.—Electron scanning photomicrographs of Arceuthobium pollen: A, Pollen of A. pusillum; B, Detail of

spines and pollen surface; C, Pollen of A. verticilliflorum; D, Detail of spines and pollen surface. Photographs

provided through the courtesy of Drs. Estella B. Leopold of the U.S. Geological Survey, and Charles Drew of the

U.S. Navy Department.

47
.».

4^ W

£TSS) • •

1^. if* /" ' *. * •

¿^ 1 *

Figure 19.—Chromosomes in Arceuthobium gillii subsp.

gillii at early meiotic metaphase. ¡Magnification ca. 950.

include two Mexican species {A. globosuyn and A.

rubrum), the Central American A. guatemalense,

and two subspecies of il. vaginatum (subsp. vaginaium

and durangense).

In the Old World, only A. oxijcedri has been

studied cytologically (Pisek 1924). This species is

apparently n = 14, but Pisek also reported the

existence of n = 13.

Although the chromosome numbers in Arceuthobium are consistently n = 14, preliminary

observations suggest that other karyotypic differences may distinguish some species. For example,

A. douglasii and A. gillii possess a bivalent significantly smaller than other members of the genome.

Both species undergo meiosis in late summer but

flower in late winter or early spring of the following

year. Initial observations of A. americanum and A.

-pusillum, which have similar patterns of meiosis and

flowering, did not show small bivalents, although

most of the meiotic cells examined did not spread

well, and a small bivalent could have gone undetected. The karyotypes of these two species

should be examined with regard to this karyological

feature.

A thorough karyotypic study of the dwarf mistletoes would be a useful contribution to our knowledge

of the group. The difficulties of obtaining mitotic

cells in large numbers, however, pose technical

problems. For example, there are no root tips, in the

usual sense, but the dwarf mistletoes have a basa.l

meristem similar to grasses, which is active primarily

in the spring. The radicular apex of the developing

embryo and axillary buds were investigated as a

possible source of mitotic cells, but mitosis was too

sporadic and occurred in numbers too small for

effective karyotypic work (Wiens 1968). The

radicular apex of germinating seeds might contain

mitotic cells, but this source has not been fully

investigated.

Physiological Characters

Phenology

Gill (1935) used time of flowering (spring vs.

summer) as a principal taxonomic character. His

"summer" flowering group (the campylopodum

group) contains some significant interspecific variation. For example, A. occidentale flowers as late as

November or December, and A. calijornicum flowers

as early as June (fig. 20).

A taxon generally flowers at given periods each

year, although normal altitudinal, latitudinal, and

seasonal variation occurs. For the summer or fall

flowering species, flowering usually begins at the

higher elevation first (Scharpf 1965) while the

opposite seems to be true for the spring flowering

taxa.

Whether meiosis occurs immediately before flowering or 5 to 8 months preceding anthesis is an important taxonomic criterion not previously utilized

in this genus. Wiens (1968) has established three

basic flowering groups in ArceutJiobium (table 15) :

Group I, spring flowering species that undergo

meiosis directly preceding anthesis; Group II, midor late-summer flowering species that undergo

meiosis directly preceding anthesis; and Group III,

spring flowering species that undergo meiosis

during the preceding late summer or early fall. Group

la was established for A. gillii and A. verticilliflorum

which undergo meiosis in late summer and flower

in early spring, a characteristic in which they

resemble the species in Group III. General floral

development and other traits, however, appear to

place the two species in a transitional position

between Groups I and II.

Time of seed dispersal and fruit maturation

periods are useful taxonomic criteria. Relatively

consistent for a given taxon, peak seed dispersal

may occur as early as July in A. globosum or as late

as November to January in A. occidentale (fig. 21).

Also, fruits 0Î A. pusillum mature in about 5 months,

while those of all other taxa require at least 12

months; fruits of A. gillii require 19 months to

reach maturity (fig. 22).

Too little data are available to attach taxonomic

significance to time of seed germination and length

of the incubation period (time from infection until

48
TABLE 15.—Flowering groups in the North American dwarf mistletoes (Wiens 1968)

Group I

(direct flowering)

February-April

Group la

(indirect flowering)

February-March

Group II

(direct flowering)

July-September

Group III

(indirect flowering)

February-May

A. ahietis-religiosae'i

A. globosum

A. vaginatum subsp. vaginatum

A, vaginatum subsp. cryptopodum

A. vaginatum subsp. durangense

A. gillii subsp. gillii

A. gillii subsp. nigrum

A. verticilliflorum

A. ahietinum f. sp.

concoloris

A. ahietinum f. sp.

magnificae

A. apachecum

A. bicarinatum

A. hlumeri

A. californicum

A. campylopodum

A. cyanocarpum

A. divaricatum

A. guatemalense

A. hondurense

A. laricis

A. microcarpum

A. occidentale

A. ruhrum

A. strictum

A. tsugense

A. americanum

A. douglasii

A. pusillum

initial shoot appearance). A. guatemalense, A.

vaginatum subsp. cryptopodum, and probably other

southern species germinate in the fall, while all

North American species in the temperate zone

germinate in late winter or spring.

Hosts

The host, important in the classification of dwarf

mistletoes, was discussed earlier. In most instances

a dwarf mistletoe will be collected on its principal

host. If a collection is made on other hosts, it is

important for identification to record the principal

host in the immediate vicinity. For example, if a

mistletoe occurs on Pinus flexilis, it is usually A.

cyanocarpum. However, if the dwarf mistletoe is a

rare crossover of A. americanum, information that

the mistletoe was rare on Pinus flexilis, but common

on associated Pinus contorta, would help to identify

the parasite. Situations are relatively rare where two

species of dwarf mistletoe are both common in a

given stand and, in such situations, crossovers are

extremely rare.

Witches' Brooms

The dwarf mistletoes cause two basic types of

witches' brooms, systemic or nonsystemic (see p. 26).

Since the type formed is usually consistent with a

host-parasite combination, this taxonomic feature

can be used to help classify the species. Five taxa

(A. americanum, A. douglasii, A. guatemalense, A.

minutissimum, and A. pusillum) consistently cause

systemic brooms. Most other taxa cause the nonsystemic type. In Guatemala, A. glohosum forms

witches' brooms, but in Mexico it does not. Witches'

broom formation is not exhibited, or is inconspicuous, in these host-parasite combinations: A. divaricatum on Pinus monophylla. A, hlumeri on Pinus

strohiformis, and A. occidentale on Pinus sahiniana.

Chemical Characters

Our analysis of the phenolic constituents of

Arceuthohium is the first comprehensive chemotaxonomic study in the mistletoes.

Greenham and Leonard (1965), who compared the

amino acids of three dwarf mistletoes (A. ahietinum

f. sp. concoloris and magnificae and A. occidentale)

with their respective host trees, reported some

similarities in amino acid composition between

host and parasite, but their results did not help

explain host specificity.

Relatively little information on the phenolics of

the mistletoes is available; hence, we are presenting

a brief review of the known literature.

ViscACEAE.—Bate-Smith (1962) made preliminary

studies of the phenolic compounds in the European

Arceuthohium oxycedri and Viscum alhum. KuangFang and Shih-Chueh (1957) recorded four flavonoids in V. alhum subsp. coloratum from China.

49
Arceufhobium

A.globosum

A. a biefi s - rellgiosae

A. douglasii

A.gillil subsp. gil Hi

A, gillii subsp. nigrum

A. vag, subsp. vaginatum

A. verficilliflorum

A. vag. subsp. durangense

A.americanum

A. pus/Hum

A. vag, subsp. crypfopadum

A.caiifornicum

A.rubrum

A. sfrief urn

A, blumeri

A. Iarlei s

A, ablet, f. s p. concoloris

A. abief, f. sp. magnificae

A,guafemalen$e

A.campylopodum

A.cyanocarpum

A, microcarpum

A, bicarinafum

A, apachecum

A, divaricafum

A fsugense

A.hondurense

A,occidentale

JAN. FEB. MAR. APR. MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC.

Figure 20.—Approximate periods of anthesis for the New World dwarf mistletoes.

LoRANTHACEAE.—Some phenoHc constituents

have been isolated from the AustraHan species

Nuytsia floribunda (Bate-Smith 1962), and from

two South American mistletoes Psittacanthus cuneifolius and Phrygilanthus flagellaris (Graziano,

Widmer, JuHani, and Coussio 1967; Widmer and

Coussio 1969). Several Asian species have also been

studied, and the flavonoid quercitin has been found

in Loranthus peniandrus and L. globosus (Wester

1921). This flavonoid and arabinoside have been

reported in Loranthus parasiticus (Kuang-Fang and

Chung-Liang 1957). Khanna, Viswanathan, Tewari,

and others (1968) found a relatively high concentration of phenolics in the mistletoe Dendrophthoe

falcata and two other parasitic angiosperms in India.

A relatively wide range of shoot colors is known in

Arceuthobium. Although there is some infraspecific

variation, shoot color is relatively constant within a

species. We attempted to determine w^iether these

color differences were associated with specific

pigments.

Prehminary analyses indicated that a dark

species {A. rubrum) contained anthocyanins, whereas

a yellow species {A, stridum) did not. These fragmentary data suggested that anthocyanins might

be positively correlated with the dark coloration

(red or black) characteristic of many species, and

negatively correlated with the lighter coloration

(greens, yellows, and oranges) of others. We then

surveyed all dwarf mistletoe taxa to determine the

50
A.globosum

A. rubrum

A. vag. s\jíbs\>. cryptopodum

A. vag. subsp. durangense

A, gillii subsp. nigrum

A.cyanocarpum

A. apachecum

A.americanum

A, vag. subsp. vaginafum

A. bicarinafum

A. blumeri

A. Iarid s

A.guatemalense

A.microcarpum

A.douglasii

A.pusillum

A.hondurense

A. abiet. f. s p. concoloris

A. campylopodum

A.californicum

A. divaricatum

A tsugense

A. strictum

A. a biet, f. sp. magnificae

A. abieti s - rellgiosae

A. verticilliflorum

A. gill il subsp. gillii

A. occidentale

JUNE JULY AUG. SEPT. OCT. NOV. DEC.

Figure 21.—Approximate periods of seed dispersal for the New World dwarf mistletoes.

relationship between anthocyanin content and

shoot coloration. In addition, we evaluated several

additional phenoHc compounds (flavones, flavonols,

and cinnamic acid derivatives) because they could be

chromatographically isolated from the original

extracts in ethyl acetate along \\dth the anthocyanins,

and appeared to be valuable taxonomic characters in

classification of Arc6i¿¿/io6mm. Alston, Rosier, Naifeh,

and Mabry (1965) found the phenolic compounds

to be of chemotaxonomic value because, at least for

the flavonoids, Mendehan mechanisms apparently

govern their qualitative differences.

Methods

The tissue utilized for extraction represented

approximately the terminal centimeter of growi^h,

and was obtained from staminate herbarium material. The internodes were ground to powder with a

mortar and pestle with the aid of glass beads. The

phenolic compounds were extracted in 5 ml. of 1

percent hydrochloric acid (HCl) in absolute

methanol (v/v), followed by four additional washes

of the same volume. The extracts were then combined and concentrated at 40°C. to approximately

3 to 4 ml. by evaporation into nitrogen. After

initial concentration, approximately 5 ml. of 0.5

percent aqueous HCl was added and the solution

then reconcentrated to about 2 ml. Carotenoids were

removed by phasing with petroleum ether (30-60°C

fraction).

Non-anthocyanin phenolic compounds were removed from the aqueous phase with ethyl acetate.

The organic phase was then back extracted with

0.5 percent aqueous HCl.

The extracts were immediately applied to Whatman #3 MAI Chromatographie paper with finely

51
Arceuthobium

A. pusi Hum

A.apachecum

A.bicarinatum

A,cyanocarpum

A.hondurense

A tsugense

A.microcarpum

A.rubrum

A.guatemalense

A. blumeri

A.campylopodum

A.occidentale

A. divaricatum

A. Iarid s

A.strictum

A, abiet. f.sp. concoloris

A. abiet. f. sp. magnificae

A.californicum

A. vag. syibsp. cryptapodum

A.globosum

A. vag. subsp. durangense

A,americanum

A, gillii subsp. nigrum

A. vag, subsp. vaginatum

A. abieti s - rellgiosae

A, douglasii

A. verticilliflorum

A, gill il subsp. gillii

8 12

MONTHS

16 20

Figure 22.—Approximate fruit maturation periods for the New World dwarf mistletoes.

drawn Pasteur pipets. For one-dimensional chromatography, the extracts were applied in strips 3 cm.

Jong. For two-dimensional development, they were

applied in spots about 2 cm. in diameter.

The anthocyanin-containing fraction was chromatographed in water: glacial acetic acid:12 N HCl in

the ratio 82:15:3 (v/v). Development time was 18

hours. The ethyl acetate fraction was chromatographed in 15 percent aqueous acetic acid (v/v)

for 6 hours. The various bands were consistently

discernible under 3440 A° ultraviolet (UV) light

both before and after drying.

Results

All species and subspecies of Arceuthobium, except

A. chineuse and A. pini, were analyzed chromatographically (table 16). These data must be considered as preliminary, however, because of the

small number of individuals analyzed in some species.

52
and because the bands are only tentatively identified.

Furthermore, the ethyl acetate fraction contained

many compounds which cochromatographed in 15

percent aqueous acetic acid, but were separable

when developed in the second dimension with

BAW (12:3:5). In the numerical analyses, however,

these paired bands were scored as unit characters,

as were the chromatographically pure bands. No

attempt was made to analyze quantitative differences, although these were sometimes obvious. The

chromatogram of A, microcarpum (fig. 23) contains

most of the flavonoid compounds presently known

in Arceuthohium, and shows the band numbering

system used in our studies.

ANTHOCYANINS.—The aqueous phase that remained after extraction with ethyl acetate contained

up to three magenta bands discernible in visible

light. After hydrolysis, the aglycon from each band

was shown to be cyanidin. This conclusion was

derived from the results of cochromatography with

authentic cyanidin in several solvent systems, and of

UV and visible spectrophotometry.

The specific distribution of cyanidin is interesting

because it occurs not only in all species with dark

coloration, which we suspected, but also in many

which are yellow, greenish yellow, or orangish in

coloration.

FLAVONOLS.—A conspicuously yellow fluorescing

band (Y), observable under UV, contains two compounds, one of which is probably quercitin, as

determined by cochromatography with known

quercitin. The second component of the band is unknown, but may be derived from kaempferol.

The unknown band YO has some flavonol-like

properties, and, in at least two species (A. campylopodum and A. microcarpum) is separable into two

compounds when developed in the second dimension

in BAW (12:3:5). The yellow fluorescing band, along

with the BG bands of putative cinnamic acid derivatives (vid infra), are of nearly universal occurrence

in the populations studied. Band Y occurred consistently in every sample except A. vaginatum subsp.

durangense. The single and rare YG band is also

tentatively placed here.

FLAVONES.—The most interesting feature of the

chromatograms viewed under UV is the occurrence of

4 (rarely 5 or 6) purple bands (A bands) that appear

to be glycosides of flavones. These bands turn yellow

with age or when fumed with ammonia. PreHminary

observations in A. microcarpum show that bands Ai

and As are separable into two compounds when

developed in the second dimension. Also, the single

D band, presently unique to A. guatemalense,

probably belongs in this group of compounds.

The flavones exhibit more discontinuous variation

and perhaps less infrataxon variation than most of

origin

»G^CD

Al

A2

A3

A4

B4

BS

B6

solvent

front

Figure 23.—Chromatogram of

Arceuthohium microcarpum

showing typical separation of

compounds soluble in ethyl

acetate.

53
O

O

Ü

0)

Ü

a

a

T3

03

O

>

03

ce

tí

o

PQ

ffl

m

o

o

PQ

o

+ +

+

+

+ ¡

' ' ' +

, , .+ ' ' '+

¡ +

+

+

+

4-4-4-4- +++^

+ ++ , + .+ ,+ , ++ + .4-^++ " + + + ++^+ '+ '4-^4-+

4- 4- 4-

4- 4-

¡ 4-

4- 4-

4- 4-

4- ,4-4-4-

'4- '4- +4-

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54
the other chemical classes; they are totally lacking

in A. hicarinatum, A. vaginatum subsp. cryptopodum

and subsp. durangense, A. oxycedri, and A. ruhrum.

This group of compounds might provide the best

possibility for further chemotaxonomic study in

Arceuihohium.

CiNNAMIC ACID DERIVATIVES.—A COUSpicUOUS

feature of most chromatograms of Arceuthohium

extracts is a series of bright blue fluorescing bands

(B bands) which are probably cinnamic acid derivatives. With two exceptions, all B bands appear to

contain single components. The exceptions are the

B5 and Be bands which are separable in the second

dimension in A. campylopodum, A. microcarpum, and

A. minutissimum. These two bands are also difficult

to separate in the single dimension, and were arbitrarily scored as B5 unless, as in the three cases

mentioned, the results of second dimension chromatograms confirmed the existence of both bands.

As with the A bands, this group of compounds

displays considerable interspecific diversity. Several

species contain none of these presumed cinnamic

acid derivatives: A. hicarinatum, A. gillii subsp.

nigrum, A. ruhrum, A. vaginatum (including all of its

subspecies), and A. verticilliflorurn; A. gillii subsp.

gillii contains only a single blue fluorescing band,

which was found in only one of four populations

analyzed.

In addition to the bands fluorescing bright blue,

two additional bands fluorescing a light blue green

under UV are tentatively placed in this class of

compounds. The most significant aspect of the BG

bands is their occurrence in every species analyzed

and in all but two of the populations sampled.

Furthermore, both of these bands are separable when

chromatographed in the second dimension.

Discussion and Conclusions

Anthocyanins were found in all the species with

dark-colored shoots, and more than half of the light

colored ones (table 16). We also found a light

magenta color pattern in the living staminate

flowers of two light-colored species that resembled

the anthocyanin bands we had isolated chromatographically from their shoots. Since the color pattern

is evident in the outer portions of living staminate

flowers oí A. americanum and A. douglasii, we suspect

its function is to visually attract pollinators (apparently Diptera in these species). We plan to study the

living flowers of all the üght-colored species to determine if a similar light magenta coloration might

be present.

CHROMATOGRAPHIC VARIATION.—Three aspects of

variation were considered : qualitative, quantitative,

and comparison of living and dead specimens.

Qualitative infrataxon variation in chromatographic patterns (presence or absence) often occurred among the populations of a particular species.

To determine the extent of such variation within a

species, 34 populations of A. americanum were

analyzed. The variables included: (1) hosts—

Pinus contorta (two subspecies), P. ponderosa, and

P. hanksiana; (2) geographical distribution—West

Coast, and the southern Rocky Mountains to

central Canada; (3) time since collection—specimens

collected a few days to over 50 years before analysis;

and (4) season of collection—most months. Analyses

showed that 30 of the 34 populations of A. americanum were qualitatively uniform (table 16). In the

four chromatograms that varied, the variation was

due to the visible anthocyanin band 2, and the bands

of phenohc compounds visible under UV (B3, B4, Ae),

with the most obvious variation in the B3 and B4

bands. The A3 and A4 bands are in close proximity,

and when small quantities of the compounds are

present interpretation becomes difficult, but some

variation was suggested in our samples.

Quantitative variation in virtually all of the bands

was a common feature of the chromatograms.

Detecting compounds at low levels of concentration

probably introduced some variability in the results.

The Y band, which contains two quantitative

phases, wus consistently detectable (except in some

samples of A. vaginatum subsp. durangense). We

thought the lighter phase might indicate the presence

of only one of the two compounds that compose this

band (see Results) but when a sample of A. americanum that contained the light phase was chromatographed in the second dimension, the Y band

still separated into two parts.

Although both qualitative and quantitative variation exists in the chromatographic patterns of

dwarf mistletoes, our experience with A. americanum

suggests that, where large samples are available,

chemical data can provide reUable taxonomic information. The data could also be more useful if two

dimensional analyses were available for each

compound.

To determine whether chromatographic variation

existed, we analyzed living and dried shoots from a

putative clone of A. americanum. The chromatograms varied little, except that the BG2 band,

apparent in all dried samples, was noticeably absent

in the live specimens, which suggests that it may be a

degradation product. Also, in the chromatogram for

the living shoots, a single band fluorescing yellow

under UV occurred between the BG2 band location

and the A band.

TAXONOMIC IMPLICATIONS.—Although the chemical data do not show discontinuous variation among

all taxa, many species are distinct chromatographically, and can be readily identified by this means.

A. americanum is the only species we intensively

55
analyzed, but our preliminary studies indicate

that comparable differences will be obvious when the

other taxa are studied.

A. hicarinatum, one of the most chromatographically distinctive of all dwarf mistletoes, lacks all

the A and B bands, and although we first confused

A. hondurense with A. hicarinatum, the Chromatographie differences we found suggested these two

taxa were specifically distinct.

Other species such as A. ahietis-religiosae, A.

ruhrum, A. stridum, and A. vaginatum subsp.

cryptopodum and durangense are similar chromatographically; they lack the A bands but are

distinctive in other ways. Although all five are Mexican species, they do not appear to form a phyletic

group, but are scattered throughout various subgenera and sections. Thus, although certain species

appear to be mutually distinct chromatographically,

no Chromatographie features seem to be associated

with the various subgeneric groupings.

NUMERICAL ANALYSES

The use of computer techniques in taxonomy

(taximetrics), essentially a development of the

1960's (Sokal and Sneath 1963; Rogers, Fleming, and

Estabrook 1967), is a powerful new tool for making

taxonomic decisions. Although some classical taxonomists view this development with suspicion, most

numerical taxonomists have emphasized that taximetric analyses will supplement but not replace

conventional systematics techniques (Rogers and

Appan 1969). How good the taximetric analyses are

for a particular group depends on the taxonomist's

ability to discern significant characters in the group

and to separate them into meaningful states. Both

the numerical taxonomist and the classical taxonomist need to thoroughly understand the group in

question before they can make systematic decisions.

Methods

The Graph Theory Clustering Program^^ is the

computer technique we found best suited to analyze

our extensive taxonomic data (table 17). We made a

prehminary taximetric analyses of the characters of

Arceuthohium (Hawks'worth, Estabrook, and Rogers

1968), then chose the following taximetric methods

for classifying the species :

1. On the basis of our field and herbarium experience with the dwarf mistletoes, we selected a number

of qualitative and quantitative characters that we

considered taxonomically significant. Because these

plants are leafless and extremely reduced morphologically, we emphasized other types of characters

in addition to morphological ones.

2. Each selected character was subdivided into 2

to 18 states, depending on the range of variation

shown in the particular character. The number of

17 Details of the Program and its application are summarized in Estabrook 19G7; Estabrook and Rogers 1966: Irwin

and Rogers 1967; Rogers and Appan 1969; and Wirth, Estabrook, and Rogers 1966.

states we selected for each character was based on the

work by Rogers and Fleming (1964) and Estabrook

and Rogers (1966).

3. We adjusted the number of states for each

character so the range of overlap (Estabrook and

Rogers 1966) was either one or zero. This overlap

value, for example, is used for shoot height (character

No. 2, table 17), where three states are based on the

relative height of the staminate and pistillate shoots :

(1) staminate shoots tallest, (2) staminate and

pistillate shoots about the same, and (3) pistillate

shoots tallest. Examination of many collections

showed that within a taxon, ratings were usually 1

and 2, but not 3; other taxa were rated as either 1, 2,

or 3; while others were 2 and 3, but not 1. Thus an

overlap value of 1 was assigned, which indicates

that only states that are more than 1 unit apart

(or in this instance states 1 and 3) are computed as

different. In many other characters, (for example.

Principal hosts, character No. 38, table 17) an overlap value of zero is used, which indicates that differences in any two states are calculated as equally

distinct.

4. The 60 characters used in this study (see table

17) are divided as follows: shoots (12), flowers (11),

pollen (5), fruit (5), phenology (4), hosts and host

reactions (3), and shoot pigments (21).

5. In the analyses we gave all characters equal

weight because of the difficulty of assigning meaningful weights to the various characters. As will be

shown, however, certain characters contain much

more taxonomic information than others and thus

contribute more to the resulting classiftcation.

6. A total of 212 specimens representing all known

28 North American taxa were examined and rated

in each of the 60 characters. The number of specimens per taxon varied from 12 to as few as 3 in some

of the seldom-collected Central American and Mexican taxa. The specimens were selected on the

following basis: (a) those that yielded the most

information, and (b) those that represented the

56
TABLE 17.—Characters and states used in taximetric studies in Arceuthobium

SHOOTS:

1. Mean plant height

(overlap—1 unit)

1. Less than 1 cm.

2. 1-2 cm.

3. 3-5 cm.

4. 6-10 cm.

5. 11-13 cm.

6. 14-18 cm.

7. 19-23 cm.

8. Over 24 cm.

2. Shoot height

(overlap—1 unit)

1. Staminate shoots tallest

2. About the same

3. Pistillate shoots tallest

3. Mean basal diameter

(overlap—1 unit)

1. 0.1 cm.

2. 0.2 cm.

3. 0.3 cm.

4. 0.4 cm.

5. 0.5 cm.

6. 0.6 cm.

7. 0.7 cm.

8. Over 0.8 cm.

4. Third internode length

(overlap—1 unit)

1. 1.0-2.9 mm.

2. 3.0-4.9 mm.

3. 5.0-6.9 mm.

4. 7.0-8.9 mm.

5. 9.0-10.9 mm.

6. 11.0-12.9 mm.

7. 13.0-14.9 mm.

8. 15.0-16.9 mm.

9. Over 17.0 mm.

FLOWERS:

13. Staminate floral buds

(overlap—none)

1. Spherical

2. Lenticular

14. Pre-flowering lateral staminate

spikes—length

(overlap—1 unit)

1. None

2. 1-2 mm.

3. 3-4 mm.

4. 5-6 mm.

5. 7-8 mm.

6. 9-10 mm.

7. 11-13 mm.

8. 14-16 mm.

9. Over 17 mm.

5. Third internode width

(overlap—1 unit)

1. Up to 1.0 mm.

2. 1.1-2.0 mm.

3. 2.1-3.0 mm.

4. 3.1-4.0 mm.

5. 4.1-5.0 mm.

6. 5.1-6.0 mm.

6. Third internode length/width ratio

(overlap—1 unit)

1. Less than 2.0

2. 2.1-4.0

3. 4.1-6.0

4. 6.1-8.0

5. Over 8.1

7. Branching

(overlap—none)

1. No accessory branches

2. Accessory branches sometimes verticillate

3. Accessory branches always flabellate

8. Sexual dimorphism

(overlap—none)

1. Branching similar

2. Branching different

9. Shoot color

(overlap—none)

1. Pistillate and staminate shoots of

same color

2. Pistillate and staminate shoots

of different color

10. Color of pistillate shoots

(overlap—1 unit)

1. Green

2. Yellow

3. 1

4. Orange

5. Red

6. Purple

7. 1

8. Brow^n

9. 1

10. Black

11. Shoot density

(overlap—none)

1. Shoots densely clustered around host

branch

2. Shoots not densely clustered around

host branch

12. Overall plant shape

(overlap—none)

1. Plants in globose masses

2. Plants not as above

15. Pre-flowering lateral staîuinate

spikes—width

(overlap—1 unit)

1. None

2. 1 mm.

3. 2 mm.

4. 3 mm.

5. 4 mm.

6. 5 mm.

16. Stajninate flowers

(overlap—none)

1. Verticillate

2. Opposite, with pedicellate

joints

3. Opposite, no pedicellate

joints

17. Number of staminate perianth segrnents

(overlap—1 unit)

1. 2 or 3 (approx. equally divided)

2. All 3's

3. Mostly 3, few 4's

4. 3 or 4 (approx. equally divided)

5. Mostly 4, few 3's

6. 3, 4, or 5 (approx. equally divided)

7. Mostly 5 or more.

18. Staminate flower diameter

(overlap—1 unit)

1. 1.0-1.4 mm.

2. 1.5-1.9 mm.

3. 2.0-2.4 mm.

4. 2.5-2.9 mm.

5. 3.0-3.5 mm.

57
TABLE 17.—Characters and states used in taximetric studies in Arceuthobium—Continued

6. Over 3.5 mm.

19. Anther diameter

(overlap—1 unit)

1. 0.3-0.4 mm.

2. 0.5-0.6 mm.

3. 0.7-0.8 mm.

4. 0.9-1.0 mm.

20. Anther distance from perianth

tip (overlap—1 unit)

1. 0.3-0.4 mm.

2. 0.5-0.6 mm.

3. 0.7-0.8 mm.

4. 0.9-1.0 mm.

POLLEN:

24. Length

(overlap—1 unit)

1. 18-19M

2. 20-21/x

3. 22-23M

4. 24-25)U

5. 26-27/i

6. 28-29M

25. Width

(overlap—1 unit)

1. 18-19M

2. 20-21M

3. 22-23/x

4. 24-25M

5. 26-27M

6. 28-29M

FRUIT:

29. Surface

(overlap—none)

1. Glaucous

2. Nonglaucous, dull

3. Nonglaucous, shiny

30. Length

(overlap—1 unit)

1. 2 mm.

2. 3 mm.

3. 4 mm.

4. 5 mm.

6. 1

7. Over 10 mm.

PHENOLOGY:

34. Peak flowering period

(overlap—1 unit)

1. March-April

2. May-June

3. July

4. August

5. September

6. October or later

35. Peak seed dispersal period

(overlap—1 unit)

1. July

2. August

3. September

4. October

5. November or later

21. Staminate perianth length

(overlap—1 unit)

1. 0.6-0.9 mm.

2. 1.0-1.3 mm.

3. 1.4-1.7 mm.

4. Over 1.8 mm.

22. Staminate perianth width

(overlap—1 unit)

1. 0.6-0.9 mm.

2. 1.0-1.3 mm.

3. 1.4-1.7 mm.

4. Over 1.8 mm.

23. Pistillate flowers

(overlap none)

1. Verticillate

2. Opposite

26. Spine length

(overlap—1 unit)

1. 0.5-0.9/1

2. 1.0-1.4M

3. 1.5-1.9M

4. 2.0-2.5M

27. Wall thickness

(overlap—1 unit)

1. 0.5-0.9M

2. 1.0-1.4/x

3. 1.5-1.9M

4. 2.0-2.4M

31. Distal portion as percent of total fruit

length (overlap—1 unit)

1. 21-25 percent

2. 26-30 percent

3. 31-35 percent

4. 36-40 percent

5. 41-45 percent

32. Sterile nodes near tips of pistillate shoots

(overlap—none)

1. Present

2. Absent

36. Mean fruit maturation period

(overlap—1 unit)

1. ca. 5 months

2. 1

3. ca. 12 months

4. ca. 13 months

5. 1

6. Over 14 months

37. TÍ7ne of meiosis in relation to flowering

(overlap—none)

1. Spring flowering, spring meiosis

2. Winter flowering, fall meiosis

3. Summer flowering, summer meiosis

4. Spring flowering, fall meiosis

28. Spine/wall ratio

(overlap—1 unit)

1. 0.3-0.7

2. 0.8-1.2

3. 1.3-1.7

4. 1.8-2.2

5. 2.3-2.7

6. 2.8-3.2

7. 3.3-3.8

33. Pedicellate fruits on main axis

(overlap^none)

1. Common

2. Occasional or rare

3. None

58
TABLE 17.—Characters and states used in taximetric studies in Arceuthobium—Continued

HOSTS AND HOST REACTIONS:

38. 39.

40.

Principal hosts

(overlap—none)

1. Abies {concolor, grandis)

2. Abies magnifica

3. Abies (other species)

4. Pseudotsuga

5. Picea {mariana, glauca)

6. Picea (engelmanniif

pungens)

7. Larix

8. Tsuga

9. Pinus lambertiana

10. Pinus (flexilis-aristata)

11. Pinus strobiformis

12. Pinus ayachauite

13. Pinus (pinyons)

14. Pinus (Leiophyllae)

15. Pinus (Australes)

16. Pinus (Ponderosae)

17. Pinus (Sabinianae and

Oocarpae)

V\. Pinus (Contortae)

CHROMATOGRAPHY OF SHOOT PHENOLICS:

41. to 43.

Anthocyanins

(overlap—none)

1. Consistently present

2. Sometimes present

3. Absent

Anthocyanin #1

Anihocyanin §2

43. Anthocyanin #3

44. to 60.

Flavones, flavonols and cinnamic acid derivatives

overlap—none

1. Consistently present

2. Sometimes present

3. Absent

44. Phenolic Yi

45. Phenolic D

46. Phenolic Bi

47. Phenolic Y2

48. Phenolic BGi

49. Phenolic YO

50. Phenolic BG2

51. Phenolic Ai

52. Phenolic YG

Witches' broom formation

(overlap—none)

1. None

2. Rare or occasional

3. Consistent

Type of witches^ brooms

(overlap—none)

1. Brooms systemic

2. Brooms nonsystemic

3. No brooms

41.

42.

53. Phenolic A2

54. Phenolic B3

55. Phenolic A3

56. Phenolic A4

57. Phenolic B4

58. Phenolic A5

59. Phenolic B5

60. Phenolic Be

^ A skipped state number indicates that the two adjacent states are calculated as distinct when the overlap value is 1 unit.

2 See table 16. Two compounds (B2 and Ae) listed in table 16 were not used in the taximetric analyses because their occurrence

was too erratic.

geographic and host range of the taxon. The specimens used in these analyses are indicated in the

specimens examined lists.

7. Data were entered on IBM cards and analyses

were made on a CDC 6400 computer at Colorado

State University.

8. A measure of similarity was then calculated

between each specimen and every other one in the

analysis. This similarity was based on the fraction

of the characters which were coded for both specimens. For example, if two specimens were rated

the same in all characters, their similarity would be

1.00.

9. The clustering program compares the overall

59
similarity (the averaged similarities of each state for

which information is given) of each specimen with

all others in the study. The specimens are then

clustered in accordance \\ith the rank of their

relative similarity. For example, specimens that

have a similarity ratio of 1.00 are separated first,

then additional specimens are clustered as the

similarity ratio is relaxed, and the program continues until all specimens are included in a single

group.

Results

A numerical analysis of the New World taxa of

Arceuthohium (fig. 24) shows the relationships of the

taxa based on average similarity of all characters

listed in table 17.

Because the characters and states used in each

taximetric study are different, the results of our

Arceuthohium study cannot be compared with those

in other groups. The taxa oí Arceuthohium, however,

were found to be unusually distinct, and all species

were formed by similarity level 0.85. The analyses

suggest that the genus consists of two major groups

plus several smaller ones, some of which contain

only a single taxon.

Arceuthohium verticilliflorurn, A. ahietis-religiosae,

and A. americanum, the most distinctive North

American species (fig. 24), share few features in

common, except all have verticillate branching of the

shoots or flowers (Kuijt 1970).

Two distinctive taxa, Arceuthohium pusillum and

A. douglasii, which form another group, are characterized by their small size, indirect spring flowering, high host specificity, and formation of systemic

witches' brooms.

A major group (the vaginatum group) consists of

A. verticilliflorum

A. abietis - rsllgiosae

A.americanum

A. pusillum

A, douglasii

A.gillH subsp. glllii

A.gillii subsp. nigrum

A.vag. subsp. vaginatum

A. vag. subsp. cryptopodum

A. vag. subsp. durangense

A. globosum

A.sfrictum

A.rutrum

A.bicarinafum

A.hondurense

A.guafema/ense

A.occidentale

A.cyanocarpum

A. blumeri

A. apachecum

A.campylopodum

A. tsugense

A. californicum

A. divaricatum

A. Iarid s

A.microcarpum

A. ablet, f. s p. concoloris

A. abiet, f. sp. magnificae

1 1 1 T 1 f 1 I

• 1

• 1

•

•

•

•

1

1 1

I—,

•

•

•

•

•

—•

-#

—•

•

1

—•

•

•

•

•

•

•

• '

-

.*

-1- - -

1—1. 1111

1.00 0.95 0.90 0.85 0.80 0.75

SIMILARITY

0.70 0.65 0.60

Figure 24.—Dendrogram showing taximetric analyses of the New World dwarf mistletoes. The scale shows similarity,

hased on combined analyses of all 60 characters used. Vertical lines indicate the level at which two taxa were

joined. Length of the horizontal line indicates the distinctness of each taxon. Dot shows the similarity level at

which all specimens of the particular taxon w^ere first united.

60
six taxa comprising three species and three subspecies: A. glohosum; A. gillii subsp. gillii and nigrum;

A, vaginatum subsp. vaginatum, cryptopodum, and

durangense. The analyses of this group illustrate our

use of subspecies in Arceuthohium. For example,

although subsp. gillii and nigrum are distinct, and

maintain their identities to about level 0.79, common

features unite them for about another 0.05 similarity

unit before the group joins with the A. vaginatum

complex. A comparable situation exists for A.

vaginatum. These are the only instances where we

believe that a subspecific category is appropriate

in Arceuthohium. This vaginatum group contains

relatively unspecialized taxa with large shoots and

broad host ranges on pines, and is centered in

Mexico. All exhibit spring flowering.

The other major group, the campylopodum

group, contains 17 taxa: A. abietinum f. sp. concoloris

and magnificae, A. apachecum, A. hlumeri, A.

californicum^ A. campylopodum, A. cyanocarpum, A.

divaricatum, A. guatemalense, A. laricis, A. microcarpum, A. occidentale, and A. tsugense; the less

closely related group A. hicarinatum, A. hondurense,

A. ruhrum; and the even more distinct A. strictum.

The 13 principal members of this group are concentrated in the western United States, but some

range from southeastern Alaska to northern Guatemala. Arceuthohium hicarinatum, A. hondurense, and

A. ruhrum form a distinct southern subgroup and

occur in Hispaniola, Honduras, and central Mexico,

respectively. The analyses suggest that A. strictum

is the most distinct member of the entire campylopodum group. Arceuthohium guatemalense is the most

distinct of the main 13-member subgroup.

The black dot in ñgure 24 indicates the similarity

level by which all specimens of a particular taxon

had been united. Thus, the distance of the dot from

the 1.00 similarity level indicates variability within

a taxon; species with dots closest to 1.00 (for example, A. ahietis-religiosae, A. strictum, A. pusillum,

A. hicarinatum, A. hondurense, A. guatemalense,

and A. hlumeri) are least variable while those with

the dot farthest from 1.00 (A. gillii subsp. gillii and

A. glohosum) are the most variable. In general, the

vaginatum group seems to be the most variable.

Perhaps this is correlated with the primitive status of

the group.

The length of the horizontal line from the dot to

the point where it joins another taxon is a direct

indication of the distinctness of the particular taxon.

For example, A. verticilliflorum, A. ahietis-religiosae,

A. americanum, A. pusillum, A. douglasii, and A.

strictum are the most distinct, and their lines range

from 0.25 to 0.33 similarity unit in length. Conversely, the/ormae spéciales of A. ahietinum are the

least distinct and their lines are only 0.03 to 0.05

unit long.

Character analyses oí Arceuthohium (Hawksworth,

Estabrook, and Rogers 1968) provide data on the

quality of information contributed by each character

toward the overall classification of the genus based

on the synthesis of all characters used. The analyses

showed that the five characters with the highest

quantity of information, that is, those that contributed the most to the classification were, in

order:

No. 39—principal hosts (most information)

No. 3—mean basal diameter of dominant

shoots

No. 38—time of meiosis in relation to time of

flowering

No. 15—length of lateral staminate spikes

No. 16—width of lateral staminate spikes.

Similarly, the five characters that contributed

least to the final classification were :

No. 27—height of pollen spines

No. 23—width of perianth segment on

staminate flowers

No. 8—sexual dimorphism of branching

No. 9—difference in color of pistillate and

staminate plants

No. 21—location of anther on perianth lobe

(least information).

COLLECTING AND CURATING TECHNIQUES

At many of the herbaria we visited, the Arceuthohium specimens were broken or fragmented.

Invariably the plants were fragmented if a large

branch with an attached dwarf mistletoe had been

pressed.

In preserving our specimens, we had little fragmentation if we pressed individual shoots rather than

preserving the entire dwarf mistletoe plant. To

include the basal shoot internodes, which we found

useful taxonomically, we cut the host branch so

that a small portion of the host bark remained

attached to each shoot. For smaller species such as

A. douglasii or A. pusillum, we pressed the entire

infected host branch if it measured less than about

one-fourth inch in diameter. Also, we retained enough

of the host branch for identification, and collected

61
both staminate and pistillate dwarf mistletoe

plants.

On many herbarium specimens we studied, collection data were meager. Information that should be

recorded includes the host, ecological data about the

site, and shoot characters such as habit size, sexual

dimorphism, and original color. Also, the presence or

absence of witches' brooms should be recorded and,

if more than one host species is involved, their

relative susceptibility noted. Any additional information that will help identify either the parasite

or its host should be recorded.

Our herbarium specimens are better preserved

when we used the following technique for mounting:

Before the plants became totally dry, we applied a

heavy coat of adhesive to the back of each specimen,

and mounted them on heavy grade herbarium

paper. Another satisfactory method is to use a

cotton-backed envelope or a Riker Mount. A

modification of the latter is to place the dried specimen directly on Tomac,i^ and file it in a large

envelope with a transparent window so it can be

observed with little handling.

Formal Taxonomy

Generic Description

ARCEUTHOBIUM M. BIEB. DWARF MISTLETOE

RazoumofskyalíoñmBin, Hort. Mosq., unpaged, 1808.

Arceuthobium M. Bieb. Fl. Taur.-Caucásica 3(IV)

Suppl.: 629, 1819. Norn. Cons. 2091.

Herbs or shrubs from 0.5 cm. to approximately

70 cm. high, parasitic on coniferous trees of the

Pinaceae or Cuppressaceae; plants glabrous, variously colored from greenish yellow to orange;

reddish and black, dioecious; stems without a central

xylem cylinder; leaves reduced to minute, opposed,

connate scales; internodes angled (at least when

young) ; flowers generally decussate or rarely whorled

on young shoots, 2-4 mm. in diameter; staminate

flowers with a central nectary, perianth segments

usually 3-4 (rarely 2 and up to 7) bearing a sessile,

uniloculate, circular anther on each perianth segment; pollen spherical with 6 alternating spiny and

smooth section; pistillate flower manifestly ep-i

gynous, with one style, perianth segments persistent,

adnate to ovary, 2-merous; ovary 1-chambered;

fruit an ovoid berry, 1-seeded, mucilaginous and

bicolored (distal and basal portions of different

shades), explosive at maturity; seeds without true

integuments, usually 3-5 mm. long, ovate-lanceolate;

containing a single, distal, cyhndrical embryo, with

copious endosperm. Basic chromosome number,

x= 14.

A genus of approximately 32 taxa mostly in the

United States and Mexico; apparently four species

in the Old World.

Type species: A. oxycedri (DC.) M. Bieb.

Subgeneric Classification and

Natural Keys to the Species

Ours is the first attempt to establish a formal

below-genus classification in Arceuthobium (fig. 25).

We recognized two subgenera, Arceuthobium and

Vaginata, based on the type of secondary branching,

verticillate and flabellate, respectively. Although

Kuijt (1970) suggested that secondary branching

could be the basis for natural subdivision of the

genus, he did not formally name the subdivisions.

We are not fully satisfied because this system emphasizes a single character (branching type), but we

have not discovered other groups of characters

that might form a better basis for natural subgenera.

Another difficulty is that some species (for example,

A. pusillum) are so reduced that their basic type of

secondary branching cannot be determined. Their

placement into subgenera, then, is based on assumed

relationships and is obviously tentative.

The subgenus Vaginata is exclusively New World,

and contains 25 taxa. This subgenus has been divided

into three sections (Vaginata, Campylopoda, and

Minuta), with the largest section Campylopoda,

further subdivided into three series: Campylopoda,

Rubra, and Stricta. We believe the three sections

represent a phyletic sequence from primitive

(Vaginata), to intermediate (Campylopoda), to

advanced (Minuta) species.

^^ Dr. W. A. Weber substituted Tomac for cotton because it

did not cling to the back of the specimen. Tomac is manufactured by the American Hospital Supply Corporation, North

Kansas City, Missouri.

62
Genus Arceuthobiunn

?;ubaenu5 Arceuthobium Subgenus Vaginata

o Ö Section Section Section

X

o

+ '

Vaginata Cannpylopoda Minuta

\I- CO Series Series Series

1

^

Cannpylopoda Rubra Stricta

-f-» H-»

O Ö

c c

CD O)

if) V)

(D 0 c

"Ö "Ö a Ö Ö o o

in X X X X X

d r^ Ö Ö Ö Ö Ö

O O +-» -+-> -h-> -H-» -H-»

U U

(D CO en ^ CM

(D 0

en en

O O

Z z

OLD WORLD NEW WORLD

Figure 25.—Taxonomic summary of the genus Arceuthobium (Hawksworth and Wiens 1970b).

The subgenus Arceuthobium contains four Old

World and three New World species. We have not

proposed formal subdivisions of this subgenus because detailed information on the Old World taxa is

lacking and there are only three New World taxa in

this group. We beheve the Old and New World

members are probably sectionally distinct, but we

hesitate to establish sections based exclusively on

geography. However, since the two groups apparently have been isolated since at least the

Miocene, it seems likely that, when the Old World

species are studied in detail, sufficient additional

differences will be found to warrant establishment of

sections. In the New World, the three members of

subgenus Arceuthobium seem to represent a phyletic

sequence similar to that shown by the sections

in subgenus Vaginata. We consider A. verticilliflorum

to be primitive, A. abietis-religiosae intermediate,

and A. americanum advanced (see fig. 15). The

three species are quite distinct and each should

perhaps be accommodated by separate series if

sectional status is estabhshed. Probably a comparable situation exists in the Old World, where A.

oxycedri is primitive, A. chinense and A. pini intermediate, and A. minutissimum advanced.

A synopsis of our classification of Arceuthobium

(see fig. 25) follows. The taxon numbers correspond

with those used in the alphabetical listing for the

New and Old World species.

I. Subgenus Arceuthobium (No formal sections

designated)

New World Species

2. A. abietis-religiosae Heil

3. A. americanum Nutt. ex Engelm.

24. A. verticilliflorum Engelm.

Old World Species

25. A. chinense Lecomte

26. A. minutissimum Hook. f.

27. A, oxycedri (DC.) M. Bieb.

28. A. pint Hawks. & Wiens

II. Subgenus Vaginata Hawks. & Wiens

A. Section Vaginata

12. A. gillii Hawks. & Wiens

12a. subsp. gillii

12b. subsp. nigrum Hawks. & Wiens

13. A. globosum Hawks. & Wiens

63
23. A. vaginatum (Willd.) Presl

23a. subsp. vaginatum

23b. subsp. cryptopodum (Engelm.)

Hawks. & Wiens

23c. subsp. durangense Hawks. & Wiens

B. Section Campylopoda Hawks. & Wiens

a. Series Campylopoda

1. A. ahietinum Engelm. ex Munz

la. f. sp. concoloris Hawks. &

Wiens

lb. f. sp. magnificae Hawks. &

Wiens

4. A. apachecum Hawks. & Wiens

6. A. hlumeri A. Neis.

7. A. californicum Hawks. & Wiens

8. A.campylopodum^ngQlxn..

9. A. cyanocarpum Coulter & Nelson

C.

10. A. divaricatum'Engelm.

14. A.guatemalenseHsiwks.&Wiens

16. A. Zanas (Piper) St. John

17. A. microcarpum (EngeJm.) Hawks.

& Wiens

18. A. occidentale Engelm.

22. A. tsugense (Rosendahl) G. N.

Jones

Series Rubra Hawks. & Wiens

5. A. bicarinatum Urban

15. A. hondurense Hawks. & Wiens

20. A. rubrum Hawks. & Wiens

Series Stricta Hawks. & Wiens

21. A. strictum Hawks. & Wiens

Section Minuta Hawks. & Wiens

11. A. douglasii Engelm.

19. A. pusillum Peck

b.

c.

Key to the Subgenera

1. Stems with at least some of the branching, or the staminate flowers, whorled.

1. Stems flabellately branched; staminate flowers never whorled.

Subgenus Arceuthohium

1. subgenus Arceuthobium

2. subgenus Vaginata

Stems with at least some of the branching, or the staminate flowers, occurring in whorls; Old and New

World.

Type species. A. oxycedri (DC.) M. Bieb.

No formal subdivision of this subgenus is made, although a practical division used here is occurrence in the

Old or New Worlds. A natural key to the New World species follows. We do not have sufficient data to construct a comparable natural key for the Old World species but we have prepared an artificial key for them

(see p. 70).

Key to the New World Species

1. Staminate flowers usually in whorls of 6; fruit at maturity about 15 mm. long; staminate inflorescences

deciduous, internodes about as long as wide.

24. A. verticilliflorum

1. Staminate flowers opposite, fruit at maturity about 4 mm. long; staminate inflorescences not deciduous

(but individual flowers deciduous) ; internodes at least 5 times longer than wide.

2. Stems with branching occasionally whorled; fruit opposite; internodes about 5 times longer than wide;

staminate buds lenticular; parasite on Abies.

2. A. abietis-religiósae

2. Stems with branching always whorled; fruit occurring in whorls; internodes about 10 times longer

than wide; staminate buds rounded; parasite on Pinus.

3. A. americanum

Subgenus Vaginata

Hawksworth & Wiens, Brittonia 22: 265, 1970.

Stems flabellately branched; staminate flowers decussate, never in whorls; New World.

Type species: A. vaginatum (Willd.) Presl

64
Key to the Sections

1. Pistillate plants usually more than 4 cm. high; shoots in individual clusters; host generally not forming

systemic brooms.

2. Plants flowering in winter or spring; internodes usually less than 4 times as long as wide.

Section 1. Vaginata

2. Plants flowering in summer and fall; internodes at least 5 times as long as wide.

Section 2. Campylopoda

1. Pistillate plants usually less than 4 cm. high; shoots scattered along the host stem near the apex; host

forming systemic brooms.

Section 3. Minuta

Section 1. Vaginata

Anthesis in winter or spring; tallest shoots usually over 15 cm. high; internodes less than 4 times as long as

wide.

Type species: A. vaginatum

Key to the Species

1. Fruit markedly whitish-blue glaucous on proximal portion; staminate plants often markedly divaricate,

pistillate plants erect; parasite on Pinus section Leiophylla^ occasionally on P. teocote.

12. A. gillii

1. Fruit not markedly glaucous on proximal portion; staminate plants and pistillate plants erect; parasite

on Pinus sections Ponderosae, Australes, and Oocarpae (rare on P. teocote).

2. Plants forming conspicuous, dense, globose clusters several dm. in diameter; stems pale to greenish

yellow; pedicels on mature fruit usually 4-5 mm. long; hosts usually not forming witches' brooms;

anthesis from February to April.

13. A. globosum

2. Plants usually scattered along the host branch, not forming dense, globose clusters; stem blackish or

orange; pedicels ca. 3 mm. long; hosts forming conspicuous witches' brooms; anthesis from April

through May.

23. A. vaginatum

Section 2. Campylopoda

Hawksworth & Wiens, Brittonia 22: 266, 1970.

Anthesis summer or fall; tallest shoots usually less than 10 cm. high; internodes generally 5 times as long

as wide.

Type species: A. campylopodum

Key to the Series

1. Staminate shoots unbranched except at base, some staminate flowers 6-or 7-merous.

Series 1. Stricta

1. Staminate shoots abundantly branched, staminate flowers 3- or 4-merous, rarely 5.

2. Tallest shoots usually over 10 cm. high; shoots dark red or redish brownish; secondary branching rare.

Series 2. Rubra

2. Tallest shoots usually under 10 cm. high; shoots yellow, green, gray or purple; secondary branching

consistent.

Series 3. Campylopoda

Series 1. Stricta

Hawksworth & Wiens, Brittonia 22: 266, 1970.

Staminate shoots unbranched in annual growth segments, usually only branched near base; staminate

flowers 3- to 7-merous.

Type species: A. strictum, the only representative

Series 2. Ruhra

Hawksworth & Wiens, Brittonia 22: 266, 1970.

Plants usually over 15 cm. tall; shoots dark, reddish or brownish; secondary branching rare.

Type species: A. ruhrum

65
Key to the Species

1. Staminate flowers ca. 1.5 mm. in diameter; shoots dark red; fruit shiny, ca. 3 mm. long, central Mexico.

1. Staminate flowers ca. 3 mm. in diameter; shoots red to brownish red; fruits dull, ca. 4-5 mm. long, Hispaniola or Honduras.

2. Shoots brownish red; internodes often twisted in dried specimens; basal diameter of older shoots

2-3 mm.; nodes not swollen on older shoots; mature fruits ca. 4 X 2 mm., reddish; nectary in staminate flower with three similar lobes; parasite on Pinus occidentalis; Hispaniola.

5. A. hicarinatum

2. Shoots olive brown to grayish green; internodes not twisted; basal diameter of older shoots 3-9 mm.;

nodes of older shoots swollen; mature fruits ca. 5.5 X 3 mm., greenish; nectary of staminate flower

with two large and one small lobe; parasite on Pinus oocarpa; Honduras.

15. A. hondurense

Series 3. Campylopoda

Plants usually under 10 cm. tall; plants variously colored but not dark reddish or dark brown; secondary

branching consistent.

Type species: A. campylopodum

Key to the Species

1. Parasites on Pinus.

2. Parasites on Pinus subgenus Haploxylon (soft or white and pinyon pines).

3. Shoots olive green to brownish; parasite on pinyon pines.

10. A. divaricatum

3. Shoots generally yellowish green to light gray or purplish; parasites on white pines.

4. Plants forming systemic witches' brooms; receptacle larger than the base of the mature

fruit; staminate flowers often 2-merous; parasite on Pinus ayacahuite.

14. A. guatemalense

4. Plants rarely forming systemic witches' brooms; receptacle the same diameter as the base

of the mature fruit; staminate flowers rarely 2-merous, usually 3- to 5-merous; never

parasitic on Pinus ayacahuite.

5. Staminate spikes in summer less than 3 mm. long, about twice as long as wide; staminate

plants less than 2 cm. high; parasitic principally on Pinus flexilis, P. aristata, or P.

alhicaulis; California, Great Basin, and Rocky Mountain Stat^tes.

9. A. cyanocarpum

5. Staminate spikes in summer over 5 mm. long, about 5 times as long as widi\; staminate

plants at least 4 cm. high; parasitic principally on Pinus lambertiana or P. strobiformis;

California, southern Oregon, Southwest, and northern Mexico.

6. Anthesis primarily in July; plants ca. 8 cm. high; shoots greenish to bright yellow;

parasitic on Pinus lambertiana; California and southern Oregon.

7. A. californicum

6. Anthesis primarily August and September; plants ca. 4 to 6 cm. high; shoots

generally yellow to light gray; parasitic on Pinus strobiformis; Arizona, New

Mexico, and northern Mexico.

7. Plants ca. 6 cm. high; staminate spikes in summer over 10 mm. long; shoots

not densely clustered; straw to light gray; extreme southern Arizona (Huachuca and Santa Rita Ranges) and in Chihuahua, Durango, and Nuevo

León, Mexico.

6. A. blumeri

7. Plants ca. 4 cm. high; staminate spikes in summer 5-7 mm. long; shoots

densely clustered, yellow; southern Arizona and New Mexico, northern

Coahuila, Mexico.

4. A. apachecum

2. Parasites on Pinus subgenus Diploxylon (hard or yellow pines).

8. Staminate spikes in summer ca. 5 mm. long, less than 3 times as long as wide; anthesis in August

and September; parasite primarily on Pinus ponderosa.

8. A. campylopodum

66
8. Staminate spikes in summer over 10 mm. long, more than 5 times as long as wide; anthesis in

October and November; parasite primarily on Pinus sahiniana.

18. A. occidentale

1. Parasites on Abies, Larix, Tsuga, or Picea.

9. Parasite on Abies; plants usually over 10 cm. high; usually yellowish; staminate buds same color as the

subtending bracts.

I. A. abietinum

9. Parasite on Larix, Tsuga, or Picea; plants usually under 8 cm. high; dark, usually purplish; staminate

buds conspicuously lighter than the subtending bracts.

10. Parasite on Larix; anthesis usually in August; terminal internodes of staminate shoots ca. 2 mm.

long.

16. A. larids

10. Parasite on Tsuga or Picea; anthesis usually in September terminal internodes of staminate

shoots over 3 mm. long.

11. Parasite on Tsuga; staminate spikes in summer over 5 mm. long; staminate flowers sometimes 4-merous; basal internodes of mature plants usually rounded.

22. A. tsugense

11. Parasite on Picea; staminate spikes in summer less than 4 mm. long; staminate flowers

rarely 4-merous; basal portions of older shoots angular.

17. A. microcarpum

Section 3. Minuta

Hawksworth & Wiens, Brittonia 22: 266, 1970.

Anthesis in spring; plants less than 4 cm. high; shoots scattered along the host stem near the apex.

Type species: A. douglasii

Key to the Species

1. Plants usually less than 2 cm. high, without secondary branching; fruit developing in one season (ca. 5

months) ; parasite on Picea; northeastern North America.

19. A. pusillum

1. Plants usually more than 2 cm. high, generally with secondary branching; fruit development requiring two

seasons (ca. 17-18 months) ; parasite principally on Pseudotsuga; Mexico and w^estern North America.

II. A. douglasii

Artificial Key to the New World Species of Arceuthobium

The color photographs will also aid in identification.

1. Mexico, Central America, or Hispaniola.

2. Hispaniola or Honduras.

3. Hispaniola; parasite on Pinus occidentalis.

5. A. bicarinatum

3. Honduras; parasite on Pinus oocarpa.

15. A. hondurense

2. Mexico, Guatemala, or British Honduras.

4. Parasites on Abies or Pseudotsuga.

5. Plants 1-3 cm. high; parasite on Pseudotsuga.

11. A. douglasii

5. Plants 10-15 cm. high; parasite on Abies.

2. A. abietis-religiosae

4. Parasites on Pinus.

6. Baja California.

7. Parasite on pinyons; plants olive green; internodes ca. 1-2 mm. wide.

10. A. divaricatum

7. Parasite on Pinus Jeffrey; or P. Coulteri plants yellow; internodes ca. 2-4 mm. wide.

8. A. campylopodum

67
6. Mainland Mexico (not Baja California) or Central America.

8. Parasites on Pinus subgenus Haploxylon (soft or white pines).

9. Guatemala (possibly in southern Mexico); parasite on Pinus ayacahuite; plants

greenish purple to purple.

14. A. guatemalense

9. Northern Mexico; parasite on Pinus sirohiformis; plants yellow or grayish.

10. Northern Coahuila; plants yellowish; ca. 4 cm. or less high.

4. A. apachecum

10. Chihuahua, Durango, Nuevo León; plants gray, usually over 6 cm. high.

6. A. blunteri

8. Parasites on Pinus subgenus Diploxylon (hard or yellow pines).

11. Parasites principally on Pinus section Leiophylla and occasionally on P. teocote.

12. Staminate plants strongly divaricate; internodes more than 3 times as long

as wide; fruit markedly bluish white glaucous.

12. A. gillii

12. Staminate plants unbranched; internodes less than twice as long as wide;

fruit slightly glaucous, but never markedly bluish white.

21. A. strictum

11. Parasites on pines other than section Leiophylla; staminate plants erect; fruit not

plants erect; fruit not markedly glaucous.

13. Plants dark reddish or blackish.

14. Plants 20 cm. or more high; blackish; basal internodes about 5 mm. wide.

23. A. vaginatum

14. Plants 15 cm. or less high, dark reddish; basal internodes about 2 mm.

wide.

20. A. ruhrum

13. Plants greenish, yellowish, brownish, or orange.

15. Staminate flowers in whorls; fruit generally over 7 mm. long; internodes

about as long as wide; plants under 10 cm. high.

24. A. verticilliflorum

15. Staminate flowers opposite; fruit 6 mm. or less long; internodes at least 3

times as long as wide; plants over 15 cm. high.

16. Plants greenish to pale yellow, generally diffusely and openly

branched; pedicels of mature fruit ca. 4-5 mm. long.

13. A. glohosum

16. Plants orangish, generally densely branched along a principal axis;

pedicels of mature fruit ca. 3 mm. long.

23. A. vaginatum

1. In the United States and Canada.

17. Parasites principally on Pinus.

18. Parasites on Pinus subgenus Haploxylon (white or soft pines or pinyon).

19. Parasite on pinyons.

10. A. divaricatum

19. Parasites on white or soft pines.

20. Parasite on Pinus strohiformis.

21. Plants usually under 4 cm. high; yellow; southern Arizona and southern New

Mexico.

4. A. apachecum

21. Plants usually over 6 cm. high; gray; Huachuca and Santa Rita Mountains,

Arizona.

6. A. hlumeri

20. Parasite on other white pines.

22. Staminate plants usually over 8 cm. high; not densely clustered around the stem;

parasite principally on P. lambertiana.

7. A. californicum

22. Staminate plants usually less than 2 cm. high, densely clustered around the stem;

parasite principally on P. flexilis, P. aristata, or P. albicaulis.

9. A. cyanocarpum

68
18. Parasites of Pinus subgenus Diploxylon (hard or yellow pines).

23. Shoots branching in whorls; parasite principally on Pinus contorta vars. murrayanna and

latifolia or P. hanksiana.

3. A. americanum

23. Shoots branching flabellately; parasites principally on other pines.

24. Arizona, Utah, and eastward.

25. Parasite on Pinus leiophylla var. chihuahuana; fruit markedly bluish white

glaucous.

12. A. gillii

25. Parasite on Pinus ponderosa vars. scopulorum and arizonica and P. engelmannii;

fruits not markedly bluish white glaucous.

23. A. vaginatum

24. Pacific Coast States, British Columbia, and Idaho.

26. Pacific Coast from Oregon northward; parasite on coastal Pinus contorta var.

contorta (generally associated with infected Tsuga, which is usually the principal

•host).

22. A. tsugense

26. California, southern Oregon and northward east of the Cascades; parasite on other

hard or yellow pines.

27. Parasite principally on Pinus ponderosa, P. ponderosa, P. jeffreyi, and P.

attenuata; California to Washington and Idaho ; staminate spikes in summer

ca. 5 mm. long, less than 3 times as long as wide; anthesis in August and

September.

8. A, campylopodum

27. Parasite principally on Pinus sdbiniana, P. radiata, and P. muricata; Califfornia; staminate spikes in summer over 10 mm. long, more than 5 times as

long as wide; anthesis in October and November.

18. A. occidentale

17. Parasites principally on Ahies, Larix, Picea, Pseudotsuga, and Tsuga.

28. Parasites on Tsuga, Larix, or Pseudotsuga.

29. Parasite on Pseudotsuga; plants less than 4 cm. high, scattered along the host stem near the

apex.

11. A. douglasii

29. Parasites on Larix and Tsuga; plants over 5 cm. high, occurring randomly in clusters.

30. Parasite on Larix.

16. A. larids

30. Parasite on Tsuga.

31. Pacific Coast from Alaska to central California, Cascades-Sierra Nevada west

of the crest; basal internodes of mature plants rounded; staminate spikes in

summer over 5 mm. long; terminal internodes of staminate shoots over 3 mm.

long ; lateral staminate flowers frequently 4-merous; anthesis usually in September; parasite on Tsuga heterophylla or T. mertensiana.

22. A. tsugense

31. Idaho and Montana; basal internodes of mature plants angular; staminate spikes

in summer less than 3 mm. long; terminal internodes of staminate shoots less

than 2 mm. long ; lateral staminate flowers rarely 4-merous; anthesis usually in

August; parasite on Tsuga mertensiana (associated with infected Larix which is

generally the principal host).

16. A. larids

28. Parasites on Ahies or Picea.

32. Parasites on A6zes.

33. Northern Arizona, southern Utah, Nevada, California, southern Oregon northward to

southern Washington east of the crest of the Cascades; plants usually over 10 cm. high,

yellowish; staminate buds same color as the subtending bracts.

I. A. ahietinum

69
33. Pacific Coast from Oregon northward to Alaska and westward to the crest of the Cascades;

plants usually less than 6 cm. high, green to purplish; staminate buds conspicuously

lighter than the subtending purplish bracts (associated with infected Tsuga which is

the principal host).

22. A. tsugense

32. Parasites on Pzcea.

34. Lake States and Saskatchewan eastward to the Atlantic; plants less than 2 cm. high,

unbranched; parasites on Picea glauca and P. mariana.

19. A. pusillum

34. Arizona and New Mexico; plants usually over 5 cm. high, secondarily branched;

.parasites on Picea engelmannii and P. pungens.

17. A. microcarpum

Artificial Key to the Old World Species of Arceuthobium

1. Parasite on Juniperus; north Africa and southern Europe from the Atlantic eastward to the Himalayas and

southward to Kenya.

27. A. oxycedri

1. Parasites on Abies, Keteleeria, or Pinus; Himalayan region and eastward to Szechwan, and Yunnan.

2. Plants usually less than 1 cm. high; host forming systemic brooms with numerous mistletoe shoots

concentrated near the apex of the host branches; parasite on Pinus griffithii; western Himalayan

region.

26. A. minutissimum

2. Plants usually over 5 cm. high; host probably not consistently forming systemic brooms, mistletoes not

concentrated near the apex of the host branches; parasites on Pinus tahulaeformis, Abies, or Keteleeria;

eastern Himalayas to southwestern China.

3. Parasite on Pinus tabulaeformis; plants ca. 10-20 cm. high; internodes ca. 10 mm. long; anthesis in

May and June.

28. A.pini

3. Parasite on Abies and Keteleeria, plants ca. 5 cm. high; internodes ca. 5 mm. long; anthesis in

August-September.

25. A. chinense

70
1. Arceuthobium abietinum í. sp. concoloris on Abies concolor. Staminate plant. California.

2. Arceuthobium abietinum f. sp. magnificae on Abies magnifica. Pistillate plant. California.

71
iÄ'.

3. Arceuthobium abietis-religiosae on ^bies religiosa.

Dried herbarium specimen. Staminate plant (center and upper left) and pistillate plant (upper

right). Note vertieillate branching. Mexico, D.F.

4. Arceuthobium americanum on Finns contorta. Staminate plant. Colorado.

72
5. Arceuthobium atnericanum on Pinus contorta. Pistillate plant. Colorado.

6. Arceuthobium apachecum on Pinus strobiformis. Staminate plant. Arizona.

73
7. Arceuthobiùtn apachecum on Pinus strobiformis.

PistUlate plant. Arizona.

8. Arceuthobium bicarinatum on Pinus occidentalis.

Staminate plant. Dominican Ret)ublic.

Arceuthobium bicarinatum on Pinus occidentalis.

Pistillate plant. Dominican Republic.

74

10. Arceuthobium blumeri on Pinus strobiformis.

Staminate plant (lower) and pistillate plant

(upper). Arizona.
11. Arceuthobium californicum on Pinus lambertiana. Staminate plant (above) and pistillate plant (below).

California.

12. Arceuthobium campylopodum on Pinus ponderosa. Staminate plant (left) and pistillate plant (right).

California.

75
13. Arceuthobiutn cyanocarpum on Pinusflexius. I'lstillate plant. Colorado.

llllllll!

76

14. Arceuthobium. cyanocarpum on Pinus flexilis.

Pistillate plant, note nectar droplets on flowers.

Colorado.
15. Arceuthobium divaricatum on Pinus edulis. Staminate plant. New Mexico.

16. Arceuthobium divaricatum on Pinus edulis. Pistillate plant. New Mexico.

77
17. Arceuthobium douglasii on Pseudotsuga menziesii.

Staminate plant. California.

19. Arceuthobium gillii subsp. gillii on Pinus leiophylla

var. chihuahuana. Staminate plant. Arizona.

18. Arceuthobium douglasii on Pseudotsuga menziesii. Pistillate plant. Idaho.

78
20. Arceuthobium gillii subsp. gillii on Pinus leiophylla var. chihuahuana. Pistillate plant. Chihuahua, Mexico.

21. Arceuthobium gillii subsp. nigrum on Pinus leiophylla. Staminate plant. Durango, Mexico.

22. Arceuthobium gillii subsp. nigrum. on Pinus leiophylla. Pistillate plant. Durango, Mexico.

79
23. Arceulhobium globosum (right) and A. vaginatum subsp. vaginatum (left) on the same branch of Pinus

cooperi. Durango, Mexico.

24. Arceuthobium guatemalense on Pinus ayacahuite. Systemic infections of pistillate plants (above) and staminate plants (below). Huehuetenango, Guatemala.

80
1

27. Arceuthobiutn laricis on Larix occidentalis. Staminate plant. British Columbia. Photograph by

courtesy of R. B. Smith.

25. Arceuthobiutn hondurense on Pinus oocarpa. Staniinate plant. Francisco Morazan, Honduras.

26. Arceuthobiutn hondurense on Pinus oocarpa. Terminal portion of pistillate plant. Francisco Morazan, Honduras.

28. Arceuthobiutn laricis on Larix occidentalis. Pistillate plant. British Columbia. Photograph by

courtesy of R. B. Smith,

81
29. Arceuthobium microcarpum on Picea pungens.

Staminate plant. Arizona.

30. Arceuthobium microcarpum on Picea pungens.

Pistillate plant. Arizona.

31. Arceuthobium occidentale on Pinus sabiniana. 32. Arceuthobium occidentale on I'iiius sabiniana.

Staminate plant. California. Pistillate plant. California.

82
333. Arceuthobium pusiUum on i'lcea glauca. Staminate plant. Manitoba. Photograph courtesy of J. G. Laut.

34. Arceuthobium rubrum on Pinus teocote. Staminate plant. Durango, Mexico.

83
35. Arceüthobium rubrum on Pinus teocote. Pistillate plant. Durango, Mexico.

36. Arceüthobium strictutn on Pinus teocote. Stami- 37. Arceüthobium strictum on Pinus leiophylla var.

nate shoots. Durango, Mexico. chihuahuana. Pistillate plant. Durango, Mexico.

84
/ / ^'

38. Arceuthobium tsugense on Tsuga heterophylla.

Staminate plant. British Columbia. Photograph

by courtesy of R. B. Smith.

39. Arceuthobium tsugense on Tsuga heterophylla.

Pistillate plant. British Columbia. Photograph

by courtesy of R. B. Smith.

85
40. Arceuthobiuni vagiiialuin subi^p. cryplopodum on Pinus ponderosa var.

scopulorum. Staminate plant. Arizona.

41. Arceuthobium vaginatum subsp. cryptopodutn on Pinus ponderosa var. scopulorum. Pistillate plant. Colorado.

86
44. Arceuthobium verticilliflorum on Pinus engelmannii. Staminate shoots in flower. Durango,

Mexico.

43. Arceuthobium vaginatum subsp. durangense on

Pinus durangensis. Pistillate plant. Durango,

Mexico.

87
45. Arceuthobium verticilliflorum in Pinus engelmannii. Pistillate plant with nearly mature fruits.

Durango, Mexico.

46. Arceuthobium minutissimum on Pinus griffithii.

Staminate shfH>ts. Kashmir.

47. Arceuthobium minutissimum on Pinus griffithii.

Pistillate shoot. Kashmir.

88
,,—-f:

•'* ^

..v>J-f//

.v v\

48. Arceuthobium oxycedri on Juniperus oxycedrus. Staminate plants. France. Photograph by courtesy of

J. Gremmen.

89
New World Taxa

ARCEUTHOBIUM ABIETINUM

FIR DWARF MISTLETOE

1. Arceuthobium ahietinum Engelm. ex Munz, Man.

South. Calif. Bot.: 114, 1935*. LECTOTYPE:

CALIFORNIA: Sierra Co.: Sierra Valley, on

Abies concolor, Lemmoîi in 1875 (MO^ Isotype

GH). Arceuthobium abietinum Engelmann in Gray,

Proc. Amer. Acad. Arts & Sei. 8: 401, 1872, 7iomen

7iudum. Arceuthobium douglasii Engelm. var.

abietinum Engelm., in Watson, Bot. Calif. 2: 106,

1880. Arceuthobium occidentale Engelm. var. abietinum Engelm. in Watson, Bot. Calif. 2: 107, 1880.

Razoumofskya douglasii (Engelm.) Kuntze var.

abietina (Engelm.) Howell, Fl. N. W. Amer. 1:

609, 1902. Razoumofskya douglasii (Engelm.)

Kuntze var. abietina (Engelm.) Piper, Contr. U.S.

Nat. Herb. 11: 223, 1906. Razoumofskya abietina

(Engelm.) Tubeuf forma párvula Tubeuf. Naturwiss. Z. Forst-u. Landwirt. 17: 219, 1919, nomen

nudum. Razoumofskya abietina (Engelm.) Tubeuf

forma magna Tubeuf, loc. cit. : 220, nomen nudum,

Razoumofskya abietina (Engelm.) Abrams. Illus.

Fl. Pac. States 1: 530, 1923. Arceuthobium campylopodum Engelm. forma abietinum (Engelm.) Gill,

Trans. Conn. Acad. Arts & Sei. 32: 195, 1935.

DESCRIPTION: Mean shoot height ca. S cm.

but up to 22 cm. Shoots yellow green to yellow,

flabellately branched. Basal diameter of dominant

shoots 1.5-6 mm. (mean 2 mm.). Third internode

4-23 mm. (mean 14) long and 1.5-4 mm. (mean 2

mm.) wide; length/width ratio ca. 7-9:1. Staminate flowers ca. 2.5 mm. in diameter; perianth

3-merous, sometimes 4, (not 7 as reported by

Munz and Keck, 1959) apex acute; same color as

shoots; segments ca. 1.2 mm. long and 1.0 mm.

wide. Mean anther diameter 0.4 mm., centered

0.7 mm. from tip of segment. Pollen polar diameter

18-23 (mean 20^); equatorial diameter 19-25

mean 22^) ; polar/equatorial diameter ratio 1:1.07;

spine height ca. 1.5-2 times greater than wall

thickness (7 collections). Mature fruit ca. 4 mm.

long and 2 mm. wide; proximal portion ca. 2.5 mm.

long. Seeds 2.8 X 1.2 mm. Meiosis in July.

Anthesis usually in August-September. Seeds

mature in September or October of the year

following pollination; maturation period averages

13-14 months, n = 14.

PRINCIPAL HOSTS: A6z6s spp.

* Dr. C. V. Morton of the U.S. National Museum has

kindly pointed out that Munz raised Engelmann's varietal

name to specific rank in 1935. Thus our pubhshed ''new"

combination of this transfer is superfluous (Brittonia 22: 268,

1970).

DISCUSSION: Parmeter and Scharpf (1963) first

reported that, on the basis of field evidence and

inoculation studies, the dwarf mistletoe on

Abies concolor will not infect A. magnifica. Conversely, the parasite of A. magnifica does not

parasitize A. concolor. Our observations in several

mixed fir stands in the Sierra Nevada confirm

Parmeter and Scharpf^s conclusions. However,

we have been unable to find morphological differences between these two mistletoes. Also, the two

are similar phenologically although, in the same

locality, seed- dispersal begins about 1 week

earlier for thé red fir parasite (Scharpf and Parmeter 1967). Greenham and Leonard (1965)

studied the amino acids of the two dwarf mistletoes, and their respective hosts, but concluded

that amino acid composition probably could not

explain the specificity of the two mistletoes'.

Since the host affinities of these two mistletoes

are distinct, and because this is of considerable

practical importance in forestry, we have designated them SiS formae spéciales in accordance with

recommendation 24B of the International Code of

Botanical Nomenclature.

Key to the formae spéciales

1. Parasitic principally on Abies concolor or A.

grandis. Southern Washington to southern

California and northern Arizona. '

, la. Arceuthobium abietinum f. sp. condoloris

1. Parasitic principally on Abies magnifica or A.

procera. Southern Oregon to central California.

' lb. Arceuthobium abietinum f. sp.,, magnificae

ARCEUTHOBIUM ABIETINUM F. SP.

CONCOLORIS,

WHITE FIR DWARF MISTLETOE

la. Arceuthobium abietinum Engelm. ex Munz f. sp.

concoloris Hawksworth & Wiens. Brittonia 22:

267, 1970.

DESCRIPTION: Mean shoot height ca. 10 cm.,

but up to 22 cm. (fig. 26). Basal diameter of

dominant shoots 1.5-6 mm. (mean 2 mm.).

Third internode 4-23 mm. (mean 13.3 ± 3.8 mm.)

long and 1.5-4 mm. (mean 2.0 mm.) wide (15

collections), length/width ratio 6.7:1. Pollen

polar diameter 18-23 (mean 20^); equatorial

diameter 19-25 (mean 22ju); polar/equatorial

diameter ratio 1:1.07; spine height ca. 1.5^

slightly greater than wall thickness—ca. l.l^u

(5 collections). Peak anthesis usually during first

half of August, with extremes from early July to

late September. Seeds usually mature in September with extremes from late August to late

October; maturation period averages 13-14

months. Seeds germinate from February through

June (Scharpf and Parmeter 1967). n = 14.

90
Figure 26.—Arceuthohium ahietinum f. sp. concoloris in summer: A^ Staminate plant; ß, Pistillate plant; C, Detail

of nearly mature fruit; D, Detail of staminate flower just prior to anthesis.

91
HOSTS : The principal liosts of this dwarf mistletoe

are Abies concolor and A. grandis. The rare

Brewer spruce, Picea breweriana, was found by

Hawksworth, Wiens and Graham (1967) to be

quite susceptible to this parasite on Flat Top

^lountain, west of Grants Pass, Oregon (see fig. 9).

On the Xorth Rim of Grand Cayon, Arizona,

Abies lasiocarpa is parasitized where this tree

grows in association with infected A. concolor.

Little is known of its occurrence on A. amabilis;

only one collection is available on this host from

the vicinity of Crater Lake, Oregon. Pinus

lambertiana, P. monticola, and P. contoria are

very rare hosts for the white fir dwarf mistletoe.

DISTRIBUTION: Arceuthobiuni abietinum f. sp.

concoloris is widely distributed from southern

Washington (Klickitat County) south through

the Cascades and Sierra Xevada to the San

Bernardino Mountains of California (fig. 27). It

also occurs along the coast ranges from Alendocino

County, California, to Curry County, Oregon. It

is known in three isolated areas: the Charleston

Mountains, Nevada; Grand Canyon National

Park, Arizona; and northwestern Kane County,

Utah.

This taxon occurs on Abies concolor throughout

most of its range in California, Nevada, Arizona,

and southern Oregon. Abies grandis is the principal host in central Oregon, southern Washington,

and in the coastal mountains of Oregon and

California. In the Siskiyou INIountains of Del

Norte County, Cahfornia, just south of the

Oregon boundary, we found the parasite commonly

infecting both Abies concolor and A. grandis in a

in a mixed stand.

Several localities for A. abietinum recorded by

Gill (1935) have since been found to be based on

occasional parasitism of Abies by other species of

Arceuthobiuni: Arizona, San Francisco Mountains,

(A. douglasii on Abies sp.) ; Washington, Longmire

and EUensburg, (A. tsugense on Abies amabilis);

Arizona, Graham Mountains, and New ^léxico,

JMogoUon, (A. douglasii on Abies lasiocarpa var.

arizonica); Utah, Spring Lake and Charles Peak,

{A. douglasii on Abies concolor); and Oregon,

McKenzie Pass, (A. tsugense on Abies lasiocarpa).

The parasite occurs from near sea level along the

California and Oregon coasts to over 8,500 feet in

the Charleston Mountains of southern Nevada.

Specimens examined: see appendix, p. 186.

DISCUSSION: In the Northwest, two other species

of Arceuthobium occur on Abies: (1) A. tsugense on

Abies amabilis, A. lasiocarpa, and A. grandis, and

(2) A. laricis on Abies lasiocarpa and A. grandis.

However, insofar as we know, neither is sympatric

with A. abietinum. For these other taxa, general

Figure 27.—Distribution of .arceuthobiuni abietinum

f. sp. concoloris (based on collections rnly). Outline

map copyrighted by Denoyer-Geppert Co., Chicago

(used by permission).

infection of pure Abies stands does not occur, but

occasional trees are infected in stands where the

principal host is parasitized (for example Larix

for Arceuthobium laricis, and Tsuga for A.

tsugense). Arceuthobium laricis is readily distinguished from A. abietinum by its smaller,

darker shoots, and small staminate spikes in

summer (2-3 mm. long) vs. 5-7 mm. long in A.

abietinum. Arceuthobium tsugense differs from A.

abietinum by its smaller, green to purple shoots

compared with the larger, yellowish shoots in A.

abietinum.

Branch dying (flagging) is one of the most

conspicuous field symptoms of A. concolor infected by dwarf mistletoe, particularly in California. Scharpf (1969c) has shown that flagging of

dwarf mistletoe infected branches is often associated with the fungus Cytospora abietis.

92
ARCEUTHOBIUM ABIETINUM F.

MAGNIFICAE,

RED FIR DWARF MISTLETOE

SP.

lb. Arceuthohium abietinum Engelm. ex Munz f.

sp. magnificae Hawksworth & Wiens, Brittonia

22: 268, 1970.

DESCRIPTION: Alean shoot height ca. 6 cm., but

up to 12 cm. Basal diameter of dominant shoots

1.5-3 mm. (mean 2 mm.). Third internode 10-22

mm. (mean 15.0 ±3.1 mm.) long and 1.5-2 mm.

(mean 1.7 mm.) wide (5 collections), length/width

ratio 8.8:1. Pollen polar diameter 19-23 (mean

21ii); equatorial diameter 19-24 (mean 22jLt);

polar/equatorial diameter ratio 1:1.07; spine

height ca. 2ß approximately twice wall thickness—

ca. 1/i (2 collections). Peak anthesis usually in late

August, with extremes from late July to late

September. Seeds mature in September or October;

maturation period averages 13-14 months, n = 14.

HOSTS AND DISTRIBUTION: This mistletoe is

known on Abies magnifica (vars. magnifica and

shastensis) from Oregon Caves National Monument in southern Oregon south to Sequoia National Park in California (fig. 28). It has been reported on Abies procera from three localities in

southern Oregon, but its distribution and hosts in

that State are poorly known, partly because of the

difficulty of distinguishing Abies magnifica from

A. procera in this area (Powells 1965). It may be

significant that the areas where dwarf mistletoe

has been reported on A. procera are in the zone

where the taxonomic status of the host is questionable. The parasite has not been found in the

"typical" A. procera stands north of Crater Lake,

Oregon. This mistletoe may occur in Nevada in

the vicinity of Lake Tahoe. Known elevational

range: 5,000 to 7,800 feet. Specimens examined:

see appendix p. 188.

DISCUSSION: As noted under the discussion of

A. abietinum, f. sp. magnificae is maintained hero

because, although it is morphologically similar to

f. sp. concoloris, it will not infect Abies concolor.

This mistletoe is a very common and serious disease agent of the A. magnifica forests of the Sierra

Nevada. In a series of sample plots in California,

this dwarf mistletoe was present on 46 percent of

103 A. magnifica plots and on 36 percent of the

trees (Calif. Forest Pest Control Action Council

1968). Scharpf (1969b) discusses infection in

young red fir stands, and gives recommendations

for control of the dwarf mistletoe.

Branch flagging is one of the most conspicuous

field symptoms of A. magnifica infected by dwarf

mistletoe. Scharpf (1969c) has shown that flagging

of dwarf mistletoe infected branches is often associated with the fungus Cytospora abietis.

Figure 28.—Distribution of Arceuthobiuin abietinum

f. sp. magnificae (based on collections only). Outline

map copyrighted by Denoyer-Geppert Co., Chicago,

(used by permission).

ARCEUTHOBIUM ABIETIS-RELIGIOSAE

2. Arceuthobium abietis-religiosae Heil, Centralbl. f.

Bakt. Abt. 2: 28, 1923. NEOTYPE: :MEXIC0:

^Mexico: Between Amecameca and Paso do Cortez,

near km. 78, below Popocatepetl and Ixtaccihuatl

Nat. Park, on Abies religiosa, Hawksworth &

Wiens 3339 in 1963 (COLO, Isotypcs F, FPF,

INIF, IMEXU, AIO, US). (See Hawksworth and

Wiens, Brittonia 17: 231, 1965).

DESCRIPTION: -Mean shoot height ca. 10 cm., but

up to 16 cm. (fig. 29). Shoots olive green, older

shoots typically with black variegations. Occasionally verticillately branched. Basal diameter of

dominant shoots 2-10 mm. (mean 4 mm.). Third

internode 8-24 mm. (mean 15.4 ± 5.3 mm.) long

and 1-4 mm. (mean 2.8 mm.) wide (3 collections),

length/width ratio 5.5:1. Staminate buds 2-4 per

node. Staminate flowers 2 mm. long and 2.4 mm.

in diameter; perianth segments mostly 3, but often

also 4, apex obtuse-acute; same color as shoots on

93
Figure 29.—Arceuthobium abietis-religiosae in March: A^ Pistillate plant; ß, Staminate plant; C, Detail of fruit;

D, Detail of staminate flower.

94
outer surface, reddish on inner surface distal to

anther; segments ca. 1.2 mm. long and 0.9 mm.

wide. Mean anther diameter 0.4 mm., centered 0.8

mm. from tip of segment. Pollen polar diameter

17-23 (mean 20M); equatorial diameter 22-26

(mean 24JLI); polar/equatorial diameter ratio

1:1.22; spine height approximately equal to wall

thickness—1.5M (2 collections). Pistillate flowers

ca. 1.0 mm. long and 0.5 mm. in diameter. Mature

fruit 3.5 mm. long and 2 mm. wide; proximal portion ca. 2.5 mm. long. Seeds 2.2 X 1.0 mm. Meiosis

in September. Peak anthesis in jNlarch and April

but also observed in September. Seeds probably

mature in October or November; maturation period apparently averages ca. 19 months, n = 14.

HOSTS: Known only on Ahies spp., including A.

religiosa and A. vejarii. It does not parasitize pines

even where these trees are associated with infected

Ahies. Probably occurs on Ahies guatemalensis at

Copainalá, Chiapis, Mexico judging by the photograph of a tree with apparent witches' brooms

published by Martinez (1963, p. 124).

DISTRIBUTION: This dwarf mistletoe is known

only in IMexico from the Distrito Federal and the

States of Mexico, Nuevo León, Jalisco, and Hidalgo, but can be expected in adjacent areas (fig.

30). Madrigal Sánchez (1967) reports that in the

mountains around the Valle de Mexico A. ahietisreligiosae generally occurs wherever Ahies religiosa

is found, and hsts these localities: D. F.: Sierra de

las Cruces; MEXICO: Sierra Nevada, Sierra de

Calpulalpan, Sierras de Monte Alto and Monte

Bajo; and HIDALGO: Sierra de Pachuca. The

known elevational range of this dwarf mistletoe is

from 8,200 to 10,600 feet. Specimens examined:

see appendix, p. 189.

DISCUSSION: The nomenclature of this distinctive Mexican dwarf mistletoe is discussed by

Hawksworth and Wiens (1965). The most distinguishing characteristics of the species are its

occasionally verticillately branched shoots (a feature shared only with A. americanum), and its exclusive occurrence on Ahies.

Additional items of interest in A. ahietis-religi-

^~- 'J /' 1 i 1 1 <r À-—-^- \ \

t^l^ ^__----T-- \ ^N

\ )

+ ,' -H -1-

/

I Í ^^-'-^^-^-^ ~"^^--. \

^ N/

Y ,.'\^ U N ( 1 T E D S ' T-"-" A \ T s /

V ( \

\ V ''\. \ \ \

\ (^^ "^ v...^ [_.

New Mexico

\ )Ivlissisji^P.',

31

Florida ^' Y -^ \

Tixos "

+ 1 N

\

Chihuahua;^g^^,lg

+

^

1 Louisionok^

^

J\A^

^ f\ *^^ \ S '^~' *

( \ jF

T\^

\ \

\ ^H'HN k ^L \ f^ j

^ \ ' V

>$Sinaloa| I -~^- f-\ / '^\S \\ 0 " ' \

F

^^\^——^

"~~——r~~~~~SL 1 V ^N. iDurango J \ Nuevo] "y !____-—— \

^'V- \t

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\

+

-f

^-^ Zacatéeos/ \ *

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T

1 c

^

\

^ \

\ -/V''' '-^

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1 "''W\

J____ o

<

VIoyant

•

1 San Luis'^>^-—1

-V„ Potos, ^.y^^V

/ \^-^ ^ ^ (Cruz >

/> Guonaiuofo^-p ^rCj"'^ ^'i {

\ [Hidalgo v;-\-^ \

ton y

V <6 ¿ .'-\ 1 7

Vo'umtona

\ /IYV]

'-^-'^^\)^,^^ ^\ ,-'Puebla ^-^

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A.abietis-

• collection

- rellgiosae ^«>/Michoacan

O C

V J Ctiiopor"^^-<7>^

Í 1

^ locality cited by Madrigal (1967) C ^^ Guerrero 1

'"^''^-X^baxaca

/ y-'^^'uty\^^ HONDURAS ^—

1 - Distrito Fede

£• ^ / i^x^"^

1,1,1 300 .00 500 «ILES

3 -Tloxcolo

4 -Queretoro

A/ ______J\^

r ' I ' 1 ' 1 ~ ., .'±

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1 ' 1 ' 1 ' i ' 1

400 600 800

1

K.OME

—1-

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6 -Colimo

es -r—^' , h

Figure 30.—Distribution of Arceuthobium abietis-religiosae.

95
osae include the occurrence of systemic witches^

brooms and verticillate staminate floral buds.

Systemic infections are not common in this species;

on those few that we found, the dwarf mistletoe

shoots had formed only at the girdles of the host

branches.

The only other case in which verticillately arranged staminate flowers occur is in ^. verticilliflorum, where the character is consistent and conspicuously developed. The discovery of this character in A. ahietis-religiosae strengthens the presumed relationship between these two species.

Most dwarf mistletoes show definite and predictable phenological periodicity; however, several

phenological characters such as flowering period

and period of seed dispersal appear to be irregular

in A. ahietis-religiosae. Our observations are based

on staminate flowers because of their larger size,

even though the staminate and pistillate flowers

appear to be synchronous.

In early April, the specimens we collected appeared to be in the peak flowering period, and the

fruits in the initial stages of development. In

September, when we made additional collections,

we expected to find seed dispersal and floral buds

in early developmental stages. Instead, we found

most of the floral buds were already formed, and

many staminate flowers already open. Only additional field observations at periodic intervals can

help clarify the habits of this species, but we suggest two possibilities. Perhaps, A. ahietis-religiosae

has two flowering periods with peaks in late September and early March, or it may be an indirect

flowering species similar to A. douglasii and A.

americanum which complete floral development in

late summer but do not flower until spring. If this

is true, unusual environmental conditions could

have induced the fall flowering of A. ahietisreligiosae, as sometimes occurs in A. douglasii and

A. americanum.

Heil (1923) stated that the fruits of A. ahietisreligiosae mature in September. The fruits we observed in mid-September 1969, near Mexico City,

however, were at least a month from maturity,

and were maturing at the higher elevations first—a

character common in most Arceuthohium species.

ARCEUTHOBIUM AMERICANUM

LODGEPOLE PINE DWARF MISTLETOE

3. Arceuthohium americanum Nuttall ex Engelmann

in Gray, Boston J. Natur. Hist. 6: 214, 1850.

TYPE: OREGON, Blue Mountains, on Pinus

contorta, Douglas in 1826 (MO!). Razoumofskya

americana (Nutt. ex Engelm.) Kuntze, Rev. Gen.

PL 2:587, 1891.

DESCRIPTION: Mean shoot height ca. 6 cm., but

up to 30 cm. (fig. 31). Shoots yellowish to olive

green, verticillately branched. Basal diameter of

dominant shoots 1-3 mm. (mean 1.5 mm.). Third

internode 6-23 mm. (mean 12.1 =t 3.0 mm.) long

and 1-2 mm. (mean 1.2 mm.) wide (20 collections),

length/width ratio 10.1:1. Staminate ñowers

borne on pedicel-like joints. Staminate flowers ca.

2 mm. long and 2.2 mm. in diameter; perianth

mostly 3-merous, sometimes 4, same color as

shoots; segments ca. 1.1 mm. long and 1.0 mm.

wide. Mean anther diameter 0.55 mm., centered 0.7 mm. from tip of segment. Pollen polar

diameter 19-28 (mean 21/x); equatorial diameter

23-30 (mean 25M) ; polar/equatorial diameter ratio

1:1.16; spine height (mean 1.8íLI) equal to or

slightly greater than wall thickness—mean 1.5/i

(8 collections). Pistillate flowers verticillate; ca.

1.5 mm. long and 1.0 mm. in diameter; 2-merous.

Mature fruit 3.5-4.0 mm. long (mean 4 mm.) and

1.5-2.5 mm. wide (mean 2 mm.); proximal portion

ca. 2.5 mm. long. Seeds 2.4 X 1.1 mm. Meiosis in

August. Anthesis usually in April and May, with

extremes from March to June. Seeds mature in

late August or September of the year following

pollination; maturation period averages about 16

months; germination begins in May in Colorado.

n = 14.

HOSTS: Principal hosts are Pinus contorta (subsp.

latifolia and murrayana) and P. hanksiana (see

table 5). Pinus ponderosa var. scopulorum is frequently parasitized in Colorado, Utah, and Wyoming primarily where this tree is associated with

infected P. contorta, but sometimes in pure stands.

Pinus ponderosa var. ponderosa is an occasional

host (Kuijt 1953) but it seems to be less susceptible

than var. scopulorum (Hawksworth 1968). Other

occasional hosts for A. americanum are Pinus

aristata, P. alhicaulis, and P. flexilis. Rare hosts

are the spruces Picea glauca, P. engelmannii, and

P. pungens (Hawksworth and Graham 1963;

Kuijt 1960b; Molnar, Harris, Ross, and Ginns

1968). Pseudotsuga menziesii is an extremely rare

host, and in the single case known (Shingle Creek,

east of Kamas, Utah), small witches' brooms

without mistletoe shoots were formed. Dissection

of twigs on brooms revealed the endophytic system

of the parasite.

Weir (1917) reports A. americanum on Pinus

attenuata at Oregon Mountain, Josephine County,

Oregon. His specimens are unquestionably A.

americanum, but when we visited the same area in

1966, we were unable to find this parasite in the

Pinus attenuata or P. contorta stands.

Weir (1918a) successfully inoculated seedlings

of the European Pinus montana with A. americanum. This mistletoe has been found on planted

96
Figure 31.—Arceuthohium americanum in spring: A, Staminate plant; ß, Pistillate plant; C, Staminate flower;

D, Pistillate flower.

97
Pinus sylvestris in Washington (Graham and

Leaphart 1961) and in Alberta (Powell 1968).

DISTRIBUTION: Arceuthohium americanum has

the largest range of any North American dwarf

mistletoe (figs. 32, 33, 34). Its latitudinal range,

which is exceeded only by A, douglasii, is from

Lake Athabasca in northern Saskatchewan south

to the southern Sierra Nevada in California and to

central Colorado. Its longitudinal distribution of

nearly 1,500 miles extends from western Ontario

to western British Columbia (Elliott, Laut, and

Brandt 1967; Kuijt 1963; Larsen and Gross 1970;

Zalasky 1956). Arceuthohium americanum occurs

to nearly 60° N. latitude; thus it is one of the

northernmost mistletoes in the world—possible

rivals are A. tsugense in Alaska and Viscum album

in Sweden (Wallden 1961). The general range of

A. americanum is centered within that of its principal host Pinus contorta, particularly the subspecies

latifolia and murrayana. An exception to this is in

southern California and Baja California where

subsp. murrayana extends about 300 miles south

of the known range of the parasite.

Arceuthohium americanum does not occur within

the range of shore pine (P. contorta subsp. contorta) except possibly in the Oregon Mountains

near the Oregon-California border. Weir (1917)

reported the parasite here on Pinus contorta, P.

jeffreyi, and P. attenuata, but when we examined

this area in 1966 we did not find A. americanum.

All Pinus contorta we found was subspecies contorta, so Weir's report must refer to this subspecies. Verification of A. americanum here is

needed because this area is approximately 60 miles

from the main range of the species.

The parasite occurs in outhers of Pinus contorta subsp. latifolia in the Cypress Hills of southeastern Alberta, in the Little Rocky Mountains

near Malta, Montana, and possibly in the Whitlash Mountains northeast of Shelby, Montana.

In central and northern Alberta, A. americanum

occurs on Pinus hanksiana, and is common on this

host in Saskatchewan and Manitoba. The reports

of A. americanum at White Otter Lake, in western

Ontario (Hord and Quirke 1956) and near Sprague

in extreme southeastern Manitoba (Zalasky 1956)

apparently were based on a rare infection of Pinus

hanksiana by A. pusillum (Laut 1967; Sippell,

Rose, and Larsen 1968). Recently, A. americanum

has been confirmed in Ontario (Larsen and Gross

1970). Its discovery in the Sioux Lookout area, is

nearly 150 miles east of the previous known eastern

limit at the southern end of Lake Winnipeg,

Manitoba.

Arceuthohium americanum only occurs in the

western part of the range of P. hanksiana so spread

of the parasite onto this tree has probably been

relatively recent. It seems likely that A. americanum evolved as a principal parasite of Pinus

contorla, then spread to P. hanksiana through central Alberta where these two trees occur together

and frequently hybridize. Yeatman (1967) suggests that P. hanksiana first became parasitized

by A. americanum after the Wisconsin glaciation,

but available evidence indicates an earlier association (Zavarin, Critchfield, and Snajberk 1969).

Arceuthohium americanum has been found on the

Pinus contorta X hanksiana hybrid near Grand

Prairie and Peace River, Alberta.^^ Also, A. americanum occurs on planted P. contorta in an infected

P. hanksiana stand at Prince Albert, Saskatchewan.^^ Arceuthohium americanum has an elevational range from about 700 feet near Lake Winnipeg in Manitoba to 11,000 feet in central Colorado. In the Rocky Mountains, the parasite occurs

at the lower limits of P. contoria, but its upper

limits are usually about 500 feet below the upper

commercial limits of the host (Hawksworth

1956b).

Arceuthohium americanum distribution maps

have been published by Kuijt (1963) for western

Canada, Baranyay (1970) for Alberta, and by

Zalasky (1956) for Alanitoba and Saskatchewan.

Specimens examined: see appendix, p. 189.

DISCUSSION: Gill (1935) discusses the typification

problem of A. americanum. Engelmann thought

that the specimen he was describing was collected

by Nuttall, and it was designated as the type

(Oregon, on Pinus). Gill states, however, that this

specimen from Nuttall's collections was actually

made by Douglas (Blue Mountains, Oregon on

Pinus contorta) and thus we have designated it as

the type specimen.

Despite the large geographic range of this parasite, we find no features that suggest ecotypic or

subspecific division. In general, the plants are

larger in the Cascades of Oregon and in northern

Idaho than elsewhere, but this is probably associated with the more vigorous host growth in these

areas. A. americanum, which induces the characteristic systemic brooms on Pinus contorta (Kuijt

1960b), causes the same type of broom on P.

ponderosa (Hawksworth 1956a, Weir 1916c), but

on Picea engelmannii, the brooms are nonsystemic

(see fig. 9).

Kuijt (1960a) noted that A. americanum does

not perpetuate itself for long periods of time on

Pinus jeffreyi or P. ponderosa var. ponderosa in

California. In northern Colorado and southern

Wyoming, however, the parasite is aggressive in

^^ Personal communication, J. A. Baranyay, Canada Dep.

Fisheries and Forestry, 1967.

20 Personal communication, J. G. Laut, Canada Dep. Fisheries and Forestry, 1967.

98
A.amerfcanum

o 200 400 600 800 1000

Scale of Miles

Figure 32.—General distribution of Arceuthohium americanum. Outline map copyrighted by DenoyerGeppert Co., Chicago (used by permission).

99
Figure 33.—Distribution of Arceuthobium americanum in the United States and the range of Pinus contorta.

Outline map copyrighted by Denoyer-Geppert Co., Chicago (used by permission).

100
A.americanum

Figure 34.—Distribution of Arceuthohium americanum on all hosts in Canada, collections and reports combined.

several pure P. ponderosa var. scopulorum stands,

some of which are several miles from the closest

infection on P. contorta. Most areas where A.

americanum occurs in pure P. ponderosa stands are

outside the range of the common ponderosa pine

dwarf mistletoe, A. vaginatum suhsp. cryptopodum

(Hawksworth 1968).

ARCEUTHOBIUM APACHECUM

APACHE DWARF MISTLETOE

4. Arceuthohium apachecum Hawksworth & Wiens,

Brittonia 22: 266, 1970. TYPE: ARIZONA: Pima

Co.: Santa Catalina Mountains near the summit

of Mt. Lemmon at 9,100 feet, on Pinus strohiformis, Hawksworth, Lightle, and Gilhertson 1110,

September 13, 1968. (US, Isotypes ARIZ, COLO,

DS, FPF, MO, RM, UC, UT).

DESCRIPTION: Mean shoot height ca. 3-4 cm.,

but up to 7 cm. (fig. 35). Shoots yellow green, or

reddish, flabellately branched and densely clustered. Basal diameter of dominant shoots 1-2 mm.

(mean 1.8 mm.). Third internode 5-10 mm. (mean

7.2 zb 2.0 cm.) long and 1-2 mm. (mean 1.5 mm.)

wide (12 collections), length/width ratio 4.8:1.

Flowers axillary. Staminate flowers 2.7 mm. in

diameter; perianth 3- or 4-merous; same color

as shoots; segments ca. 1.3 mm. long and 0.9

mm. wide. Mean anther diameter 0.45 mm., centered 0.7 mm. from tip of segment. Pollen polar

diameter 16-23 (mean 19)Lt); equatorial diameter

18-23 (mean 21jLt); polar/equatorial diameter ratio

1:1.11; spine height 1.5^ slightly greater than wall

thickness—ca. l.lju (4 collections). Mature fruit

4 mm. long and 2.5 mm. wide; proximal portion

ca. 2.5 mm. long. Seeds 2.8 X 1.2 mm. Meiosis in

July. Anthesis usually in early September. Seeds

usually mature in September; maturation period

averages ca. 12 months, n = 14.

HOST : Known only on Pinus strohiformis.

DISTRIBUTION: This species occurs from eastcentral Arizona (near Springerville) and New

Mexico (Mangas, San ]\Iateo, Magdalena, and

Capitán Mountains) south to the Santa Catalina,

Rincón, and Chiricahua Mountains of Arizona

and to the Sierra del Carmen in northern Coahuila,

Mexico (fig. 36). Elevational range: 7,700 to 9,800

feet. Specimens examined: see appendix, p. 195.

DISCUSSION: GilPs Arceuthohium campylopodum

f. hlumeri is here considered to comprise three nonsympatric species: A. apachecum, A. hlumeri, and

A, californicum. These differ in many characters

including morphology, hosts, phenology, and distribution. Our numerical analyses confirm that

these taxa should be regarded as species (see fig.

24).

The exclusive occurrence of two dwarf mistletoes on a single host species {Pinus strohiformis) is

unique in Arceuthohium. In our early studies of

A. hlumeri and A. apachecum, we thought they

might represent a single, variable taxon. However,

101
h 2 mm

<c^o>

Figure 35.—Arceuthobium apachecum in September: A, Staminate plant; B, Detail ofstaminate flower; C, Pistillate

plant; D, Detail of mature fruit.

102
Figure 36.—Distribution of Arceuthobium apachecum (based on collections only). Outline map copyrighted by

Denoyer-Geppert Co., Chicago (used by permission).

when further investigation revealed that the

morphological differences were geographically consistent, we decided that separate taxonomic status

was warranted. Also, under common conditions in

a greenhouse at Fort Collins, Colorado, each species maintained its distinct morphological features.

We have not found the two species in the same

mountain range, although they occur about 40

miles apart in southern Arizona. Arceuthobium

apachecum is abundant north and east from the

Chiricahua, Graham, Santa Catahna, and Rincón

ranges, and A. blumeri is south from the Huachuca

and Santa Rita ranges.

A comparison of the two taxa is given in table

18. We do not have precise flowering data for A.

apachecum, but its peak period of anthesis seems

to be somewhat later (early September) than that

of A. blumeri (mid-August?). Fruits of both taxa,

however, mature in September.

TABLE 18.—Comparison ofArceuthobium apachecum

and A. blumeri

Characters A. apachecum A. blumeri

Shoots;

Size, mean

(maximum) 3-4 (7) cm. 6-7 (18) cm.

Color Yellowish Gray

Habit Densely clustered Not densely

around host clustered

branch

Lateral staminate

spikes in summei 6 X 1 mm. 11X2 mm.

Witches' brooms Consistent Rare

103
ARCEUTHOBIUM BICARINATUM

HISPANIOLAN DWARF MISTLETOE

5. Arceuthobium hicarinatum Urban, Symb. Antillanae 7: 204, 1912. TYPE: DOMINICAN REPUBLIC: Constanza, on Pinus occidentalis, von

Turckheim 3241 in 1910 (Z! Isotypes BM, F, ILL,

MO, NY, US). Razoumofskya hicarinata (Urban)

Tubeuf, Naturwiss. Z. Forst-u. Landwirt. 17: 195,

1919.

DESCRIPTION: Mean shoot height ca. 10 cm., but

up to 17 cm. (fig. 37). Shoots dark brownish red,

terminal branches of Hving plants conspicuously

whitish-blue glaucous, not glaucous when dried;

shoots frequently twisted when dry, flabellately

branched. Basal diameter of dominant shoots 2-4

mm. (mean 3 mm.). Third internode 6-14 mm.

(mean 10.5 ±2.1 mm.) long and 1.5-4 mm. (mean

2.0 mm.) wide (6 collections), length/width ratio

5.2:1. Staminate flowers ca. 3 mm. in diameter;

perianth 3- (sometimes 4-) merous, apex acute,

reddish; segments ca. 1.5 mm. long and 1.0 mm.

wide. Mean anther diameter 0.5 mm., centered 1.0

mm. from tip of segment. Pollen polar diameter

21-25 (mean 23M); equatorial diameter 25-28

(mean 26jLt); polar/equatorial diameter ratio

1:1.14; spine height (ca. Zß) about 3 times wall

thickness—ca. 1/x (3 collections). Mature fruit ca.

4 mm. long and 2 mm. wide; proximal portion ca.

2.5 mm. long. Seeds 2.5 X 1.2 mm. Meiosis in

August. Anthesis usually in September. Seeds

mature in late August or September of the year

following pollination; maturation period averages

ca. 12 months, n = 14.

HOSTS: Known only on Pinus occidentalis on the

island of Hispaniola (Haiti and the Dominican

Republic).

DISTRIBUTION: Arceuthobium bicarinatum is

common in both Haiti and the Dominican Republic (fig. 38). In Haiti, the dwarf mistletoe is

common in the Morne la Selle and the Morne des

Commissaires, in the southeastern part of the

country.

Dominican Republic

A, bicarinatum

In the Dominican Republic, A. bicarinatum is

damaging on all three major mountain ranges

where Pinus occidentalis is found: The Cordillera

Central, the Sierra de Neiba, and the Sierra de

Baoruco. It is estimated that more than half of the

pine forests in the Dominican Republic are infested, and damage is particularly severe in Santiago and San Juan Provinces.^i In our studies of

this parasite, mostly in La Vega Province, we observed A. bicarinatum commonly between 3,000

and 8,500 feet. Specimens examined: see appendix,

p. 195.

DISCUSSION: This is a distinctive species restricted to Hispaniola (see fig. 12). Distinguishing

features are the dark brownish-red color, sterile

nodes near the tips of the pistillate shoots, and

glaucous terminal internodes. Internodes on dried

specimens often show a twist of about 1/4 turn,

but this trait is not discernible in living plants.

Some characteristics that distinguish this species

from the related A, hondurense of Honduras are

summarized in table 19.

The witches' brooms caused by A. bicarinatum

are mostly nonsystemic (as illustrated by MarieVictorin 1943). In one area near Valle Nuevo in

the Cordillera Central, however, some witches'

brooms were anomalous : they appeared to be systemic but dwarf mistletoe shoots were not regularly produced on the infected branches as is

normally the case for systemic infections in other

host-parasite combinations. Detailed studies are

needed on the witches' brooms induced by A.

bicarinatum.

At higher elevations in Haiti and the Dominican

Republic a leafy mistletoe, Dendropemon pycnophyllus Krug & Urban, is common on Pinus

occidentalis and frequently occurs on trees infected

by A. bicarinatum.

ARCEUTHOBIUM BLUMERI

6. Arceuthobium blumeri A. Nelson, Bot. Gaz. 56:

65, 1913. HOLOTYPE: ARIZONA: Cochise Co.:

Huachuca Mountains., on Pinus strobiformis,

Blumer in Oct., 1910 (RM no. 78604!). Razoumofskya blumeri Standley, Proc. Biol. Soc. Wash. 29:

86, 1916. Arceuthobium campylopodum Engelm.

forma blumeri (Engelm.) Gill, Trans. Conn. Acad.

Arts & Sei. 32: 207, 1935 (in part).

DESCRIPTION: Mean shoot height ca. 6-7 cm.,

but up to 18 cm. in Durango, Mexico (fig. 39).

Shoots gray to straw or light green, flabellately

branched. Basal diameter of dominant shoots 1-3

Figure 38.—Distribution of Arceuthobium hicarinatum

in Hispaniola (based on collections only).

21 We wish to thank Anthony F. Gasbarro, Peace Corps

Volunteer in the Dominican Republic during 1962 and 1963,

for this information.

104
Figure 37.—Arceuthohium hicarinatum in September: A, Pistillate shoot; B, Detail of fruit; C, Staminate shoot;

D, Detail of staminate flowers.

105
TABLE 19.—Comparison of Arceuthobium bicarinatum of Hispamola and

A. hondurense of Honduras

Characters A. bicarinatum

Host

Shoots:

Basal diameter

Nodes on old shoots

Color

Mature fruits:

Size, mean

Color

Stigmas

Staminate flowers:

Perianth shape

Nectary shape

Pinus occidentalis

2-3 mm.

Not swollen

Brownish red

2.0 X 4.0 mm.

Reddish, nonglaucous

Not exserted

Sides parallel, pointed tip

Three lobes the same size

A. hondurense

Pinus oocarpa

3-9 mm.

Swollen

Olive brown to grayish green

3.0 X 5.5 mm.

Greenish, glaucous

Markedly exserted

Rounded to tip

One lobe markedly smaller

than the other two

mm. (mean 2.1 mm.). Third internode 5-14 cm.

(mean 9.1 db 2.5 cm.) long and 1-2 mm. (mean 1.6

mm.) wide (8 collections), length/width ratio

5.5:1. Staminate flowers ca. 2.5 mm. long and

2.5-3 mm. in diameter; perianth 4-, 5-, or 6-

merous, apex acute; segments ca. 1.3 mm. long and

1.0 mm. wide. Mean anther diameter 0.6 mm.,

centered 0.4 mm. from tip of segment. Few pollen

grains available for study, but those found had a

polar diameter of ca. 19/^ and an equatorial diameter of ca. 20/x; spine height and wall thickness

each ca. l/x (1 collection). Mature fruit ca. 4 mm.

long and 2.5 mm. wide; proximal portion ca. 2.5

mm. long. Seeds 2.7 X 1.0 mm. Meiosis in July.

Anthesis not precisely known but presumably in

August. Seeds mature in September of the year

following polUnation; maturation period averages

ca. 13 months, n = 14.

HOST : Known only on Pinus strobiformis.

DISTRIBUTION: This mistletoe ranges southward

from the Huachuca and Santa Rita ranges in

southern Arizona through the Sierra Madre Occidental to southern Durango, and on to Cerro

Potosí in Nuevo León in Mexico (fig. 40). Not yet

collected in Sonora but we expect it to be found

there. Known elevational range: 7,100 to 9,800

feet. Specimens examined: see appendix, p. 195.

DISCUSSION: The parasitism of Arceuthobium

blumeri and A. apachecum on Pinus strobiformis is

discussed under A. apachecum and the two taxa

compared in table 18.

Unusual features of A. blumeri include its gray

shoots, 4- to 6-merous staminate flowers, and rare

formation of witches' brooms.

ARCEUTHOBIUM CALIFORNICUM

SUGAR PINE DWARF MISTLETOE

7. Arceuthobium californicum Hawksworth & Wiens,

Brittonia 22: 266, 1970. TYPE: CALIFORNIA:

MARIPOSA CO.: Fish Camp, 0.25 mi. W of

State Route 41 on Summit Camp rd., on Pinus

lambertiana, Hawksworth cfc Hawksworth 1147,

November 6, 1968 (US, Isotypes ARIZ, COLO,

DS, FPF, MO, RM, UC, UT). Arceuthobium

campylopodum Engelm. var. crytopodum (Engelm.)

Jepson, Man. Fl. PL Calif. 284. 1925.

DESCRIPTION: Mean shoot height ca. 8 cm., but

up to 12 cm. (fig. 41). Shoots greenish to bright

yellow, turning brown at base of older shoots,

flabellately branched. Basal diameter of dominant

shoots 1.5-4 mm. (mean 2 mm.). Third internode

6-16 mm. (mean 10.5 zt 2.9 mm.) long and 1-2

mm. (mean 1.5 mm.) wide (8 collections), length/

width ratio 7.0:1. Flowers axillary. Staminate

flowers 3.3 mm. in diameter; perianth 3- or 4-

merous, segments ca. 1.5 mm. long and 1.1 mm.

wide. Mean anther diameter 0.5 mm., centered

0.7 mm. from tip of segment. Pollen polar diameter 18-23 (mean 20ju); equatorial diameter 19-25

(mean 23/x); polar/equatorial diameter ratio

1:1.12; spine height ca. 1.6^ approximately twice

wall thickness—ca. 0.8/x (4 collections). Mature

fruit 4 mm. long and 2.5 mm. wide; proximal portion ca. 2.0 mm. long. Seeds 3.2 X 1.2 mm. Meiosis

in July. Anthesis usually in July, with extremes

from late June to early August. Seeds usually

mature about mid-September, with extremes from

late August to early October; maturation period

averages ca. 13-14 months, n = 14.

106
c m

Figure 39.—Arceuthobium blumeri in winter: A, Staminate plant; B, Pistillate plant; C, Immature fruit; D, Terminal portion of staminate shoot.

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Figure 40.—Distribution of Arceuthobium blumeri in Mexico and Arizona (based on collections only).

HOSTS AND DISTRIBUTION: The principal

host is sugar pine, Pinus lamhertiana. The parasite

ranges from southern Oregon south through the

Sierra Nevada to the Cuayamaca Mountains of

San Diego County and in the Coast Range south

to Lake County, California (fig. 42). A, californicum has been reported on Pinus monticola, the

only other common host, in the Plumas National

Forest, California, and in the Siskiyou Mountains,

southern Oregon. The only other known host is the

rare Picea hreweriana in southern Oregon, but in

this area the distribution of sugar pine mistletoe

is poorly known. Also, A. californicum was observed in several CaHfornia localities during disease surveys22 (Calif. Forest Pest Control Action

Council 1968). GilFs (1935) report of this species

on Pinus monticola at Rhododendron, in northern

Oregon, is based on A. tsugense. The known eleva22 Personal communication, D. R. Miller, U.S. Forest

Service, 1968.

tional range of this mistletoe is from 2,000 to

6,500 feet. Specimens examined: see appendix,

p. 196.

DISCUSSION: Jepson's (1925) variety cryptopodum^ which was intended for the parasites of

Pinus lamhertiana and Abies grandis, is invalid because it is based on a misapplied name A. cryptopodum Engelm., a Rocky Mountain parasite of

Pinus ponderosa.

As discussed under A. apachecum, GilPs A.

campylopodum f. hlumeri includes three distinct

species: A, apachecum, A. hlumeri, and A. californicum. A californicum differs from A. hlumeri

and A. apachecum in its larger size, earlier meiosis

and anthesis, distinct hosts, and a more westerly

range. A. californicum is distinguished from A.

cyanocarpum, the only other western dwarf mistletoe that parasitizes the white pines, by its much

larger size, shoots not densely clustered, earlier

meiosis and flowering, larger staminate spikes, and

its parasitism of P. lamhertiana. Arceuthohium

108
Figure 41.—Arceuthohium californicum in summer: A^ Pistillate plant; ß, Staminate plant; C, Pistillate flower

(left) and nearly mature fruit (right); D, Terminal portion of staminate shoot.

109
Figure 42.—Distribution of Arceuthobium californicum. Outline map copyrighted by Denoyer-Geppert

Co., Chicago (used by permission).

cyanocarpum did not attack P. lambertiana in the

only case we have seen where the mistletoe was

associated with this pine (in the San Jacinto

Mountains of southern California).

Gill (1935) reported that "infected trees are infrequent and were found only in presence of other

infected species." Our studies indicate that neither

statement is applicable to A. californicum, because this dwarf mistletoe is very damaging in

California (Scharpf and Hawksworth 1968). Results of surveys show that the parasite was present

on 22 percent of 274 Pinus lambertiana plots in

CaUfornia (Calif. Forest Pest Control Action

Council 1968). We have seen many instances where

A. calif ornicum was the only dwarf mistletoe present in a stand. Sometimes it is sympatric with A.

campylopodum, however, and this may have been

the basis for Gill's statement.

This mistletoe induces large compact witches'

brooms in Pinus lambertiana (see fig. 10).

ARCEUTHOBIUM CAMPYLOPODUM

WESTERN DWARF MISTLETOE

8. Arceuthobium campylopodum Engelmann in Gray,

Boston, J. Natur. Hist. 6: 214, 1850. TYPE:

"Oregon," on Pinus ponderosa, Geyer 677 in 1843

(GH! Isotype US). Arceuthobium campylopodum

Engelm. var. macrarthron Engelm. loc. cit.

Razoumofskya campylopoda (Engelm.) Kuntze,

Rev. Gen. PI. 2:587, 1891. Arceuthobium campylopodum Engelm. forma typicum Gill, Trans. Conn.

Acad. Arts & Sei. 32:185,1935 (in part).

DESCRIPTION: Mean shoot height ca. 8 cm., but

up to 13 cm. (fig. 43). Shoot olive-green to yellow,

flabellately branched. Staminate plants brownish,

and pistillate plants greenish, particularly in

northern populations. Basal diameter of dominant

shoots 1.5-5 mm. (mean 3 mm.). Third internode

7-22 mm. (mean 11.3 ± 3.8 mm.) long and 1.5-2.5

mm. (mean 2.0 mm.) wide (27 collections), length/

width ratio 5.6:1. Staminate ñowers 3.0 mm. in

diameter; perianth 3- (occasionally 4-) merous,

segments ca. 1.4 mm. long and 1.0 mm. wide.

Mean anther diameter 0.5 mm., centered 0.8 mm.

from tip of segment. Pollen polar diameter 18-25

(mean 20/1); equatorial diameter 20-27 (mean

24AI) ; polar/equatorial diameter ratio 1:1.18; spine

height ca. 2.2/LI about 2.5 times wall thickness—ca.

0.8/i (5 collections). Mature fruit 5.0 mm. long

and 3.0 mm. wide. Meiosis in July. Peak anthesis

usually during mid-September, with extremes

from early August to late October. Seeds usually

mature in late September, with extremes from

late August to late November; maturation period

averages about 13 months, n = 14.

HOSTS: The principal and commonly infected hosts

are Pinus ponderosa var. ponderosa and P. jeffreyi.

Other trees frequently infected, particularly when

associated with the above hosts, are P. coulteri and

P. attenuata. In the Charleston Mountains, Nevada, Pinus ponderosa var. scopulorum is a common host, but this is the only record where A.

campylopodum occurs naturally on this tree. In

Oregon, however. Roth (1967) inoculated varieties

scopulorum and ponderosa with A. campylopodum,

and found both to be equally susceptible.

Occasional hosts ior A. campylopodum are Pinus

contorta vars. latifolia and murrayana. Dr. T. W.

Childs^' informs us that Pinus lambertiana is very

rarely infected by this mistletoe in the vicinity of

Hammer Butte in the Deschutes National Forest

in northern Klamath County, Oregon.

Weir (1918a) successfully inoculated seedlings

of Pinus sylvestris, P. montana, P. resinosa, Picea

abies, Abies concolor, A. grandis, and Larix occidentalis. The latter three trees are commonly asso-

' Personal communication, U.S. Forest Service, 1964.

110
Figure i^.—Arceuthobium campylopodum in winter: A^ Pistillate plant with young fruits; B, Staminate plant

with remnants of flowers; C, Old flower.

Ill
ciated with A. campylopodum on Pinus ponderosa,

but we have never seen the parasite on any of the

three. Kuijt (1960b) found the European Pinus

pinaster infected naturally at the Institute of

Forest Genetics Arboretum at Placerville, California. J. R. Weir collected A. campylopodum on

Pinus sylvestris near Hayden Lake, Idaho (see

specimens cited).

Pinus washoensis is probably susceptible to A.

campylopodum, but we know of no collections on

this host. We found none on the extensive populations of P. washoensis in the vicinity of Patterson

Guard Station in the southern end of the Warner

Mountains, Alodoc County, California. Similarly,

Dr. R. S. Peterson^* reports no A. campylopodum

in the type locality for P. washoensis on Mount

Rose, near Reno, Nevada.

DISTRIBUTION: This dwarf mistletoe occurs

from northern Washington and eastern Idaho,

south through Oregon and California (but not

known in the southern coast range) to the Sierra

de San Pedro Mártir in Baja California, Mexico.

In Nevada, it is found in the vicinity of Lake

Tahoe and in the Charleston Mountains of Clark

County (fig. 44). Not known from British Columbia, A. campylopodum occurs within about 20 miles

of the International Boundary near Kettle Falls,

Washington. Gill's (1935) reports of this species

in western Montana are based on occasional parasitism of P. ponderosa by the larch dwarf mistletoe,

A. laricis. The wide elevational distribution of A.

campylopodum ranges from about 100 feet along

the Columbia River near the mouth of the Hood

River in Oregon to 7,800 feet in the Charleston

Mountains of southern Nevada. Specimens examined: see appendix, p. 196.

DISCUSSION: The type locality is Hsted as "Oregon," although Piper (1906) says that the specimen must have come from northern Idaho or

northeastern Washington because Geyer did not

collect in the area that is now Oregon.

A. campylopodum is a serious pathogen in the

Pinus ponderosa and P. jeffreyi forests. According

to Wagener (1965), P. jeffreyi is more susceptible

than P. ponderosa, but both species suffer considerable damage and are classed here as principal

hosts. Our observations suggest that damage is

more severe in southern parts of the hosts' range;

the most seriously damaged stands were seen in the

Laguna Mountains in San Diego County, California. In the northern Sierra Nevada, damage is

more conspicuous on the drier east-side than along

the moister west-side forests. Roth (1966) reports

that drooping-needled races of ponderosa pine in

Oregon are less frequently infected by dwarf

mistletoe.

Daubenmire (1961) found A. camylopodum in

only two of the seven habitat types with ponderosa

pine in eastern Washington and northern Idaho.

The two habitat types with dwarf mistletoe—

Pinus/Agropyron and Pinus/Purshia—were the

driest of the seven recognized. The presence or absence of Arceuthobium was consistent enough to

be used as one of the key characters to determine

ponderosa pine habitat types (Daubenmire and

Daubenmire 1968).

Arceuthobium campylopodum f. typicum of Gill

(1935) is here considered to comprise two species,

A. campylopodum and A. occidentale, separable on

the basis of phenology, morphology, geographic

distribution, and hosts (fig. 45, table 20).

G. G. Hedgcock (unpublished manuscript dated

1914) first noted some differences between these

two mistletoes and, although he suggested that

they warranted separate taxonomic status, he did

not publish descriptions of them. In addition to

the differences mentioned in table 20 the two

" Personal communication, U.S. Forest Service, 1966.

Figure 45.—Comparative distribution of Arceuthobium

campylopodum and A. occidentale. Outline map

copyrighted by Denoyer-Geppert Co., Chicago (used

by permission).

112
Figure 44.—Distribution of Arceuthobium campylopodum on all hosts and the range of Pinus ponderosa var.

ponderosa (based on collections only). Localities for the mistletoe outside the range of this host are on Pinus

ponderosa var. scopulorum (southern Nevada) and on Pinus jeffreyi (Baja California). Outline map copyrighted

by Denoyer-Geppert Co., Chicago (used by permission).

113
TABLE 20.—Comparison of Arceuthobium campylopodum and

A. occidentale

Characters A. campylopodum A. occidentale

Principal hosts

Shoots

Lateral staminate

spikes in summer :

Length

Length/width ratio

Peak flowering time

Peak seed dispersal time

Distribution

Pinus ponderosa

Pinus jeffreyi

Pinus attenuata

Glaucous; shoot

usually open

clusters

Less than 10 mm.

Three or less

September or earlier

October or before

Washington and Idaho to

Baja California

Pinus sahiniana

Pinus radiata

Pinus muricata

Nonglaucous or only slightly

so; shoots in dense, globose

clusters

Over 10 mm.

Five or more

October or later

November or later

California

dwarf mistletoes differ in germination characteristics and host reactions. The seeds of A. campylopodum require an afterripening period (Beckman

and Roth 1968, Wicker 1965), while those of A,

occidentale do not (Scharpf and Parmeter 1962).

Witches^ brooms caused by A. campylopodum (see

fig. 14) are larger than those caused by A. occidentale (see fig. 8).

Host data for A. campylopodum and A. occidentale, based on observations of infection in

stands where at least two species of pines occur,

are given in fig. 46. The known natural hosts for

A. campylopodum are Pinus ponderosa, P. jeffreyi,

P. attenuata, P. coulteri, P. contorta (subsp. latifolia, murrayana, and contorta) and, very rarely,

P. lamhertiana. The hosts of A. occidentale are P.

sahiniana, P. radiata, P. muricata, P. attenuata,

P. coulteri, and P. contorta subsp. holanderi. Pinus

attenuata and P. coulteri may be attacked by either

dwarf mistletoe, depending on the locality where

the hosts occur.

Arceuthohium campylopodum (on P. ponderosa)

and A. occidentale (on P. sahiniana) are sympatric

in many areas of the Sierra Nevada and north

Coast ranges. Despite intensive searches in several

such areas, we found no evidence of cross infection

between these two hosts. Neither did we observe

sympatry between these two dwarf mistletoes on

other hosts.

ARCEUTHOBIUM CYANOCARPUM

LIMBER PINE DWARF MISTLETOE

9. Arceuthohium cyanocarpum Coulter & Nelson,

New Man. Bot. Cent. Rocky Mts.: 146, 1909.

LECTOTYPE: WYOMING: Carbon Co.: Ferris

Mountains, on Pinus flexilis. Nelson 4959 in 1898

(RM! Isotypes NY, MO). Razoumofskya cyanocarpa A. Nelson in Rydberg, Colo. Agr. Exp. Sta.

Bull, 100: 101, 1906 nomen nudum. Arceuthohium

cyanocarpum (A. Neis.) Abrams, lUus. Flora

Pacific Coast States I: 531, 1923. Arceuthohium

campylopodum Engelm. forma cyanocarpum (A.

Neis.) Gill, Trans. Conn. Acad. Arts & Sei. 32:

204, 1935.

DESCRIPTION: Mean shoot height ca. 3 cm., but

up to 5 cm. (fig. 47). Shoots yellow green, flabellately branched, densely clustered. Basal diameter

of dominant shoots 1-2 mm. (mean 1.4 mm.).

Third internode 2-14 mm. (mean 5.2 d= 2.0 mm.)

long and 1-1.5 mm. (mean 1.1 mm.) wide (15

collections); length/width ratio 4.7:1. Staminate

flowers 3.0 mm. in diameter; perianth 3- (rarely

4-) merous, apex acute; same color as shoots; segments ca. 1.4 mm. long and 1.0 mm. wide. Mean

anther diameter 0.4 mm., centered 0.7 mm. from

tip of segment. Pollen polar diameter 15-21 (mean

19M); equatorial diameter 20-25 (mean 22/i);

polar/equatorial diameter ratio 1:1.19; spine

114
contorta latifolia contorta nnurrayana

ponderosa

contorta contorta jeífreyi

attenuate ^ coulteri

sabmiana

contorta bolanderi radiata

muricata

Dwarf mistletoe

Crossinfection

No crossinfection

Arceuthobium campylopodum

Arceuthobiunn occidentale

Figure 46.—^Natural hosts of Arceuthobium campylopodum and A, occidentale.

height (ca. l.öju) approximately twice wall thickness—ca. 0.8/i (6 collections). Mature fruit 3.5

mm. long and 2.0 mm. wide; proximal portion ca.

2.0 mm. long. Seeds 2.0 X 0.9 mm. Meiosis in

July. Anthesis usually in late August, with extremes from mid-July to early September. Seed

mature in late August or early September; maturation period averages ca. 12 months. Germination

mostly in June, n = 14.

HOSTS: The principal host is Pinus flexilis. Pinus

aristata and P. alhicaulis appear to be equally susceptible, but these trees are not as common within

the range of the mistletoe, and relatively few collections on them have been made. In northern

California, A. cyanocarpum also occurs on Pinus

montícola and rarely on P. halfouriana (Miller and

Bynum 1965). Rare hosts include Pinus ponderosa

var. scopulorum and Pinus contorta subsp. latifolia

in the Rocky Mountains (Hawksworth and Peterson 1959).

Tubeuf (1919) mentions a single infection on

Picea engelmannii from Pikes Peak, Colorado,

which Hawksworth and Graham (1963) surmised

was this taxon.

Inoculations of seedlings by A. cyanocarpum in

a greenhouse have been successful on Pinus strohus

and P. strohiformis (Hawksworth, unpublished

data). The latter is of interest because, although

115
Figure 47,—ArceuthobiuTn cyanocarpum in September: A, Pistillate plant; ß, Detail of mature fruit; C, Pistillate

flowers just pollinated; D, Staminate plants; E, Mature staminate flower.

116
this tree is naturally exposed to A. cyanocarpum

on the San Francisco Peaks, Arizona, it is not infected there (Hawksworth 1965a). This emphasizes

that conclusions on host relations based on inoculations under artificial conditions must be viewed

with caution.

Arceuthobium cyanocarpum is not sympatric

with A. hlumeri, and the former does not attack

P. strohiformis in the only known area where this

tree and the mistletoe occur together in the San

Francisco Peaks in northern Arizona (Hawksworth 1965a).

DISTRIBUTION: This dwarf mistletoe occurs

from southern Montana (Brown 1970) and northern Colorado southwestward to northern CaUfornia; down the east side of the Sierra Nevada

and in San Bernardino and San Jacinto Mountains of southern California (fig. 48). Isolated occurrences are known from the vicinity of Pikes

Peak, and in the Sangre de Cristo Mountains near

Coaldale, Colorado (on P. flexilis), La Sal Mountains near the Utah-Colorado border {on P.

flexilis), San Francisco Peaks, Arizona* (on P.

aristata), and the Mount Shasta area, California {on

P. alhicaulis and P. monticola). Known now from

only three areas in Idaho and two in Nevada, it

seems likely that A. cyanocarpum will be found in

other P. flexilis localities in these States. For

example, Linsdale, Howell, and Linsdale (1952)

illustrate a broomed Pinus flexilis from the

Toiyabe Mountains (Nye County, Nevada) which

appears to be infected by A. cyanocarpum. The

specimens listed by Gill (1935) as A. campylopodum f. cyanocarpum on Pinus alhicaulis from

the Crater Lake region of Oregon are A. tsugense.

This mistletoe may occur in the mountains of

southern Oregon, however, and we have tentatively listed a collection on P. alhicaulis from

''Obsidian Cliff,'' Oregon, as this taxon.

The known elevational range of this mistletoe is

from 5,200 feet in southern Montana to 9,800 feet

in central Colorado. Specimens examined: see appendix, p. 201.

DISCUSSION: Weir made several collections of A.

cyanocarpum in the vicinity of Anaconda, Montana, in 1913 and 1914, but we were unable to

find it on Pinus flexilis in that area in 1966. Possibly the sulfur dioxide fumes that damaged the

vegetation so severely there have eUminated the

parasite. Scheffer and Hedgcock (1955) who studied the area in the late 1920's, noted that Pinus

* 1970 and 1971 studies of the San Francisco Peaks population on Pinus aristata, which we tentatively class as A.

cyanocarpum, revealed four previously unrecorded hosts:

Picea pungens and P. engelmannii are occasional hosts and

Abies lasiocarpa var. arizonica and Pinus strohiformis rare

hosts. Taxonomic studies of this population are continuing.

flexilis and dwarf mistletoe were relatively rare in

the fume-damaged area.

This characteristic dwarf mistletoe of Pinus

flexilis and associated high-altitude white pines is

easily recognized by its small, densely clustered

shoots. In many cases shoot mortality is excessive

(cause unknown but probably a fungal disease),

and only dense clusters of shoot bases about 1 mm.

high remain.

Arceuthohium cyanocarpum causes heavy mortality in Pinus flexilis in the Rocky Mountains

and in P. alhicaulis on Mount Shasta, California.

In the latter area, Cooke (1955) noted that it

causes death of trees over large areas resulting in

'^ghost forests.^^

Witches' brooms induced by A. cyanocarpum

are typically small and compact. In many instances, however, infection is so general throughout the crown of the tree that witches' brooms are

not distinct.

ARCEUTHOBIUM DIVARICATUM

PINYON DWARF MISTLETOE

10. Arceuthohium divaricatum Engelmann in U.S.

Geogr. Surv. W of 100th Merid. (Wheeler Rep.)

6: 253, 1878. LECTOTYPE: ARIZONA: County

unknown. Salt River Valley, on Pinus edulis,

Gilhert 116 in 1873 (MO! Isotype US). Arceuthohium gracile Engelmann in Gray, Mem. Amer.

Acad. N.S. 4: 59, 1849, nomen nudum. Razoumofskya divaricata Coville, Contr. U.S. Nat. Herb. 4:

192, 1892. Arceuthohium campylopodum Engelmann forma divaricatum (Engelm.) Gill, Trans.

Conn. Acad. Arts & Sei. 32:193,1935.

DESCRIPTION: Mean shoot height ca. 8 cm. but

up to 13 cm. (fig. 49). Shoots olive green to brown,

flabellately branched. Basal diameter of dominant

shoots 1.5-4 mm. (mean 2 mm.). Third internode

6-15 mm. (mean 9.8 dz 2.4 mm.) long and 1-2

mm. (mean 1.6 mm.) wide (19 collections), length/

width ratio 6.1:1. Staminate flowers 2.5 mm. in

diameter; perianth 3-merous; segments ca.^ 1.1

mm. long and 0.9 mm. wide. Mean anther diameter 0.45 mm., centered 0.5 mm. from tip of segment. Pollen polar diameter 18-26 (mean 20M);

equatorial diameter 21-30 (mean 24^); polar/

equatorial diameter ratio 1:1.17; spine height 1.6/x

ca. 1.5 times wall thickness—ca. 1.0^ (8 collections). Mature fruit 3.5 mm. long and 2.0 mm.

wide; proximal portion ca. 2.0 mm. long. Seeds

2.0 X 0.9 mm. Meiosis in July. Peak anthesis

usually in early September, with extremes from

early August to late September. Seeds usually

mature in late September with extremes from early

September to early November in the year following poUination; maturation period average ca. 13

months, n = 14.

117
Figure 48.—Distribution of Arceuthobium cyanocarpum (based on collections only). Outline map copyrighted by

Denoyer-Geppert Co., Chicago (used by permission).

HOSTS : This mistletoe, known only on the pinyons,

is common on Pinus edulis and P. monophylla, but

rare on P. quadrifolia and P. cembroides. The

latter two appear to be equally susceptible, but the

parasite occurs infrequently within their ranges.

Although mistletoe-infected pinyons are seldom

associated with other pines, the associated pines

(JP. ponderosa and P. jeffreyi) were not parasitized

in the few situations we observed. Thus the pinyon

dwarf mistletoe is not only restricted to pinyons.

118
Figure 49.—Arceuthobiuin divaricatum in summer: A, Pistillate plant; B, Detail of fruit; C, Staminate plant.

119
but also is the only dwarf mistletoe that parasitizes pinyon in the United States. In a single

locality in Mexico, however, we observed rare

parasitism of P. culminicola by A. vaginatum

subsp. vaginatum (Hawksworth and Wiens 1965).

DISTRIBUTION: Pinyon dwarf mistletoe occurs

in eastern and southern California (east side of

Sierra Nevada, Mount Pinos area, San Bernardino

Mountains), the southern two-thirds of Nevada,

southwestern Colorado, Arizona (except southwest). New Mexico (except northeast), and south

to the Davis Mountains of western Texas (fig.

50). In Baja CaUfornia, Mexico, A. divaricatum is

known in the Sierra Juárez and is probably in the

Sierra de San Pedro Mártir.^^ J. R. Weir (unpublished manuscript, 1920) reported A. divaricatum

on P. monophylla in extreme southern Idaho, but

we have seen no Idaho collections of this taxon.

The principal host in CaUfornia and Nevada is

P. monophylla; in Colorado, New Mexico, and

Texas it is P. edulis; but both species are parasitized in Arizona and Utah. The distribution map

(see fig. 50) suggests this mistletoe may be more

abundant on Pinus edulis than on P. monophylla,

but probably specimens have been more intensively collected within the range of P. edulis. The

mistletoe has been collected on P. cuadrifolia only

in Baja CaUfornia, (Hawksworth, Lightle, and

Scharpf 1968) but it is to be expected on this tree

in southern California. A. divaricatum is known

on P. cembroides only in the extreme northern part

of its range where this tree is associated with P.

edulis. The only coUections that we have seen are

from the Davis Mountains, in western Texas,

where Little (1966) determined the host as P.

cembroides, but noted that it was near the range

of P. edulis and may be intermediate. Pinyon

dwarf mistletoe also occurs on ^'typicaF' P. edulis

in this range. Elevational range of A. divaricatum

is from 5,300 feet in Baja CaUfornia to 8,900 feet

in the San Mateo Mountains of New Mexico.

Specimens examined: see appendix, p. 202.

DISCUSSION: The type locality for this taxon is

given by Gill (1935) as Salt '^Run'^ VaUey. The

original label, however, is handwritten and scarcely

legible, but it appears that Salt ^'River'' VaUey

was intended.

Kuijt (1960a) points out the spotty distributions of this parasite in California. We feel that

A. divaricatum is probably more widely distributed

than coUections indicate, but may be less fre25 Although we have seen no specimens from the Sierra de

San Pedro Mártir, Mr. Phillipe Meling, lifelong resident of

the area, informed us in 1965 that dwarf mistletoe occurs on

pinyon (presumably P. quadrifolia) on the north slopes of

this range.

120

quently coUected than other species of Arceuthobium. Reasons for its paucity in herbaria may be

that (1) it often does not form witches' brooms so

it is less conspicuous than most dwarf mistletoes,

and (2) its hosts are not commercial timber trees

so little attention has been paid to their pathology.

The pinyon dwarf mistletoe does cause witches'

brooms, but they are often poorly developed and

not conspicuous because of the stunted habit of

healthy trees. Our observations suggest that

brooming is more consistent in P. edulis than in

P. monophylla.

Shoots of the pinyon dwarf mistletoe are long

and slender, often divaricate. The staminate

plants have relatively few flowers per shoot.

ARCEUTHOBIUM DOUGLASII

DOUGLAS-FIR DWARF MISTLETOE

11. Arceuthobium douglasii Engelmann in U.S.

Geogr. Surv. W of 100th Merid. (Wheeler Rep.)

6: 253, 1878. LECTOTYPE: NEW MEXICO:

Santa Fe Co.: Santa Fe River, on Pseudotsuga

menziesii, Rothrock 69 in 1874 (MO! Isotypes F,

US). Razoumofskya douglasii (Engelm.) Kuntze

Rev. Gen. PI. 2:587,1891.

DESCRIPTION: Mean shoot height ca. 2 cm., but

up to 7 cm. (fig. 51). Shoots olive green, flabellately branched. Basal diameter of dominant

shoots 1-1.5 mm. (mean 1 mm.). Third internode

2-6 mm. (mean 3.6 ± 1.2 mm.) long and ca. 1.0

mm. wide (29 coUections), length/width ratio

3.6:1. Flowers usuaUy axiUary in pairs, occasionally borne on pedicel-like segments as in ^. americanum. Staminate flowers 2.0 mm. long and 2.3 mm.

in diameter; perianth mostly 3-(occasionally 4-)

merous; segments rounded at apex without a keel;

inner surface reddish to purple, lower surface same

color as shoots; segments ca. 1.0 mm. long and

1.0 mm. wide. Mean anther diameter 0.4 mm.,

centered 0.6 mm. from tip of segment. PoUen

polar diameter 16-25 (mean 21^); equatorial diameter 20-28 (mean 24^) ; polar/equatorial diameter ratio 1:1.16; spine height approximately equal

to waU thickness—1-1.2At (8 collections). PistiUate

flowers ca. 1.5 mm. long and 1.5 mm. in diameter.

Mature fruit olive green 3.5-4.5 mm. long (mean

4 mm.) and 1.5-2 mm. wide (mean 2 mm.),

obovate; proximal portion ca. 2.5 mm. long. Seeds

2-4 X 1.1 mm. Staminate meiosis in September,

pistillate meiosis in April (Jones and Gordon

1965). Anthesis usuaUy in March and Apnl, with

extremes from February to May. Seeds usually

mature in September of the year following pollination; maturation period averages 17-18 months.

Seeds germinate in March (Wicker 1967a). n = 14.
Figure 50.—Distribution of Arceuthobium divaricatum within the composite range of the pinyons (based on collections only). Outline map copyrighted by Denoyer-Geppert Co., (used by permission).

121
2 mm

Figure 5l.—Arceuthobium douglasii in spring: A, Pistillate (left) and staminate (right) plants on the same host

twig; ß. Detail of nearly mature fruit; C, Staminate shoot; D, Staminate shoot with mature buds and open

flower (left) and detail of open flower (right); E, Staminate shoots showing characteristic rounded buds.

122
Figure 52.—General distribution of Arceuthobium douglasii. Outline map copyrighted by Denoyer-Geppert Co.,

Chicago (used by permission).

123
HOSTS: The principal and only commonly infected

host is Pseudotsuga menziesii; both var. menziesii

(in Washington, Oregon, and California) and var.

glauca (from British Columbia to central Mexico)

are parasitized. The following trees are sometimes

infected when they are associated with infected

Douglas-fir: Abies concolor, A. grandis, A. lasiocarpa vars. lasiocarpa and arizonica, Picea engelmannii, and P. pungens. Of these secondary hosts,

only A. lasiocarpa var. arizonica seems to be

severely parasitized. We concur with Kuijt (1960a)

that reports of this mistletoe on Pseudotsuga

macrocarpa (Jepson 1925, Munz 1935) are probably erroneous.

DISTRIBUTION: Douglas-fir dwarf mistletoe

ranges as far north as Lytton and Sicamous in

southern British Columbia,^^ about 130 miles

north of the International Boundary. These areas

are somewhat north of the distribution of the

parasite given by Kuijt (1963). In Washington

and Oregon, it occurs east of the crest of the Cascades, but is common on the Pacific slope from the

Umpqua River drainage south to about latitude

40° N. in northern Cahfornia. It is found in eastern

Idaho, western Montana, western Wyoming,

Utah, the southern two-thirds of Colorado, on

Wheeler Peak in extreme eastern Nevada, and

throughout the mountainous areas of Arizona and

New Mexico (figs. 52, 53). In Mexico, it is known

in only three isolated localities—Coahuila, Durango, and Nuevo León (fig. 54)—but probably

occurs throughout the Sierra Madre Occidental in

Chihuahua, Sonora, and Durango.

Also shown on the distribution map (see fig. 53)

are several localities for the Douglas-fir dwarf

mistletoe in Cahfornia obtained during field work

for State-wide disease surveys (Calif. Forest Pest

Council 1968).

The distribution of this species shows some interesting pecuHarities. It does not occur west of

the Cascades in British Columbia, Washington, or

northern Oregon except for a few instances where

it is found within 4 to 10 miles west of the crest

at White Pass, Washington (Wicker 1969) and in

Clackamas and Linn Counties, Oregon.^^ However,

Wicker (1969) successfully inoculated seedlings of

coastal Pseudotsuga menziesii in western Washington with this dwarf mistletoe, and he suggests

that isolational factors have precluded its occurrence on the western slope of the Cascades. This

26 Personal communication, Dr. R. B. Smith, Canada Dep.

Fisheries and Forestry, 1966.

2^ Personal communications, J. L. Stewart and D. M.

Knutson, U.S. Forest Service, 1969-70.

124

dwarf mistletoe is also absent in a number of other

areas where P. menziesii is common, such as the

central Sierra Nevada of California, central Montana, and northern Colorado. The altitudinal

range of this mistletoe is correlated with latitude,

and it occurs as low as 900 feet near Lytton in the

Thompson River Valley in southern British

Columbia to as high as 10,000 feet on Cerro

Potosí in Nuevo León, Mexico. Specimens examined: see appendix, p. 205.

DISCUSSION: Although Gill (1935) questioned

the validity of this species, we consider ^. douglasii

to be one of the most distinctive and stable members of the genus. It has a latitudinal range of over

2,000 miles, yet has no characteristics that warrant

ecotypic or subspecific separation.

Much of the confusion in the literature on this

species is due to its supposed affinities with various

members of the A. campylopodum complex (Gill

1935). Weir (1918a) discussed the occurrence of

''a small purple-flowered form on Picea'' in the

Inland Empire which he, and we, consider to be

merely 4. douglasii on an unusual host.

This dwarf mistletoe severely damages Douglasfir (see fig. 8B).

ARCEUTHOBIUM GILLII

12. Arceuthobium gillii Hawksworth & Wiens, Brittonia 16: 55, 1964. HOLOTYPE: ARIZONA:

Cochise Co.: Huachuca Mountains, 0.5 mi. E of

Reef, on Pinus leiophylla var. chihuahuana,

Hawksworth & Lightle 236, in 1962 (COLO., Isotypes FPF, MO, ARIZ, DS).

DESCRIPTION: Shoots up to 25 cm. long, greenish yellow or dark brown to almost black; pistillate

plants usually erect, but the staminate plants often

strongly divaricate; flabellately branched, internodes 0.4-1.9 cm. long and 2-8 mm. wide; staminate flower ca. 3-3.5 mm. long and up to 3.5 mm.

in diameter; mostly 3-merous, perianth segments

ca. 1 4 mm. long and ca. 1.0 mm. wide, apex

rounded-obtuse; pistillate flower ca. 1.5 mm. long,

anthers 0.7 mm. in diameter, centered 0.9 mm.

from tip of segment, and ca. 1.0 mm. in diameter;

fruit ca. 4-5 mm. (mean 4.5) long and 2-3 mm.

(mean 2.5) wide, proximal portion 2-3 mm. long,

elliptical-obovate, conspicuously whitish-blue

glaucous on proximal portion; anthesis from approximately mid-February through April and

possibly May.

Parasitic on pines of the section Leiophylla and

P. teocote. This species comprises two subspecies

differentiated by shoot color, internode thickness,

geographic ranges, time of anthesis, and host

affinities (Hawksworth and Wiens 1965).
A. doug/asít

• collection

+ report

\h

100 200 300

Scale of Miles

Figure 53.—Distribution of Arceuthobium douglasii on all hosts in the United States and in British Columbia,

Canada (based on collections and reports). The range oîPseudotsuga menziesii is shown by the solid line. Outline

map copyrighted by Denoyer-Geppert Co., Chicago (used by permission).

125
Figure 54.—Distribution of Arceuthobium douglasii in Mexico (based on collections only).

Key to the Subspecies

1. Plant greenish to brownish yellow; parasitic

almost exclusively on Pinus leiophylla var. chihuahuana; anthesis from mid-February through midApril; fruit maturity in November; Sierra Madre

Occidental of Sonora and Chihuahua, Mexico

north to southern Arizona.

12a. A. gillii suhsp. gillii.

1. Plants dark brown to almost black; parasitic

on Pinus leiophylla vars. leiophylla and chihuahuana, P. lumholtzii and P. ieocote; anthesis from

mid-March through mid-April or later; fruit

maturity in September; Sierra Madre Occidental

of Durango to Hidalgo, Oaxaca, and Veracruz,

Mexico.

12b. A. gillii subsp. nigrum.

ARCEUTHOBIUM GILLII SUBSP. GILLII

CHIHUAHUA PINE DWARF MISTLETOE

DESCRIPTION: Mean shoot height ca. 11 cm. but

up to 25 cm. (fig. 55). Shoots greenish brown.

Basal diameter of dominant shoots 2.5-8 mm.

(mean 4 mm.). Third internode 5-18 mm. (mean

10.7 db 3.4 mm.) long and 2-4.5 mm. (mean 2.8

mm.) wide (9 collections), length/width ratio 3.8:1.

Staminate flowers 3.5 mm. long and 2.5-4 mm.

(mean 3.2 mm.) in diameter; pollen polar diameter

19-25 (mean 22jLt); equatorial diameter 24-28

mean 26M) ; polar/ equatorial diameter ratio 1:1.19;

spine height (1.7M) slightly greater than wall

thickness—1.3/x (4 collections). Pistillate flowers

ca. 1.5 mm. long and 1 mm. in diameter. Proximal

portion of fruit conspicuously glaucous. Seeds 3.1

X 1.4 mm. Meiosis in September. Anthesis usually

in March and April. Seeds mature in October^^

of the year following pollination; maturation

period averages ca. 19 months, the longest in the

genus, n — 14.

12a. Arceuthobium

subsp. gillii.

gillii Hawksworth & Wiens

28 Dr. R. L. Gilbertson of the University of Arizona found

that seed dispersal in 1967 occurred between October 13 and

November 5 in the Santa Catalina Mountains near Tucson.

126
Figure 55.—Arceuthobium gillii subsp. gillii in spring: A, Staminate plant in anthesis; ß, Detail of a mature staminate flower; C, Pistillate plant; D, Detail of fruit showing characteristic glaucousness of the proximal portion.

127
HOSTS : The principal and only commonly infected

host is Pinus leiophylla var. chihuahuana. Pinus

lumholtzii and P. leiophylla var. leiophylla seem to

be equally susceptible to the parasite, but are not

common in the areas where A, gillii occurs. In

western Chihuahua we found this mistletoe rarely

on Pinus ponderosa vars. scopulorum and arizonica,

but we have not found these trees to be infected

in Arizona although both are exposed to A. gillii

there.

DISTRIBUTION: This dwarf mistletoe occurs in

southeastern Arizona (Santa Catalina, Rincón,

Santa Rita, Huachuca, and Chiricahua Mountains), eastern Sonora, western Chihuahua, and

Sinaloa (fig. 56). It is to be expected in the Animas

Mountains in extreme southwestern New Mexico.

We have examined several other Pinus leiophylla

var. chihuahuana stands in New Mexico (on Bear

Mountain near Silver City, Mule Mountains, and

Peloncillo Mountains) without finding the parasite. Also the following Arizona localities of this

pine have been examined and found to be free of

A, gillii: Graham Mountains, the Final Mountains, and below the Mogollón Rim near Colcord

Mountain, Gila County.^^

Arceuthohium gillii ranges from 5,600 feet in

Arizona to 8,700 feet in southern Chihuahua.

Specimens examined: see appendix, p. 210.

DISCUSSION: This subspecies had long been confused with A. vaginatum subsp. cryptopodum but

it differs in several respects (Hawksworth and

Wiens 1964), some of which are summarized in

table 21.

29 Personal communication, Dr. P. C. Lightle, U.S. Forest

Service, 1969.

In Arizona, the two species are not sympatric

because they are usually separated by at least

1,000 feet in elevation. In an area about 36 miles

southwest of La Junta in Chihuahua, Mexico,

however, we found these two taxa growing together, but intermediates between them were not

evident.

This dwarf mistletoe causes open, nonsystemic

brooms similar to those induced by A. gillii subsp.

nigrum (see fig. 10).

ARCEUTHOBIUM GILLII SUBSP. NIGRUM

12b. Arceuthohium gillii subsp. nigrum Hawksworth

& Wiens, Brittonia 17: 233, 1965. HOLOTYPE:

MEXICO: Durango, 32 mi. E of El Salto on

Route 40, on P. teocote, Hawksworth & Wiens 304

in 1963 (COLO. Isotypes FPF, MEXU, MO,

US).

DESCRIPTION: Mean shoot height ca. 15-20 cm.,

but up to 45 cm. (fig. 57). Shoots dark brown to

black. Basal diameter of dominant shoots 3-8 mm.

(mean 5 mm.). Third internode 5-19 mm. (mean

10.8 ± 3.8 mm.) long and 2.5-6 mm. (mean 3.7

mm.) wide (6 collections), length/width ratio

2.9:1. Staminate ñowers ca. 3 mm. long and 3.5

mm. in diameter. Pollen poorly known and only a

few grains available for study in one collection:

polar diameter ca. 20^, equatorial diameter ca.

23M; spine height ca. 2.0/i, ca. twice wall thickness—l.Oju. Mature fruit 6-9 mm. (mean 7 mm.)

long and ca. 3.5 mm. wide, proximal portion ca.

2-3 mm. Seeds 3.5 X 1.3 mm. Meiosis in August

and September. Anthesis usually in March

or April. Seeds mature in September of the year

following pollination; maturation period averages

17-18 months, n = 14.

TABLE 21,—Comparison o/Arceuthobium gillii and A. vaginatum suhsp.

cryptopodum

Characters

Principal hosts

Shoot color

Fruits

A. gillii

A. vaginatum subsp.

cryptopodum

Peak flowering time

Seed maturation period

Pinus leiophylla var. chihuahuana

Greenish yellow

Proximal portion of fruit

bluish-white glaucous;

distal portion (including

perianth segments) accounting for ca. 40 percent of fruit length

February to early April

ca. 19 months

Pinus ponderosa vars. arizonica and scopulorumf P.

engelmannii

Yellow to orange

Fruit essentially nonglaucous;

distal portion (including

perianth segments) accounting for ca. 30 percent

of fruit length

May to June

14-15 months

128
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Figure 56.—Distribution of the two subspecies of Arceuthobium gillii (based on collections only).

HOSTS: Principally on pines of section Leiophyllae

{Pinus leiophylla vars. leiophylla and chihuahuana

and P. lumholtzii) and Teocote (P. teocote). Rare

on P. montezumae.

DISTRIBUTION: Presently known from the Mexican States of Durango, Hidalgo, Veracruz, and

northern Oaxca, but is to be expected in other adjacent States (see fig. 56). Known elevational

range is 7,400 to 8,800 feet. Specimens examined:

see appendix, p. 210.

DISCUSSION: This subspecies is distinguished

from subsp. gillii by its darker color, more southerly range, markedly divaricate staminate shoots,

and larger plants. Plants from Cofre de Perote in

Veracruz, tentatively referred to this subspecies,

are morphologically similar to subsp. nigrum from

other localities but somewhat lighter in color.

Witches' brooms caused by subsp. nigrum are

illustrated in figure 10.

ARCEUTHOBIUM GLOBOSUM

13. Arceuthobium glohosum Hawksworth & Wiens,

Brittonia 17: 223, 1965. HOLOTYPE: MEXICO:

DURANGO: 1.5 mi. E of El Salto on Route 40,

on Pinus cooperi, Hawksworth & Wiens 34-14 i^i

1963 (COLO, Isotypes DS, F, FPF, INIF, MEXU,

MO, US).

DESCRIPTION: Mean shoot height ca. 20-30 cm.,

but up to 70 cm. in Guatemala (fig. 58). Shoots

yellow to greenish, flabellately branched. Basal

diameter of dominant shoots 3-44 mm. (mean 12

mm.). Third internode 4-35 mm. (mean 20.5 dz

8.4 mm.) long and 2-23 mm. (mean 5.5 mm.)

wide (6 collections), length/width ratio 3.7:1.

Staminate flowers ca. 3.5 mm. long and 3-3.5

mm. in diameter; perianth 3- or 4-merous; same

color as shoots; segments ca. 1.3 mm. long and

1.0 mm. wide. Mean anther diameter 0.6 mm.,

129
Figure 57. Arceuthobium gillii subsp. nigrum in spring: A, Staminate shoot; B, Pistillate shoot.

130
Figure 58.—Arceuthobium globosum in March: A^ Staminate shoot; ß, Detail of staminate flower; C, Pistillate

shoot; D, Detail of fruit.

131
centered 0.9 mm. from tip of segment. Pollen

polar diameter 19-23 (mean 22ju); equatorial diameter 24-28 (mean 26/x) ; polar/equatorial diameter ratio 1:1.18; spine height (ca. 1.9fjL) approx.

twice wall thickness—ca. I.OJLí (4 collections).

Pistillate flowers ca. 1.5 mm. long and 1.5 mm. in

diameter; 2-merous. Mature fruit ca. 6-7 mm.

long, and ca. 4 mm. wide; proximal portion ca.

3.5 mm. long. Seeds 5X2 mm. Pedicels 4-5 mm.

long. Time of meiosis unknown. Anthesis usually

in ]\iarch-April. Seeds mature in June-July in

northern and central aMexico and September in

Guatemala; maturation period averages 15-16

months, n = unknown.

HOSTS: This species has the broadest known host

range of any of the dwarf mistletoes; it occurs on

at least 14 pines as well as Cupressus. Hawksworth

and Wiens (1965), in the original description of

the species, listed 11 hosts: Pinus cooperi, P.

durangensis, P. engelmannii, P. hartwegii, P. lawsonii, P. michoacana, P. montezumae, P. ponderosa

var. arizonica, P. pseudostrohus, P. rudis, and P.

tenuifolia, all from Mexico. Three additional hosts

have been noted in Michoacán, Mexico: Pinus

douglasiana, P. pringlei, and P. teocote, but the

dwarf mistletoe was not identified (Valdivia

1964). We believe the parasite is A. glohosum because: (1) we have examined many of Valdivia's

specimens, (2) his description fits that of this

species, and (3) this is the only species known from

Michoacán. Valdivia (1964) gives the hosts in

northeastern Michoacán, in decreasing order of

susceptibility, as: Pinus tenuifolia, P. pseudostrohus, P. pseudostrohus f. protuherans, P. montezumae, P. michoacana, P. durangensis, P. douglasiana, P. lawsonii, P. pringlei, P. teocote, and P.

rudis. Pinus leiophylla and Ahies religiosa were

recorded as immune.

Hunt (1962) reported a dwarf mistletoe (as ^'A.

vaginatum^') on Pinus carihaea from Mountain

Pine Ridge, British Honduras; we have examined

his specimens and determined that they are A.

glohosum. Etheridge (1968) noted that A. glohosum

was damaging in two localities on Mountain Pine

Ridge.

The hosts of A. glohosum in Guatemala are

poorly known because many collections recorded

the host only as Pinus sp. However, at least P.

pseudostrohus, P. tenuifolia, P. rudis, and Cupressus are infected there. This is the only known instance of Arceuthohium on Cupressus in the Western Hemisphere (Standley and Steyermark 1946).

Parasitism of Cupressus is apparently rare, however, because we did not observe infection of this

tree where it was associated with A. glohosum on

Pinus rudis in the Sierra Cuchumatanes of Guatemala.

DISTRIBUTION: This is one of the most widely

distributed dwarf mistletoes in ^Mexico and Guatemala, and is the only species known from British

Honduras (fig. 59). In ^léxico it occurs from

western Chihuahua and eastern Sonora south

through the Sierra Madre Occidental to Michoacán; east to Veracruz and south through Oaxaca

and Chiapas, where it crosses the border into the

highlands of Guatemala. It is known from a single

area in west-central British Honduras.

Arceuthohium glohosum is usually a high-altitude

species; in central Mexico it ranges from 10,000

feet to as high as 13,000 feet in Nevado de Toluca.

It occurs as low as about 3,000 feet in the Alaya

Mountains in British Honduras. Specimens examined: see appendix, p. 210.

DISCUSSION: One of the most distinctive dwarf

mistletoes, A. glohosum is characterized by globose

masses, 30-100 cm. in diameter, of bright yellow

to greenish shoots. The shoots on older plants may

exceed 4 cm. in diameter and 70 cm. in height.

The form of the species in the Sierra Cuchumatanes of Guatemala differs from the populations

in northern and central ^Mexico. For example, in

Guatemala the plants are larger and consistently

dark greenish yellow, while those in ^Mexico are

smaller and yellow. Guatemalan plants produce

witches' brooms; the ^lexican populations do not.

In Mexico, the peak seed dispersal period is apparently in June and early July, but in Guatemala, the fruits were just maturing in mid-September. In view of the differences between the

two population systems, subspecific separation

might be desirable if the differences are further

confirmed. Detailed field studies are also needed

in southern ^Mexico to determine whether the differences between central ?^Iexican and Guatemalan

populations are discontinuous.

Valdivia (1964), who surveyed a million acres

of pine forest in northeastern ^lichoacán, Mexico,

found A. glohosum on nearly 40 percent of the

forest area.

ARCEUTHOBIUM GUATEMALENSE

14. Arceuthohium guatemalense Hawksworth &

Weins, Brittonia 22: 267, 1970. HOLOTYPE:

GUATEMALA: DEBT. HUEHUETENANGO,

Sierra Cuchumatanes, on road from Huohuetenango to Santa Eulalia at Km 142 ; 54 km north

of Huohuotonango or 10 km south of San Juan

Ixcoy, on Pinus ayacahuite, Hawksworth & Weins

1221, Sept. 11, 1969. (US, Isotypos COLO, DS,

EAP, FPF, MO, UT).

DESCRIPTION: Mean shoot height ca. 1-3 cm.

on systemic brooms, but shoots on nonsystemic

infections up to ca. 7 cm. high (fig. 60); shoots

132
1 1 1 1

_J

+ 1 -h

,-'\^ u N 1 T E D

V ^• ? \^ \.. New Mexico

Figure 59.—Distribution of Arceuthobium globosum in Mexico, Guatemala, and British Honduras.

greenish to purple, yellow to brown when dried,

flabellately branched; basal diameter of dominant

shoots 2-2.5 mm. Third internode 8-15 (mean

11.4 ± 2.8 mm.) long and 1.5-2 (mean 1.7) mm.

wide; length/width ratio 6.7:1 (3 collections).

Staminate flowers ca. 2 mm. in diameter; perianth

2- or 3-merous, segments 0.9 mm. long and 0.7

mm. wide. Mean anther diameter 0.5 mm. Few

pollen grains available for study; mean polar diameter 21jLt and mean equatorial diameter 24jit;

polar/equatorial diameter ratio 1:1.14; spine

height—ca. 1.5/x, wall thickness—ca. 1.0/z (1 collection). Mature fruit ca. 3.7 mm. long and 1.8

mm. wide; distal portion ca. 1.2 mm. long; dark

green, nonglaucous, with a slightly swollen ring

at the base of the fruit where it joins the receptacle

of the pedicel (fig. 60). Seed 2.0 X 0.8 mm. Time

of meiosis unknown. Anthesis apparently in August and early September. Fruits mature in September so the fruit maturation period is from

12-13 months. Germination in September, n =

unknown.

HOSTS AND DISTRIBUTION: This parasite of

Pinus ayacahuite is known only from the high

mountains of Guatemala (Departments of Huehuetenango and Totonicipán), but it probably

also occurs on P. ayacahuite in southern Mexico.

We collected it in the Sierra Cuchumatanes, and

along the Pan American Highway between Nahualá and Quetzaltenango. It is quite common in the

vicinity of Santa Eulalia, north of Huehuetenango.

The elevational distribution of this species is

poorly known, but we observed it between 8,300

and 10,100 feet. No range map was prepared for

this species because its distribution is too poorly

known. Specimens cited: see appendix, p. 212.

DISCUSSION: The consistent formation of systemic witches^ brooms is a distinctive characteristic of this species. The brooms are sometimes

10 to 15 feet in diameter. An unusual feature of

the brooms is that mistletoe shoots are consistently

formed on host twigs as young as 1 year old and,

in some cases, we found buds on the current year's

growth (in September).

133
2mm

Figure 60,—Arceuthobium guatemalense in September: A, Staminate shoots; ß, Detail of staminate shoot; C,

Pistillate shoot; D, Detail of mature fruit showing typical enlargement at base.

134
This species is quite damaging and causes considerable mortality (see fig. IID).

ARCEUTHOBIUM HONDURENSE

15 Arceuthohium hondurense Hawksworth & Wiens,

Brittonia 22: 267, 1970. HOLOTYPE: HONDURAS: DEPT. FRANCISCO MORAZAN,

Piedra Herrada, 22 km. southeast of Tegucigalpa

on road to Escuela Agrícola Panamericana, on

Pinus oocarpa, Hawksworth, Wiens, and Molina

1203, Sept. 4, 1969 (US, Isotypes EAP, COLO,

DS, F, FPF, MO, UT).

DESCRIPTION: Mean shoot height ca. 14 cm.,

but up to 21 cm. (fig. 61) ; shoots olive brown to

grayish green, markedly glaucous; flabellately

branched; basal diameter of dominant shoots 3-9

(mean 5) mm.; nodes of older shoots swollen;

lateral branches of staminate plants at nearly

right angle to axis of the main shoot; third internode 7-12 (mean 9.1 dz 1.5 mm.) long and 2.5-4

(mean 3.2 mm.) wide; length/width ratio 6.1:1

(2 collections). Staminate flowers ca. 2.5 mm. in

diameter; inner surface reddish, lower surface the

same color as shoots; perianth usually 3- (sometimes 2- or 4-) merous, segments ca. 1.2 mm. long

or 0.8 mm. wide; nectary with 2 large lobes and

1 small lobe. Pollen polar diameter 18-23 (mean

20/x) ; equatorial diameter 21-25 (mean 24JLI) ; polar/

equatorial diameter ratio 1:1.21; spine height (ca.

2.5/X) about twice wall thickness—1.1 M (2 collections). Pistillate flowers with stigmas exserted ca.

0.5 mm., copious nectar secretion. Mature fruit

ca. 5.5 mm. long and 3.0 mm. wide, greenish

glaucous; proximal portion ca. 4.0 mm. long;

stigma exserted. Seeds ca. 3.1 X 1.5 mm. Meiosis

in August or early September. Anthesis and seed

maturation in September so the seed maturation

period averages ca. 12 months, n = 14.

HOSTS AND DISTRIBUTION: This species has

been collected only on Pinus oocarpa, although it

has been seen on Pinus pseudostrohus.^^ The distribution of this species is poorly known, and the

only three collections that have been made are

from the same general area near Piedra Herrada,

southeast of Tegucigalpa, Honduras, on the road

to Escuela Agrícola Panamericana. Roadside surveys totaling 140 miles through P. oocarpa stands,

mainly in the Departments of Francisco Morazán,

El Paraiso, and Comayagua, failed to reveal additional populations of the parasite. The indiscriminate cutting of the original forests has apparently

reduced the occurrence of A. hondurense in these

areas, which are now either agricultural lands or

30 Personal communication, Antonio Molina R., Escuela

Agrícola Panamericana, Tegucigalpa, Honduras, 1969.

second-growth forests. Unless the species occurs

in areas less subject to severe disturbance, A.

hondurense faces extinction. No range map was

prepared for this species because its distribution

is so poorly known. The few populations observed

were between 4,000 and 5,300 feet in elevation.

Specimens cited: see appendix, p. 212.

DISCUSSION: Originally we believed A. hondurense (see fig. 12) to be conspecific with A. hicarinatum, a species endemic to Hispaniola. Observations of living plants of both dwarf mistletoes,

however, demonstrated discontinuities in many

characters (see table 19). A. hondurense appears

most closely related to A. ruhrum and A. hicarinatum, which are slender, dark-colored species in

the southern range of the genus. In fact, A.

hicarinatum and A. hondurense, occupy the distributional limits of the genus in the southeastern

and southern parts of the New World, respectively.

ARCEUTHOBIUM LARICIS

LARCH DWARF MISTLETOE

16. Arceuthohium laricis (Piper) St. John, Flora SE

Wash. & Adj. Idaho: 115, 1937. LECTOTYPE:

WASHINGTON: Kittitas Co.: N of EUensburg,

on Larix occidentalis, Brandegee 1071 in 1883

(US! Isotypes GH, PH, UC). Razoumofskya

douglasii (Engelm.) Kuntze subsp. laricis Piper,

Contr. U. S. Nat. Herb. 11: 223, 1906, nomen

nudum. Arceuthohium douglasii Engelm. var.

laricis M. E. Jones, Univ. Mont. Bull. 61 (Biol.

Ser. 15): 25, 1910, nomen nudum. Razoumofskya

laricis Piper, in Piper and Beattie, Flora SE

Wash. & Adj. Idaho: 80, 1914. Arceuthohium

campylopodum Engelm. forma laricis (Piper) Gill,

Trans. Conn. Acad. Arts & Sei. 32: 202, 1935.

DESCRIPTION: Mean shoot height ca. 4 cm. but

up to 6 cm. (fig. 62). Shoots mostly dark purple,

flabellately branched. Basal diameter of dominant

shoots 1.5-3 mm. (mean 2 mm.). Third internode

5-14 mm. (mean 8.0 ± 2.0 mm.) long and 1-2.5

mm. (mean 1.3 mm.) wide (12 collections), length/

width ratio 6.1:1. Staminate flowers 2.7 mm. in

diameter; perianth mostly 3- (sometimes 4-)

merous; segments ca. 1.4 mm. long and 1.1 mm.

wide. Mean anther diameter 0.5 mm., centered

0.5 mm. from tip of segment. Pollen polar diameter 18-25 (mean 20M); equatorial diameter 21-27

(mean 23/i); polar/equatorial diameter ratio

1:1.16; spine height (ca. 2jLt) about twice wall

thickness—ca. 1.0M (3 collections). Pistillate

flowers ca. 1 mm. long and 1 mm. in diameter.

Mature fruit 4.5 mm. long and 2.5 mm. wide;

proximal portion ca. 2.5 mm. long. Meiosis in

June. Peak anthesis usuafly in early August, with

extremes from late July to early September. Seeds

135
3 mm

Figure 61-Arceuthobiuni hondurense in September: A, Staminate shoot; B, Detail of terminal portion of staminate shoot showing unopen flowers; C, Open staminate flower; D, Pistillate shoot; E, Detail of mature fruit.

136
B

1 cm

Figure 62,—Arceuthobium laricis in August: A, Staminate shoot in anthesis; B, Detail of staminate flowers; C,

Pistillate shoot; D, Nearly mature fruit.

137
usually mature in September, with extremes from

early August to early October. Maturation period

averages 13-14 months, n = 14.

HOSTS: The principal and only commonly infected host is Larix occidentalis. Other trees that

are sometimes infected when they grow in association with infected L. occidentalis are: Abies lasiocarpa, A. grandis, Picea engelmannii, Pinus alhicaulis, P. contorta, P. monticola, P. ponderosa, and

Tsuga mertensiana. The status of this mistletoe

on subalpine larch, Larix lyallii, is poorly known,

and only two early collections are available from

this host (by J. R. Weir, from the Bitterroot and

Cabinet Mountains of Montana). Exotic trees infected naturally include Pinus hanksiana (Graham

1959a), P. resinosa (U. S. Department of Agriculture 1962), P. sylvestris (Graham and Leaphart

1961), and Picea ahies (U. S. Department of Agriculture 1963).

Weir (1918a) has shown that, on the basis of

seedling inoculations, the European larch, Larix

europaea, and Japanese larch, L. leptolepis, are

susceptible to the larch dwarf mistletoe.

The occurrence of A. laricis on Tsuga mertensiana in Idaho and Montana (see fig. 13B) is of

interest, because this mistletoe will not infect T.

heterophylla. Our observations in many mixed

Larix-Tsuga stands in the Cascades in Oregon and

Washington, and in Idaho and Montana confirm

that T. heterophylla is not susceptible to A. laricis.

In mixed Tsuga heterophylla-T. mertensiana stands

in the the Coeur d' Alene National Forest, Idaho,

only the latter was infected. We do not know

whether A. laricis is associated with T. mertensiana

in the Cascades.

DISTRIBUTION: The larch dwarf mistletoe occurs generally throughout the range of its principal host, Larix occidentalis, in southern British

Columbia, east of the Cascades in Washington

and northern Oregon, north and central Idaho,

and western Montana (fig. 63). Known elevational

range is 2,300 to 6,500 feet.

Arceuthohium laricis extends to Salmon Arm

and north to Kootenay Lake,^^ which is somewhat

farther north than the range Kuijt (1963) gives

for this taxon. Specimens examined : see appendix,

p. 212.

DISCUSSION: A. laricis is a serious pathogen of

Larix occidentalis (Weir 1916a). For example, surveys show that about two-thirds of the western

larch stands are infested on the Coeur d^Alene

National Forest, Idaho, and in the Kootenai

National Forest, ]\Iontana (Graham 1959b, 1959c).

On the Colville National Forest and adjacent

private lands in northeastern Washington, infestation was 86 percent (Graham and Frazier

1962).

In nature, selective parasitism is the easiest

diagnostic feature that separates A. laricis from

A. tsugense: A. laricis infects Larix occidentalis

but not Tsuga heterophylla; A. tsugense attacks T.

heterophylla but not L. occidentalis. In greenhouse

and plantation inoculations, however, R. B. Smith

has infected Larix occidentalis with A. tsugense.^^

Arceuthohium tsugense is generally larger and more

profusely branched than A. laricis (Kuijt 1970).

Table 22 gives some other distinguishing features

for these two Northwestern taxa (see fig. 13).

ARCEUTHOBIUM MICROCARPUM

WESTERN SPRUCE DWARF MISTLETOE

17. Arceuthohium microcarpum (^ngQ\vû.2inii) Hawksworth & Wiens, Brittonia 22: 268, 1970. LECTOTYPE: ARIZONA: Apache Co.: Sierra Blanca,

^^ Personal communication, Dr. R. B. Smith, Canadian

Dep. Fisheries and Forestry, 1967.

TABLE 22.—Comparison of Arceuthohium laricis and A. tsugense

Characters A. laricis A. tsugense

Principal hosts

Shoots:

Base of older shoots

Terminal internodes

on main axis of

staminate shoots

Length of lateral staminate spikes in summer

Staminate flowers

Peak flowering time

Peak seed dispersal time

Larix

Angular

Ca. 2 mm. long

2-3 mm.

Rarely 4-merous

August

September

Tsuga

Rounded

Over 3 mm. long

5-6 mm.

Sometimes 4-merous

September

October

138
Figure 63.—Distribution of Arceuthobium. lands on all hosts in the United States and in British Columbia, Canada

(based on collections and reports). Also shown is the range of Larix occidentalis. Outline map copyrighted by

Denoyer-Geppert Co., Chicago (used by permission).

on Picea engelmannii, Gilbert 112 in 1873 (MO!

Isotype ILL, \j^). Arceuthobiumdouglasii'EngelTa..

var.? microcarpum Engelm. in U.S. Geogr. Surv.

W of 100th Meridian (Wheeler Rep.) 6: 253, 1878.

Razoumofskya microcarpa (Engelm.) Wooton &

Standley, Contr. U.S. Nat. Herb. 19: 179, 1915.

Razoumofskya douglasii (Engelm.) Kuntze var.

microcarpa (Engelm.) Tubeuf, Naturw. Zeitschr.

Forst, u. Landw. 17: 216, 1919. Arceuthobium

campylopodum Engelm. forma microcarpum

(Engelm.) Gill, Trans. Conn. Acad. Arts & Sei. 32:

209, 1935.

DESCRIPTION: Mean shoot height ca. 5 cm.,

but up to 10 cm. (fig. 64). Shoots green to purple.

139
Figure 64.—Arceuthobium microcarpuní in August: A, Staminate shoot; B, Pistillate shoot; C, Detail of young

pistillate flower; D, Nearly mature fruit.

140
flabellately branched. Basal diameter of dominant

shoots 1.5-3 mm. (mean 2 mm.). Third internode

5-16 mm. (mean 9.3 ± 2.2 mm.) long and 1-2 mm.

(mean 1.5 mm.) wide (9 collections), length/width

ratio 6.2:1. Staminate flowers 2.3 mm. in diameter;

perianth mostly 3-, rarely 4-merous; segments

ca. 1.2 mm. long and 1.0 mm. wide. Mean anther

diameter 0.4 mm., centered 0.6 mm. from tip of

segment. Pollen polar diameter 18-21 (mean 19M);

equatorial diameter 21-23 (mean 22/z); polar/

equatorial diameter ratio 1:1.16; spine height

(1.2M) approximately equal to wall thickness—1.4^

(1 collection). Pistillate flowers ca. 1 mm. long and

1 mm. in diameter. Mature fruit 3.5 mm. long and

2.0 mm. wide; proximal portion ca. 2.5 mm. long.

Seeds 2.4 X 1.0 mm. Meiosis in July. Anthesis

usually in late August or early September. Seeds

mature in September. Maturation period averages

12 to 13 months, n = 14.

HOSTS: Common and serious only on Picea

engelmannii and P. pungens. Gill (1935), who

collected one specimen on Ahies lasiocarpa var.

arizonica near Columbine Ranger Station in the

Graham Mountains of Arizona, noted that A.

microcarpum rarely infected this host; our searches

in the same area failed to reveal any infection on

Ahies, although the parasite was common on Picea.

Also, we have examined Ahies in many other

stands where A, microcarpum is abundant, but

have found no infection on it.

DISTRIBUTION : This dwarf mistletoe, which has

been found only in Arizona and New Mexico

(fig. 65), has one of the smallest ranges of any

species of Arceuthohium. In Arizona, the parasite

has been observed in the Apache National Forest,

the Graham Mountains, the North Rim of Grand

Canyon, and the San Francisco Peaks and nearby

Kendrick Peak,^^ ^^t not in the Chiricahua

Mountains.22 j^ New Mexico, the only infested

area known is in the Mogollón Mountains,

although we suspect the parasite is more widely

distributed in the mountains of southwestern New

Mexico. We found no spruce dwarf mistletoe,

however, in northern, central, and south-central

New Mexico, or in the commercially valuable

spruce forests from southern Utah and Colorado

northward. Known elevational range is 7,900 to

10,000 feet. Specimens examined: see appendix,

p. 213.

Apparently, the rare Picea chihuahuana is not

parasitized, because no infections have been

reported from the Sierra Madre Occidental of

Chihuahua or Durango, Mexico. Gordon (1968)

did not observe the parasite in his ecological study

of the tree near El Salto, Durango.^^ If P. chihuahuana is parasitized by dwarf mistletoe, A.

microcarpum would be the most likely taxon.

DISCUSSION: The original specimens cited by

Engelmann are Gilbert 100 and 102. The only

specimen now at the Missouri Botanic Garden is

Gilhert 112 which is labeled in Engelmann's handwriting as the type. The original label for specimen 100 is in the same packet. Presumably,

Gilhert 102 in the original citation should have

been 112. In any case, the collections are now

combined so Gilhert 112 is our choice for a

lectotype.

This localized southwestern species is characterized by its almost exclusive occurrence on

Picea. Gill (1935) gave a much larger distribution

for this taxon (to California, Idaho, and Montana).

Hawksworth and Graham (1963), however, have

shown that GilPs records are based on occasional

parasitism of spruce by other dwarf mistletoes

{A. laricis. A. douglasii, A. americanum, and

possibly A. tsugense and A. cyanocarpum).

This species induces small, dense witches'

brooms (see fig. 9). Heavily infected trees may

bear hundreds of brooms. Although this dwarf

mistletoe causes heavy mortality in Picea engelmannii and P. pungens in Arizona and New

Mexico, the spruces constitute only a minor part

of the total forest volume. The dwarf mistletoe

is not found in the main commercial range of

P. engelmannii.

ARCEUTHOBIUM OCCIDENTALE

DIGGER PINE DWARF MISTLETOE

18. Arceuthohium occidentale Engelmann, U.S. Geogr.

Surv. W of 100th Meridian (Wheeler Rep.) 6: 375,

1878 and Bot. Calif. 2: 107, 1880. LECTOTYPE:

CALIFORNIA: Kern Co.; Walker's Basin, on

Pinus sahiniana, Rothrock 429 in 1875 (MO!

Isotype US). Razoumofskya occidentale (Engelm.)

Kuntze, Rev. Gen. PI. 2: 587, 1891.

DESCRIPTION: Mean shoot height ca. 8 cm.,

but up to 17 cm. (fig. 66). Shoots yellowish,

glaucous, flabellately branched. Basal diameter of

dominant shoots 1.5-5 mm. (mean 2 mm.). Third

internode 7-18 mm. (mean 12.7 ± 2.0 mm.)

long and 1.5-3.5 mm. (mean 1.8 mm.) wide (11

collections), length/width ratio 7.1:1. Staminate

flowers 3.0 mm. in diameter; perianth 3- or 4-

merous; segments ca. 1.5 mm. long and 1.0 mm.

wide. Mean anther diameter 0.5 mm., centered

0.6 mm. from tip of segment. Pollen polar diameter

18-23 (mean 20M); equatorial diameter 21-25

32 Personal communication, Dr. John R. Jones, U.S. Forest

Service, 1968.

33 Personal communication, Dr. A. G. Gordon, Canada Dep.

Fisheries and Forestry, 1969.

141
Figure 65.—Distribution of Arceuthobium microcarpiim. Outline map copyrighted by Denoyer-Geppert Co.,

Chicago (used by permission).

142
Figure 66,—Arceuthobium occidentale in March: A, Staminate shoot: B, Remnant of staminate flower; C, Pistillate

shoot; D, Young fertilized fruit.

143
(mean 23yii); polar/equatorial diameter ratio

1:1.16; spine height (ca. 2ju) approximately twice

wall thickness—1.1M (3 collections). Mature fruit

4.5 mm. long and 3.0 mm. wide; proximal portion

ca. 3.0 mm. long. Seeds 2.6 X 1.0 mm. Meiosis

in August. Peak anthesis usually during late

October, with extremes from early September to

late November. Seeds usually mature in November, with extremes from early October to January;

maturation period averages ca. 13 months.

n = 14.

HOSTS: Principal hosts are Pinus sabiniana, P.

radiata, and P. tnuricata. Pinus coulteri and P.

attenuata are sometimes infected in the coastal

mountains of central California. Pinus contorta

var. bolanderi is occasionally attacked where this

tree is associated with infected P. muricata.

A. occidentale will not infect P. ponderosa, although

it is commonly associated with this tree on P.

sabiniana in the Sierra Nevada.

Hedgcock and Hunt (1917) successfully inoculated A. occidentale on seedlings of Pinus banksiana, P. bungeana, P. caribaea, P. pinea, and P.

virginiana as did Kuijt (1960a) for P. torreyana.

DISTRIBUTION: This dwarf mistletoe occurs

throughout the foothills and low mountains

surrounding the Central Valley of California and

along the coastal mountains from the Mount

Pinos area (Ventura County) north to Mendocino

County (fig. 67). It is found in two of the three

populations of Pinus radiata: Monterey and Cambria but not Swanton (Offord 1964a). One of the

few dwarf mistletoes to become established outside

its natural range, A. occidentale has been collected

on P. radiata at Stanford University (Peirce 1905)

and at Berkeley, California (Offord 1964b).

In both cases, the source of infection presumably

was diseased trees transplanted from the Monterey

area.

This mistletoe ranges from near sea level on P.

radiata and P. muricata to an elevation of about

4,700 feet in the southern Sierra Nevada on P.

sabiniana. Specimens examined: see appendix, p.

214.

DISCUSSION: Engelmann's 1878 report specifically

lists only the specimen Rothrock 1^29 on Pinus

sabiniana, but also mentions that it occurs on P.

ponderosa. No specimens are cited in Engelmann's

1880 report. Engelmann thus considered A.

occidentale to include both the parasite on Pinus

ponderosa (our A. campylopodum) and P. sabiniana

(our A. occidentale). Engelmann's writings are not

clear as to what he considered to be the relationship of A. occidentale to his previously described A.

campylopodum (in Gray 1850). Traditionally, the

name A. campylopodum has been associated with

Figure 67.—Distribution of Arceuthobium occidentale

in California. Outline map copyrighted by DenoyerGeppert Co., Chicago (used by permission).

the ponderosa pine parasite, and we maintain this

usage.

Several characteristics that distinguish A.

occidentale from A. campylopodum (see table 20)

are discussed under the latter taxon. Also, the type

of witches' brooms formed differs considerably.

Frequently, A. occidentale does not form witches'

brooms in Pinus sabiniana (see fig. 8c), but those

that are formed are small and open compared with

the large, dense brooms that A. campylopodum

causes on P. ponderosa. Witches' brooms are more

144
frequent in Pinus radiata (Offord 1964a) and P.

muricata than in P. sahiniana. Swellings at the

point of infection are particularly conspicuous on

P. radiata.

Shoot color in A. occidentale varies with the

species of host parasitized: shoots are light brown

to straw colored on P. sahiniana, olive green on

P. radiata, and dark brown on P. muricata. These

color differences may be due to hosts, or to local

populations of the parasite in areas where the

three main hosts occur.

Populations of A. occidentale on its three principal hosts {Pinus sahiniana, P. radiata, and P.

muricata) are usually geographically separated,

although there are a few instances of cross infection between them (see fig. 46). Pinus muricata

is attacked by A. occidentale on Huckleberry Hill

near Monterey where the parasite is common on

P. radiata. However, infection in P. muricata is

most common in the north coastal area of California where P. contorta var. holanderi is the only

other infected tree. Pinus radiata and P. sahiniana do not occur together naturally, but P.

radiata has been planted within the range of P.

sahiniana. An example is at Mount Hamilton

Observatory where planted P. radiata near mistletoe-infected P. sahiniana was also generally

infected. Apparently Pinus muricata and P.

sahiniana are not sympatric.

Scharpf (1969a) successfully inoculated A.

occidentale from Pinus sahiniana on P. radiata;

Parmeter and Cobb (1967) grew this mistletoe

from P. muricata on P. radiata.

Even though Pinus sahiniana typically occurs

in very open forests, A. occidentale is widely

distributed in these stands. The distribution

pattern suggests that vectors (possibly birds),

in addition to explosive seed dispersal, are important in disseminating this mistletoe.

ARCEUTHOBIUM PUSILLUM

EASTERN DWARF MISTLETOE

19. Arceuthohium pusillum Peck, Trans. Albany

Inst. 7: 191, 1872. HOLOTYPE: NEW YORK:

Rensselaer Co.: Sandlake, on Picea mariana.

Peck in 1871 (Isotype MO!) Arceuthohium

minutum Engelmann, Bull. Torrey Bot. Club 2:

43, 1871, nomen nudum. Arceuthohium oxycedri

(DC.) M. Bieb. var. ahigenium Wood, Amer.

Botanist and Florist: 446, 1871. Arceuthohium

ahigenium Wood, Class-Book of Botany : 832,1881.

Razoumofskya pusilla (Peck) Kuntze, Rev. Gen.

PI. 2: 587, 1891. Razoumofskya minuta (Engelm.)

Kuntze, Loc. Cit.

DESCRIPTION: Mean shoot height ca. 1 cm., but

up to 3 cm. (fig. 68). Shoots green to brown, without secondary branching. Basal diameter of

dominant shoots ca. 1.0 mm. Third internode 1-4

mm. (mean 1.9 zt 0.8 mm.) long and 0.5-1.5 mm.

(mean 1.0 mm.) wide (17 collections), length/

width ratio 1.9:1; segments often markedly wider

at top than at base. Pistillate shoots often longer

than staminate. Staminate flowers 1.7-2.2 mm.

(mean 1.8 mm.) in diameter; perianth mostly

3-merous, sometimes 2- or 4-; segments ca. 0.8 mm.

long and 0.7 mm. Avide. Mean anther diameter

0.4 mm., centered 0.5 mm. from tip of segment.

Pollen polar diameter 19-25 (mean 22^); equatorial diameter 23-28 (mean 26/x) polar/equatorial

diameter ratio 1:1.16; spine height (ca. 2.2/x) ca.

1.5 times wall thickness—1.4^t (4 collections).

Mature fruit ca. 3.0 mm. long and 1.25-1.75 mm.

wide (mean 1.5 mm.); proximal portion ca. 2.0

mm. long. Seeds 2.0 X 0.9 mm. Staminate

meiosis in September and pistillate meiosis in May

(Tainter 1968). Anthesis usually in April or May,

with extremes from late March to June (Fernald

1950, Tainter 1968). Seeds mature in September

or early October of the same year as pollination;

maturation period averages about 5 months.

Germination mostly in May and June (Tainter

1968). n = 14.

HOSTS: Arceuthohium pusillum is mainly parasitic

on Picea mariana, and to a somewhat lesser

extent on P. glauca; its occurrence on P. glauca

is somewhat variable. This tree is heavily attacked

along the Maine Coast (Nash 1955), and in the

pure P. glauca stands near Glenboro, Manitoba.^^

Occasional hosts are Picea ruhens and Larix

laricina (Kliejunas 1969; Tainter and French 1967,

1968). Rare hosts are Picea pungens (U.S. Department of Agriculture 1960), Pinus hanksiana

(Laut 1967; Pomerleau 1942; Sippell, Rose, and

Larsen 1968), Pinus resinosa (Krotkov 1940, Kuijt

1955), and Pinus strohus (House 1935).

DISTRIBUTION: Eastern dwarf mistletoe occurs

in Canada from the Hudson Bay and Cumberland

areas in eastern Saskatchewan to southern Manitoba, southern Ontario, Quebec, the Maritime

Provinces and Newfoundland. In the United States

it is found in the northern parts of Minnesota,

Wisconsin, and Michigan, northeastern Pennsylvania, extreme northwestern New Jersey and all

six New England States (figs. 69, 70, 71). Distribution maps for A. pusillum have been prepared for

New England (Eaton 1931, Eaton and Dow 1940);

Maine (Brower 1960); Minnesota (Tainter and

French 1969a) ; New Brunswick, Nova Scotia, and

Prince Edward Island (Forbes, Underwood, and

Van Sickle 1968) ; and for Manitoba and Saskatchewan (Kuijt 1963, Zalasky 1956). In Ontario and

Quebec the northern limits of this mistletoe are

^"^ Personal communication, J. G. Laut, Canada Dep. of

Fisheries and Forestry, 1968.

145
5 mm

Figure 68.—Arceuthobium pusillum: A^ Staminate plant with mature flowers, May; ß, Pistillate plant with immature fruit, midsummer; C, Pistillate plant with mature fruit, September. Drawn from photographs provided

by F. H. Tainter, University of Minnesota.

146
Figure 69.—General distribution of Arceuthobium pusillum. Outline map copyrighted by DenoyerGeppert Co., Chicago (used by permission).

147
poorly known; also, the report from White Cloud

in West Central Michigan (Martin 1926) should

be verified, because no other collections are

known within about 50 miles of this locality. This

dwarf mistletoe ranges from sea level in Maine and

the Maritime Provinces to 2,700 feet on Mount

Katahdin, Maine (Brower 1960). Specimens

examined: see appendix, p. 216.

DISCUSSION: Woodys (1871) variety ahigenium

was based on Peck's specimen from Sandlake,

New York. Since it was proposed as a variety,

it cannot replace Peck's (1872) validly published

specific name pusillum.

The discovery of A. pusillum at Warrensburg,

New York, in 1871 triggered a flurry of activity

which eventually showed the parasite to be

widely distributed in the spruce forests of the Lake

States, New England, and southern Canada. The

plant, no longer considered a curiosity, is recognized as an important damaging agent of spruce in

many parts of its range. For example, mortality

is severe in Picea glauca along the Maine Coast

(Nash 1955), and the parasite is considered to be

the most serious disease agent of Picea mariana

in the Lake States (Beckwith and Anderson 1956).

Eastern dwarf mistletoe shoots average about 1

cm. high. Only A. minutissiynum of the Himalayas

has smaller shoots.

The witches' brooms induced by A. pusillum

have been the subject of several investigations.^^

Witches' brooms appear to be mostly of the

systemic type (see fig. 14) ; thus shoot formation is

regular and progressive. Shoots usually first appear

in late summer or autumn as small eruptions in the

bark of twigs two seasons old; during the third

season the shoots mature; they flower the following

spring; the fruits mature the next autumn. Thus,

shoots with mature fruits usually occur only on

4-year-old host growth. Thomas (1954) stated that

the type of brooms formed on Picea mariana is

dense and dark, while that on P. glauca is light and

loose. The brooms he observed possibly were in

different stages of development, because Anderson

and Kaufert (1959), in a study of the parasite on

P. mariana showed that brooms grow rapidly for

the first 5 to 10 years and then abruptly decline

in vigor.

Brower (1960) and Nash (1955) suggest that the

dwarf mistletoe on Picea glauca along the Maine

Coast is distinct from the inland mistletoe characteristically found in spruce bogs. We have not

studied this possibility in the field, but the herbarium specimens we examined from hosts on the

two sites showed no morphological differences.

35 Von Schrenk 1900, Gill 1935, Pomerleau 1942, Anderson

and Kaufert 1959, Kuijt 1960b.

Although P. glauca appears to be more severely

infested along the Maine Coast, the occurrence of

A. pusillum on this host is not unique to this area.

It occurs commonly on P. glauca in the Mackinac

Straits area of Michigan and in the Sprucewoods

area of southwestern Manitoba, and sporadically

on this host throughout the range of the parasite

(see specimens cited).

That A. pusillum occurs in P. mariana bogs is

well known, but ecology of the parasite has not

been adequately investigated. Only complete

studies will help explain the unusual distribution

patterns reported for A. pusillum: In Quebec, it is

restricted to within 1 mile of lakes or rivers

(Bernard 1957) ; in Maine on P. glauca to within Ji

mile of the Coast (Nash 1955). If, as Bonga (1969a)

suggests on the basis of New Brunswick studies, A.

pusillum requires an uninterrupted period of high

atmospheric humidity in the spring, its absence in

inland-upland sites except bogs might be explained.

When Eaton (1931) noted an 80- to 100-mile

gap in the distribution of A. pusillum in southern

New England, only one collection was known between central Vermont and New Hampshire and

central Massachusetts. Later, A. pusillum was

recorded from two other localities, and, since the

mistletoe also occurs in New York just west of the

Massachusetts-Vermont boundary (fig. 70), the

gap is probably more apparent than real (Eaton

and Dow 1940).

Several islands where A. pusillum infections

occur are isolated 20 miles or more from the

parasite's main range; for example, Monhegan

Island off the Maine Coast and Beaver Island in

Lake Michigan.

ARCEUTHOBIUM RUBRUM

20. Arceuihohium ruhrum Hawksworth & Wiens,

Brittonia 17: 233, 1965. HOLOTYPE: MEXICO:

DURANGO: 29 mi. E of El Salto on Route 40,

on Pinus teocote, Hawksworth & Wiens 3490 in

1963. (COLO, Isotypes FPF, INIF, MEXU, MO,

US).

DESCRIPTION: Mean shoot height ca. 10 cm.,

but up to 15 cm. (fig. 72). Shoots reddish brown,

flabellately branched. Staminate plants usually

taller than pistillate plants. Basal diameter of

dominant shoots 2-4 mm. (mean 2.4 mm.). Third

internode 4-12 mm. (mean 6.9 ± 2.7 mm.) long

and 2-3 mm. (mean ca. 2.0 mm.) wide (5 collections), length/width ratio 3.4:1. Staminate flowers

1.0-1.5 mm. in diameter; mostly 3-merous; segments ca. 0.6 mm. long and 0.6 mm. wide. Pollen

polar diameter 23-26 (mean 24^1) ; equatorial diameter ratio 1:1.05; spine height (ca. LO/x) slightly

148
A. pusi Hum

Figure 70.-Distribution of Arceuthohiumpusillx^m in Eastern North America (based on collections and reports).

The dashed line shows the approximate southern limits of ?icea mariana.

greater than wall thickness—0.8M (1 collection).

Mature fruit ca. 3.5 mm. long and 2.0 mm. wide.

Seeds 2.0 X 1.0 mm. Meiosis in June? Anthesis

usually in July. Seeds mature in mid-July to

August of the year following pollination; maturation period averages 12-13 months, n = unknown.

HOSTS: Known on Finu^ teocote, P. cooperi, P.

engelmannii, P. herrerai and P. durangensis, all

of which appear to be highly susceptible.

DISTRIBUTION: This rare species is known only

from the Sierra Madre Occidental in Durango and

Sinaloa, Mexico. No range map was prepared for

this taxon because its distribution is so poorly

known. Known elevational range is 8,200 to 8,900

feet. Specimens examined: see appendix, p. 219.

DISCUSSION: This distinctive slender, red mistletoe is apparently widespread in the mountains of

Durango (see fig. 10 B). As the plants dry, red

turns to dull brown, and the apical portion of each

segment becomes golden yellow, which gives

dried specimens a characteristic banded appearance. The shiny fruits, a unique character that

readily distinguishes this species from all other

dwarf mistletoes, remain shiny when dried but the

bright red color turns dark brown.

Another distinctive characteristic of this taxon

is the very small, scarcely opened staminate

flowers. In all July collections examined, the

staminate flowers were still not expanded, although

they were past maturity and nearly all pollen

had been shed. Instead, the perianth segments

formed a nearly closed cup with only a slight

separation between the segments. The plants must

be examined closely to tell whether they are

undergoing anthesis.

ARCEUTHOBIUM STRICTUM

21 Arceuthobium stridum Hawksworth & Wiens,

Brittonia 17: 234, 1965. HOLOTYPE: MEXICO:

DURANGO: 16 mi. S of city of Durango on road

to La Flor, on Pinus leiophylla var. chihuahuana,

Hawksworth & Wiens 3465 in 1963 (COLO,

Isotypes DS, FPF, INIF, MEXU, MO, US).

DESCRIPTION: Mean shoot height ca. 7 cm., but

up to 13 cm. (fig. 73). Pistillate shoots generally a

149
tsAao\^

DaKota

\~Ä\^^^'

so^o

A, pusi Hum

\]g\sc'pos'^^ \J\\C:^NO^^

Figure 71.—Distribution of Arceuthohiunt pusillum in central Canada and the Lake States (based on collections

and reports). The dashed line indicates the approximate southern limits of spruce.

greenish yellow brown, flabellately branched.

Staminate shoots brownish, seldom branching.

Pistillate plants usually taller than staminate

plants. Basal diameter of dominant shoots 2.5-4

mm. (mean 3.1 mm.). Third internode 1-8 mm.

(mean 3.6 i 2.0 mm.) long and 1.5-3.5 mm.

(mean 2.3 mm.) wide (5 collections), length/width

ratio 1.6:1. Staminate flowers ca. 3 mm. in

diameter, perianth 3-, 4- or 5-, rarely 6- or 7-

merous (Kuijt 1970); segments ca. 1.5 mm. long

and 1 mm. wide. Pollen polar diameter 18-19

(mean 18M); equatorial diameter 20-25 (mean

22/x); polar/equatorial diameter ratio 1:1.19;

spine height ca. l^u, approximately equal to wall

thickness (1 collection). Mature fruit ca. 4 mm.

long and ca. 2.5 mm. wide. Seeds 2.5 mm. X 1.0

mm. Meiosis in July. Anthesis from late July to

October, with peak in September. Seeds mature

from mid-September to October of the year

following pollination; maturation period averages

ca. 13 months, n = 14.

HOSTS: Chihuahua pine, Pinus leiophylla var.

chihuahuana is the principal host, P. teocote an

occasional host, and P. engelmannii a rare host.

DISTRIBUTION: This rare dwarf mistletoe is

known only in the Sierra Madre Occidental south

and west of Durango, State of Durango, Mexico.

No range map for this species was prepared

because its distribution is too poorly known.

Elevational range is 7,300 to 8,200 feet. Specimens

examined: see appendix, p. 219.

DISCUSSION: The most distinctive feature of this

mistletoe is its branching pattern (Hawksworth

and Wiens 1965, Kuijt 1970). The staminate

shoots seldom branch; instead they may become

single spikes 4 to 6 cm. long (see fig. 73). The

pistillate plants, however, have abundant secondary branching with dense clusters of small, yellowgreen fruits. This species is damaging to Pinus

leiophylla var. chihuahuana (see fig. 10).

150
I—I p

2mm w

Figure 12.-Arceuthobium rubrunt in July: A, Staminate shoot; B, Pistillate shoot; C, Detail of staminate flower.

151
Figure 73-Arceuthobium strictum in July: A, Staminate plant showing characteristic lack of branching in the

annual growth segments; B, Pistillate plant. urantning m tne

152
ARCEUTHOBIUM TSUGENSE

HEMLOCK DWARF MISTLETOE

22. Arceuthohium tsugense (Rosendahl) G. N. Jones,

Univ. Wash. Publ. Biol. 5: 139, 1936 (as A.

tsugensis). HOLOTYPE: CANADA: BRITISH

COLUMBIA: Vancouver Island, Port Renfrew

on Tsuga heterophylla, Rosendahl 826 in 1902,

(MIN. Isotypes COLO, DS, FPF, ILL, MO, RM).

Razoumofskya tsugensis Rosendahl, Minn. Bot.

Stud. 3: 272, 1903. Razoumofskya douglasii

tsugensis (Rosendahl) Piper, Contr. U.S. Nat.

Herb. 11: 222, 1906. Arceuthohium douglasii

Engelm. var. tsugensis (Rosendahl) M. E. Jones,

Univ. Mont. Bull. 61 (Biol. Ser. 15): 25, 1910.

Arceuthohium camplyopodum Engelm. forma tsugensis (Rosendahl) Gill, Trans. Conn. Acad. Arts

& Sei. 32:200,1935.

DESCRIPTION: Mean shoot height ca. 5 cm.,

but up to 13 cm. (fig. 74). Shoots greenish to

reddish, darker in winter, ñabellately branched.

Basal diameter of dominant shoots 1.5-4 mm.

(mean 2.0 mm.). Third internode 4-16 mm.

(mean 9.2 i 2.5 mm.) long and 1-2 mm. (mean

1.5 mm.) wade (15 collections), length/width ratio

6.1:1. Staminate flowers 2.8 mm. in diameter;

perianth 3- or 4-merous, segments ca. 1.2 mm.

long and 1.0 mm. wide. Mean anther diamter 0.5

mm., centered 0.6 mm. from tip of segment. Pollen polar diameter 19-21 (mean 20/x); equatorial

diameter 23-25 (mean 24/x); polar/equatorial

diameter ratio 1:1.19; spine height (2.4/x) approximately 3 times wall thickness—O.S/x (4 collections).

Pistillate flowers ca. 1 mm. long and 1 mm. in

diameter. Mature fruit 3 mm. long and 2 mm.

wide; proximal portion ca. 2.0 mm. long. Meiosis

in July. Peak anthesis usuaUy in late August, with

extremes from early August to late September, or

even December in British Columbia (Baranyay

1962). Seeds mature from late September to early

November (Smith 1966a). Maturation period

averages ca. 13-14 months. Germination usually

from February to May in British Columbia (Smith

1966b). n = 14.

HOSTS: The hemlocks, Tsuga heterophylla and

T. mertensiana, are the principal hosts, although

several other trees are attacked, particularly when

they grow in association with infected hemlocks:

Ahies lasiocarpa, A. amahilis, A. grandis, A.

procera, Pinus contorta subsp. contorta, P. alhicaulis, and P. monticola. A. tsugense is fairly

common on Ahies amahilis; many reports from

Washington should show A. tsugense as the parasite on this host rather than A. ahietinum. Very

rarely A. tsugense is found on Picea sitchensis in

Alaska (Laurent 1966) and in British Columbia

(Molnar, Harris, Ross, and Ginns 1968). Also, we

found it on P. engelmannii in the Oregon Cascades.

Arceuthohium tsugense rarely occurs on Picea hreweriana in northern California.

Along the south coastal areas of British Columbia and on Oreas Island, Washington, a dwarf

mistletoe is common on Pinus contorta subsp.

contorta and on P. monticola. We classify this as A.

tsugense although, as mentioned by Kuijt (1956),

the pines are often infected in areas where there is

little or no current infection on Tsuga.

Eastern hemlock, Tsuga canadensis, was shown

to be susceptible on the basis of seedling inoculations (Weir 1918a). Larix europea became infected

when planted under infected Tsuga heterophylla

on Vancouver Island, British Columbia (Kuijt

1964), but it apparently was not a compatible host

because swellings formed but no shoots developed.

On the basis of artificial inoculations of potted

seedlings. Smith (1965) showed the following as

susceptible to A. tsugense; Tsuga heterophylla

from south coastal and interior British Columbia,

Pinus contorta subsp. latifolia from Montana,

Picea glauca from central British Columbia, and

Picea ahies from Europe. Smith has also successfully infected Picea engelmannii (1970b) and, with

difficulty, Larix occidentalis (1970a). In the latter

host-parasite combination, most infections resulted

only in swellings, but in one instance a few shortlived shoots were produced. Of the several mixed

Larix occidentalis-Tsuga heterophylla stands in the

Oregon Cascades we examined, where Tsuga grew

in close association with A. laricis on Larix, we

found no infected Tsuga. Based on greenhouse and

plantation inoculations in British Columbia, Pinus

ponderosa, P. radiata, and P. sylvestris were susceptible to A. tsugense (Smith and Craig 1968).

DISTRIBUTION: This dwarf mistletoe ranges

from near Haines, Alaska^^ to central California

(figs. 75, 76, 77). It is found throughout the coastal

hemlock forests of Alaska, British Columbia^^ (Kuijt

1963), and Washington. In Oregon, A. tsugense is

common on the west slope of the Cascades and in

the coastal mountains, and occurs in California at

least as far south as Alpine County in the Sierra

Nevada and along the coast to Mendocino County.

This taxon apparently is widespread in the coastal

mountains of Oregon and California, although

few specimens have been collected in these areas.

Hulten (1968) mentions that this plant is '^much

overlooked;^^ this observation is emphasized by his

distribution map of A. tsugense in southeastern

Alaska, which indicates only two Alaska localities.

^^ Personal communication, T. W. Laurent, U.S. Forest

Service, 1965.

2^ Personal communication, Dr. R. B. Smith, Canadian Dep.

Fisheries and Forestry, 1969.

153
Figure 74.—Arceuthobium tsugense in August: A, Pistillate plant; ß, Staminate plant; C, Immature staminate

flower; D, Nearly mature fruit.

154
Our field investigations did not confirm the

reports of A. tsugense in Idaho. Although Weir

(1916c) reports and illustrates infected Tsuga

heterophylla on Marble Creek on the St. Joe

National Forest, the host probably is T. mertensiana.^^ In intensive mistletoe surveys in northern

Idaho, western Montana, and eastern Washington,

D. P. Graham^^ found no infection on T. heterophylla in Marble Creek or elsewhere in this region.

Graham (19o9b) reports mistletoe on Tsuga

mertensiana on the Coeur d'Alene National Forest

and our studies there in 1966 identified the parasite

involved as A. laricis (see fig. 13c). Tsuga heterophylla was not infected even though several

individuals grew in close association with infected

T. mertensiana, Larix occidentalism and Ahies

lasiocarpa.

The specimens that Weir collected in Idaho,

reportedly on Tsuga heterophylla (see specimens

cited), are so fragmentary that specific identification of the mistletoe is impossible. Thus, whether

A. tsugense does attack T. heterophylla in Idaho

will be in doubt until populations of the parasite

can be rediscovered and studied. INIarble Creek

and nearby areas have been extensively logged

during the 50 years since Weir recorded his observations, and it is remotely possible that the

parasite has been eradicated. Although cutting the

timber might be expected to reduce the amount of

mistletoe in a stand, we know of no instances

where logging has eliminated a species.

The known elevational range of A. tsugense is

from sea level in Alaska, British Columbia, and

Washington, to 8,100 feet in the central Sierra

Nevada of California. Specimens examined: see

appendix, p. 220.

DISCUSSION: The name tsugensis was published

as a species of Razoumofskya; however, the Botanical Rules of Nomenclature require that the gender

of generic and specific names agree, so with the

transfer to Arceuthohium the correct name became

tsugense. This mistletoe has been variously

treated as a distinct species, or as alHed to A.

douglasii or A. campylopodum (Gill 1935). Its

phenology and general appearance certainly relate

it to the campylopodum group. A,, tsugense and

A. laricis are compared in table 22.

We have not observed A. tsugense growing with

other summer-flowering dwarf mistletoes. However, it occurs within about 34 niile of A. abietinum

f. sp. concoloris and A. occidentale near Fort

Figure 75.—General distribution of Arceuthohium

tsugense in Canada and the United States. Outline

map copyrighted by Denoyer-Geppert Co., Chicago

(used by permission).

38 Personal communication, Dr. E. F. Wicker, U.S. Forest

Service, 1967.

39 Personal communication, U.S. Forest Service, 1965.

155
Figure 76.—Distribution of Arceuthobium tsugense in

the United States (based on collections only). Outline

map copyrighted by Denoyer-Geppert Co., Chicago

(used by permission).

Bragg, California, and within a few miles of A.

abietinum f. sp. concoloris and A. lands in the

Oregon and Washington Cascades.

Since the dwarf mistletoes are sensitive to light,

this may account for the scarcity of shoots on

most lower infections under the dense shade of

coastal hemlock forests (Smith 1969). In such

situations vigorous shoots are often found only

along margins of stands, on young trees in openings, or in higher branches of older trees. Witches'

brooms caused by this taxon in Tsuga mertensiana

are illustrated in figure 13D.

ARCEUTHOBIUM VAGINATUM

23. Arceuthobium vaginatwn (Willdenow) Pre si in

Berchtold, O Pfirozenosti Rostlin aneb Rostlináf

2:28,1825.

<i^X

•»îp

Figure 77.—Distribution of Arceuthobium tsugense in

British Columbia and Alaska (based on collections

and reports).

DESCRIPTION: Shoots up to approximately 30

cm. long, but rarely reaching 50 cm., bright

orange to dark brown or black, usually densely

branched and erect, but large older plants sometimes become pendulous; flabellately branched;

internodes ca. 1-3 cm. long, 0.2-0.4 cm. wide;

staminate flower up to 3.5 mm. long and up to 3.5

mm. in diameter, mostly 3-, sometimes 4-merous,

perianth segments up to 2.0 mm. long and up to

1.5 mm. wide, apex acute to obtuse, pistillate

flower up to 2.5 mm. long and up to 1.5 mm. in

diameter; fruit 4-6 mm. long and 2-3 mm. wide.

156
elliptical to obovate; anthesis from approximately

late March through May. Parasitic on Dipoxylon

pines.

Our studies in Mexico (Hawksworth and Wiens

1965) and the United States show that A. vaginaturn is composed of three subspecies distinguishable on the basis of shoot color, size, staminate

flowers, fruit, time of anthesis, and geographic

distribution (figs. 78, 79).

Usually the three subspecies are distinct,

although some intermediates were found in a

transition zone between subsp. cryptopodum and

vaginatum southwest of Matachic, in western

Chihuahua (Hawksworth and Wiens 1965).

Key to the Subspecies

1. Plants dark brown to black; staminate flowers

usually greater than 3.0 mm. long and wide;

anthesis from approximately March through

April; Sierra Madre Occidental from central

Chihuahua southward to the Central Cordillera,

the Sierra Madre Oriental, and the mountains

of Nuevo León and Coahuilla.

23a. A. vaginatum suhsp. vaginatum

1. Plants dull orange to bright reddish orange;

staminate flowers usually less than 3.0 mm.

long and wide; anthesis from approximately

April through May; Sierra Madre Occidental of

northern Chihuahua and Sonora, or central

Durango; southwestern and central Rocky

Mountain States.

2. Mature plants generally bright reddish

orange, often greater than 20 cm. long;

mature internodes usually greater than 1.5

cm. long; fruit ca. 5 mm. long, the proximal

portion ca. 4 mm. long; the stigmas of

mature fruit usually slightly exceeding the

perianth segments (under magnification);

staminate flowering peduncles 1 to 1.5 cm.

long, unbranched; known only from several

localities near the crest of the Sierra Madre

Occidental in Durango and Sinaloa on the

road between Mazatlán and Durango.

23c. A. vaginatum subsp. durangense

2. Mature plant generally orange to dull orange,

not bright, usually less than 20 cm. long,

internodes usually less than 1.5 cm. long,

fruit ca. 4 mm. long, the proximal portion

ca. 3 mm. long; the stigmas on mature

fruit generally about equal to the perianth

segments (under magnification) ; staminate

flowering peduncles usually under 0.5 cm.

long, branched; Sierra Madre Occidental of

northern Chihuahua and Sonora; central

Coahuila and west Texas north to central

Utah and northern Colorado.

23b. A. vaginatum subsp. cryptopodum

ARCEUTHOBIUM VAGINATUM

SUBSP. VAGINATUM

23a. Arceuthobium vaginatum (Willdenow) Presl

subsp. vaginatum in Berchtold, O Pfirozenosti

Rostlin aneb Rostlináf 2: 28, 1825. TYPE:

MEXICO: VERACRUZ: Cofre de Perote, on

Pinus sp., Humboldt & Bonpland in 1804 (Isotype

MO!). Viscum vaginatum Willdenow, Sp. PI. ed. 4,

4: 740, 1806. Razoumojskya mexicana Hoffman,

Hort. Mosquensis (unpaged), 1808. Viscum vaginatum (Willd.) Kunth. Nov. Gen. et Spec. 3: 445,

1820. Arceuthobium vaginatum (H.B.K.) Eichler,

in Mart. Fl. Bras. 5(2); 105, 1868. Razoumojskya

vaginata (Willd.) Kuntze, Rev. Gen. PI. 2: 587,

1891.

DESCRIPTION: Mean shoot height ca. 20 cm.,

but up to 50 cm. (fig. 80). Shoots dark brown to

black. Basal diameter of dominant shoots 4-20

mm. (mean 7 mm.). Third internode 5-30 mm.

(mean 14.4 ± 6.0 mm.) long and 2.5-8.5 mm.

(mean 5.0 mm.) wide (11 collections), length/

width ratio 2.9:1. Staminate flower segments ca.

1.6 mm. long and 1.1 mm. wide. Mean anther

diameter 0.6 mm. centered 0.8 mm. from tip of

segment. Pollen polar diameter 19-23 (mean 21jLt);

equatorial diameter 22-26 (mean 25/x); polar/

equatorial diameter ratio 1:1.17; spine height

1.7M ca. 1.5 times wall thickness—1.2/i (3 collections). Mature fruit ca. 5.5 mm. long and 3.5 mm.

wide. Meiosis in February. Anthesis usually in

March-April. Seeds mature in August of the year

following pollination; maturation period averages

16-17 months, n = unknown.

HOSTS : This dwarf mistletoe is common on Pinus

montezumae, P. hartwegii, P. herrerai, P. lawsonii,

P. rudis, P. cooperi. P. ponderosa vars. scopulorum

and arizonica, P. durangensis and P. engelmannii,

all of which appear to be highly susceptible. It

rarely infects Pinus culminicola on Cerro Potosí,

Nuevo León (Hawksworth and Wiens 1965).

DISTRIBUTION: This subspecies, one of the most

widely distributed dwarf mistletoes in Mexico,

occurs in the Sierra Madre Occidental from

western Chihuahua south through Durango,

Jalisco, and into the Central Cordillera of Mexico

and Puebla; southeastward in the Sierra Madre

Oriental from Coahuila and Nuevo León to

Veracruz (see fig. 79). Known elevational range

is from 7,000 feet in Nuevo León to 12,800 feet on

Nevado de Toluca near Mexico City. Specimens

examined: see appendix, p. 222.

DISCUSSION: The shoots of A. vaginatum subsp.

vaginatum range up to 50 cm. high in the vicinity

of Mexico City.

The hosts and ecological requirements of A.

vaginatum subsp. vaginatum and A, globosum are

similar; the two mistletoes frequently occur to157
^subsp. vaginatum

^subsp. crypfopodum

subsp. durangense

Figure 78.—General distribution of the three subspecies of Arceuthobium vaginatum. Outline map

copyrighted by Denoyer-Geppert Co., Chicago (used by permission).

158
• A. vaginatum subsp. vaginatum

^ A, vaginatum subsp. durangenseo

- A. vaginatum subsp. cryptapodum I I -Distrito Fe

2-Morelos

100 200 300 400 500 MILES

600 800 KILOMETERS

5 - Aguascolientes

Figure 79.—Distribution of the three subspecies of Arceuthobium vaginatum in Mexico.

gether, sometimes on the same trees (Hawksworth

and Wiens 1965).

ARCEUTHOBIUM VAGINATUM

SUBSP. CRYPTOPODUM

SOUTHWESTERN DWARF MISTLETOE

23b. Arceuthobium vaginatum subsp. cryptopodum

(Engelm.) Hawksworth & Wiens, Brittonia 17:

230, 1965. HOLOTYPE: NEW MEXICO: Santa

Fe Co., Santa Fe, on Pinus ponderosa, Fendler 283

in 1847 (MO! Isotypes GH, K, NY). Arceuthobium

robustum Engelmann in Gray, Mem. Amer. Acad.

N.S. 4: 59, 1849, nomen nudum. Arceuthobium

cryptopodum Engelmann in Gray, Bost. J. Nat.

Hist. 6: 214, 1850. Razoumofskya robusta (Engelm.)

Kuntze, Rev. Gen. PI. 2: 587, 1891. Razoumofskya

cryptopoda (Engelm.) Coville, Contr. U.S. Nat.

Herb. 4: 192, 1893. Arceuthobium vaginatum

(Willd.) Presl forma cryptopodum (Engelm.) Gill,

Trans. Conn. Acad. Arts & Sei. 32: 178, 1935.

DESCRIPTION: Mean shoot height ca. 10 cm.,

but up to 27 cm. (fig. 81). Shoots orange to reddish

brown. Basal diameter of dominant shoots 2-10

mm. (mean 4 mm.). Third internode 4-16 mm.

(mean 7.8 d= 3.2 mm.) long and 2-4.5 mm. (mean

3.1 mm) wide (25 collections), length/Avidth ratio

2.5:1. Staminate flowers 2.5-3.0 mm. (mean 2.7

mm.) in diameter; perianth segments ca. 1.3 mm.

long and 1.0 mm. wide. Mean anther diameter 0.5

mm., centered 0.5 mm. from tip of segment. Pollen

polar diameter 18-28 (mean 22/x); equatorial

diameter 23-30 (mean 26M); polar/equatorial

diameter ratio 1:1.15; spine height (ca. 1.5ju) or

1.5 times wall thickness—1.0M (5 collections).

Mature fruit 4.5-5.5 mm. long (mean 5.0 mm.)

and 2.0-3.0 mm. wide (mean 2.5 mm.) ; proximal

portion ca. 3.5 mm. long. Seeds 2.7 X 1.1 mm.

Meiosis in April. Anthesis usually in ]\Iay-June.

Seeds mature in late July or early August of the

year following pollination; maturation period

averages 14-15 months. Germination in August to

September, n = 14.

HOSTS: Principal hosts are Pinus ponderosa vars.

scopulorum and arizonica, P. engelmannii, and the

159
Figure SO,—Arceuthobium vaginatum subsp. vaginatum in March: A, Staminate plant; ß, Pistillate plant; C,

Staminate flower; D, Immature fruit.

160
Figure 81.—Arceuthobium vaginatum subsp. cryptopodum in July: A^ Staminate plant; ß, Pistillate plants; C,

Detail of nearly mature fruit.

tree known as P. arizonica var. stormiae Martinez

in the Sierra del Carmen, Coahuila, Mexico;

occasional hosts are P. contorta var. latifolia

(Hawksworth 1956a) and P. aristata (Hawksworth

1965a); rare hosts are Pinus flexilis and P.

strohiformis.

DISTRIBUTION: Southwestern dwarf mistletoe is

widely distributed throughout the Pinus ponderosa

forests from central Utah* (Sevier County) and

* Robert C. Loomis (Personal communication, 1971) of the

Southwestern Region of the U.S. Forest Service, reports that

A. vaginatum subsp. cryptopodum is common on Navajo

Mountain, San Juan County, Utah.

northern Colorado (Larimer County) to Arizona,

New Mexico, west Texas (Guadalupe and Davis,

but not Chisos Mountains),'^^ at least as far south

as the Sierra del Pino in central Coahuila and into

the Sierra Madre Occidental in Sonora and

Chihuahua (figs. 79, 82).

Found as far west as the Hualapai Mountains

near Kingman, Arizona, this taxon occurs in

nearly every southwestern mountain range where

P. ponderosa grows. Andrews and Daniels (1960)

^0 Personal communication, Dr. P. C. Lightle, U.S. Forest

Service, 1966.

161
Figure 82.—Distribution of Arceuthobium vaginatum

subsp. cryptopodum in the United States and northern Mexico and the range of Pinus ponderosa var.

scopulorujn. Outline map copyrighted by DenoyerGeppert Co., Chicago (used by permission).

recorded the parasite on 36 percent of some 3,000

P. ponderosa plots in Arizona and New Mexico.

In 1969, when we examined several P. ponderosa

localities in northwestern Arizona (Mount Dellenbaugh, Black Rock Mountain, and Mount

Trumbull) we found no dwarf mistletoe, although

an infection has previously been reported from

Mount Trumbull.^' Known elevational range is

5,500 to 9,700 feet although this species is found

mainly between 6,600 and 8,500 feet in southwestern United States. Specimens examined: see

appendix, p. 222.

DISCUSSION: This mistletoe, common on P.

ponderosa pine in the central Rocky Mountains

and Southwestern States, is characterized by

thick, orange-colored shoots. While the shoots are

usually some shade of orange, certain populations

show various gradations from yellow to red, and

rarely may be purple or nearly black. Plants

growing in deep shade tend to be greenish, as do

those on Pinus contorta subsp. latifolia.

Some writers (for example, Kuijt 1955) have

questioned whether this taxon is distinct from A.

campylopodum, the parasite of Pinus ponderosa

along the Pacific Coast. Some additional features

that distinguish these two taxa are summarized in

table 23.

Southwestern dwarf mistletoe is particularly

damaging to P. ponderosa in south central New

Mexico (Lincoln National Forest, Mescalero

Apache Reservation), central Arizona, and along

the Front Range in Colorado. For reasons yet to

be explained, the parasite, although common, is

not as damaging to P. ponderosa in southwestern Colorado or southeastern Utah.

" Personal communication, John Churches, U.S. Forest

Service, 1956.

TABLE 23.—Comparison of Arceuthobium vaginatum subsp. cryptopodum

and A. campylopodum

Characters A. vaginatum subsp.

cryptopodum.

A. campylopodum

Shoots:

Third segment

Basal diameter

Color

Fruits

Meiosis

Peak flowering time

Peak seed dispersal time

Seed germination

Ca. 8 mm. long and 3 mm.

wide; length/width ratio

ca. 2-3

Up to 10 mm.

Usually orange

Orange, nonglaucous

April

May-June

Late July-August

Late summer or fall of year

of dispersal

Ca. 11 mm. long and 2 mm.

wide; length/width ratio

ca. 5-6

Under 5 mm.

Olive green to brownish

Greenish

July

August-September

September

Spring following year of dispersal

162
The witches^ brooms induced by this taxon

(Hawksworth 1961a; see fig. 11A,B)C) are similar

on all hosts except Pinus contorta subsp. latijolia.

On this species, witches' brooms are smaller and

swellings at the point of infection more pronounced

(often two to three times the diameter of healthy

parts of the branch) than in P. ponderosa.

ARCEUTHOBIUM VAGINATUM

SUBSP. DURANGENSE

23c. Arceuthohium vaginatum subsp. durangense

Hawksworth & Wiens, Brittonia 17: 230, 1965.

HOLOTYPE: MEXICO: DURANGO: 36 mi. W

of El Salto on Route 40, on Pinus durangensis,

Hawksworth & Wiens 3507 in 1963 (COLO.

Isotypes DS, F, FPF, INIF, MEXU, MO, US).

DESCRIPTION: Mean shoot height ca. 20-30 cm.

but up to 50 cm. (fig. 83). Shoots bright orange.

Longer shoots becoming pendulous. Basal diameter

of dominant shoots 4-8 mm. (mean 6 mm.).

Third internode 9-22 mm. (mean 14.9 dz 4.1 mm.)

long and 3.5-6 mm. (mean 4.5 mm.) wide (3

collections), length/width ratio 3.3:1. Internodes

often slightly swollen at base. Staminate ñowers

ca. 2.5 mm. long and 2.5 mm. in diameter, segments ca. 1 mm. long and 1 mm. wide. Pollen

polar diameter 18-19 (mean 18/x); equatorial

diameter 20-21 (mean 21/x); polar/equatorial

diameter ratio 1:1.15; spine height 1.5/x about 1.5

times wall thickness—1.0/x (1 collection). Mature

fruit ca. 7 mm. long and 3.5 mm. wide, bluish;

proximal portion ca. 4 mm. long. Seeds 4 X 1.5

mm. Time of meiosis unknown, probably February. Anthesis usually in April. Seeds mature

from mid-July to September of the year following

pollination; maturation period averages 15-18

months, n = unknown.

HOSTS: Pinus durangensis, P. montezumae, and P.

herrerai.

DISTRIBUTION: This rare dwarf mistletoe,

known only in extreme western Durango and adjacent Sinaloa on the western edge of the escarpment near the crest of the Sierra Madre Occidental

on Highway 40 between Durango and Mazatlán

(see figs. 78, 79), probably occurs in other areas

along the west side of this mountain range. Known

elevational range is 5,900 to 8,900 feet. Specimens

examined: see appendix, p. 227.

DISCUSSION: Although its distribution is poorly

known, this taxon is distinct and readily recognizable. It differs from subsp. vaginatum by its bright

orange shoot color and from subsp. cryptopodum

by its larger size, brighter color, branching pattern

of the staminate spikes, and geographical

distribution.

The branching pattern of the staminate plants

on the populations on Pinus herrerai in extreme

western Durango near the Sinaloa boundary seem

to differ slightly from nearby populations on P.

montezumae. We believe that both populations are

the same taxon, but we will need further field

studies to confirm this.

ARCEUTHOBIUM VERTICILLIFLORUM

24. Arceuthohium verticilliflorum Engelmann, Bot.

Calif. 2: 107, 1880. TYPE: MEXICO: DURANGO: Sierra Madre, (on Pinus sp.?), Seemann

2138 in 1852 (MO! Isotype K, ILL). Razoumojskya verticillata (Engelm.) Kuntze, Rev. Gen.

PI. 2:587,1891.

DESCRIPTION: Mean height ca. 7 cm., but up to

11 cm. (fig. 84). Shoots mostly yellowish green to

purple, without secondary branching, shoots

lightly glaucous when young. Basal diameter of

dominant shoots 2.5-5 mm. (mean 3.6 mm.).

Third internode 2-7 mm. (mean 3.0 ±1.2 mm.)

long and 2.5-4.5 mm. (mean 3.2 mm.) wide (4

collections), length/width ratio 0.9:1. Staminate

flowers 3.5-4.5 mm. (mean 4.0 mm.) in diameter;

perianth mostly 4-, sometimes 3-merous; verticillate, with 5-6 flowers per whorl; segments 1.8 mm.

long and 1.2 mm. wide. Mean anther diameter 1.0

mm., centered 0.8 mm. from tip of segment. Pollen

polar diameter 21-25 (mean 24/x) ; equatorial diameter 26-30 (mean 28/x) ; polar/equatorial diameter

ratio 1:1.18; spine height (ca. 1.0/x) or ca. % the

wall thickness—1.5^ (2 collections). Mature fruit

ca. 15 mm. long and 10 mm. wide. Seeds ca. 11 X 6

mm., embryos 4X1 mm. Meiosis in SeptemberOctober. Anthesis usually in March-April. Fruits

mature in September and October of the year

following pollination; maturation period averages

18-19 months, n = 14.

HOSTS: This dwarf mistletoe is known only on

Pinus engelmannii and P. cooperi; Pinus leiophylla

vars. leiophylla and chihuahuana and P. teocote

associated with these infected pines were not

parasitized.

DISTRIBUTION: This poorly known species has

been collected only west of Durango along the

Durango-El Salto highway at elevations between

7,900 and 8,900 feet. The location of Seemannes

type collection is uncertain but it is presumably

somewhat south of the above area. Specimens

examined: see appendix, p. 227.

DISCUSSION: This species, although first described by Engelmann in 1880, had received no

serious consideration until we showed that it was

indeed a valid, and very distinctive, species

(Hawksworth and Wiens 1965). The staminate

shoots are characterized by the thick spikes (4-6

mm. in diameter) with verticillate 4-merous

flowers, mostly 6 in a whorl; the staminate spikes

163
Figure 83.—Arceuthobium vaginatum subsp. durangense in July: A, Pistillate shoot; B, Detail of nearly mature

fruit; C, Staminate shoot; D, Detail of staminate floral remnant.

164
Figure 84.—Arceuthobium verticilliflorum in March: A^ Distal portion of pistillate shoot; ß. Distal portion of

staminate shoot; C, Pistillate shoot; D, Immature fruit; E, Staminate shoot.

165
are deciduous after flowering. The species has

fruits larger than any other dwarf mistletoe. These

averaged 3X6 mm. in March, 7 X 14 mm. in

July, and 10 X 15 mm. in September, when they

were nearly mature. A few fruits were expelled

in early September, but the peak period of seed

dispersal is presumably in October.

In all taxa except this one, the pedicels elongate

so that the distal ends of the fruits are oriented

downward, and the seeds at maturity are expelled

upward. By contrast, an apparently unique

feature of A. verticilliflorum is that the pedicels do

not elongate at maturity and the seeds seem to be

expelled more or less at random. This feature, plus

apparent evidence of bird feeding, suggests that

the taxon may be evolving away from explosive

seed dispersal, perhaps in favor of bird dispersal.

Other observations that also suggest bird dispersal

are the common occurrence of the mistletoe in

very open stands and the almost random distribution of infection in tree crowns. All other dwarf

mistletoes are concentrated in the lower portions

of the tree crowns during the early stages of

infection.

Kuijt (1970) concluded that the branching

pattern of A. verticilliflorum was basically verticillate. This dwarf mistletoe causes massive witches'

brooms (see fig. lOA).

Old World Taxa

ARCEUTHOBIUM CHÍNENSE

25. Arceuthohium chinense Lecomte, Notulae Systematicae 3: 170, 1915. TYPE: CHINA: YUNNAN: Bois audessus de Ta pin tze; parasite on

Abies, Delavay s. n., not dated.

DESCRIPTION: Plants 2-5 cm. high, color of

living plants unknown; secondary branching

verticillate (Kuijt 1970). Basal diameter of dominant shoots ca. 1.5 mm. Third internode 4-6 mm.

(mean 5 mm.) long and 1-1.5 mm. (mean 1.1 mm.)

wide (4 collections); length/width ratio 4.4:1.

Staminate flowers ca. 2 mm. in diameter, 4-,

rarely 3-merous; segments ca. 1 mm. long and

1 mm. wide; perianth segments tapered to tip,

widest at base. Pollen polar diameter 14-18

(mean 16^); equatorial diameter 16-19 (mean

17/x); polar/equatorial diameter ratio 1:1.09;

spine height and wall thickness ca. l.O/z (2 collections). Mature fruit ca. 3-3.5 mm. long and 1.5-2

mm. wide, distal portion ca. 2 mm. long, proximal

portion nonglaucous (?). Time of meiosis unknown.

Anthesis usually in September but beginning in

July in one specimen, fruits mature in September,

so fruit maturation period is ca. 11-12 months.

n = unknown.

HOSTS AND DISTRIBUTION: On Abies and

Keteleeria in Yunnan and Szechwan Provinces in

southwestern China at 8,200 to 13,000 feet in

elevation. Handel-Mazzetti (1929) gives the lower

elevational range of ''A. chinense'' as 1,600

meters (5,250 feet) but it is uncertain whether

this refers to A. chinense sensu stricto or A. pini.

The dwarf mistletoe reported on Abies pindrow

in India may be this taxon (Rajagopaliengar 1955).

Specimens examined: see appendix, p. 227.

DISCUSSION: Information on this very poorly

known taxon is available only from a few reports

(Anonymous 1939, Danser 1936, Handel-INIazzetti

1929, and Lecomte 1915) but these papers do not

distinguish A. chinense from the recently described A, pini. Some features of these two parasites are compared in table 24.

Type of witches' brooms formed by A. chinense

has not been reported, but the specimen Schneider

2951 (at KewO on Keteleeria exhibits a systemic

type of parasitism. Handel-Mazzetti (1929) reports a young Keteleeria stand in Yunnan Province

that was completely killed, presumably by A.

chinense.

TABLE 24:.—Comparison of Arceuthohium chinense and A. pini

Characters A. chinense

Hosts

Mean plant height

Shoots:

Third internode

Basal diameter of

dominant shoots

Staminate flowers:

Lobes

Perianth segments

Abiesj Keteleeria

2-5 cm.

5X1 mm.

Ca. 1.5 mm.

A. pini

Pinus tahulaeformis

10-20 cm.

12 X 2 mm.

2.5-3 mm.

Mostly 4-merous Mostly 3-merous

Rounded; widest near mid« Tapered to tip; widest at base

die

Peak time of flowering August-September May-June

166
ARCEUTHOBIUM MINUTISSIMUM

HIMALAYAN DWARF MISTLETOE

26. Arceuthohium minutissimum J. D. Hooker, Fl.

Brit. India 5: 227, 1886. TYPE: KUMAON

HIMALAYA, alt. 10,700 feet, on Pinus griffithii

(as P. excelsa), Duthie s.n., not dated. Razoumojskya minutissima (Hook.) Tubeuf, Naturw.

Zeitschr. Forst, u. Landw. 17:195,1919.

DESCRIPTION: Plants ca. 5 mm. high, but up to

10 mm. (fig. 85), yellow to yellow green; primary

branching multiple from basal cups, but without

secondary branching (Kuijt 1970). Basal diameter

of dominant shoots ca. 1 mm. Third internode

0.5-1.4 mm. long (mean 0.8 db 0.3 mm.) and

0.3-1.0 mm. (mean 0.4 mm.) wide (3 collections);

length/width ratio 1.5:1. Staminate flowers 2-2.5

mm. in diameter; perianth mostly 4-, sometimes

3-merous; segments ca. 0.8 mm. long and 0.8 mm.

wide. Anther diameter ca. 0.5 mm. centered ca.

0.4 mm. from tip of segment. Pollen polar diameter

16-19 (mean 18^); equatorial diameter 19-21

(mean 21/x); polar/equatorial diameter ratio

1:1.16; spine height about the same as wall

thickness—O.SjLi (3 collections). Mature fruit 2-2.5

mm. long and 1-1.5 mm. wide. Time of meiosis

unknown. Anthesis, reported to be from August

to October (Datta 1956), was in September in all

flowering specimens that we examined. Seed

dispersal also in September, so fruit maturation

period averages ca. 12 months, n = unknown.

HOSTS: The only certain host is Pinus griffithii

(syn. P. excelsa Wall., P. wallichiana A. B. Jacks),

but reported indirectly on Abies pindroiu in a

description of a land shell said to occur in basal

cups of A. minutissimum on this host in the Simla

Hills, East Punjab, India (Rajagopaliengar 1955).

DISTRIBUTION: This dwarf mistletoe is known

only from the western Himalayas in West

Pakistan, Kashmir, India, and Nepal. Brandis

(1907) reports it at elevations of from 8,000 to

11,000 feet. Bagchee (1952) gives its distribution

in India as the dry zone of the Himalayas in

Upper and Lower Bashahr in Himachal Pradesh,

Kulu in East Punjab, and the four divisions of

Kashmir. Specimens examined: see appendix, p.

227.

DISCUSSION: Hooker (1886) described A. minutissimum as the smallest known dicotyledonous

plant. While it does have very small aerial shoots,

which average less than 5 mm. high, the plant is

much more extensive because of its endophytic

system within the host tissues. Thus, it is difficult

to compare the size of the mistletoes with that of

autotrophic plants.

Whether A. minutissimum is dioecious (as are

all the other members of the genus) or monecious

has not been resolved (Danser 1931, Datta 1951,

and Hooker 1886), because intermixed pistillate

and staminate shoots frequently occur. This does

not mean, however, that this species is monecious.

Perhaps mixed infections occur more frequently in

A. minutissimum than in the North American

dwarf mistletoes. Datta (1951) described a single

hermaphroditic flower in A. minutissimum, and

Bhandari and Nanda (1968) discuss the embryology and morphology of this distinctive taxon.

Kuijt (1960b) suggests that fruits of A, minutissimum, as in the North American A. pusillum,

may mature in a single growing season (ca. 5

months). However, the specimens that we have

examined and our field studies in Kashmir clearly

indicate that peak periods for both anthesis and

seed dispersal occur in September; thus, a maturation period of abour 12 months is indicated.

The parasite is a serious pathogen of P. griffithii,

particularly on the drier sites where it causes severe

mortality (Gorrie 1929). A. minutissimum induces

systemic witches' brooms (Gorrie 1929, Kuijt

1960b; see fig. 14C).

ARCEUTHOBIUM OXYCEDRI

JUNIPER DWARF MISTLETOE

27. Arceuthohium oxycedri (DC.) M. Bieb., Flora

Taurico-Caucasia III: 629, 1819. TYPE: details of

type unknown but Komarov (1936) says that it is

in Paris. Viscum oxycedri DC, Fl. Fr. 4: 274, 1805.

Razoumofskya caucásica Hoffman, Hort. Mosqu.

(unpaged), 1808. Razoumofskya oxycedri (DC.)

Schultz, ex Nym. Consp.: 320, 1853. Viscum

caucasicum Steud. No. ed 1: 888, 1891. Arceuthohium juniperi Bubani, Flora Pyrenacea: 131,

1897. Arceuthohium juniperi-procerae Chiovenda,

Ann. di Botánica (Rome) IX (fase. 2): 134, 1911.

Arceuthohium juniperorum Reynier, Bull. Soc.

Bot. France 66: 97, 1919. Arceuthohium oxycedri

(DC.) M. Bieb. ySLYCupressii Zefirov, Akad. Nauk

SSSR. Bot. Inst. 17:110,1955.

DESCRIPTION: Shoots mostly 5-10 cm. high, but

up to 20 cm.; yellow green; verticillately branched

(Kuijt 1970). Basal shoots 1.5-2 mm. in diameter.

Third internode 5-9 mm. long and ca. 1 mm. in

diameter; shoots terete; segments often markedly

wider at the top than at the base. Staminate

flowers 1.5-2.5 mm. in diameter, perianth mostly

3-, but sometimes 2- or 4-merous. Pollen polar

diameter 17-19 (mean 18ß), equatorial diameter

19-22 (mean 21^); polar/equatorial diameter

ratio 1:1.17; spine height (1.5/x) slightly greater

than wall thickness—1.0^ (2 collections). Mature

fruit ca. 3 mm. long and 1.5-2 mm. wide. Anthesis

in September and October (Heinricher 1915a).

Seed dispersal in autumn (Heinricher 1915a).

n = reported to be 13 or 14 (Pisek 1924) but

167
2 mm

Figure SS.—Arceuthobium niinutissimum in September: A, General habit of shoots on a piece of host bark; B,

Pistillate shoot; C, Staminate shoot with two flowers. Other drawings illustrate various shoot details (same

scale as B and C).

168
Wiens (1968) suggests that it is probably n = 14,

as are all North American taxa of the genus

counted to date.

HOSTS: Parasitic mainly on Juniperus oxycedrus,

but also on several other species of Juniperus

(table 25).

DISTRIBUTION: Described as 'Very local'' (Tutin,

Heywood, Burges, and others 1964:), Arceuthohium

oxycedri is the most widely distributed dwarf

mistletoe. The most comprehensive accounts of its

distribution (Turrill 1920, 1926) give a 6,000-mile

range from the Azores, throughout the Mediterranean area of Europe and Africa, Asia Minor,

and east to India, in addition to the Aberdare

Mountains of Kenya, the only known locality

for Arceuthohium in the southern hemisphere.

Turrill (1920), and others as noted, have

reported A. oxycedri from the following countries :

EUROPE: Azores, Portugal (Palhinha 1942),

Spain, France, Corsica (specimen at US), Italy?

(Tutin, Heywood, Burges, and others 1964),

Yugoslavia, Hungary, Albania, Greece, Bulgaria,

U.S.S.R. (Crimea); AFRICA: Algeria, Ethiopia,

Kenya; ASIA: Turkey, Lebanon, Syria, Iran,

U.S.S.R. (Armenia, Georgia, Turkmen S.S.R.,

Tadzhik S.S.R.) (Komarov 1936) and India.

DISCUSSION: The occurrence of A. oxycedri over

such an extremely wide geographical area suggests

that detailed field study may reveal segregates of

the species (Wiens 1968). For example, the

apparent outlier in east central Africa has been

accorded specific rank as A. juniperi-procerae

(Chiovenda 1911). Also Zefirov (1955), who

described a new variety (var. cupressi) on an

introduced tree (Cuppressus macrocarpa), said

the variety could be distinguished from the typical

form on Juniperus by its longer, thinner, greenish

shoots. We consider such erection of taxonomic

entities based on their occurrence on introduced

hosts as spurious.

ARCEUTHOBIUM PINI

28. Arceuthohium pini Hawksworth & Wiens, Brittonia 22: 267, 1970. HOLOTYPE: CHINA: YUNNAN Prov, E flank of Lichiang Range, Lat. 27°

35' N, elevation 12,000 feet, on PinuSy Forrest

10169 in 1913 (BM, Isotype K).

DESCRIPTION: Plants 10-22 cm. high, olive to

dull green; no secondary branching observed in 5

specimens examined. Basal diameter of dominant

shoots 2.5-3 mm. Third internode 5-14 mm.

(mean 12) mm. long and 1.5-2.5 (mean 2 mm.)

wide (5 collections); length/width ratio 6.2:1.

Staminate flowers ca. 2 mm. in diameter; perianth

mostly 3-, rarely 4-merous; segments ca. 1 mm.

long and 1 mm. wide; perianth segments rounded,

widest near the middle. Pollen polar diameter

15-18 (mean 16jLt); equatorial diameter, 16-18

(mean 18/x); polar/equatorial diameter ratio

TABLE 25.—Hosts of Arceuthohium oxycedri

Host Locality Reference

Juniperus oxycedrus (in- General Turrill 1920

cluding /. rufescens)

J. communis do. do.

J. drupácea do. do.

J. sabina do. do.

J. hrevifolia Azores do.

J. procera Kenya do.

Do. Eritrea Chiovenda 1911

/. Phoenicia France Leveillé 1904

J. macropoda India Brandis 1907

J. excelsa U.S.S.R. Komarov 1936

J. semiglohosa do. do.

J. seravschanica do. do.

J. turcomanica do. do.

Cupressus macrocarpa do. Zefirov 1955

Chamaecyparis thyoides^ Inoculations Heinricher 1930

Thuja Orientalis 2 Beer 1951

^ Heinricher (1930) listed this tree as Chamaecyparis sphaeroides var. péndula, although it is now known as C. thyoides. The same tree was erroneously recorded earlier

as Cupressus péndula (Heinricher 1920).

2 Transmitted vegetatively to Thuja orientalis scions on infected Juniperus communis

stock.

169
1:1.13; spine height 1.5fx, wall thickness—ca. 1.0^

(3 collections). Mature fruit ca. 3-3.5 mm. long

and ca. 2 mm. wide, distal portion probably

glaucous. Time of meiosis unknown. Anthesis

usually in April-June, but one specimen collected

in July had a few flowers still with pollen. Fruit

maturation time not precisely known, but apparently in August or September; fruit maturation

period presumably averages ca. 16 months, n =

unknown.

HOST: Known only on Pinus tahulaeformis.

DISTRIBUTION: This dwarf mistletoe is known

only from Yunnan and Szechwan Provinces in

Southwestern China and in Tibet. Known elevational range is from 9,000 to 12,000 feet. HandelMazzetti (1929) gives the lower altitudinal range

range of "A. chínense'' as 1,600 meters (5,250 feet),

but it is not known whether this refers to this

species or A. chínense sensu stricto. Specimens

examined: see appendix, p. 227.

DISCUSSION: Some characteristics that serve to

distinguish this little-known taxon from the

equally poorly known A. chínense were given in

table 24.

Nothing has been published on the witches'

brooms formed by this dwarf mistletoe. The type

specimen {Forrest 10169) at Kew has one branch

that is systemically infected. This specimen, which

was collected in June 1913, is described below:

1913 and 1912 growth: no shoots

1911 growth: dwarf mistletoe buds ca.

1 mm. high at proximal

and distal ends of segment.

1910 and 1909 growth: shoot bases throughout

segment, shoots broken

off.

We do not know whether A. píní consistently

forms this type of broom. The British Museum

specimen of Forrest's same collection and four

other specimens examined did not show systemic

brooming. Handel-Mazzetti (1929) discusses swellings associated with the parasite on pines, but he

does not mention witches' brooms.

Rejected Species

1. Arceuthohium honaniae Grisebach, Cat. Plant.

Cuben.: 121, 1866 = Dendrophthora honaniae

(Griseb.) Eichler, Flora Brasil. 5 (2) : 103, 1868.

2. Arceuthohium cupressoides (Macf.) Grisebach,

Flora Brit. West Indies: 315, 1864. Razoumojskya cupressoides (Macf.) Kuntze, Rev. Gen.

2: 587, 1891 = Dendrophthora cupressoides

(Macf.) Eichler, Flora Brasil. 5 (2): 103, 1868.

3. Arceuthohium dacrydii Ridley, J. Fed. Malay

States Mus. 6:170, 1915 = Korthalsella dacrydii

(Ridley) Danser, Rec. Trav. Bot. Neer. 31:

759,1934.

4. Arceuthohium domingensis (Spreng.) Grisebach,

Cat. Plant. Cuben: 121, 1866 = Dendrophthora domingensis (Spreng.) Eichler, Flora

Brasil. 5 (2) : 103,1868.

5. Arceuthohium epiviscum Grisebach, Mem. Amer.

Acad. Arts and Sei. N.S., 8: 192, 1861 =

Dendrophthora epiviscum (Griseb.) Eichler, Flora

Brasil. 5 (2) : 103,1868.

6. Arceuthohium glauca Grisebach, Cat. Plant.

Cuben.: 121, 1866 = Dendrophthora glauca

(Griseb.) Eichler, Flora Brasil. 5 (2) : 103, 1868

7. Arceuthohium gracile (DC.) Grisebach, Flora

Brit. West Indies: 315, 1864 (non Engelmann,

Mem. Amer. Acad. Arts, and Sei., N.S., 4 (4):

59, 1849). Razoumofskya gracilis (DC.) Kuntze,

Rev. Gen. 2: 587, 1891 = Dendrophthora

flagelliformis (Lam.) Krug & Urban, Ber.

Deutsch Bot. Ges. 14: 287,1896.

8. Arceuthohium mancinellae Grisebach, Cat. Plant.

Cuben.: 121, 1866 = Dendrophthora mancinellae

(Griseb.) Eichler, Flora Brasil. 5 (2): 103, 1868.

9. Arceuthohium opuntioides (L.) Grisebach, Flora

Brit. West Indies: 315, 1864. Razoumofskya

jamaicensis Hoffman, Hortus ]Mosquensis (unpaged), 1808. Razoumofskya opuntioides (L.)

Kuntze, Rev. Gen. 2: 587, 1891 = Dendrophthora opuntioides (L.) Eichler, Flora Brasil. 5 (2) :

102,1868.

MAJOR CONCLUSIONS

Some of the major conclusions of this work are:

1. Arceuthohium is a well-defined genus of the

Viscaceae parasitic on conifers. The genus is

probably of early Tertiary origin in Eastern

Asia. Korthalsella, which occurs from Hawaii

throughout the South Pacific area, is probably

the closest related genus.

2. Arceuthohium presumably migrated to the New

World in pre-Miocene times. An intensive

adaptive radiation occurred into the Pinaceae

of the New World, and this area is now the

center of species diversification. Of the 32 taxa

recognized in this paper, 28 are in the New

World, and 4 in the Old World.

170
3. Most of the Pinaceae of western North America

and Mexico have been colonized by Arceuthohium. In the western United States, all the

species of Ahies, Picea^ Tsuga, and Larix are

parasitized to some degree. In addition, about

80 percent of the Pinus and 50 percent of the

Pseudotsuga species are dwarf mistletoe hosts.

The few species that are not known to be parasitized are mostly local endemics.

4. Arceuthobium has a base chromosome number of

X = 14. No hybridization or polyploidy is

known in the genus. The absence of hybridization and polyploidy has apparently led to a

dendritic pattern of evolution that resulted in

clearly demarcated taxa.

5. Although taxa are clearly demarcated, the differences may be cryptic and apparent for only

portions of the life cycle. The genus has undergone extreme morphological reduction due

to the parasitic habit. Species and subspecies

differ only in the number of discontinuous

variables. With the possible exception of A.

vaginatum subsp. vaginatum and cryptopodum,

even subspecies do not intergrade.

6. Most taxa are sympatric with other members of

the genus somewhere in their geographic range.

In addition, the flowering periods of sympatric

species often overlap. If the taxa are compatible,

gene exchange should occur, but does not. Even

though the differences among taxa are less

pronounced than in some autotrophic groups,

each should be regarded as a separate species.

All dwarf mistletoe taxa are relatively true

breeding, so cannot be considered as ecotypes

or geographic races.

SUGGESTIONS FOR FURTHER RESEARCH

Research into the host-parasite relationships of the

dwarf mistletoes should include investigation of the

genetic and physiological basis for host specificity.

Studies such as Greenham and Leonard's (1965) on

the amino acid components of dwarf mistletoes and

their hosts should be continued. Research similar to

the study of the ultrastructure of Arceuthohiwn

pusillum in compatible and noncompatible hosts

(Tainter and French 1969b) should be expanded

because it could provide much-needed basic information on the host-parasite relationship.

Attempts to grow dwarf mistletoes in vitro

(Blakely 1959, Bonga and Chakraborty 1968) should

be continued. Studies should be conducted on the

comparative evolution of the dwarf mistletoes and

their hosts, and crossing experiments between

various taxa attempted.

Dwarf mistletoe pollination, pollinators, and

effective distances of pollen transport should be

experimentally investigated.

Local dispersal of dwarf mistletoes by their

explosive fruits has been studied in some detail

(Hinds and Hawksworth 1965), but httle is known

of the long-range dispersal of these plants. Presumably birds are involved, but experimental data

are lacking.

Specific areas suggested for study are:

1. Complete chromosome counts for New World

species not yet studied {A. glohosmn, A. guatemalense, A. ruhrum), two subspecies of A.

vaginatum, and all the Old World species.

Make comparative karyotype studies of all

members of the genus.

2. Study all the Old World species to determine

whether they can be ranked into the three

stages of evolutionary divergence we propose

for the New World species.

3. Conduct detailed field and experimental studies

on Arceuthohiwn oxycedri populations throughout its vast range (from the Azores to India to

Kenya) to determine whether geographic segregates are discernible.

4. Study the hosts, biology, morphology, and life

history of the very poorly known A. chinense

and A. piniin southeastern Asia.

5. Determine by field and morphological studies

whether the central Mexican (Durango) and

southern Mexican populations (Hidalgo, Veracruz, Oaxaca) of A. gillii subsp. nigrum are, in

fact, the same taxon.

6. Determine whether the marked differences we

found between the Guatemalan and central

Mexican populations of A. glohosum are continuous or discontinuous by making field

studies in southern Mexico.

7. Determine with certainty the taxonomic status

of the dwarf mistletoe on Pinus herrerai in

Durango, Mexico, which we tentatively classed

as A. vaginatum subsp. durangense.

8. Determine more fully in the field the host and

geographical distributions of the Mexican and

central American dwarf mistletoes.

9. Determine, particularly in Ontario and Quebec,

the northern limits of Arceuthobium pusillum,

10. Conduct field and experimental studies to

determine the consistency of differences observed in populations of A. occidentale on its

171
three principal hosts: Pinus sabiniana, P.

radiata, and P. muricata.

11. Clarify the status of the dwarf mistletoes on

Pinus montícola in northern California and

southern Oregon, where Arceuthohium cyanocarpum, A. calif ornicum, and A. tsugense all

parasitize this tree. We consider some of our

identifications to be tentative, until field

studies are completed on the parasites on P.

rnonticola.

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APPENDIX

Glossary

ADAPTIVE RADIATION. The evolution of species

into a diversity of previously unoccupied

ecological niches.

ADVANCED TAXA. Plants that have derived

characters not present in their ancestors (see p

38).

ALLOPLOID. An individual of hybrid origin whose

sets of chromosomes are derived from different

parental species.

ANISOPHASIC WITCHES' BROOM. See nonsystemic witches' broom.

ANTHESIS. The period of flowering.

ARCHESPORIUM. A cell or group of cells from

which spores (including pollen) are derived.

AUTOPLOID. An individual which has one or more

exact duplications of the diploid set of

chromosomes.

BIVALENT. An homologus pair of chromosomes in

the synapsed, or paired, state during prophase

of the first meiotic division.

CHIASMA. The microscopically visible, apparent

exchange of chromatid parts between homologous chromosomes during prophase of the

first meiotic division.

CHROMATOGRAPHY. Separation of chemical

compounds by allowing a solution of them to

seep through an adsorbent material (paper in

our studies) so that the different compounds

become adsorbed in separate colored layers.

CONNATE. A uniting of parts, particularly similar

structures united to form a single organ.

CONVERGENCE. An evolutionary phenomenon

whereby similar forms or functions occur in

groups which are not closely related.

CROSS-OVER. Parasitism by a dwarf mistletoe on a

tree other than its principal host.

DECUSSATE. An arrangement of shoots in that the

parts alternate in pairs at right angles (see fig. 1).

DENDRITIC EVOLUTION. A pattern of evolution

in which distinct hneages are evident.

DICHASIUM. A type of inflorescence, which in its

basic form consists of an older apical flower

subtended laterally by two younger flowers of

axillary origin.

DIMORPHISM. The occurrence of two forms, often

associtated with morphological differences relating to sex.

DIOECIOUS. The situation in which each individual

of a species possesses only a single sex.

DIPLOID. An individual which has twice the number of chromosomes which characterizes a

gamete; i.e. the presence of both members of

homologous pairs of chromosomes.

DIPLOXYLON. The hard or yellow pines, one of the

two major groups of the genus Pinus.

DIRECT FLOWERING. A plant with uninterrupted development of the floral buds from

initiation to anthesis.

DISTAL. Part of a structure farthest from the point

of attachment (cf. proximal).

DIVARICATE. Parts spreading widely away from

one another.

ENDOPHYTIC SYSTEM. The ^^root-like'^ parts

of dwarf mistletoe within the host tissues. The

endophytic system consists of ''strands'' within

the cortex and ''sinkers" which are embedded in

successive layers of xylem.

EPIGYNOUS. Originating from above the ovary;

generally used to describe situations where the

ovary is situated below the perianth.

182
EPIPHYTE. A plant growing on another plant

from which it derives no benefit other than

support, such as various orchids or bromeliads.

EXTRA-LIMITAL HOSTS. Infection of trees that

do not occur within the natural range of a

particular dwarf mistletoe. Susceptibility of

such trees may be established either by inoculations or by introducing trees into areas where the

dwarf mistletoe occurs.

FLABELLATE. Fan shaped, a branching pattern

produced by the continued development of

superposed axillary buds.

GENOME. The haploid (n) set of chromosomes.

GIRDLE. The point on a vegetative branch or main

stem of a conifer between two annual growth

segments. This is frequently, but incorrectly,

referred to as a ''node."

GLABROUS. Bald, without hairs or trichomes.

GLAUCOUS. A usually delicate whitish waxy

coating which is easily rubbed away.

HAPLOID. A situation where each nucleus contains

only one member of each homologous pair of

chromosomes; the n number of chromosomes.

HAPLOXYLON. The white or pinyon pines; one of

the two major groups of the genus Pinus.

HOLDFAST. A disc-like swelling at the distal end of

the radicle through which infection of the host

takes place.

HOLOTYPE. The single specimen designated by an

author of a taxonomic name to which that name

is permanently attached.

INDIRECT FLOWERING. The intervention of a

rest period between the initiation of a floral bud

and anthesis.

INFLORESCENCE. A flower cluster or mode of

bearing fllowers.

ISOPHASIC WITCHES' BROOMS. See systemic

witches' broom.

ISOTYPE. A specimen collected at the same place

and time and by the same collector as the

holotype.

KARYOTYPE. The morphological and quantitative

characterization of chromosome set, including

such features as size, centromere position, arm

length, satellites and chromosome member.

LECTOTYPE. A specimen selected from the

original material on which a taxon is based,

when no holotype was designated originally, or

when the holotype is missing or destroyed.

LENTICULAR. Shaped Hke a double-convex lens;

not spherical.

MEDELIAN. A type of inheritance which is

allelomorphic, i.e. exhibits alternate forms,

usually with respect to dominance and

recessiveness.

MEGASPOROGENESIS. A meiotic nuclear division in plants which produces megaspores, which

in turn give rise to mega- or female gametophytes. The megagametophyte in flowering

plants (embryo sac) eventually produces the

egg and associated cells at maturity.

MEIOSIS. A type of cell division which produces

gametes (in animals) or. spores (in plants) which

are characterized by having the reduced or

haploid (n) chromosome number, i.e. one member of each homologous chromosome pair.

MICROSPOROGENESIS. A meiotic nuclear division in plants which produces microspores

(pollen grains) which in turn produce a microor male gametophyte. The microgametophyte

in flowering plants (pollen tube) eventually

produces the sperm nuclei.

NEOTYPE. A specimen designated as the type

when the original type and all specimens on

which the name Avas based has been either

destroyed or are missing.

NONSYSTEMIC WITCHES' BROOM. Witches'

broom in which the dwarf mistletoe shoots

remain concentrated near the original point of

infection (compare with systemic broom, p. 26).

PALEOBOTANY. The study of fossil plants.

PALYNOLOGY. The study of pollen.

PARASITE. An organism, such as mistletoe, that

obtains sustenance from another organism.

PEDICELLATE JOINT. The point of attachment

of the flower or fruit to the pedicel.

PERIANTH. A collective term for the sepals and

petals.

PHENOLOGY. The relationship between seasonal

and periodic biological phenomena, e.g. flowering and fruiting times.

PHOTOPERIOD. A light period, usually the daylight portion of a 24-hour period at a particular

time of the year.

PHYLAD. A natural group of any rank, usually

considered with respect to its evolutionary

origin or history; any evolutionary line of development.

PHYLETIC. The evolutionary history of a group of

organisms.

POLYMORPHISM. The occurrence of two or more

forms within a population.

POLYPLOIDY. The exact dupUcation of entire

cromosome sets.

PRIMARY BRANCHING. The basic, decussate

type of branching exhibited by all species of

Arceuthohiu77i.

PRIMITIVE TAXA. Plants that have characters

also possessed by their ancestors (see p. 38).

PRINCIPAL HOSTS. The main, or most susceptible

hosts, for a particular dwarf mistletoe (see p. 16).

183
PROXIMAL. Part of a structure closest to the point

of attachment (cf. distal).

RADICLE. The growing tip of the hypocotyl in a

germinating seed.

RECOMBINATION INDEX. The haploid number

of chromosomes plus the chiasma frequency.

RECOMBINATION POTENTIAL. The possibility for expressing genetic variability which is

determined by the genetic system (cf. recombination index).

RELICTUAL. A population or species occurring in

only a small geographical portion or portions of

its previous distribution.

RETICULATE EVOLUTION. A netHke structure

of relationships, normally resulting from hybridization and alloploidy.

SECONDARY BRANCHING. Branching that

develops in addition to primary branching. It

may be either flabellate or verticillate.

SEGMENT. The single years' growth in length of a

vegetative branch or main stem of a conifer.

This is often, but incorrectly, referred to as an

'^internode.''

SYMPATRY. Occurring together; used specifically

in this study to denote situations where temporal and spatial consideration should permit

hybridization between taxa unless they are

reproductively isolated.

SYSTEMIC WITCHES' BROOMS. Witches'

brooms in which the endophytic system keeps

pace with the growth of the infected host branch.

Dwarf mistletoe shoots are scattered along the

branch (seep. 26).

TAXIMETRICS. Analyses associated with numerical taxonomy.

TAXON (plural Taxa). A taxonomic unit of any

rank (species, subspecies, form, etc.).

TETRAPLOIDY. A polyploid condition charaacterized by four sets of chromosomes.

TRICÓME. An epidermal appendage or hair.

TYPE. A specimen which is the basis of a taxonomic

name, usually a holotype, but sometimes also

lectotypes or neotypes.

VERTICILLATE (whorled). The occurrence of

more than two leaves or branches at a single

node.

VISCOUS. Sticky, generally used in this study to

indicate the material around the seed of dwarf

mistletoes.

WHORLED. See verticillate.

Scientific and Common Tree Names

Most names are from Little (1953) or

Critchfleld and Little (1966)

Abies amaUlis (Dougl.) Forbes—Pacific silver fir

Ahies halsamea (L.) Mill.—balsam fir

Ahies hracteata D. Don—bristlecone fir

Ahies concolor (Gord. & Glend.) Lindl.—white fir

Ahies durangensis Martinez

Ahies grandis (Dougl.) Lindl—grand fir

Ahies guatemalensis Rehd.

Ahies hickeli Flous & Gaussen

Ahies lasiocarpa (Hook.) Nutt.

var. lasiocarpa—subalpine fir

var. arizonica (Merriam) Lemm.—corkbark fir

Ahies magnifica A. Murr.

var. magnifica—California red fir

var. shastensis Lemm.—Shasta red fir

Ahies mexicana Martínez

Ahies oaxacana Martinez

Ahies pindrow Royale—Pindrow fir

Ahies procera Rehd.—noble fir

Ahies religiosae (H.B.K.) Schlecht. & Cham.

Ahies vejarii Martinez

Chamaecyparis thyoides (L.) B.S.P.—Atlantic whitecedar

Cupressus macrocarpa Hartw.—Monterey cypress

Juniperus hrevifolia Ant.

Juniperus communis L.

Juniperus drupácea Labill.

Juniperus excelsa Bieb.

Juniperus macropoda Boiss.

Juniperus oxycedrus L.

Juniperus phoenicia L.

Juniperus procera Höchst.

Juniperus rufescens Link

Juniperus sahina L.

Juniperus semiglobosa Regel

Juniperus seravschanica Komarov

Juniperus turcomanica B. Fedtsch

Larix europae DC.—European larch

Larix laricina (Du Roi) K. Koch—eastern larch

Larix leptolepis (Sieb. & Zuce.) Gord.—Japanese

larch

Larix lyallii Pari.—subalpine larch

Larix occidentalis Nutt.—^western larch

184
Picea abies (L.) Karst.—Norway spruce

Picea hreweriana S. Wats.—Brewer spruce

Picea chihuahuana Martínez

Picea engelmannii Parry—Engelmann spruce

Picea glauca (Moench) Voss—^white spruce

Picea mariana (Mili.) B.S.P.—black spruce

Picea mexicana Martínez

Picea pungens Engelm.—blue spruce

Picea ruhens Sarg.—red spruce

Picea sitchensis (Bong.) Carr.—Sitka spruce

Pinus alhicaulis Engelm.—^whitebark pine

Pinus aristata Engelm.—bristlecone pine

Pinus attenuata Lemm.—knobcone pine

Pinus ayacahuite Ehrenb.—Mexican white pine

Pinus halfouriana Grev. & Balf.—foxtail pine

Pinus hanksiana Lamb.—jack pine

Pinus hungeana Zuce.—lacebark pine

Pinus carihaea Morelet—Caribbean pine

Pinus cemhroides Zuce.—Mexican pinyon

Pinus chiapensis (Martínez) Andresen

Pinus contorta Dougl. ex Loud.—lodgepole pine

subsp. contorta—shore pine

subsp. bolanderi (Pari.) Critchf.

subsp. murrayana (Balf.) Critchf.

subsp. latifolia (Engelm. ex Wats.) Critchf.

Pinus cooperi C. E. Blanco—Cooper pine

Pinus coulteri D. Don—Coulter pine

Pinus culminicola Andresen & Beaman—Potosí pine

Pinus douglasiana Martinez—Douglas pine

Pinus durangensis Martinez—Durango pine

Pinus edulis Engelm.—pinyon

Pinus engelmannii Carr.—Apache pine

Pinusflexilis James—limber pine

Pinus greggii Engelm.—Gregg pine

Pinus griffithii McClelland—blue pine

Pinus hartwegii Lindl.—Hartweg pine

Pinus herrerai Martinez

Pinus Jeff reyi Grev. & Balf.—Jeffrey pine

Pinus lamhertiana Dougl.—sugar pine

Pinus lawsonii Roezl—Lawson pine

Pinus leiophylla Schiede &Deppe

var. leiophylla

var. chihuahuana (Engelm.) Shaw—Chihuahua

pine

Pinus lumhoUzii Robins. & Fern.—Lumholtz pine

Pinus maximartinezii Rzedowski—Martinez pine

Pinus michoacana Martinez—Michoacân pine

Pinus monophylla Torr. & Frém.—singleleaf pinyon

Pinus montezumae Lamb.—Montezuma pine

Pinus montícola Dougl.—western white pine

Pinus mugo Turra—Swiss mountain pine

Pinus muricata D. Don—bishop pine

Pinus nelsonii Shaw—Nelson pinyon

Pinus occidentalis Sw.—West Indian pine

Pinus oocarpa Schiede

Pinus palustris Mill.—longleaf pine

Pinus patula Schiede & Deppe—Mexican weeping

pine

Pinus pinaster Ait.—maritime pine

Pinus pinceana Gord.—Pince pinyon

Pinus pinea L.—Italian stone pine

Pinus ponderosa Laws.—ponderosa pine

var. ponderosa

var. scopulorum Engelm.

var. arizonica (Engelm.) Shaw—Arizona pine

Pinus pringlei Shaw—Pringle pine

Pinus pseudostrobus Lindl.

Pinus quadrifolia Pari.—Parry pinyon

Pinus radiaia D. Don—Monterey pine

Pinus resinosa Ait.—red pine

Pinus rudis Endl.

Pinus rzedowskii Madrigal and Caballero

Pinus sahiniana Dougl.—Digger pine

Pinus stroUJormis Engelm.—southwestern white pme

Pinus strobus L.—eastern white pine

Pinus sylvestris L.—Scotch pine

Pinus tabulaeformis Carr.—Chinese pine

Pinus tenuifolia Benth.

Pinus teocote Schiede & Deppe

Pinus torreyana Parry—Torrey pine

Pinus virginiana Mill-Virginia pine

Pinus washoensis Mason & Stockwell—Washoe pine

Pseudotsuga macrocarpa (Vasey) Mayr—bigcone

Douglas-fir

Pseudotsuga menziesii (Mirb.) Franco—Douglas-fir

var. menziesii

var. glauca (Beissn.) Franco

Thuja orientalis L.—Oriental arborvitae

Tsuga canadensis (L.) Carr.—eastern hemlock

Tsuga heterophylla (Raf.) Sarg.—western hemlock

Tsuga mertensiana (Bong.) Carr.—mountain hemlock

185
Specimens Examined

Herbaria

The dwarf mistletoe specimens that were examined

for these studies are tabulated in the following lists:

the host, location, collector, year of collection, and

herbarium where the collection is housed is given

for each specimen. The herbaria in which specimens

were examined and their abbreviations are listed

below. An asterisk indicates an abbreviation not in

the 1964 edition of Index Herbariorum (Lanjou and

Stafleul964).

ARIZ—University of Arizona, Tucson

ßLH—Cranbrook Institute of Science, Bloomfield Hills, Michigan

BM—British Museum, London, England

CAS—California Academy of Sciences, San

Francisco

COLO—University of Colorado, Boulder

CFB—Canadian Dep. Forestry, Calgary, Alberta

CS—Colorado State University, Fort Collins

DAVFP—Canadian Dep. Forestry, Victoria,

British Columbia

DS—Dudley Herbarium, Stanford University

EAP—Escuela Agrícola Panamericana, Tegucigalpa, Honduras

F—Field Museum of Natural History, Chicago

FPB*—U.S. Forest Service, Forest Pathology

Herbarium, Berkeley, California

FPF*—U.S. Forest Service, Forest Pathology

Herbarium, Fort Collins, Colorado

PPT*—Canadian Dep. Forestry, Forest Pathology, Toronto, Ontario

GH—Gray Herbarium, Harvard University

ILL—University of Illinois, Urbana

INIF*—Instituto Nacional de Investigaciones

Forestales, Mexico D. F.

JEPS—Jepson Herbarium, University of California, Berkeley

K—Royal Botanic Garden, Kew, England

MEXU—Institute de Biología, Mexico D. F.

MICH—University of Michigan, Ann Arbor

MIN—University of Minnesota, Minneapolis

MNA*—Museum of Northern Arizona, Flagstaff

MO—Missouri Botanical Garden, St. Louis

MSC—Michigan State University, East Lansing

NY—New York Botanical Garden, New York

OSC—Oregon State University, Corvallis

PH—Academy of Natural Sciences, Philadelphia, Pennsylvania

POM—Pomona College, Claremont, California

RM—Rocky Mountain Herbarium, University

of Wyoming, Laramie

RSA—Rancho Santa Ana, Claremont,

California

SRSC—Sul Ross State College, Alpine, Texas

UC—University of California, Berkeley

US—U.S. National Museum, Washington, D.C.

UT—University of Utah, Salt Lake City

UTC—Utah State University, Logan

WINF—Canadian Department of Forestry,

Winnipeg, Manitoba

WIS—University of Wisconsin, Madison

WSP—Washington State University, Mycological Herbarium, Pullman

WTU—University of Washington, Seattle

Z—Universität Zurich, Switzerland

Three collector abbreviations are used in the

specimens cited hst: (Hawksworth = H, Lightle = L,

and Wiens = W). An asterisk (*) after the collection

denotes a specimen used in the taximetric analyses.

The following abbreviations are used in the collection citations:

mountain = mtn.

National = Nat.

not dated = n.d.

range = R.

Reservation = Res.

route = Rte.

road = rd.

section = Sec.

Station = Sta.

township = T.

boundary = bdy.

canyon = can.

creek = cr.

collector unknown = col?

county = CO.

Experimental = Exp.

Forest = For.

junction = Jet.

miles = mi.

mount = mt.

Monument = Mon.

New World Taxa

la. ARCEUTHOBIUM ABIETINUMf. sp.

CONCOLORIS

Specimens Examined: on Abies concolor except as

noted.

UNITED STATES

ARIZONA

COCONINO CO. : Kaibab Plateau, Richards in 1938

(UC) and on A. lasiocarpa, Richards in 1938 (WTU).

Grand Can. Nat. Park, North Rim: Cape Royal rd.,

Gill FP 68297'' in 1934 (FPF); Fairview Point,

Peterson 37-61 in 1961 (FPF) and H 252 in 1962

(FPF) and on A. lasiocarpa, Peterson 39-61 in 1961

(FPF); Bright Angel Point, H 254 in 1962 (FPF);

Inspiration Point, W, 3180 in 1962 (COLO). Grand

Can. Nat. Park, South Rim: 1 mi. E of Grandview

Point, H & L 184 in 1962 (FPF) and H & Scharpf

7Ö^in 1964 (FPF).

186
CALIFORNIA

ALPINE CO.: 8 mi. SW of Silver Cr. on Rte. 4,

H & Scharpf 667 in 1964 (FPF). BUTTE CO.:

Jonesville, Copeland P4 in 1930 (ARIZ, CAS, MO,

POM, RM, RSA, UC, US, Z) and Copeland in 1929

(UC). CALAVERAS CO. : 12 mi. SW of Tamarack on

Rte. á,H& Scharpf 662 in 1964 (FPF). DEL NORTE

CO.: 0.25 mi.S of Oregon bdy. on O'Brien-Happy

Camp rd., H & Hinds SQQ"" in 1966 (FPF) and on A.

grandis, H & Hinds 998 in 1966 (FPF). ELDORADO

CO.: US 50 near Camino, Kuijt 1272 in 1957 (UC);

Tahoma, Kuijt 1340 in 1957 (UC) and on Pinus

contorta, rare, Kuijt 1341 in 1957 (UC). FRESNO

CO.: 2 mi.S of summit on Shaver Lake—Dinkey Cr.

rd., Quick 53-27 in 1953 (CAS); Huntington Lake,

Wall in 1919 (CAS); Bubbs Cr. and Charlotte Cr.,

Howell 15674 in 1940 (CAS). GLENN CO.: Bear

TFallow rd. near Alder Springs, Boyce 1906 in 1930

(FPF); Plaskett Ranger Sta. Boyce 1905* in 1930

(FPF). HUMBOLDT Co.: Lasseck's Peak, Kildare

2634 in 1926 (DS, UC) ; Humboldt Mill, vicinity of

Eureka, on Abies grandis, Tracy 1060 in 1901 (UC).

KERN CO. : Portuguese Pass, 7 mi. N of Summit

Guard Sta., W. 3610 in 1964 (FPF). LASSEN CO.:

Crater Mtn., 15 mi. W of Eagle Lake, Meinecke in

1914 (FPF) and Whitney 1703 in 1934 (UC).

MADERA CO. : 1 mi. E of Fish Camp, H 30 in 1954

(FPF); Bass Lake, Gill & Wright FP 68161 in 1932

(FPF) ; Mile High Point, 27 mi. from North Fork on

Mammoth Pool rd., H 937'' in 1966 (FPF); EUis

(now Benedict) Meadows, Zieman in 1917 (FPF).

MODOC CO.: Warner Mtns.: Cedar Pass, Alava in

1957 (UC); 2.5 mi. W of Patterson Guard Sta.,

H & Hinds 1002"" in 1966 (FPF). MARIPOSA CO.:

Yosemite, Hedgcock & Meinecke FP 4823 in 1910

(FPF) ; Fish Camp, Hedgcock & Meinecke FP 4834

in 1910 (FPF); 1 mi. E of Fish Camp, /Í* in 1963

(FPF). MENDOCINO CO.: Van Damme State

Park, on Abies grandis, Kuijt 1216 in 1957 (UC) and

H & Scharpf 855 in 1966 (FPF). NEVADA CO.:

2 mi. N of Truckee, W 3228 in 1962 (COLO, FPF);

Hobart Mills, Boyce FP 15981 in 1914 (FPF); Bear

Valley [6 mi. SE of Sierra City], Jepson in 1898

(JEPS); Gaston Ridge, 5 mi. N of Washington,

Howell 2459 n.d. (CAS). PLACER CO. : Trail to Mt.

ElHs, above Homewood, Schreiber 891 in 1933 (UC) ;

CarneUan Bay, Lake Tahoe, Blakely 3793 in 1960

(CAS). PLUMAS CO.: Chester, Meinecke FP 10754

in 1914 (FPF) ; Bucks Lake, Scharpf in 1963 (FPF) ;

4 mi. SE of Almanor Jet. on US 89, W 3232 in 1962

(COLO, FPF); 3 mi. SE of Almanor Jet. on US 89,

W 3233* in 1962 (COLO, FPF); Lassen Buttes,

Brown 675 in 1897 (MO, US) ; Taylorsville, Clemens

in 1919 (CAS). SAN BERNARDINO CO.: San

Bernardino Mtns.: 10 mi. N of Big Bear Lake,

Gill & Wright FP 68225 in 1931 (FPF); Barton

Flat, 3 mi. above Camp Ángelus, W. 3210 in 1962

(COLO, FPF) ; Lost Creek, Mum & Johnston 8576

in 1924 (POM); Barton Flats, Ross 1917 in 1937

(POM); Inspiration Point, Perkins in 1917 (ILL).

SHASTA CO.: 22 mi. N of Lassen Nat. Park

entrance on Rte. 89, Kuijt 1365 in 1957 (UC); 4 mi.

N of Lassen Nat. Park entrance on Rte. ^^,H &W

648 in 1964 (FPF); Rte. 89 at Pondosa Jet., on

Pinus lambertiana, Kuijt 1369 in 1957 (UC).

SIERRA CO.: Sierraville, Meinecke & Boyce FP

17193 and FP 17194 in 1915 (FPF). SISKIYOU

CO.: Mt. Shasta, Sisson South Trail, Cooke 16132

in 1941 (COLO, MO, PH); Can. Cr., Hedgcock FP

1858 in 1917 (FPF); N side Cascade Gulch, Mt.

Shasta, Cooke 17722 in 1947 (WTU); Bartle-Whitehorse rd., Newcomb 156 in 1957 (UC); Sisson South

Trail, Cooke 13574 in? (DS); Shackelford Cr. [8 mi.

W of Greenview], Butler 270 in 1908 (UC); Joe Cr.,

10 mi. S of Copper, Oregon, H & W 635 in 1964

(FPF); 5 mi. E of US 99 on Mt. Shasta rd., H &W

641 in 1964 (FPF) ; Along trail on N side of Black

Butte, H & W 643 in 1964 (FPF); Base of Mt.

Shasta, Engelmann in 1880 (MO); Trestle rd. to

Hilts, N of Oak Knoll Ranger Sta., Quick 60-21 in

1960 (CAS). TEHAMA CO.: 2 mi. N of Hole-inGround Campground, S of Mineral, Kuijt 1503 in

1958 (UC). TULARE CO.: Hollow Log Camp,

[near General Grant], Dudley in 1900 (DS); Sequoia

Nat. Park, 1 mi. N of Lodgepole, W. 3613 in 1964

(FPF). TUOLUMNE CO.: Cow Cr., Gill FP 68060

in 1932 (FPF), Quick in 1966 (FPF) and on Pinus

lambertiana, Quick in 1966 (FPF); Strawberry,

Wright FP 68052 in 1931 (FPF); Long Barn,

Wright FP 68115 and FP 68116 in 1932 (FPF);

Pinecrest, Gill FP 68224 in 1932 (FPF) and on

Pinus lambertiana, Gill FP 68224 in 1932 (FPF);

Tuolumne Grove, Peterson 63-139 in 1963 (FPF); 7

mi. E of Long Barn, W 3223 in 1962 (COLO, FPF);

5 mi. E of Long Barn, Kuijt 1431 in 1957 (UC);

5.1 mi. E of Nat. For. bdy. on Rte. 120, Kuijt 1393

in 1957 (UC); Yosemite Nat. Park, Tuolumne Can.,

CZemensin 1919 (CAS).

NEVADA

CLARK CO.: Charleston Mtns.: Little Falls,

Clokey 5433 in 1935 (CAS, DS, FPF, RM, UC, US,

WTU) and 5434 in 1935 (CAS, DS, FPF, RM, US,

WTU) ; Charleston Park, W 3204 and 3033 in 1962

(COLO, FPF) ; Deer Cr. Grade, 3 mi. E of Lee Can.,

Train 2154 in 1918 (ARIZ, UC); Lee Can., Heller

11060 in 1913 (DS, MO, PH, UC); Deer Cr. Campground, Breedlove 1109 in 1961 (DS); without

locality, Jaeger in 1926 (POM) ; Lee Can., 5 mi. W of

Deer Cr. Jet., H & Scharpf 674 in 1964 (FPF);

Kyle Can., 2.5 mi. W of Ranger Sta., H & Scharpf

676* in 1964 (FPF). WASHOE CO.: Vicinity of

Reno, Hitchcock 559 in 1913 (US).

187
OREGON

CURRY CO.: S of Humbug Mtn., on Abies grandis,

Ferris 7828 in 1929 (DS); Grizzly Mtn., 0.5 mi. Eof

Gold Beach, on Abies grandis, Theisenin 1965 (FPF),

DESCHUTES CO.: McKenzie Pass, on A. grandis,

Gill FP 68186* in 1932 (FPF); 9 mi. W of Sisters,

on A. grandis, W 32U in 1962 (COLO, FPF) ; 7.5 mi.

W of Sisters on Rte 242, on A. grandis, H & W 613

in 1964 (FPF) and H & Hinds 990 in 1966 (FPF) ; 14

mi. NW of Sisters on US 126, on A. grandis, H &W

615 in 1964 (FPF). JACKSON CO.: 10 mi. E of

Butte Falls, Graham in 1964 (FPF) : Long John Cr.

[SSW of Ashland], Wheeler 3005 in 1934 (CAS, POM,

WTU) ; 11 mi. SW of Ashland on Copper rd., H&W

637 in 1964 (FPF); Summit of Siskiyou Mtns., near

Siskiyou Sta., on A, grandis, Abrams 12130 in 1927

(DS); W Fork of Evans Cr., 18 mi. NE of Grants

Pass, Graham in 1966 (FPF); 8 air mi. S of Ashland,

Graham in 1965 (FPF); 18 air mi. E of Ashland,

Howard in 1964 (FPF); Fish Lake rd., 12 mi. SE of

Butte Falls, Graham in 1965 (FPF). JEFFERSON

CO.: 3 mi. E of Santiam Pass on US 20, on A.

grandis, H & Seharpf 1257 in 1969 (FPF). JOSEPHINE CO. : Flat Top Mtn., 24 air mi. W of Grants

Pass, Grahmn in 1963 (FPF), H & W 632 in 1964

(FPF), and Bijnum in 1967 (FPF) and on Pieea

breweriana, Graham in 1963 (FPF), H óc W 631 in

1965 (FPF), and Bynumm 1967 (FPF). KLAMATH

CO.: Hamner Butte, [S of Davis Lake] on A.

grandis, Childs 28 in 1939 (OSC) and Englerth FP

91031 in 1939 (OSC); 9 mi. N of Fort Klamath on

Rte. Çi2, H & W 626 in 1964 (FPF); 31 mi. W of

Klamath Falls on Rte. ^^, H & W 629 in 1964

(FPF); vicinity of Crater Lake, Weir 2^29 in 1916

(FPF, ILL) and on A. amabilis. Weir 2^23 in 1916

(FPF, ILL). LAKE CO.: Silver Lake, Jaenicke FP

21285 in 1916 (FPF); W Fork of Silver Cr., 12 mi.

SW of Silver Lake, Stewart in 1968 (FPF) ; 1 mi. N

of Bly Summit, 8 mi. SW of Beatty on Rte. 140,

^cfcTF574inl966(FPF).

UTAH

KANE CO. : 8 mi. S of Navajo Lake on Zion Park rd.,

W 4122 m 1966 (FPF, UT).

WASHINGTON

KLICKITAT CO.: Falcon Valley, on A. grandis,

Suksdorf 2246 in 1893 (FPF, MO, UC, US); Snowden, ca. 12 mi. NE of White Salmon, on A. grandis,

Graham'' in 1964 (FPF) ^nd H & W 592 in 1964

(FF'F).

lb. ARCEUTHOBIUM ABIETINUM f. sp.

MAGNIFICAE

Specimens Examined: on Abies magnifica (including

var. shastensis) except as noted.

UNITED STATES

CALIFORNIA

AMADOR CO.: E of Lower Bear Res., Quick 5^-87

in 1954 (CAS). BUTTE CO.: Summit of Humboldt

rd. near Jonesville, Copeland in 1928 (UC); '^Big

Summit,'' Jonesville, Copeland in 1931 (UC).

CALAVERAS CO. : 10 mi. above Dorrington, Wright

FP 68050 in 1931 (FPF) and FP 68123 in 1932

(FPF); 8 mi. SW of Tamarack on Rte. 4:, H &

Scharpf 663 in 1964 (FPF). EL DORADO CO.: S

end of Echo Lake, Howell 22902 in 1946 (CAS).

HUMBOLDT CO.: Trinity Summit, E of Corral

Prairie, Tracy 10571 in 1932 (UC) ; South Fork Mtn.,

near Blake Lookout, Tracy 8950 in 1930 (JEPS,

WTU). FRESNO CO.: Huntington Lake, Miller

FP 98112 in 1965 (FPB). MADERA CO.: Haskell

Meadow, Miller FP 97956 in 1910 (FPB). KERN

CO.: Greenhorn ]Mtns., N side Sundry Peak,

Howell 388U in 1962 (CAS). MARIPOSA CO.:

5 mi. E of Fish Camp, H in 1963 (FPF). NEVADA

CO.: Hobart Mills, Boyce FP 15982 in 1914 (FPF,

ILL) ; 5 mi. E of Hobart Mills, Gill FP 68223 in 1931

(FPF). PLUMAS CO.: Grizzly Summit, SW of

Bucks Lake, Scharpf FP 98107 in 1964 (FPB);

Claremont Peak near Quincy, Weaiherby 1674 in 1955

(CAS, RM, RSA). SHASTA CO.: Lassen Nat.

Park on main rd., 10 mi. E of N entrance, H&W

650'' in 1964 (FPF). SISKIYOU CO.: near head of

E Fork of Indian Cr., Evans in 1912 (FPB); 0.5 mi.

E of Siskiyou Nat. For. bdy. on Happy CampO'Brien rd., 14 air mi. NNW of Happy Camp,

H & Hinds 1000* in 1966 (FPF); Kidder Cr. [ca. 11

mi. WSW of Greenview], Gill & Sargent FP 68181"^

in 1932 (FPF). TRINITY CO.: South Fork .Altn.,

near Hyampom, Boyce 30 in 1914 (FPF); South

Fork Mtn., Parks cfc Tracy 11532 in 1941 (RM,

RSA, UC, US, WTU). TULARE CO.: Bald Mtn.,

Kaweah River Valley, Dudley 1395 in 1896 (DS);

Roaring River, Meinecke in 1910 (FPB); Mosquito

Cr. near Mineralking, Hopping in 1913 (FPB); 17

mi. NW of Lodgepole, W 3615* in 1964 (FPF).

TUOLUMNE CO.: Pinecrest, Scharpf* in 1963

(FPF); Yosemite Nat. Park, 7 mi. E of Crane Flat

Jet. on Tioga Pass rd., H 938* in 1966 (FPF).

COUNTY UNCERTAIN: Lassen Nat. Park,

Boldenbeckin 1910 (DS).

OREGON

JOSEPHINE CO.: Flat Top Mtn., ca. 24 air mi. W

of Grants Pass, on A. procera; Bynum in 1967 (FPF) :

Oregon Caves Nat. Mon., Sike Mtn. Trail, Root &

Goading FP 68274* in 1932 (FPF) ; near Bolen Mtn.,

S of Oregon Caves, Theisen in 1966 (FPF).

KLAIMATH CO. : Bay on W side of Upper Klamath

Lake, on A. procera. Peck 9525 in 1920 (WTU);

near Crater Lake, on A. procera. Weir 2427 in 1916

188
(FPF, ILL). COUNTY UNCERTAIN: AshlandKlamath Falls wagon rd., Weir 2^26 in 1916 (ILL).

2. ARCEUTHOBIUM ABIETIS-RELIGIÖSAE

Specimens Examined:

MEXICO

JALISCO: NW Slopes of Nevado de Colima, on

Abies sp., McVaugh & Wilher 10130 in 1949 (MICH).

^MEXICO: Ixtaccihuatl, on Abies religiosa, Purpus

6491 in 1912 (MO, UC, US) ; Popocatepetl, on Abies

religiosa, Balls B4228 in 1938 (BM, K, UC, US);

17 km E of Amecameca, on Abies religiosa, H & W

SrS"" in 1963 and 1229 in 1969 (FPF); District

Temascaltepec, vicinity of Mesón Viejo, on Abies

religiosa, Hinton 3278 in 1933 (K, US) ; Sultepec rd.

10 mi. S of Nevado de Toluca Jet., on Abies religiosa,

Peterson 68-91* in 1968 (FPF). NUEVO LEON:

Cerro Potosí, on Abies vejarii, H & W 394"^, 396"",

3359 in 1963 (COLO, FPF); Municipio de Derrumbadero, in moister canyons above Santa Elena,

San Juanito to San Miguel and E as far as Los Toros,

on Abies sp., Mueller in 1935 (MO).

3. ARCEUTHOBIUM AMERICANUM

Specimens Examined : Host is Pinus contorta except

as noted. For these records the varieties murrayana and latifolia are not distinguished

because these can be determined by the geographic area involved (Critchfield 1957).

CANADA

ALBERTA

N fork of N branch of Saskatchewan River, Broten

795 in 1908 (PH); Kananaskis For. Exp. Sta.,

Muir 64-327 in 1964 (FPF) and on Pinus sylvestris,

Powell in 1968 (CFB, FPF); Banff Nat. Park

Bankhead, Brown 794 in 1906 (MO,PH, US). 2 mi

E of Lake Louise, H & Baranyay 90 in 1961 (FPF)

4 mi. W of Lake Louise, Petty 64-568 in 1964 (FPF)

Eisenhower Jet. ; on Picea glauca, H óc Baranyay 92

in 1961 (FPF); 1.5 mi. E of Eisenhower Jet., H &

Baranyay 91 in 1961 (FPF); Cypress Hills, W of

Spring Cr. Ranger Sta., Breitung 5586 in 1947 (MO) ;

Cypress Hills, Elkwater, Lawrence in 1967 (FPF,

WINF); Jasper Nat. Park, Snaring River, on

Picea glauca, Bourchier in 1954 (CFB, FPF);

7 mi. W of Cold Lake, on Pinus banksiana, Wilkinson in 1960 (CFB, FPF) ; 38 mi. SE of Conklin, on

Pinus banksiana, Wilkinson in 1961 (CFB, FPF);

20 mi. NW of Slave Lake Town, on Pinus banksiana,

Smüh in 1962 (CFB, FPF); 2 mi. W of Fort Assiniboine, on Pinus banksiana, Emond & Laijton

in 1966 (CFB, FPF); 23 mi. SW of Clearwater

Ranger Sta., Smüh in 1966 (CFB, FPF); 3 mi. SW

of Clearwater Ranger Sta., Smith in 1965 (CFB,

FPF) ; 2 mi. N and 2 mi. E of St. Vincent, on Pinus

banksiana, Layton in 1966 (CFB, FPF) ; 4 mi. NW of

Rocky, Smith in 1966 (CFB, FPF).

BRITISH COLUMBIA

Guichon Cr., 13 mi. S of Savona, Hitchcock &

Martin 7425 in 1941 (COLO, DS, MO, POM, RM,

RSA, UC, WTU); 10 mi. N of Clinton, Calcler,

Parmalee & Taylor 16644 m 1956 (COLO, US);

Prince George, Calcler, Savile & Ferguson 14363 in

1954 (WTU); Kleena Kleen P. O., 50° 57^N, 124°

52' W, Calder, Parmalee & Taylor 19159 in 1956

(DS, WTU); 3 mi. S of Ta Ta Cr., N of Cranbrook,

Calder & Savile 11374 in 1953 (UC, US); near

Allenby, McCallum in 1940 (UC); Settlers rd.

Kootenny Nat. Park, Morf 64-86 in 1964 (FPF)

20 mi. NW of Kelowna, Stewart in 1965 (FPF)

2.5 mi. SW of Telkwa, Calder, Savile & Ferguson

15273 in 1954 (US); Bonaparte River, Macoum in

1889 (US); Salmon Arm, Weir 8444 m 1913 (ILL)

25 mi. SE of Kelowna, Kuijt in 1954 (DAVFP, FPF)

Perry Cr., Kuijt 598 in 1954 (DAVFP, FPF)

Moyie River, Kuijt in 1954 (DAVFP, FPF)

between Yahk and JNIoyie Lake, Kuijt in 1954

(DAVFP, FPF); 33 mi. N of Rock Cr., Kuijt in 1954

(DAVFP, FPF) ; 1 mi. S of Beaverdell, Kuijt in 1954

(DAVFP, FPF) ; Carmi, Kuijt 591 and 592 in 1954

(DAVFP, FPF); McCulloch, Kuijt 589 in 1954

(DAVFP, FPF); Thompson River, Kuijt in 1954

(DAVFP, FPF); 8 mi. N of Kimberly, Kuijt 528

in 1954 (DAVFP, FPF) ; 45 mi. S of Radium, Kuijt

in 1953 (DAVFP, FPF) ; Windermere, Kuijt 535 in

1953 (DAVFP, FPF); Kootenay Park, Kuijt in

1954 (DAVFP, FPF) ; 12 mi. N. of Clinton, Kuijt

in 1954 (DAVFP, FPF); Jesmond, Kuijt in 1954

(DAVFP, FPF); Pavilion Mtn., Kuijt in 1954

(DAVFP, FPF); Eholt, Kuijt in 1954 (DAVFP,

FPF); Balfour, Kuijt in 1954 (DAVFP, FPF);

Manning Park, 10 mi. E of Allison Summit, Kuijt

in 1954 (DAVFP, FPF); Manning Park, Sunday

Summit, Kuijf" in 1954 (DAVFP, FPF); Moyie

Lake, Kuijt in 1954 (DAVFP, FPF); Mt. Robson

Park, Fraser River, Kuijt in 1954 (DAVFP, FPF) ;

Mt. Robson Park, Yellowhead, Kuijt 536 in 1953

(DAVFP, FPF); lOO-^NIile House, Simms in 1955

(DAVFP, FPF); :Manning Park, near E bdy,

Sugden in 1960 (DAVFP, FPF); Alexis Cr., Simms

in 1955 (DAVFP, FPF).

MANITOBA

Near Cowan, on P. banksiana, Riley 47-208 in 1947

(MO, RM); Victoria Beach, on P. banksiana,

Jackson in 1924 (ILL) and Love & Love 5740 in 1953

(US); Belaire Provincial For., on P. banksiana,

Lawrence in 1967 (FPF, WINF); Iskwasum Lake,

on Pinus banksiana. Laut in 1968 (FPF, WINF);

Black Island, Lake Winnipeg, on Pinus banksiana,

Laivrence in 1968 (FPF, WINF).

189
SASKATCHEWAN

Prince Albert, on planted P. contorta, Beveridge

in 1968 (FPF, WINF) ; MacDowall, on P. hanksiana,

Blumer 5177 in 1913 (FPF); Lac la Ronge, on P.

banksiana, McLeocl in 1965 (FPF, WINF); Cumberland House on P. banksiana, Craivford in 1967

(FPF, WINF); S shore of Lake Athabaska, on P.

banksiana, McLeod in 1966 (FPF, WINF); Lac la

Loche, on P. banksiana, Rentz in 1967 (FPF, WINF);

Mile 10.7 on Cumberland rd., on P. banksiana, Craivford in 1968 (FPF, WINF); Mile 15 on Turner Lake

rd., on P. banksiana, Laut in 1968 (FPF, WINF);

He a' la Crosse, on P. banksiana, Laut in 1968 (FPF,

WINF).

UNITED STATES

CALIFORNIA

ALPINE CO.: Sandy Flat, N Stanislaus River

[4 mi. NE of Alpine], Stanford 629 in 1927 (RM);

5 mi. W of Alpine Co. bdy. on Rte. 4, on P. ponderosa, Kuijt 1411 in 1957 (UC); 0.5 mi. W of grade

summit on Rte. 4, on P. ponderosa, Kuijt 1412 in

1957 (UC). CALAVERAS CO. Rte. 4 near Alpine

Co. bdy., Peterson 6'5-134* in 1963 (FPF); 3 mi. SW

of Tamarack on Rte. 4:, H & Scharpf 664 in 1964

(FPF). EL DORADO CO.: US 50 at Pyramid Cr.,

Kuijt 1513 in 1958 (UC); Lily Lake, Glen Alpine

Can., Abrams 12753 in 1903 (DS, MO, POM, UC);

16 mi. S of Tahoe City, on P. ponderosa, Kuijt 1335 in

1957 (UC); 1.1 mi. N of Meyers on US b{), H & W

838 in 1966 (FPF); 9 mi. E of Kyburz on US 50,

H&W839 in 1966 (FPF). FRESNO CO. : Huntington Lake, Solbrig 2420 in 1957 (UC) and Perkins in

1920 (ILL); House Meadows, N fork of Kings

River [5 mi. W of Wishon Dam], Hall & Chandler 426

in 1900 (UC) ; VermilHon Valley, Quibell & Quibell

2641 in 1953 (RSA). KERN CO.:Havileah [7 mi. S

of Bodfish], Coville & Funston 1596 in 1891 (FPF;

US). LASSEN CO.: Bridge Cr., 7-8 mi. SW of

Eagle Lake, Whitney 1717'm 1934 (UC). MADERA

CO. : Red's Meadow to Rainbow Falls, Raven 3678

in 1951 (RSA); Devils Post Pile Nat. Mon., H &

Scharpf 670 in 1964 (FPF). MARIPOSA CO.:

Tenaya Lake, Root in 1919 (FPF, ILL) and H 939

in 1966 (FPF) ; Little Yosemite Valley, Bolander 5095

in 1866 (MO, UC, US), Rodin 877 in 1946 (UC);

Merced Lake Trail, Yosemite, Schreiher 1948 in 1935

(UC); Upper Yosemite Valley, Lenimon in 1878

(MO). MONO CO.: 4.3 mi. W of Jet. of Rtes. 120

and 395, Kuijt 1390 in 1957 (UC) ; 2 mi. W of Lee

Vining, on P. jeffreyi, Peterson 63-129 in 1963 (FPF) ;

W of Lee Vining on Tioga Pass rd., Krebill & Nelson

468 in 1967 (FPF). NEVADA CO.: 4 mi. from

Truckee, Wright FP 68107 in 1932 (FPF); Hobart

Mills, Gill FP 68063 in 1931 (FPF); Donner State

Park, H in 1963 (FPF); Donner Lake, Dudley in

1893 (DS) and Soniie in 1898 (US); near Truckee,

Sonnem 1898 (UC), Wright FP68048m 1931 (FPF),

Gill FP 68227 in 1931 (FPF). PLACER CO.: Near

Cisco, Boyce 40 in 1919 (FPF); 7 mi. W of Soda

Springs, W 3226 in 1962 (COLO, FPF); N Tahoe

Meadow, Lake Tahoe, Schreiber 776 in 1932 (UC) ;

3.5 mi. N of Tahoe City on Rte. 89, Kuijt 1332

in 1957 (UC); 8.5 mi. W of Soda Springs, on P.

ponderosa, Kuijt 1330 in 1957 (UC) ; 10 mi. N of

Tahoe City on Rte. 89, on P. ponderosa, Kuijt 1331

in 1957 (UC); 9.2 mi. N of Tahoe City on Rte. 89,

on P. jeffreyi, Kuijt 1343 in 1957 (UC). PLUMAS

CO.: Chester, Meinecke FP 17053 in 1914 (FPF);

18 mi. N of Greenville, Kuijt 1350 in 1957 (UC);

Graegle Cr. near Lake Center Camp, Mason 1079

in 1924 (JEPS, UC); 8 mi. W of Rte. 89 on Webber

Lake rd., H & W 657 in 1964 (FPF); Buck^s Ranch,

Meinecke in 1920 (FPF); Little Grizzly Valley, on

P. jeffreyi, Boyce in 1915 (FPF). SHASTA CO.:

Thousand Lake Basin, Peirson 10136 in 1932 (RSA);

Lassen Nat. Park on main rd. 10 mi. E of north

entrance, H & W 651 in 1964 (FPF) ; headwaters of

Hat Cr., Eggleston 7459 in 1911 (MO, US). SIERRA

CO.: Gold Lake, Gill FP 68O46 in 1931 (FPF); 9.5

mi. N of Truckee, Kuijt 1344 in 1957 (UC) ; Webber

Lake, ''Loran'', n.d., (UC). SISKIYOU CO.:

Military Pass, NE side Mt. Shasta, Cooke 16034 in

1941 (DS, UC) ; Mt. Shasta, Sisson, Pendleton in 1919

(FPF, ILL); 3 mi. S of Tennant, Boyce in 1914

(FPF) ; N fork of Sacramento River, Raven 10456 in

1959 (CAS). TEHAMA CO.: Mineral, Meinecke

FP 97934 in 1911 (FPB); Mineral Campground,

near Ranger Sta., Kuijt 1358 in 1957 (UC) ; Deer Cr.,

1.5 mi. S of Jet. of Rtes. 32 and 36, Kuijt 1501 in

1958 (UC); Deer Cr., 1 mi. E of Rtes. 89 and 36 on

Rte. 32, Waters 242 in 1958 (JEPS); 13 mi. W of

Rtes. 36 and 89 at Lake Almanor, on P. ponderosa,

Kuijt 1354 & 1355 in 1957 (UC) ; 7 mi. W of Rtes.

36 and 89 at Lake Almanor, on P. ponderosa, Kuijt

1352 in 1957 (UC). TULARE CO.: Long Meadow,

Peterson 63-149* in 1963 (FPF) ; Bakeoven Meadows, S fork Kern River, Howell 27038 in 1950 (DS,

UC, US) ; Jet. Meadows, Kern River, Raven 8349 in

1955 (UC); Sequoia Nat. Park; 13 mi. NW of

Lodgepole, W 3614 in 1964 (FPF) ; Funston Camp

[11 mi. SE of Mineral King], Peirson 1720 in 1919

(RSA); ridge between Monache and Bakeoven

Meadows, Mum 15220in 1950 (RSA). TUOLUMNE

CO. : Pinecrest, Wright FP 68114 in 1932 (FPF) and

Gill FP 68229 in 1931 (FPF). COUNTY UNCERTAIN: N California border, Salmon Nat. For.,

GalbreathFP 1691 in 1909 (FPF).

190
COLORADO

BOULDER CO.: 3 mi. S of Ward on Rte. 160,

H & Gill in 1958 (FPF); 1.5 mi. W of Rte. 160

on Brainard Lake rd., H 1,18 in 1963 (FPF);

Allenspark, Johnston & Thompson FP 21,881 in 1917

(FPF) ; Niwot Hill, near Ward, Schmoll 498 in 1922

(COLO); Pine Glade School, Ramaley 3754 in 1907

(COLO): Ward, Newman in 1922 (COLO); near

Nederland, W 2966 in 1961 (COLO) ; Allenspark, on

P. ponderosa, H 828 in 1965 (FPF). CHAFFEE CO. :

Alpine, Branclegee in 1880 (FPF) ; 8 mi. W of Buena

Vista, H 445 in 1963 (FPF) ; Forest City, Branclegee

12201 in 1880 (MO). CLEAR CREEK CO.: Silver

Plume, Bethel FP 27111 in 1918 (FPF); 0.5 mi. SE

of Alice, H &W 283 in 1963 (FPF); 4.4 mi. E of

Berthoud Pass, Greene & Richter 112 in 1962

(COLO, FPF); Clear Cr. Valley, Engelmann in 1874

(MO, UC); Empire, Patterson 297 in 1892 (MO, UC,

US) ; 3 mi. W of Bergen Park, H 566 in 1964 (FPF) ;

above Empire, Engelmann in 1874 (MO); Bergen

Park, Greene in 1873 (MO); Empire, Torrey in 1872

(MO); Georgetown, Wootson 23 in 1873 (US).

DOUGLAS CO.: Rampart Range rd., 5 mi. S of

Long Hollow Jet., H 205 in 1962 (FPF). EAGLE

CO.: Mt. of the Holy Cross, Coulter 7655 in 1873

(MO, PH, US) and Hedgcock FP 624 in 1909 (FPF) ;

Red Cliff, Gayman FP 26295 in 1917 (FPF) ; 5 mi. E

of Vail on US 6, H 947 in 1966 (FPF). GILPIN CO. :

Rollinsville, Wheeler 330 in 1901 (COLO, RM);

Rollinsville, on P. ponderosa, Hedgcock FP 22598 in

1916 (FPF); Tolland, Bethel 317 in 1921 (CS),

Gayman & Hinson in 1917 (FPF), Hedgcock FP 22592

in 1916 (FPF, UC) ; Lake Eldora, Hedgcock FP 22549

in 1916 (FPF). GRAND CO. : Granby, Bethel in 1906

(CS), Benson 4907 in 1933 (POM); Fraser, Hedgcock

& Johnston FP 26068 in 1917 (FPF); Idlewild,

Hedgcock FP 1649 in 1909 (FPF); 6 mi. W of

Fraser, H 81 in 1961 (FPF) ; 18 mi. S of Parshall

on Leal rd., Hinds & H in 1960 (FPF); 8.3 mi.

N of Fraser on US 40, Greene & Richter 113 in

1962 (COLO, FPF) ; 11.3 mi. N of Granby on US 34,

Greene & Richter 114 in 1962 (COLO, FPF); Grand

Lake, col? in 1881 (MO) and Shear & Bessey 5317

in 1898 (US). GUNNISON CO.: 5 mi. E of Taylor

Park on Cottonwood Pass rd., H 945 in 1966 (FPF) ;

2.5 mi. W of Monarch Pass on US 50, Hinds 63-13

in 1963 (FPF). JACKSON CO.: North Park, Barber

in 1874 (US); E side of Buffalo Pass, H 414 in 1963

(FPF); Silver Cr., 4 mi. S of Gould, H 415"" in 1963

(FPF); Silver Cr., 4 mi. SE of Gould, on Picea

engelmannii, H 934 m 1966 (FPF); 1 mi. N of

Willow Cr. Pass, H 4I6 in 1963 (FPF); Big Cr.

Lake, H 830 in 1965 (FPF) ; near Wyoming line on

Rte. 127, Davidson & Gill FP 89991 in 1952 (FPF).

JEFFERSON CO.: Redskin Cr., Hill FP 68279 in

1932 (FPF). LAKE CO.: Leadville, Clokey 2684 m

1916 (DS, FPF, ILL, RM, UC, US) and Schedin &

Schedin, n.d. (RM). LARIMER CO.: Longs Peak

Inn, Cooper 23 in 1908 (RM) ; 9.8 mi. W of Pingree

Park rd. on Crown Point rd., H 4IÏ in 1963 (FPF);

Rocky ^Mountain Nat. Park, Glacier Basin, Smith

in 1937 (CS); 5 mi. W of Redfeather Lakes, H 190

in 1960 (FPF) ; 4 mi. W of Redfeather Lakes, H 407*

in 1963 (FPF) ; Longs Peak Valley, Kiener 5686 in

1937 (COLO); Buckhorn Mtn., Stove Prairie

Summit, H 553 in 1964 (FPF) ; 2 mi. W of Buckhorn

Ranger Sta., H 560 in 1964 (FPF); Bear Gulch, 6

mi. W of Masonville, H 711 in 1964 (FPF); Stove

Prairie Summit, on P. ponderosa, H 192 in 1961

(FPF); 3 mi. E of Creedmore Lakes rd., on P.

ponderosa, H 261 in 1962 (FPF) ; Boulder Ridge, on

P. ponderosa, H 263 in 1962 (FPF), Tie SidingCherokee Park rd., on P. ponderosa, H 286 in

1962 (FPF); 1 mi. W of Redfeather lakes, on P.

ponderosa, H 4O8 in 1963 (FPF); Devils Cr. on

Sand Cr. Pass rd., on P. ponderosa, Schacht in 1962

(FPF); 0.5 mi. E of Buckhorn Ranger Sta., on P.

ponderosa, H 555 in 1963 (FPF); Pennock Cr.,

2 mi. S of Pinge Park rd., on P. ponderosa, H 563 in

1964 (FPF) ; Bear Gulch rd., 6 air mi. W of Masonville, on P. ponderosa, H 712 in 1964 (FPF) ; 4 mi.

W of Redfeather Lakes, on P. fiexilis, H 258 in 1962

(FPF) ; 3 mi. W of Redfeather Lakes, on P. fiexilis, H

409 in 1963 (FPF); 4.8 mi. W of Pingree rd. on

Crown Point rd., on P. fiexilis, H 412 in 1963 (FPF) ;

Pennock Cr. 2 mi. S of Pingree Park rd., on P. fiexilis,

H 561 in 1964 (FPF) ; 5 mi. W of Redfeather Lakes,

on Picea engelmannii, H 189 in 1961 (FPF) ; 4 mi. W

of Redfeather Lakes on Picea engelmannii, H 4O6 in

1963 (FPF) ; SE of Chambers Lake, on Picea engelmannii, Schacht in 1962 (FPF); N Fork Poudre

River, ca. 7 mi. W of Redfeather Lakes on Picea

engelmannii, H 559 in 1964 (FPF); 1 mi. E of

Buckhorn Ranger Sta., on Picea pungens, H 554 in

1964 (FPF). MOFFAT CO.: Cold Spring Mtn.,

0.25 mi. E of Utah and 1.5 mi. S of Wyoming, Brown

in 1965 (FPF). PARK CO.: Fairplay, Gill FP 68140

in 1932 (FPF) ; 3 mi. NW of Fairplay, H 447 in 1963

(FPF) ; Summit of Kenosha Pass on US 285, H 448

in 1963 (FPF); 4.4 mi. N of Grant on Geneva Basin

rd., H 450 in 1964 (FPF) and on P. aristata, H 449

in 1963 (FPF) ; 0.3 mi. E of Santa Maria, H 451 in

1963 (FPF) and on P. ponderosa, H 452 in 1963

(FPF); Payne Gulch [near Bailey], on P. fiexilis,

Hill FP 68236 in 1932 (FPF). ROUTT CO. : Walton

Cr. Campground, Hinds 63-21 in 1963 (FPF);

Steamboat Springs, Galpin in 1908 (FPF); Hahn's

Peak, Edmonston FP 188 in 1908 (FPF) and on

Picea engelmannii, Edmonston FP 199 in 1908 (ILL) ;

0.25 mi. S of Trout Cr. Campround (T. 3 N., R. 86

W., Sec. 18), H, Andrews & Buchanan in 1961 (FPF) ;

9 mi. E of Clark, H 191 in 1961 (FPF); 4 mi. SE of

Clark, H 413 in 1963 (FPF) ; Seedhouse, E of Clark,

Weber 6725 in 1951 (COLO, WTU); 8 mi. E of

191
Clark, Weber 6093 in 1951 (COLO); near Steamboat

Springs, Greene & Richter 115 in 1962 (COLO).

SUMMIT CO. : Breckenridge, Seay FP 33403 in 1917

(FPF), Mackenzie 78 in 1901 (MO, PH, RM, US);

Dillion, Shear 3764 in 1896 (RM); near Dillon, W

3260 in 1962 (COLO); Mt. Baldy near Breckenridge,

Henderson in 1897-1909 (MO). COUNTY UNCERTAIN: Sawatch Range, Brandegee in 1880

(PH, UC); South Park, Wolf & Rothrock 70 in 1873

(PH) and Hall & Harbour 574 in 1862 (MO);

Middle Park, Patterson in 1875 (PH) and Torrey in

1872 (MO) ; Latitude 39-41°, Parry 574 in 1862 (US).

IDAHO

ADAMS CO. : Evergreen, Hedgcock FP 1908 in 1909

(FPF). BEAR LAKE CO.: 7 mi. NW of Liberty on

Rte. 36, H & Hinds 1007 in 1966 (FPF). BENEWAH

CO.: 1 mi. S of Emida, Owenbey & Oiuenbey 2023 in

1940 (ARIZ, DS, MO, POM, UC, US, WTU); S end

of Coeur d' Alene Lake, col.? in 1891 (UC). BLAINE

CO. : Near source of Wood River, Henderson 3581 in

1895 (US). BOISE CO.: Idaho City, Weir 8456 in

1917 (ILL). BONNER CO.: Priest Lake, Hedgcock

& Weir 8456 in 1917 (ILL). BONNER CO.: Priest

Lake, Hedgcock & Weir FP 9489 in 1911 (FPF) and

Piper 3701 in 1901 (US). BONNEVILLE CO.: SW

of Victor, Peterson 62-14 in 1962 (FPF); 11 mi. NE

of Swan Valley on Rte. 31, H & Hinds 1120 in 1968

(FPF). CAMAS CO.: Boise River, 10 mi. NE of

Elmore Co. bdy., Peterson 62-51 in 1962 (FPF).

CASSIA CO.: 14 mi. W of Oakley on Rogerson rd.,

H & Hinds 1005 in 1966 (FPF). CARIBOU CO.:

Diamond Cr., 18 mi. NE of Soda Springs, Mentz in

1914 (ILL); 4 mi. W of Freedom, Wyoming on Rte.

34, H & Hinds 1119 in 1968 (FPF). CUSTER CO.:

Redfish Lake, Macbride & Payson 3654 in 1916

(MO, RM); Marsh Cr., 25 mi. NW of Stanley,

Cronquist 2829 in 1941 (MO). FRANKLIN CO.:

Franklin Basin, Peterson 63-361 in 1963 (FPF).

FREMONT CO.: Eccles [near Pineview], Hedgcock

FP 879 in 1909 (FPF) ; Big Springs, Hedgcock FP 888

in 1909 (FPF); Buffalo For. Camp on Buffalo River,

Cronquist 1613 in 1929 (MO). IDAHO CO.: Warren

Divide, 4 mi. from Warren, Stillinger & Root in 1920

(ILL) ; 14 mi. S of Salmon River on US 95, Peterson

62-47 in 1962 (FPF) ; 7 mi. N of Burgdorf on Salmon

River rd., H 972 in 1966 (FPF). KOOTENAI CO.:

Coeur d^ Alene, Weir 8442* in 1917 (FPF, RM, US)

and Weir 9829 in 1918 (FPF, ILL), and on P. ponderosa, Weir 2387, n.d. (FPF, ILL); Fishhook Cr.,

Gill FP 68200* in 1932 (FPF); Rathdrum, Sandberg,

MacDougal & Heller 892 in 1892 (DS, US); Hayden

Lake, Weir 8440 in 1917 (FPF, ILL); without

locaHty, Sandberg 9358 in 1892 (MO, RM, WTU)

and Lieberg in 1892 (PH). LEMHI CO.: Gibbonsville, Wolpert in 1917 (ILL); 4 mi. N of Gibbonsville

on US 93, H & Hinds 1124 in 1968 (FPF) ; 20-25 mi.

mi. N of Salmon on US 93, Gill & Hinds FP 89981

in 1951 (FPF); Parker Mtn., ca. 18 mi. WNW of

Challis, [listed as Custer Co.], Macbride & Payson

3270 in 1916 (MO, RM, US) ; SW of Salmon, between

WilHams Cr. and Moccasin Cr., Krebill 127 in 1964

(FPF); 4 mi. E of Gibbonsville, on P. ponderosa,

Peterson 63-295 in 1963 (FPF). MADISON CO.:

Hawley Ranger Sta., Peterson 62-17 in 1962 (FPF).

TETON CO.: Packsaddle Can., W of Driggs,

Peterson 62-10 in 1962 (FPF). TWIN FALLS CO.:

15 mi. E of Rogerson on Oakley rd., H & Hinds 1004

in 1966 (FPF); 30 mi. W of Oakley, Benlow in 1917

(FPF, ILL). VALLEY CO.: Deadwood Basin,

Korstian in 1918 (FPF, ILL); 3 mi. NW of McCall

on shore of Payette Lake, H 976 in 1966 (FPF) ; Smith

Ferry, Benloio in 1917 (ILL) ; 13 mi. E of Cascade, on

P. ponderosa, Jones 5076 in 1934 (UC, WTU).

COUNTY UNCERTAIN: Middle Fork of Salmon

River, Trey in 1941 (US); Ross, Weir 8389 in 1915

(ILL); Lemhi Nat. For., on P. flexilis. Weir in 1917

(ILL).

MONTANA

BEAVERHEAD CO.: Pioneer Mtns., 11 mi. S of

Dewey on Can. Cr. rd., H & Wicker 949 in 1966

(FPF); 2.5 mi. E of Lemhi Pass, H & Hinds 1121

in 1968 (FPF). CARBON CO.: Woohoe Cr., near

Red Lodge, Languor in 1917 (ILL). CASCADE CO. :

Neilhart, Languor n.d. (ILL). DEERLODGE CO.:

3 mi. N of US lOA on Warm Springs rd., H & Wicker

951 in 1966 (FPF); Warm Springs Ranger Sta.,

Peterson 62-18 in 1962 (FPF). FLATHEAD CO.:

Glacier Nat. Park: Bowman Lake, Jones in 1910

(POM) and McDowell Lake, Standley 16335 in 1919

(US). GALLATIN CO. : West Yellowstone, Payson &

Payson 1937 in 1920 (MO, RM) and Hopkins 426 in

1922 (US); Hyalite Can., Weber 1957 in 1941

(COLO); Shoefelt Ranger Sta. [9 mi. SE of Bozeman], Hedgcock FP 4338 in 1910 (FPF). GLACIER

CO.: 7.8 mi. S of St. Mary on US 89, H 89 in 1961

(FPF); 0.7 mi. S of Going-to-Sun Chalets, Gill FP

68205 in 1932 (FPF). GRANITE CO.: Emerine

Lookout rd., on Piuus albicaulis, H & Wicker 952 in

1966 (FPF); Skalkaho Pass, Peterson 62-60* in 1962

(FPF); Garnet, Scheuber in 1902 (UC, US); Stumptown Cr. near Philhpsburg, Hughes in 1917 (FPF,

ILL); Georgetown, Wolpert in 1917 (FPF, ILL).

LINCOLN CO.: Libby, Smith in 1915 (ILL).

MEAGHER CO.: Little Belt Mtns.; Sheep Cr.,

Languor in 1918 (ILL) ; Adams Cr., 16 mi. N of White

Sulfur Springs, Gill FP 89984 in 1948 (FPF).

MINERAL CO.: St. Regis, Weir 7473 in 1918

(ILL) ; 6 mi. W of St. Regis on US 10, H efe Wicker

954 in 1966 (FPF) ; east of Savenac Nursery [near

Haugan], Hubert in 1915 (ILL); De Borgia, Hughes

in 1917 (FPF, ILL). MISSOULA CO.: Lolo Hot

Springs, Barkley & Barkley 3505 in 1938 (MO, UC,

192
US); Seeley Lake, Weir 9780 in 1918 (ILL). PARK

CO. : Specimen Cr., Sioingle in 1933 (CS, RM, WTU).

PHILLIPS CO.: Little Rocky Mtns., Midvale,

Umhach 594 in 1903 (ARIZ, DS, RAI, US).

RAVALLI CO.: Darby, on P. ponderosa, Weir 2S82

in 1916 (ILL) and on P. alhicaulis, Weir 2379 in 1916

(ILL); Alta, Jones in 1909 (US). SANDERS CO.:

Thompson Falls, Hedgcock & Weir FP 9U^ in

1911 (FPF). SILVER BOW CO.: Highland Mtns., 5

mi. SE of US 91 on Highland lookout rd., i/ cfc

Wicker 950 in 1966 (FPF). COUNTY UNCERTAIN: Big Adrian Gulch near Helena, Starz in 1893

(DS).

NEVADA

COUNTY UNCERTAIN: Without locality, Wheeler

in 1872 (US).

OREGON

BAKER CO. : Jim Cr. fork of Eagle Cr., 30 mi. NE of

Baker, Graham in 1965 (FPF); Horse Cr., 20 mi. W

of Baker, Graham in 1965 (FPF). CLACKAMAS

CO.: Lemiti Butte, SE corner of co., Thompson in

1968 (FPF). CROOK CO.: N Big Meadow, Nelson

845 in 1905 (RM); Laidlow area, Whited 3171 in

1906 (US). DESCHUTES CO.: 18 mi. W of Bend,

Ferris & Duthie 467 in 1919 (DS, RM); Head of

Fall River [9 mi. NW of La Pine], Peck 14322 in 1925

(DS, PH) ; near Pistol Butte, ca. 9 mi. N of La Pine,

on P. ponderosa, H & Scharpf 1261 in 1969 (FPF) ;

10 mi. NW of Sisters on US 126, H &W6I4 in 1964

(FPF); 21 mi. S of Bend on US 97, H & W 618 in

1964 (FPF); Pringle Butte, H 988 in 1966 (FPF); 4

mi. E of McKenzie Pass on Rte. 242, H & Hinds

993 in 1966 (FPF). DOUGLAS CO. : Diamond Lake,

Henderson 11501 in 1929 (PH); 20 mi. NE of Union

Cr. on Rte. 230, H 1143 in 1968 (FPF). GRANT

CO.: Whitman Nat. For., Bates, Boyce FP 40120 in

1920 (OSC) and Boyce 703 in 1920 (FPF); Dixie

Butte rd., 5 mi. N of US 26, H 979 in 1966 (FPF) ; 10

mi. N of John Day, Graham^ in 1964 (FPF); 15 mi.

N of John Day, Graham in 1964 (FPF) ; John Day

Exp. For. [9 mi. E of Bates], Childs 245 in 1955

(OSC); Dixie Pass on Rte. 26, H & W 581 in 1964

(FPF); Dry Meadow, ca. 24 air mi. E of Seneca,

H & Scharpf 1269 in 1969 (FPF). HARNEY CO.:

[was Crook Co.], Black Butte [27 mi. NW of Burns],

Cusick 2691 in 1901 (MO, POM, RM, UC, US).

JACKSON CO. : 22 mi. E of Ashland on Lake of the

Woods rd., H&W871 in 1966 (FPF). JEFFERSON

CO.: Metolius area. Gill FP 68190 in 1932 (FPF).

JOSEPHINE CO.: Waldo [7 mi. S of Cave Jet.], on

P. jeffreyi. Weir 2384 in 1916 (FPF, ILL); Oregon

Mtn. [14 mi. SW of Cave Jet.], on P. attenuata,

Weir 2376 in 1916 (FPF, ILL) and 2374 (ILL).

KLAMATH CO.: Crater Lake Nat. Park, Gill

FP 68185 in 1932 (FPF); T. 23 S., R. 7 E. [NE of

Crescent], Overhay FP 68282 in 1932 (FPF); 15 mi.

SW of Crescent Lake, Childs 67 in 1937 (OSC);

Crescent, Peck 9594 in 1928 (DS, MO, PH); Crescent, Boyce FP 40198 in 1921 (OSC); 7 mi. S of

Crescent, Peck 21639 in 1943 (UC); 6 mi. S of Rte.

58 on Windigo Pass rd., H & W 620 in 1964 (FPF);

5 mi. W of US 97 on Rte. 230, H & W 623 in 1964

(FPF) ; S of Sand Cr. [5 mi. N of Yamsey Ranger

Sta.], Childs 14 in 1947 (OSC); Fort Klamath, Walpole 234 & 323 in 1899 (US) and Meinecke in 1915

(FPF) ; Buck Lake, Applegate 2478 in 1898 (US) ; 7

mi. N of Fort Klamath, Meinecke in 1913 (FPF);

Crater Lake, on P. ponderosa, Dachnowski-Stokes in

1913 (US). LAKE CO. : Dairy Cr. Guard Sta. [13 mi.

NE of Quartz Pass], Steward & Steward 7471 in 1958

(DS, US). LINN CO. : 0.5 mi. W of Rte. 22 near Rte.

20, Steward 6605 in 1953 (RSA, WTU). UMATILLA

CO.: Langdon Lake, 20mi. Eof Westonon Rte. 204,

H &W591 in 1964 (FPF). UNION CO. : 3 mi. SE of

Kamela, H & W 586 in 1964 (FPF). WALLOWA

CO. : 6 mi. SE of Wallowa, Sampson & Pearson 21 in

1907 (US) ; Billy Meadows [30 mi. NNE of Enterprise,] Coville 2487, 2488, 2489, 2490, and 2491 in 1907

(US). WASCO CO.: Bear Springs, Hanshrough FP

68270 in 1931 (FPF); 20 mi. NW of Warm Springs

on US 26,H&W 612 in 1964 (FPF). COUNTY UNCERTAIN: Big Meadows, Deschutes River, Leiberg

520 in 1894 (DS, MO, POM, RM, US); Blue Mtns.,

Nuttalln.d. (MO, FR).

UTAH

CACHE CO.: Jet. of Rte. 89 and Beaver Mtn. rd.,

Mielke in 1952 (UC). DAGGETT CO.: 45.5 mi. S

of Manila on Rte. 44, Greene & Richter 116 m 1962

(COLO, FPF); 19 mi. S of Manila on Rte. 4:4:,H568

in 1964 (FPF) and on P. ponderosa, H 569 in 1964

(FPF). SUMMIT CO.: Kamas, Hedgcock FP 9339

in 1911 (FPF) and FP 22718 in 1916 (FPF); 8 mi. E

of Kamas, on P. ponderosa, Hedgcock FP 22719 in

1916 (FPF, UC); 20 mi. E of Kamas on Rte. 150,

Peterson 62-4 in 1962 (FPF) and on P. ponderosa,

Peterson 62-5 in 1962 (FPF); Kamas Canyon at

Shingle Cr., Peterson 65-90 in 1965 (FPF), on

Pseudotsuga menziesii, Peterson 65-88 in 1965 (FPF)

and on Picea pungens, Peterson 65-89 in 1965 (FPF) ;

Uinta Mtns., 6 mi. S of Wyo. bdy. on Rte. 150, H

1188 in 1969 (FPF). COUNTY UNCERTAIN:

Uinta Mtns., Greeman & Greeman 4^50 in 1931

(MO); Little Bush Cr. Knob, Ashley Nat. For.,

[Uintah Co.?], Harrison & Larson 7859 in 1936

(COLO).

193
WASHINGTON

CHELAN CO.: Washington For. Reserve, head of

Twenty-five-mile Cr. [W side of Chelan Lake near

Stormy Mtn.], Gorman 576 in 1897 (US). FERRY

CO.: Sherman Cr. near Growden, Wicker in 1961

(WSP); Boulder Cr., Eggleston 10599 in 1914

(US); 1.5 mi. W of Sherman Pass on Rte. 30, H^

Stewart & Thompson 1131 in 1968 (FPF). KITTITAS

CO.: Ellensburg, Brandegee 1069 in 1883 (US).

KLICKITAT CO. : Near Swauk Pass, on US 97, 18

mi. N of US 10, Wicker in 1966 (FPF); Falcon

Valley, Suksdorf in 1883 (PH, US). OKANOGAN

CO.: 8 mi. E of Methow, Grahmn in 1964 (FPF).

PEND OREILLE CO. : 3 mi. SW of Newport on US

2, H & Wicker 1126 in 1968 (FPF). SKAMANIA

CO.: SW of Mt. Adams, col.? in 1915 (ILL).

SPOKANE CO. : 21 mi. SW of Newport on US 2,

H & Wicker 1127 in 1968 (FPF); Little Spokane

River, Hedgcock FP 67206 in 1934 (FPF) ; 5 mi. SW

of Deer Park on US 395, H, Stewart & Thompson 1128

in 1968 (FPF). STEVENS CO.: Northport,

Hedgcock FP 1^.7157 and FP 47756 in 1928 (FPF);

13 mi. E of Northport, Hedgcock FP Ii.95U m 1929

(FPF); 12 mi. E of Northport, Hedgcock FP 1^7529

in 1928 (FPF); Alladin, [15 mi. NE of Colville],

Hedgcock FP 54116 in 1931 (FPF); Colville, Hedgcock FP 54116 in 1931 (FPF); Colville, Hedgcock

FP 59245 in 1933 (FPF). YAKIMA CO.: Big

Klickitat River, N of Mt. Adams, Henderson 2539 in

1892 (WTU); 7 mi. E [as ''W'] of Chinook Pass,

Thompson 15105 in 1940 (DS, MO, PH, RSA, UC,

US, WTU); Tieton Cr., 2 mi. SW of Tieton Lake,

H &W 599 in 1964 (FPF); 1 mi. W of Rimrock, N

of Tieton Lake, Steivart in 1968 (FPF). COUNTY

UNCERTAIN: Cascade Mtns., Tweedy in 1882

(MO).

WYOMING

ALBANY CO.: Medicine Bow Mtns.: Centennial,

Hedgcock FP 15834 in 1914 (FPF, UC); Lake Cr.,

Hedgcock FP 15849 in 1914 (FPF); Fox Park,

Hedgcock FP 15965 in 1914 (FPF, RSA, UC),

Hedgcock FP 15848 in 1914 (FPF, UC), Evans FP

17832 in 1915 (FPF), Hedgcock & Johnston FP

26425 in 1917 (FPF), Evans FP 17843 in 1915

(FPF); Libby Lodge, Owenby 875 in 1935 (RM);

Mtn. Home, Osterhout 2523 in 1897 (RM); 6 mi.

W of Centennial on Rte. 130, Greene & Richter

126 in 1962 (COLO, FPF); Libby Cr., Williams

2402 in 1935 (MO, US). Laramie Mtns.: East

of Laramie, Weir 9874 in 1918 (FPF, ILL);

head of Pole Cr., Nelson & Nelson 6833 in 1899

(RM); Sherman, Letterman in 1884 (PH); Boulder

Ridge, 7 mi. SW of Tie Siding, //. 285 in 1962 (FPF) ;

east of Laramie, on P. ponderosa, Weir 9875 in 1918

(FPF, ILL). Laramie Mtns., 5 mi. E of Interstate 80

on Happy Jack rd., H & Bailey 1009 in 1966 (FPF)

and, on P. ponderosa, H & Bailey 1010 in 1966 (FPF).

Pole Mtn. : on P. ponderosa, Simmons in 1917 (ILL) ;

near Happy Jack, on P. ponderosa, H 193 in 1960

(FPF) and Porter 8756 in 1961 (RM). BIGHORN

CO.: Bighorn Mtns., Meadowlark Lake, H 265'' in

1962 (FPF). CARBON CO.: Encampment, Edmonston FP 51 in 1908 (FPF) ; 2 mi. E of Medicine Bow

Lodge, Greene & Richter 125 in 1962 (COLO, FPF).

CONVERSE CO.: Laramie Mtns., Cold Springs,

Simmons in 1917 (ILL). FREMONT CO.: Popo

Agie Campground, ca. 17 air mi. SSW of Lander,

Stewart in 1967 (FPF); Fossil Hill, ca. 12 air mi. SW

of Lander, Steivart in 1967 (FPF) : 6 mi. E of Togwotee Pass on US 287, H 272 in 1962 (FPF) ; Horse Cr.,

14 miles NW of Dubois, H 273 in 1962 (FPF);

Wiggins Fork, 16 mi. N of Dubois, H 275 in 1962

(FPF); Long Cr., 21 mi. WNW of Dubois, H 276 in

in 1962 (FPF); 1 mi. SE of Sheridan Guard Sta., H

278 in 1962 (FPF) ; T Cross Ranch, Dubois, Cox FP

68228 in 1931 (FPF); Wiggins Fork, Shoshone Nat.

For., on P. flexilis, H 274 in 1962 (FPF); 23 mi.

WNW of Dubois, on P. flexilis, Stewart in 1964

(FPF). JOHNSON CO.: Bighorn Mtns.: Duck Cr.

Burn, ca. 20 mi. W of Buffalo on US 16, ^ 264 in

1962 (FPF) ; 2 mi. E of Elgin Park, on P. ponderosa,

Stewart in 1965 (FPF). LINCOLN CO. : 28 mi. W of

La Barge on La Barge Can. id., H & Hinds 1118 in

1968 (FPF). NATRONA CO.: N Entrance Casper

Mtn. Park, Peterson 125-61 on 1961 (FPF). PARK

CO.: Cody Can. at Mormon Cr., H 266 in 1962

(FPF); Cody Can., Pahaska Teepee, on P. flexilis,

Gill & Davidson in 1954 (FPF). SHERIDAN CO.:

Bighorn Mtns. : Burgess Ranger Sta., 27 air mi. W of

Sheridan, Stewart in 1966 (FPF); E fork of South

Tongue River, 20 air mi. W of Big Horn, Stewart in

1966 (FPF). SUBLETTE CO. : 8.8 mi. N of Pinedale

on Faler Cr. rd., H 545 in 1963 (FPF) ; Pine Cr., near

inlet to Fremont Lake, Krehill 538 in 1967 (FPF);

The Rim, 13.4 mi. SE of Bondurant on US 187, H

546 in 1963 (FPF) and on P. flexilis, H 547 in 1963

(FPF). 12.3 mi. E of US 89 on US 187, H 548 in

1963 (FPF). TETON CO.: Jackson Hole, Reed 1831

in 1948 (RM); Jackson Hole, Coulter Bay Jet., H

271 in 1962 (FPF) ; 10.7 mi. NE of Jackson, H 549

in 1963 (FPF) ; 4.9 mi. N of Moose, H 550 in 1963

(FPF); 11.2 mi. W of Togowotee Pass on US 287,

H 551 in 1963 (FPF) ; 3 mi. S of Jenny Lake, Greene &

Richter 118 in 1962 (COLO, FPF); 1 mi. N of Teton

Nat. Park on US 287, Greene & Richter 119 in 1962

(COLO, FPF); Double Diamond Ranch near

Moose, Williams in 1963 (RM). UINTA CO.: Black

Fork Camp, SE of Evanston, Haugl in 1916 (ILL).

YELLOWSTONE NAT. PARK: Fire Hole Basin,

(Hayden Expedition) in 1872 (PH); Fountain

Geyser, Oleson 328 in 1904 (RM); Upper Basin,

Cooper 106y in 1906 (RM); Sylvan Pass, H 267 in

194
1962 (FPF) and on P. fiexilis, H 268 in 1962 (FPF) ;

Yellowstone Lake Lodge, H 269 in 1962 (FPF)

Lewis River, 7 mi. N of S entrance on US 287, H 270

in 1962 (FPF) : Upper Madison, Rydberg & Bessey

3938 in 1897 (PH, RM, US); Madison River,

Nelson & Nelson 5515 in 1899 (R^I, MO, UC, US);

Mammoth Hot Springs, Greene & Richter 121 in 1962

(COLO, FPF) ; 8 mi. W of Can., Greene & Richter 122

in 1962 (COLO, FPF) ; 1 mi. E of Fishing Bridge,

Greene & Richter m in 1962 (COLO, FPF); Geyser

Basin, Firehole River, Jones 5372 in 1934 (WTU) ;

Upper Geyser Basin, Setchell in 1905 (UC); Yellowstone Lake, Hayden in 1871 (PH); Cache Cr.,

Tioeedy U^ in 1885 (UC) ; Lake Fork, Hayden in 1860

(MO); (without locahty) Forwood in 1882 (MO)

and Mearns 4513 in 1902 (US). COUNTY

UNCERTAIN: Hebron, Hedgcock & Johnston FP

26433 in 1917 (FPF); near E entrance, Yellowstone

Nat. Park, Greene & Richter 123 in 1962 (COLO) ;

Cooperton, Garrett in 1918 (ILL); Copper City,

Garrea n.d. (UT).

4. ARCEUTHOBIUM APACHECUM

Specimens Examined: All on Pinus strohijormis.

UNITED STATES

ARIZONA

APACHE CO.: 4 mi. S of Eagar on Big Lake rd.,

H & L210 in 1962 (FPF) ; Escudilla Mtn., Andresen

& SÍ7npson 1787* in 1962 (FPF). COCHISE CO.:

Chiricahua Mtns. : Rustler Park, Andresen & Miles

631 in 1960 (FPF); Onion Saddle, Gilbertson in 1968

(FPF) and H & L 156 in 1962 and 1107 in 1968

(FPF). GRAHAM CO.: Graham INItns.: Ellis FP

89431 in 1939 (FPF) ; Treasure Park, Stouffer & Gill

FP 68292 and FP 68293 in 1934 (FPF) and Gill FP

89313 in 1936 (FPF); 0.5 mi. E of Hospital Flat,

H & L 228 in 1962 (FPF) and 1105 in 1968 (FPF).

GREENLEE CO. : Blue Summit, Gill FP 68306 in

1934 (FPF); 1 mi. S of Mogollón Rim on US 666,

L 64-21 in 1964 (FPF). PIMA CO.: Santa Catalina

Mtns.: (without locality), Hedgcock FP 9782 in 1911

(FPF, ILL) and Shreeve in 1908 (ARIZ); Mt. Lemmon, Peterson 62-145"" in 1962 (FPF); Soldiers

Camp, Goodding in 1934 (RM) and Gill FP 68128 in

1932 (FPF); head of Marshall Gulch, Gill FP 68130

in 1932 (FPF); Bear Wallow, H & L 167 in 1962

(FPF). Rincón Mtns., Mica Mtn., Ela in 1968

(FPF).

NEW MEXICO

CATRON CO. : 6 mi. S of Luna on US 260, H & L

212 in 1962 (FPF) ; 7 mi. E of Mogollón on Rte. 78,

H & L 220 in 1962 (FPF); Mangas Mtn., // ¿c

Scharpf696* in 1964 (FPF) ; 1 mi. E of Fox Mtn., ca.

20 mi. N of Apache Cr., H & L 929 in 1966 (FPF).

GRANT CO. : Hillsboro Peak, Metcalfe 1173 in 1904

(CAS, FPF, MO, PO:\I, UC, US); Emory Pass,

H & L 121 in 1962 (FPF) and 1102 in 1968 (FPF);

24 mi. N of Mimbres on Rte. 61, L 65-40 in 1965

(FPF) ; Mimbres Valley and its east can. tributary,

Holzinger in 1911 (US) ; 4 mi. E of Rte. 25 on Signal

Peak rd., ca. 13 mi. N of Silver City, H & L 891 in

1966 (FPF). LINCOLN CO.: Capitán Mtns.: E

side Capitán Peak, Martin 902 in 1945 (WTU);

Seven Cabins Can., Gill, Ellis & Hackleman"* FP

89490 and Ellis FP 89492 in 1938 (FPF) ; 1-2 mi. E

of Capitán Gap, H & L105,. 107, 109 in 1962 (FPF)

and 1098 in 1968 (FPF). SOCORRO CO.: San

Mateo Mtns. : Hughes Sawmill, (4 mi. SW of Mt.

Withington), Long FP 21128* in 1915 (FPF) ; on Rte.

52, 12 mi. S of US 60, L 65-36 in 1965 (FPF) and 13

mi. S,H & L 1117 in 1968 (FPF). ]Magdalena iNItns.:

North Baldy Mtn., L 66-19in 1966 (FPF).

MEXICO

COAHUILA: Sierra del Carmen, Ocampo, H, L &

Muñoz 1032in 1967 (FPF).

5. ARCEUTHOBIUM BICARINATUM

Specimens Examined: All on Pinus occidentalis.

DOMINICAN REPUBLIC

AZUA PROV.: Las Cañitas, Fuertes 1923 in 1912

(ILL, US, Z) and Gasharro* in 1963 (FPF); San

Jose de Ocoa, Bejucal, Ekman 12024 in 1929 (US).

BARAHONA PROV.: Between Pedernales and

Aceital, Howard & Howard 8129 in 1946 (B:\I, US).

LA VEGA PROV.: Constanza, Allard 17432 in 1947

(US) SiudH & W 1193 in 1969 (FPF); 26 km. S of

Constanza on Valle Nueva rd., H & W 1199 in 1969

(FPF). SAN JUAN PROV. : Sabana Nueva, Howard

& Howard 9125 in 1946 (US); 10 km. N of Hato

Nueva, Gasbarro* in 1963 (FPF). SANTIAGO

PROV.: Pico des Rubio, Jijninez 1069 in 1946 (US).

HAITI

DEPT. DE L^ OUEST: Vicinity of Furcy, Leonard

4690 in 1920 (US) ; Massif de la Selle, EkiJian 1313 in

1924 (US) and Proctor 10771* and 10772 in 1955

(US); ^Nlormes des Commissiares, Holdrige 1328 in

1942(BM,F,MO,UC,US).

6. ARCEUTHOBIUM BLUMERI

Specimens Examined: All on Pinus strohijormis,

UNITED STATES

ARIZONA

COCHISE CO.: Huachuca Mtns., Carr Can.,

Goodding 309 in 1909 (ARIZ); 1.2 mi. W of Reef,

H &L 235* in 1962 (FPF) and 1109* in 1968 (FPF).

195
SANTA CRUZ CO.: Santa Rita Mtns.: Upper

Florida Can., Graham in 1939 (DS); Mt. Wrightson

Trail, 4 mi. from end of rd. in Madera Can., H 796

inl965(FPF).

MEXICO

CHIHUAHUA: 36 mi. SW of La Junta, H & W 296

in 1963 (FPF); 13 mi. W of El Vergel, H & W 314

in 1963 (FPF); 13 mi. SE of Mesa Huacan on Chico

Yd.,H&W 475 in 1963 (FPF); 65 mi. SW of Matachic on Ocampo rd., H & W 493'' in 1963 (FPF);

'^Sierra Madre,'' Giguieliat in 1912 (ILL). DURANGO: 15 mi. W of El Salto on Rte. 4.0, H & W

350 and 550* in 1963 (FPF) ; 48 mi. S of Durango on

La Flor rd., H & W 516 in 1963 (FPF) ; 19 mi W of

Santiago Papasquiaro, H &W 536* in 1963 (FPF).

NUEVO LEON: Cerro Potosí, Andresen & Steinhoff

A2047 in 1962 (FPF) and J^ cfc 17 392 in 1963 (FPF).

7. ARCEUTHOBIUM CALIFORNICUM

Specimens Examined: All on Pinus lamhertiana

except as noted.

UNITED STATES

CALIFORNIA

BUTTE CO.: Big Bar Mt. Ridge, E of Pulga,

Quick 53-32 in 1953 (CAS). DEL NORTE CO.: 0.5

mi. E of Sourdough Jet., ca. 10 air mi. N of Gasquet,

on P. monticola, Miller in 1969 (FPF). KERN CO.:

Greenhorn Mtns.: 4 mi. N of Summit Guard Sta.,

W 3609'' in 1964 (FPF); near Tiger Flat, Howell

38805 in 1962 (CAS). LAKE CO.: 23.5 mi. N of

Upper Lake on Pillsbury Lake rá., H & W 862 in

1966 (FPF). LOS ANGELES CO.: Between Wrightwood and Kratka Ridge, Embree in 1956 (UC);

0.5 mi. E of Vincent Gulch Divide, Krebill 247 in

1956 (FPF); Little Rock-Bear Cr. Divide, Ewan

10110 in 1936 (MO). MADERA CO.: Ellis (now

Benedict) Meadow, Meinecke FP 17062 in 1914

(FPF) ; Fish Cr. on Rock Cr. rd., H 9 m 1953 (FPF) ;

Soquel-Bass Lake rd.. Gill & Wright FP 68160"" in

1932 (FPF) 5 mi. N of Bass Lake on Soquel rd.,

H & H 1146 in 1968 (FPF). MARIPOSA CO.:

Mariposa Grove, Hedgcock & Meinecke FP 4822 in

1910 (FPF); Fish Cr. Camp, Hedgcock & Meinecke

FP 4833 in 1910 (FPF, ILL); Willow Cr., Hedgcock

& Meinecke FP 4838 in 1910 (FPF); Fish Camp,

Hedgcock & Meinecke FP 4827 in 1910 (FPF); 1 mi.

E of Fish Camp, H* in 1963 (FPF); 0.5 mi. SW of

Fish Camp, H 660 in 1964 (FPF). NEVADA CO.:

Graniteville, Boyce 32 in 1917 (FPF). PLUMAS

CO.: Massack, Gill & Wagener FP 68O4I in 1931

(FPF) ; Silver Lake, near Quincy, Scharpf* in 1963

(FPF); Slate Cr., 5.5 mi. W of Quincy, W 3230* in

1962 (COLO, FPF) and H & W 654 in 1964 (FPF);

Meadow Valley, Weatherhy 1667 in 1955 (CAS, RM,

RSA, UC) ; Feather River Exp. For. near Quincy,

Meinecke n.d. (FPF). RIVERSIDE CO.: San

Jacinto Mtns., Idyllwild, Meinecke FP 20151 in 1915

(FPF, ILL), Cooper & Silva 923 in 1943 (ARIZ,

RSA); 8 mi. N of Idyllwild, H 740 in 1965 (FPF).

SAN BERNARDINO CO.: San Antonio Mtns.,

Coldwater Fork of Lytle Cr., Johnston 1688 in 1917

(DS, UC). San Bernardino Mtns.: Inspiration

Point, Perkins in 1917 (ILL); Baldy Notch-Stockmans Flat, W 2808 in 1961 (COLO, FPF); W slope

Job Peak, Eioan 3564 in 1929 (POM, UC) ; ridge E of

Foxesee Cr., Peirson 2818 in 1920 (RSA) ; 1 mi. E of

Lake Gregory, H 730 in 1965 (FPF). SAN DIEGO

CO.: Cuyamaca Mtns., Cuyamaca Peak, Cox in

1967 (FPF, UT). SHASTA CO. : 0.5 mi. S of Siskiyou

Co. bdy. on Rte. 89, W 3238* in 1962 (COLO, FPF) ;

head of Tom Neal Cr. (ca. 3 mi. SE of Castella),

Wyckoff & Randall in 1920 (ILL). SIERRA CO.:

Between Downieville and Forest, BoTjce 31 in 1916

(FPF). SISKIYOU CO.: SE of Bartle on Rte. 89,

H &W644 in 1964 (FPF); Dillon Mt. rd., T. 13 N,

R. 6 E, Sec. 5, 12 mi. N of Somes Bar, Miller FP

98034 in 1962 (FPB). TEHAMA CO.: 2 mi. N of

Hole-in-Ground Campground, S of Mineral, Kuijt

1502 in 1958 (UC). TULARE CO.: Sequoia Nat.

Park, Crystal Cave Jet., W 3611 in 1964 (FPF).

TUOLUMNE CO.: 5.1 mi. E of Nat. For. bdy. on

Rte. 120, Kuijt 1394in 1957 (UC).

OREGON

CURRY CO.: Saddle Mtn., T. 37 S., R. 12 W., Sec.

20, ca. 15 air mi. ESE of Gold Beach, on P. monticola, Bynum, in 1967 and 1968 (FPF) and on Picea

hreweriana, Bynum in 1967 and 1968 (FPF). JOSEPHINE CO.: Oregon Mt. (14 mi. SW of Cave Jet.),

Weir 3191 in 1916 (FPF, ILL) and Theisen in 1964

(FPF) and on P. monticola, Weir 3240 in 1916 (FPF,

ILL); 6.5 mi. SSW of O'Brien on Oregon Mtn. rd.,

on P. monticola, H & Hinds 994 in 1966 (FPF).

COUNTY UNCERTAIN: Ashland-Klamath Falls

wagon rd.. Weir 3288 in 1916 (FPF, ILL) and on P.

monticola, Weir 3239in 1916 (FPF, ILL).

8. ARCEUTHOBIUM CAMPYLOPODUM

Specimens Examined: on Pinus ponderosa var.

ponderosa except as noted.

MEXICO

BAJA CALIFORNIA: Sierra de San Pedro Mártir:

Low hills NW of La Encantada, on P. jeffreyi,

Wiggins & Demaree 5018 in 1930 (DS, F, POM,

RSA, US); without locality, as Pinus sp., probably

P. jeffreyi, Brandegee in 1893 (UC) ; 0.25 mi. N of La

Encantada, on P. jeffreyi, H & Scharpf 765 in 1965

(FPF) ; La Grulla, on P. jeffreyi, H & Scharpf 767

in 1965 (FPF). Sierra Juárez: 30 mi. S of La Rumo196
rosa on Laguna Hanson rd., on P. jeffreyi, L & Gill

64-35 in 1964 (FPF) ; 8 mi. W of Laguna Hanson, on

P. jeffreyi H & Scharpf783 in 1965 (FPF).

UNITED STATES

CALIFORNIA

ALPINE CO.: Silver Cr. Campground, E side

Ebbetts Pass, on P. jeffreyi, Munz 21347 in 1955

(RSA); 5 mi. SW of Silver Cr. on Rte. 4, on P.

jeffreyi, H & Scharpf 668 in 1964 (FPF). BUTTE

CO. : Jonesville, on P. jeffreyi, Copeland in 1931 (UC).

CALAVERAS CO.: 15 mi. SW of Tamarack on

Rte. 4:, H & Scharpf 661'' in 1964 (FPF); Angels

Camp, Eggelston 9718 in 1913 (US). DEL NORTE

CO.: Near Gasquet, on P. attenuata, Tracy 16464 in

1939 (DS, JEPS, UC, US, WTU); Smith River,

between bridges 1 and 2, on P. attenuata, Kildare

10484 in 1931 (DS: Gordon Mtn., on rd. from

Gasquet to Big Bar, on P. attenuata, Newcomh

165 in 1957 (UC) ; Grassy Flat Guard Sta. [5 mi. E of

Gasquet], on P. attenuata, Wright FP 91563 in 1947

(OSC) ; Bear Wallows, 2 mi. N of Sänger Peak, on

P. attenuata, Kildare 8785 in 1929 (DS); Gasquet

Mtn., on P. attenuata, Eastwood 12138 in 1923 (CAS) ;

0.5 mi. N of Gasquet, H & W 863 in 1966 (FPF) ; 4

mi. NE of Gasquet on old toll rd., on P. attenuata,

H &W 865 in 1966 (FPF) ; 6 mi. NE of Gasquet on

old toll rd., H & W 867 in 1966 (FPF) and on P.

contorta subsp. contorta, H & W 866 in 1966 (FPF) ;

Ship Mtn. trail, on P. jeffreyi, Weir 3189 in 1916

(ILL); 18 Mile Cr. Can., Elk Camp Ridge, on P.

contorta subsp. contorta. Parks & Parks 24063 in 1930

(UC). ELDORADO CO.: 3 mi. S of Meyers on U.S.

50, on P. jeffreyi, Peterson 63-315"" in 1963 (FPF);

Kyburz, Rossbach 237 in 1955 (UC); Lake Tahoe,

Emerald Bay, Boyce FP 97944 m 1913 (FPB) ; 8 mi.

E of Kyburz on US bÇ^, H & W 840 in 1966 (FPF) ;

3 mi. NW of Placerville on Rte. 49, i/ cfc If 842 in

1966 (FPF); Fallen Leaf Lake, on Pinus contorta

subsp. murrayana, Copeland 1425 in 1931 (POM);

Lily Lake, Glen Alpine Can., on P. jeffreyi, Ahrams

12752 in 1930 (UC) ; 9 mi. S of Camp Richardson on

Fallen Leaf Lake rd., on P. jeffreyi, Pickle 57-15

in 1957 (UC); 0.5 mi. S of Camp Richardson on

Fallen Leaf rd., on P. jeffreyi, Kuijt 1337 in 1957

(UC). FRESNO CO.: 1.3 mi. N of Mono Hot

Springs, on P. jeffreyi, H in 1963 (FPF) ; between

Brown Cone and Dualton rd. [near Kaiser Diggings],

on P. jeffreyi, Quihell 280 in 1951 (DS, RSA) ; Camp 6,

Big Creek, Meinecke FP 97948 in 1917 (FPB). Kings

Can. Nat. Park: Cedar Grove, on P. jeffreyi, W 3616

in 1964 (FPF) ; Copper Cr. Can. Trail, on P. jeffreyi,

Howell 34204 in 1958 (CAS); Simpson Meadow,

middle fork Kings River, Hoiuell 34093 in 1958

(CAS). KERN CO. : 6 mi. W of Summit Guard Sta.,

Glenville-Sierra Alta rd., W 3606 in 1964 (FPF); 3

mi. N of Summit Guard Sta., on rd. to Tiger Flat,

W 3607 in 1964 (FPF); N of Tiger Flat, Howell

38824 in 1962 (CAS); Indian Cr., on P. jeffreyi,

Root in 1919 (ILL); rim of Kern Can., SW of Pine

Flat, on P. jeffreyi, Tivisselmann 8681 in 1963 (CAS) ;

Mt. Pinos region, on P. jeffreyi. Wheeler in 1931

(CAS). LAKE CO.: Elk Mtn. [8 mi. N of Upper

Lake], Tracy 2355 in 1905 (UC, US, WTU); Boggs

Mtn. State For., Scharpf in 1965 (FPF); 16 mi. N of

Upper Lake on Pillsbury Lake Yd.,H&W 860 in 1966

(FPF) ; 20 mi. N of Upper Lake on Pillsbury Lake

rd., on P. attenuata, H & W 861 in 1966 (FPF);

Upper Lake to Elk Mtn., on P. attenuata, Mason

2590 in 1926 (UC) ; near Lucerne, on P. attenuata,

Sutliffe in 1939 (CAS). LOS ANGELES CO.: San

Gabriel Mtns. : Chilao, Peirson 2322 in 1921 (JEPS) ;

Prairie Fork, San Gabriel River, Johnston 1720 in

1917 (POM, US, WTU); Pine Flat, Wheeler 1115 in

1932 (UC) and on P. coulteri, Sloan FP 97943 in 1914

(FPB); Crystal Lake, Martindale in 1937 (POM);

Cedar Can. off Mescal Cr., Eioan 9924 in 1936 (MO) ;

Little Rock-Bear Cr. Divide, on P. coulteri, Ewan

10111 in 1936 (MO); Barley Flat, on P. coulteri,

Perkins in 1919 (ILL); 0.5 mi. W of Big Pines, on

P. jeffreyi, H 728 in 1965 (FPF) and on P. jeffreyi,

KreUll 244 in 1965 (FPF). MADERA CO.: Bass

Lake, Gill & Wright FP 68162 in 1932 (FPF); 2 mi.

NW of North Fork, H 659 in 1964 (FPF) and i^*

in 1964 (FPF) ; 10 mi. SE of North Fork on Mammoth Pool rd., H 848 in 1966 (FPF). MARIPOSA

CO.: Yosemite, Hedgcock & Meinecke FP 4788 and

FP 4789 in 1910 (FPF) and Bolander 6380 in 1866

(US); Fish Camp, Hedgcock FP 4835 in 1910 (FPF)

and on P. jeffreyi, Hedgcock & Meinecke FP 4836

in 1910 (FPF) ; Wawona, Posey in 1919 (ILL) ; Crane

Flat, on P. jeffreyi, Kuijt 1407 in 1957 (UC) and

Rose 60113 in 1960 (RSA); Snow Cr., on P. jeffreyi.

Hall in 1911 (UC). MENDOCINO CO.: Without

locality, on P. jeffreyi. Brown 94I in 1898 (FPF,

MO, US). MODOC CO.: 3 mi. N of Crowder Flat

Guard Sta., Peterson 64-IOO in 1964 (FPF). MONO

CO. : 3 mi. E of Sonora Pass, on P. jeffreyi, Kuijt

1432 in 1957 (UC). NAPA CO.: Conn Valley,

Jepson in 1893 (JEPS) ; near Pacific Union College,

Angwin, Boyce 1908 in 1930 (FPF); 2 mi. N of

Angwin on Pope Valley rá., H & W 858 in 1966

(FPF). NEVADA CO.: Between Truckee and Lake

Tahoe, Gill FP 68243 in 1931 (FPF) and on P.

jeffreyi, Gill FP 68237* in 1931 (FPF); 1 mi. N of

Lake Tahoe, W 3227 in 1962 (COLO, FPF) ; 5 mi. W

of Truckee, on P. jeffreyi, Wright FP 68109 in 1932

(FPF). PLACER CO. : 1.5 mi. E of Tahoe State Park

on Rte 28, on P. jeffreyi, Kuijt 1339 in 1957 (UC);

1 mi. N of Tahoe City on Rte 89, on P. jeffreyi,

Kuijt 1333 in 1957 (UC). PLUMAS CO.: Chester,

on P. jeffreyi, Boyce FP 15983 in 1914 (FPF); 1 mi.

E of Rte. 89 on Rte. 40A, on P. jeffreyi, H & W 656

in 1964 (FPF); Quincy Jet., Gill & Wagener FP

197
68040 in 1931 (FPF); Quincy, Feather River Exp.

For., Boyce in 1913 (FPF) and Hunt FP 25071 in

1919 (FPF); Chester, Meinecke FP 17100 in 1914

(FPF); Slate Cr., 5.5 mi. W of Quincy, W 3231 in

1962 (COLO, FPF); 2 mi. SE of Cromberg, W 3229

in 1962 (COLO, FPF); near CHo, Meinecke in 1915

(FPF); Quincy, Boyce FP 97955 in 1913 (FPB);

21.5 mi. N of Sierraville, Kuijt 1346 in 1957 (UC);

7.5 mi. W of Blairsden, Kuijt 1347 in 1957 (UC) ; 6.5

mi. E of Jet. of Rtes. 40 & 89, near Indian Falls,

Kuijt 1348 in 1957 (UC) ; 5 mi. W of Quincy, H &W

655 in 1964 (FPF); 5 mi. W of Meadow Valley,on

P. contorta subsp. inurrayana, Gill & Wagener FP

68039 in 1931 (FPF). RIVERSIDE CO.: San

Jacinto Mtns.: Spenser 1018 in 1919 (POM); Grant

1113 in 1901 (ARIZ, US) ; and on P. jeffreyi, Mum &

Johnston 8705 in 1924 (POM). Idyllwild-Banning,

Clokey & Anderson 6574 in 1935 (POM, RM, RSA,

UC, WTU); Idyllwild, Perkins in 1916 (ILL); 8 mi.

N of Idyllwild on Rte. 74, H 739 in 1965 (FPF) ;

11 mi. N of Idyllwild on Rte. 74, on P. coulteri, H 738

in 1965 (FPF); 2 mi. SE of Idyllwild, on P. jeffreyi,

H 742 in 1965 (FPF); Tahquitz Ridge, on P.

jeffreyi, Meyer 688 in 1929 (JEPS); Pine Cove,

Cooper 1547 in 1944 (RSA); Strawberry Valley,

Wheeler 242 in 1931 (DS, POM) and on P. coulteri,

Grand n.d. (DS) ; 0.6 mi. SE of Keen Camp Summit

on Rte. 74, on P. coulteri, Kuijt 1497 in 1958 (UC) ;

Tripp Mill, on P. coulteri. Meinecke FP 20152 in

1915 (FPF). Santa Rosa Mtns.: Peak rd. 9 mi. from

Rte. 74, on P. jeffreyi, H 745 in 1965 (FPF). SAN

BERNARDINO CO.: San Bernardino Mtns.:

Engelmann in 1880 (MO) ; on P. coulteri. Parish in

1892 (JEPS) and Parish & Parish 966 in 1884

(MO, US) ; Big Bear Lake, Gill & Wright FP 6824O*

in 1931 (FPF) and W 2444 in 1959 (RSA, UC, US)

and 2741 in 1960 (RSA); Camp Ángelus, W 3211

in 1962 (COLO, FPF); Wrightwood, W3212 in 1962

(COLO, FPF); S fork of Santa Ana River, Munz

6260 in 1922 (POM) and on P. jeffreyi, Peirson 1721

in 1909 (RSA) ; 2 mi. E of Bluff Lake, Munz 10566 in

1926 (POM); Bear Valley, on P. jeffreyi, Ahrams

2818 in 1902 (FPF, MO, POM, WTU, Z) ; 5 mi. W of

Arrowbear, on P. jeffreyi. Raven 16874 in 1961 (RM,

RSA); Mill Cr. rd. near Big Bear, on P. jeffreyi,

Howell 386A in 1927 (RSA); Snow Valley W of Big

Bear, on P. jeffreyi, H 731 in 1964 (FPF) ; 2 mi. W of

Snow Valley, on P. coulteri, H 734 in 1965 (FPF) ; 3

mi. SW of Running Springs on Rte. 30, on P.

conlteri, H 736 in 1965 (FPF) and, on P. attenuata,

H 735 in 1965 (FPF); City Cr. rd., on P. attenuata,

Johnston in 1924 (POM). SAN DIEGO CO.: Without locality, on P. coulteri. Weir in 1913 (ILL);

JuHan, on P. coulteri. Bethel FP 24449 in 1918 (FPF) ;

4 mi. W of JuHan, on P. coulteri, Wiggins 2106 in 1926

(DS, UC); 4.1 mi. S of Julian on Rte. 79, on P.

coulteri, Kuijt 1471 in 1958 (UC); 4 mi. SW of

JuHan, on P. coulteri, H 751 in 1965 (FPF) ; 5 mi. SW

of Pine Valley, H & Scharpf 789 in 1965 (FPF).

Cuyamaca Mtns. : Palmer in 1875 (UC) ; Cuyamaca

Lake, Roos I24O in 1937 (POM); Base of Stonewah

Peak, on P. coulteri, Wiggins 2725 in 1926 (DS, UC).

Laguna Mtns.: Huey in 1946 (RSA); Laguna,

Schoenfeldt 3600 in 1894 (UC) ; 5 mi. N of Laguna

Mtn. Lodge, on P. jeffreyi, H 752 in 1965 (FPF);

Laguna Campground, on P. coulteri, Wiggins 2817

in 1927 (DS) and on P. coulteri, Munz 8350 in 1924

(POM). SANTA BARBARA CO.: Los Olivos, on

P. jeffreyi. Hartley & Weir in 1912 (ILL). SHASTA

CO. : 3 mi. N of north entrance to Lassen Nat. Park

on Rte. 89, W 3236 in 1962 (COLO, FPF); Sisson

Tavern Park, col.? FP 97945 in 1913 (FPB); Jet. of

Rtes. 89 and 299, Kuijt 1366 in 1957 (UC) ; 12 mi. N

of Rte. 299 on Rte. S^, H & W 646 (FPF) ; Lassen

Nat. Park, 8 mi. E of N entrance, H & W 649 in

1964 (FPF); Long Valley, SW of Burney Falls, on

P. jeffreyi, Scharpf FP 98114 in 1965 (FPB) ; between

Oak Run and Whitmore, Anderson in 1949 (CAS).

SIERRA CO.: Sierraville, Lemmon in 1875 (MO);

Lemon Can., Sierraville, on P. jeffreyi. Meinecke

& Boyce FP 17196 in 1915 (FPF);'20.5 mi. N

of Truckee, on P. jeffreyi, Kuijt 1345 in 1957 (UC) ;

2 mi. W of Bassetts, on P. contorta subsp. murrayana,

Peterson 63-166 in 1963 (FPF). SISKIYOU CO.:

Mt. Shasta, Engelmann in 1880 (FPF, MO) ; Yreka,

Hedgcock FP 1897 in 1909 (FPF); Scotts River,

Pond FP 193 n.d. (FPF); Mt. Shasta, Brown

580 in 1897 (FPF, MO, US); Sisson, Hedgcock,

Meinecke & Long FP 9668 in 1911 (FPF); Big

Carmen Lake, Scott Mtn. rd., [6 mi. E of Callaham],

Barhe 314 in 1955 (RSA, UC); Mt. Shasta, Sisson

South Trail, Cooke 11593 in 1938 (DS, UC); Gravely

Ridge, W of Craggy Mtn., NW of Yreka, Owenhy

& Brown 2428 in 1941 (UC); Can. Cr. [ca. 12 mi.

SSW of Hamburg], Hedgecock FP 1873 in 1909

(FPF) ; 3 mi. NW of McCloud, Scharpf & Miller

in 1965 (FPF), and on P. attenuata, Scharpf &

Miller in 1965 (FPF); Russian Cr., on P. attenuata,

Butler 273 in 1908 (UC); 3 mi. N of Branch Guard

Sta., N of Happy Camp, on P. attenuata, Kuijt

1277 in 1957 (UC); 16 mi. N of Happy Camp,

on O'Brien rd., on P. attenuata, H & Hinds 1001 in

1966 (FPF); Scott Mtn., on P. jeffreyi. Engelmann

in 1880 (MO). TEHAMA CO.: Childs Meadow, 2

mi. W of Morgan Springs on Rte. 36, on P. contorta

subsp. murrayana, Scharpf FP 38026, n.d. (FPB);

15 mi. W of Rte. 36 on Rte. 89, Kuijt 1356 in 1957

(UC) and on P. contorta subsp. murrayana, Kuijt 1357

in 1957 (UC). TRINITY CO.: Trinity, Munns FP

9440m 1911 (FPF);PlummerSpring, [nearPeanut?],

Meinecke in 1912 (FPF); Trail to Granite Peak,

Baker 209 in 1926 (DS); Scott Ranch, Cantielow

1659 in 1936 (RSA). TULARE CO.: Jordan Hot

Springs, Zeile in 1924 (JEPS) ; Peppermint Valley,

198
Dudley 758 in 1895 (DS); Whitaker For.-, Benson

5705 in 1933 (POM) ; Mineralking, on P. jeffreyi,

Coville & Funston U60 in 1891 (DS) and U82 in

1891 (US); Sequoia Nat. Park, 1 mi. N of Lodgepole

Campground, on P. jeffreyi, W 3612 in 1964 (FPF);

Mt. Sillman [ca. 3 mi. N of Lodgepole], on P.

jeffreyi, Hopping 391 in 1905 (UC); Little Kern

River, between Lion Meadow and Trout River, on

P. jeffreyi, Dudley 1978 in 1897 (DS). TUOLUMNE

CO.: Clark Fork of Tuolumne River, 1.5 mi. below

Arnott Cr., on P. jeffreyi, Wiggins 9322 in 1939

(DS, POM, UC, WTU) ; Crocker Sta., on P. jeffreyi,

col? FP 97946 in 1915 (FPB); 2.8 mi. NE of Long

Barn, on P. jeffreyi, Thomas 10262 in 1963 (FPF,

RSA) ; near Kennedy Meadows on Rte. 108, on P.

jeffreyi, Kuijt 1423 in 1957 (UC); Long Barn, Gill

FP 68087 and FP 68097 in 1932 (FPF), Wright FP

68117 in 1932 (FPF), and Gill & Wright FP 68072

in 1932 (FPF) ; 6 mi. E of Long Barn, W. 3222 in 1962

(COLO, FPF); 1 mi. E of Twain Harte, Kuijt 1428

in 1957 (UC) ; Cottonwood Meadows, E of Mather,

Clausen 1777 in 1939 (DS); Pate Valley, Yosemite,

Clemens in 1919 (CAS). VENTURA CO.: Middle

Fork, Mt. Pinos, Hall 6642 in 1905 (UC) ; Mt. Pinos,

Rothroch 213 m \Slb{V^).

IDAHO

ADAMS CO.: Evergreen, Hedgcock FP 1909 in 1909

(FPF); Bear, Barr in 1915 (ILL); 10 mi. N of

Council on US 95, H 978 in 1966 (FPF). BOISE CO. :

S fork of Payette River, Garden Valley, Miles FP

15108 in 1913 (FPF); Karney Cr., Peterson 62-49

in 1962 (FPF). IDAHO CO.: N bank of Salmon

River, 11 mi. E of Riggins, H 970'' in 1966 (FPF);

MacKay Bar, on Salmon River 2 mi. E of Jet. of S

fork of Salmon River, W 4127h in 1966 (FPF, UT).

KOOTENAI CO.: Without locality, Sandherg in

1892 (US); Coeur d' Alene, Weir FP 17047 in 1914

(FPF) ; Coeur d^ Alene City Park, Gill FP 68203* in

1932 (FPF) ; 5 mi. W of Coeur d^ Alene, W 3254 in

1962 (COLO, FPF) ; Spokane Bridge, Heller 925 in

1892 (DS, MO) ; E side Coeur d' Alene Lake, 4 mi. N

of inlet of Coeur d' Alene River, on P. contorta subsp.

latifolia, Krebill 154 in 1964 (FPF); Spirit Lake, on

P. contorta subsp. latifolia, Dodge in 1916 (ILL);

Fourth of July Can., on P. contorta subsp. latifolia,

Weir 8252 in 1914 (ILL); Fernán Lake, H & Wicker

966* in 1966 (FPF) and on P. contorta subsp.

latifolia. Weir 8243 in 1916 (ILL) ; Hayden Lake, on

P. sylvestris. Weir in 1920 (ILL). LEWIS CO.:

Nezperce, Amson in 1914 (ILL).

NEVADA

CLARK CO.: Charleston Mtns: All on P. ponderosa

var. scopulorum; Clokey 5431 in 1935 (CAS, FPF,

JEPS, WTU) and 5432 in 1935 (CS, FPF, JEPS,

WTU) ; Charleston Park, W 3032 and 3205 in 1962

(COLO, FPF) ; Kyle Can., 2.5 mi. W of Ranger Sta.,

H & Scharpf675in 1964 (FPF); Kyle Can., Peterson

64-144 in 1964 (FPF). DOUGLAS CO.: Glenbrook,

on P. jeffreyi, Baker 1006 in 1902 (MO, POM, UC);

E of Spooner Jet. on US 50, on P. jeffreyi, Peterson

65-403 in 1965 (FPF) ; 5 mi. E of Spooner Jet. on

US 50, on P. jeffreyi, H & W 836 in 1966 (FPF).

ORMSBY CO. : 2 mi. N of US 50 near Carson City,

on P. jeffreyi, Peterson in 1960 (FPF). WASHOE

CO.: Franktown, col? n.d. (PH); Hunter Cr. rd.,

Kennedy in 1907 (CAS) ; Incline Camp [N shore of

Lake Tahoe, 3 mi. E of California bdy.], on P.

jeffreyi, Wright FP 68111 in 1932 (FPF); Bowers

Mansion, 26 mi. S of Reno, on P. jeffreyi. Archer

6396 in 1938 (MO, UC) ; Mtns. W of Bowers, on P.

jeffreyi, Heller 10660 in 1912 (CAS, DS, MO);

N shore of Lake Tahoe, Crystal Bay, on P. jeffreyi,

Peterson65-395m 1965 (FPF).

OREGON

CURRY CO. : 12 mi. NW of Agness, on P. attenuata,

Graham* in 1963 (FPF). BAKER CO.: 7 mi. S of

Baker, Peterson 63-224 in 1963 (FPF); Dooley

Mtn. area, 12 mi. S of Baker, Graham in 1965 (FPF) ;

Sumpter, Hedgcock FP 998 in 1909 (FPF) and on

P. contorta subsp. latifolia, Weir 3200 in 1913 (ILL) ;

23 mi. S of Baker on Rte. 1, H &W 583 in 1964

(FPF); Buck Mtn., E Oregon [near Unity?], Cusick

2701 in 1901 (MO, POM, RM, UC, US); Halfway,

Miller n.d. (ILL); Squaw Cr., 28 mi. ESE of Prairie

City, Graham in 1965 (FPF). CROOK CO.: Prineville, Harvey FP 4159 in 1910 (FPF); Gerow Butte

rd., Ochoco Nat. For., Childs 52 in 1949 (OSC)

Marks Cr. rd., Ochoco Nat. For., Childs 61 in 1949

(OSC). DESCHUTES CO.: Deschutes River near

Bend, Nelson 866 in 1906 (RM) ; 10 mi. S of Sisters,

Henderson 14130 in 1931 (PH);Pringle Butte, [6 mi.

NW of La Pine] H 989 in 1966 (FPF) and Steward &

Sowder 6803 in 1954 (RSA) ; 9 mi. S of Bend on US

97, H & W 617 in 1964 (FPF). DOUGLAS CO.:

Big Camas Ranger Sta., Kern in 1915 (ILL).

GRANT CO.: Strawberry Mtns., Graham Cr.,

Blue Mtn. Hot Springs, [8 mi. SSE of Prairie City],

Ferris & Duthie 857 in 1919 (DS); 16 mi. S of John

Day on Rte. 395, H & W 577 in 1964 (FPF) ; 10 mi.

NE of Prairie City on Rte. 2^, H & W 580 in 1964

(FPF). HARNEY CO.: T. 24 S., R. 34 E., Sec.

34 [ca. 30 mi. ESE of Burns], Overhy FP 68281 in 1932

(FPF); Calamity Cr., [near Van], Griffith & Morris

803 in 1901 (US); Crow Flat Ranger Sta. [ca. 18 mi.

mi. N of Burns], Porter in 1915 (ILL); Lee Cr., 25

air mi. ESE of Seneca, H efe Scharpf 1265 in 1969

(FPF). HOOD RIVER CO.: Hood River, W 3245

in 1962 (COLO, FPF) and Gorman in 1891 (WTU);

without locality, Henderson 527 in 1923 (MO).

199
JACKSON CO.: 18 mi. SW of Ashland, Graham

in 1963 (FPF); Rogue Elk [near ]McLeod], Stillinger

in 1919 (FPF, ILL, US) ; 12 mi. S of Ruch on Copper

rd., H &W 633 in 1964 (FPF). JEFFERSON CO.:

Warm Springs Indian Res. Chilcls FP 68272 in 1931

(FPF); Shuttle Lake, Metolius Area, Gill FP 68191

in 1932 (FPF) ; Montgomery Ranch, Lower Metolius

River, [13 mi. W of Metolius], Boyce 2121 in 1931

(FPF) and Chilcls 68 in 1931 (OSC). JOSEPHINE

CO.: Waldo [7 mi. S of Cave Jet.], Lewis in 1917

(ILL); on P. contorta subsp. contorta, Weir 3187 in

1916 (ILL); and on P. jeffreyi, Lewis in 1917 (FPF,

ILL). Grants Pass, Weir 3206 in 1916 (FPF, ILL)

and on P. attenuata, Weir 9868 in 1916 (FPF, ILL);

5 mi. W of Grants Pass, on P. jeffreyi, Graham in

1964 (FPF); 5 mi. N of Grants Pass, Ahrams 8677

in 1922 (DS); 6.5 mi. SSW of O'Brien on Oregon

Mtn. rd., H & Hinds 996 in 1966 (FPF) and on P.

attenuata, H & Hinds 995 in 1966 (FPF); Oregon

Mtn. [14 mi. SW of Cave Jet.], Weir 3194 in 1916

(ILL), on P. contorta subsp. contorta, Theisen in 1965

(FPF), and on P. attenuata, Weir 5755 in 1916 (FPF,

ILL) and Bynum in 1967 (FPF); 2 mi. E of Cave

Jet., Theisen in 1965 (FPF); near Lookout Gap, 20

air mi. S of Gahce, onP.jeffreyi, Graham'm 1965(FPF) ;

Hobson Horn Lookout, 25 air mi. NW of Grants Pass,

on P. attenuata, Graham in 1965 (FPF); Bain Sta., 2

mi. W of Oregon Mtn., on P. attenuata, Howard in

1964 (FPF); Oregon Mtn., on P. jeffreyi, Howard in

1964 (FPF); Wonder, on P. jeffreyi, Mitchell FP

68212 in 1932 (FPF); Mt. Peavine, near Gahce, on

P. attenuata, Childs FP 91615 in 1950 (OSC):

Crescent City-Grant Pass rd., on P. attenuata,

Goodding & Root in 1932 (UC); vicinity of Oregon

Caves, Weir 3193 in 1916 (ILL); 4 mi. E of Cave

Jet. H & W 869 in 1966 (FPF). KLAMATH CO.:

Near Jet. of Rtes. 230 and 97, W 3239 in 1962

(COLO); Ringo Butte, Childs 10 in 1946 (OSC);

Brookside Ranch, Swan Lake Valley, "EJ.'' 3187

in 1904 (DS); Hamner Butte, Bedwell & Childs

FP 91033 and FP 91034 in 1939 (OSC) and on P.

contorta subsp. murrayana, Childs 22 in 1939 (OSC)

and Bedwell FP 91035 in 1939 (OSC); 11 mi. NW of

Rte. 97 on Rte. 58, H & W 619 in 1964 (FPF); 14

mi. W of Klamath Falls on Rte. m, H óc W 627 in

1964 (FPF); Applegate Ranch, Swan Lake Valley,

Walpole 417 in 1899 (US); 19 mi. NW of Klamath

Falls on Rte. 140, H & W 872 in 1966 (FPF) ; 14 mi.

SW of Beatty on Rte. 140, H & W 873 in 1966

(FPF). LAKE CO.: Winter Rock Mtns. near

Summer Lake, Ferris & Duthie 422 in 1919 (DS,

RM); Quartz Mtn. Pass, Betting 4255 in 1940

(UC); Warner Mtns., 1 mi. W of Warner Can. Ski

Area on Rte. 140, H (È W 875 in 1966 (FPF).

WALLOWA CO.: Wallowa Mtns., between Bear

Cr. and Minam River, Sheldon 8845 in 1897 (MO,

US). WASCO CO: Between Friend and head of

Fifteen Mile Cr., Milburge 1328 in 1936 (WTU);

Friend, Boyce FP 40179 in 1921 (OSC) ; Bear Springs,

H &W 611 in 1964 (FPF) and Boyce 1907"" in 1930

(FPF). WHEELER CO.: 1 mi. N of Nat. For. bdy.

on Derr Meadows rd., ca. 12 mi. SSE of Mitchell,

H 983"" in 1966 (FPF); 1 mi. N of Ochoco Divide,

11 air mi. W of Mitchell, H 989 in 1966 (FPF);

26 mi. S of Hardman on Rte. 207, H & Scharpf 1271

in 1969 (FPF). COUNTY UNCERTAIN: Umpqua

Mtns., Wilkes Exped. 1185 n.d. (US); Mtns. of

eastern Oregon, Cusick 2324 in 1899 (MO).

WASHINGTON

CHELAN CO.: 8 mi. NW of Chelan, Hedgcock FP

48627 in 1929 (FPF); Rainbow Trail, 1 mi. from

Stehekin River, Ward 595 in 1946 (WTU) ; Peshastin,

Sandherg &. Leiberg 593 in 1893 (MO, UC, US);

Stehekin, Chelan Lake, Jones in 1911 (POM, UC)

and Graham in 1964 (FPF); Antwine Cr. [Antoine

Cr. ?, 8 mi. N of Chelan], Harris FP 68283 in 1932

(FPF); Leavenworth, Umback in 1901 (US) and

Otis 1011 and 1021 in 1920 (CAS, UC). FERRY

CO.: 10 mi. E of Nespelem, Peterson 63-225 in 1963

(FPF). KITTITAS CO.: 8 mi. N of Teanaway on

US 97, Wicker in 1966 (FPF). KLICKITAT CO.:

Trout Lake, W 3246 in 1962 (COLO, FPF); W

Khckitat Co., Suksdorf 1364 in 1892 (MO, UC, US);

1 mi. N of Satus Pass on Rte. 97, H & W 594* in

1964 (FPF); Khckitat, Hoivell 929 in 1881 (US).

OKANOGAN CO.: E of Omak, Fiker 695 in 1932

(DS, MO, US, WTU); Squaw Cr., 5 mi. SW of

Methow, Graham, in 1964 (FPF) ; Pateros, Mitchell

in 1915 (ILL). PEND ORIELLE CO.: Newport, on

P. contorta subsp. latifolia. Weir 3202 in 1916 (ILL).

SPOKANE CO.: 3 mi. N of Spangle, Wicker in 1962

(WSP); Spokane, Weir 8475 in 1917 (FPF, RM,

US) and FP 29660 in 1918 (FPF); Sandberg 925 in

1892 (xMO, US, WTU), and Hubert FP 91292 in 1917

(OSC). 12 mi. S of Spokane, Hansbrough, Mielke &

Joy FP 68273 in 1931 (FPF); Riverside State Park,

NW of Spokane, W 3253 in 1962 (COLO, FPF);

near Spokane, Milburge 253 in 1932 (WTU); Horseshoe Lake (NE part of co.), Yocom in 1947 (WTU);

Tekoa Mtn. [near Latah], St. John 3051 in 1921

(UC); Medical Lake, Elmer 1246 in 1898 (US).

STEVENS CO.: 3.5 mi. S of Gifford, Hedgcock FP

68059 in 1931 (FPF); 4 mi. S of Kettle Falls, Hedgcock FP 49442 in 1929 (FPF) ; S of Gifford, Hedgcock

FP 54990 and FP 54991 in 1931 (FPF); N of Clark

Lake, near Cedonia, Dennis in 1946 (WTU). YAKIMA CO.: Upper Naches River region. Grant in

1938 (WTU); 23 mi. E of White Pass on Rte. 14, 2

mi. W of Nat. For. bdy., H & W 595 in 1964 (FPF).

COUNTY UNCERTAIN: Colville Indian Res.,

Sanpoil drainage, Childs 53 in 1948 (OSC) and

FP 91575 in 1948 (OSC); Cascade Mtns., Tweedy in

1882 (MO).

200
9. ARCEUTHOBIUM CYANOCARPUM

Specimens Examined: on Pinus flexilis unless as

noted.

UNITED STATES

ARIZONA

COCONINO CO.: San Francisco Peaks: On P.

aristata, Leiherg 5884 in 1901 (US) and Fremont

Pass rd., 6 mi. N of Schultz Pass rd., on P. aristata,

H & L 247 in 1962 (FPF).

CALIFORNIA

DEL NORTE CO.: Ship Mtn., ca. 12 air mi. SE of

Gasquet, on P. montícola, Reynolds in 1916 (ILL).

INYO CO. : 17 mi. W of Bishop on Sabrina Lake rd.

H & Scharpf 671* in 1964 (FPF). MONO CO.

Saddle above Convict Lake, Kuijt IJ^löm 1957 (UC)

2 mi. ESE of Lundy Lake, on Pinus sp., probably

flexilis, Hendrix 616 in 1937 (RSA, UC). RIVERSIDE CO.: San Jacinto Mtns., Tahquitz Peak,

Andresen & Cochrane 1729 in 1962 (FPF); Saddle

N of Tahquitz Peak, W 3617 in 1964 (FPF, UT).

SAN BERNARDINO CO.: T. 1 N, R. 3 E, Sec. 6

(4 mi. S of Tiptop Mtn.), Miller FP 98023 in 1962

(FPB). SIERRA CO.: 0.25 mi. N of Hawley Lake

(3 mi. W of Gold Lake), on P. monticola, Menzies

in 1961 (CAS). SISKIYOU CO.: Summit of Black

Butte, on P. alhicaulis. Root & Goodding FP 68275 in

1932 (FPF), Mielke & Zentmeyer FP 89238* in

1937 (FPF) SindH &W 642 in 1964 (FPF) ; N fork

of Molly Cr., on P. alhicaulis, Butler 272 in 1908

(UC); N slope of Mt. Shastina on P. alhicaulis,

Cooke 11576 in 1938 (DS, UC); NW side of Mt.

Shastina, Cooke 16212 in 1941 (MO, PH, UC);

Scott Mtns., Toad Lake (3 mi. S of Mt. Eddy) on

P. monticola. Engelmann in 1880 (ILL). TRINITY

CO.: High Camp Cr., 5 mi. WNW of Mt. Eddy:

On P. monticola. Miller & Bynum* in 1964 (FPF),

Scharpf & Miller in 1965 (FPF), and Broiun in

1965 (FPF), on P. halfouriana. Miller & Bynum

in 1964 (FPF), Scharpf & Miller in 1965 (FPF) and

Brown in 1965 (FPF). COUNTY UNCERTAIN:

Tenderfoot Lake, Siskiyou Nat. For., on P. monticola, Leivisin 1916 (ILL).

COLORADO

BOULDER CO.: 3 mi. S of Ward, H & Gill 195 in

1958 (FPF); Brainard Lake rd. Jet. at Ward, H

417 in 1963 (FPF); Ward, Bethel, Schmoll & Clokey

4314 (MO) and 4^15 in 1921 (CAS, CS, COLO,

JEPS, MO, POM, RM, UC, US, WTU), Bethel

in 1921 (CS), and W 2926 in 1961 (COLO) ; Alienspark, Johnston & Thompson FP 24883 in 1917

(FPF); 1.3 mi. E of Eldora, H 689* in 1964 (FPF);

3 mi. S of Ward on Rte. 160, on P. contorta, H &

Gill 194 in 1958 (FPF) and on P. ponderosa, H &

Gill 198 in 1958 (FPF) ^nà H & W 565 in 1964

(FPF). CLEAR CREEK CO. : Hamlin Gulch rd. 1.4

mi from Fall River rd., ca. 7 air mi. NW of Idaho

Springs, H 1057 and 1058 on P. ponderosa, in 1967

(FPF). EL PASO CO. : Ridge rd., Christ 1877 in 1935

(CS); Rock Cr. (T. 15 S, R. 67 W, Sec. 1), Hill

FP 68276 in 1932 (FPF) ; 5 mi. S of Bruin Inn on

Gold Camp rd., H 54O* in 1963 (FPF); Pikes Peak,

Lake Moraine, Eioan 15154 in 1943 (COLO); Pikes

Peak, Hedgcock FP I6I4 in 1909 (FPF) and FP 19185

in 1911 (FPF); Machride 2677 in 1913 (J\IO);

Fremont Exp. For. (2 mi. W of Manitou Springs),

Hedgcock FP 22706 and 22707 in 1916 (FPF) and

Hedgcock, Payson & Johnston FP 24852 in 1917

(FPF); Pikes Peak, jNIinnehaha, Hedgcock FP 635

in 1909 (FPF); Pikes Peak For. Reserve, Flintham

in 1903 (US); Pikes Peak, Halfway, Hartley FP

1774 in 1909 (FPF) and Hedgcock FP 15909 in 1914

(FPF, RSA); Rampart Range rd., 3 mi. SE of

Woodland Park, Schacht in 1964 (FPF). Crystal

Park, Gill FP 68235 in 1932 (FPF). GILPIN CO.: 1

mi. SW of Tolland, H 827 in 1965 (FPF). (FPF).

FREMONT CO: Sangre de Cristo Range, 8 mi. SW

of Coaldale on Hayden Pass va., H óc Laut 1317 in

1970 (FPF). LARIMER CO.: Eaton Reservoir,

Douglass in 1957 (CS) ; 4 mi. W of Eaton Reservoir,

Hinds* in 1962 (FPF) ; Estes Park, Cooper 65 in 1904

(RM) ; Boulder Ridge, 0.5 mi. S of Sand Cr. rd., H 82

in 1961 (FPF); 1.6 mi. E of Pingree Park on main

rd., H 1055 in 1967 (FPF); Rocky Mtn. Nat. Park,

Gem Lake Trail, Ashton 66 in 1930 (RMNP) ; Rocky

i\Itn. Nat. Park, Glacier Gorge, Loll FP 11832 in

1924 (FPF); Boulder Ridge, H 196 in 1961 (FPF);

Boulder Ridge, 0.5 mi. S of Wyoming bdy., H 84

in 1961 (FPF) ; 6.5 mi. S of Estes Park on Rte. 7,

H & Staley 288 in 1962 (FPF); Pennock Cr., 2 mi. S

of Pingree Park rd., H 562 in 1964 (FPF); Medicine

Bow Mtns., 4.3 mi. S of Wyoming bdy. on Roach

rd., H829 in 1965 (FPF). COUNTY UNCERTAIN:

La Sal Mtns., col.? in 1915 (FPF, ILL).

IDAHO

BEAR LAKE CO. : Ridge S of Bloomington Lake,

W of Bloomington, Peterson 65-370 in 1965 (FPF).

BUTTE CO.: Craters-of-the-Moon Nat. Mon.,

Peterson 62-53* in 1962 (FPF) and Mahoney in 1965

(FPF). LEMHI CO.: Near Salmon, Wolpert in 1917

(FPF, ILL).

MONTANA

BEAVERHEAD CO.: Big Hole Valley, on P.

alhicaulis, Weir 3216 n.d. (FPF, ILL). BIG HORN

CO.: Crow Res., Bighorn Mtns., Rotten Grass

drainage, Gregory in 1969 (FPF). CARBON CO.:

Rock Cr. Ranger Sta., Hedgcock FP III47 in 1911

201
(FPF); 5 mi. W of Red Lodge on Rock Cr. rd.,

W 3865 in 1965 (FPF, UT). Pryor Mtns., Crooked

Cr., Brown in 1969 (FPF). DEERLODGE CO.:

French Gulch near Anaconda, Weir 321^6 in 1913

(FPF, ILL) ; Mt. Haggin near Anaconda, Weir 3218

in 1914 (FPF, ILL) ; near Anaconda, Weir 825^ in

1914 (ILL). GALLATIN CO.: Rocky Can., near

Bozeman, Blankinship 436 in 1905 (POM, RM,

US) : 7 mi. E of Bozeman, W 3257* in 1962 (COLO,

FPF): Bozeman, Brandegee in 1883 (UC) and

Langohr in 1917 (ILL). COUNTY UNCERTAIN:

Mt. Powell, Weir 8345in 1914 (ILL).

OREGON

COUNTY UNCERTAIN: Obsidian Cliff, on P.

albicaulis, Gorman 1753in 1903 (US).

NEVADA

CLARK CO.: Charleston Mtns.: On P. aristata,

Coville & Funston 311 in 1891 (US); head of Tractor

Lane, Lee Can. Train 2150 in 1938 (UC). Peak

Trail, Clokeij 5430 in 1935 (CAS, DS, FPF, RM,

RSA, UC, US, WTU), and on P. aristata, Clokey

5429 in 1935 (CAS, DS, FPF, JEPS, UC, WTU)

and Clokey 5492 in 1935 (CAS, DS, JEPS, RM, RSA,

WTU). ELKO CO.: Ruby Mtns.: (kover in 1958

(FPF) and Lamoille Can., 12 mi. SE of Lamoille,

i7*TF575*in 1964 (FPF).

UTAH

CACHE CO.: Logan Can., near Jardine Juniper,

Krebill 454 in 1967 (FPF); Tony Grove Lake,

Maguire & Maguire 20377 in 1939 (RM, WTU) and

20378 in 1939 (RM); 0.25 mi. NW of Tony Grove

Lake, Peterson 62-75 in 1962 (FPF) ; Mt. Naomi, 1

mi. from Tony Grove, Snell 1047 in 1938 (UC,

WTU). DAGGETT CO.: 2 mi. W of Utah, Wyo.,

and Colo, corner on Loop rd. from Clay Basin,

Bleazard in 1965 (FPF, UT). DUCHESNE CO.:

Indian Can., 25.5 mi. SW of Duchesne on Rte. 33,

H 571 in 1964 (FPF). GARFIELD CO.: 10 mi. W

of Bryce Can. Lodge, Dorn in 1947 (POM); near

Escalante, Peterson 52-61 in 1961 (FPF); Bryce

Can. Nat. Park; Headquarters area. Gill FP 68144

in 1932 (FPF) and on P. aristata, FP 68143 in 1932

(FPF); Garrett FP 38107"" in 1921 (FPF) and on P.

aristata, FP 38106 in 1921 (FPF); H 255 in 1962

(FPF) and on P. aristata, 256 in 1962 (FPF) ; Bryce

Point, W 3034 and 3178 in 1962 (COLO, FPF),

Howell & Eastwood 7204 n.d. (CAS). IRON CO.:

16.5 mi. E of Cedar City on Rte. 14, H 679 in 1964

(FPF) and on P. aristata, H 680 in 1964 (FPF) ; 1 mi.

W of Summit on Rte. 14, W 4'i-27 in 1966 (FPF, UT)

and on P. aristata, W 4125 in 1966 (FPF, UT).

KANE CO.: 1 mi. N of Navajo Lake on Rte. 14,

W 4124 in 1966 (FPF, UT). UTE CO.: 8 mi. S of

Belknap Guard Sta., above Deep Cr., Peterson

65-339 in 1965 (FPF). COUNTY UNCERTAIN:

Charles Peak, southern part of State, Siler in 1879

(MO).

WYOMING

ALBANY CO. : 1.0 mi. N of Albany rd. on Cinnabar

Park rd., H & Hinds 197 in 1961 (FPF); Centennial

Mt., Nelson in 1915 (RM); Laramie Mtns., Nelson

8247 in 1901 (ARIZ, MO, RM, US) : Pilot Knob,

Nelson in 1905 (MO, RM); Boulder Ridge, 7 mi.

SW of Tie Siding, H 85 in 1961 (FPF); Sheep Mtn.,

Nelson 3320 in 1897 (RM, US); Hermosa, Garrett

FP 38174 in 1921 (FPF); Centennial, Mullin Cr.,

Garrett FP 15833 in 1914 (FPF); Dale Cr. Bridge

near Sherman, col.? n.d. (MO); Laramie Mtns.,

Weir 9872 in 1918 (FPF, ILL), 2 mi. SW of Pilot Hill,

H & Bailey 1008 in 1966 (FPF); Boulder Ridge, 10

feet N of Colorado bdy., 7 mi. SW of Tie Siding, on

P. ponderosa, H 284 m 1962 (FPF). CARBON CO.:

Medicine Bow Mtns.: 3 mi. S of Pennock Mtn.,

Landgrafin 1965 (FPF) and Cooper Cr., 12 air mi. N

of Centennial, Landgrafin 1965 (FPF); Ferris Mtns.,

NE slope of Whiskey Gap, Dunder in 1969 (FPF);

Shirley Mtns., Lower Prior Flat, Dunder in 1969

(FPF); N side of Elk Mtn., Rattlesnake Cr.,

Dunder in 1969 (FPF). CONVERSE CO.: Laramie

Mtns., Cold Springs, Simmons in 1917 (ILL).

FREMONT CO.: Slate Cr., SE Lander District,

Peterson in 1960 (FPF) ; Fossil Hill, ca. 10 air mi.

SW of Lander, Steioart in 1967 (FPF); W fork of

Long Cr., 21 mi. WNW of Dubois, H 277 in 1962

(FPF) ; 4 mi. WNW of South Pass City, Landgraf &

Davel in 1965 (FPF); Green Mtns., Crook Gap,

H & Landgraf 279* in 1962 (FPF). LINCOLN CO.:

Alpine, along Snake River, Payson & Armstrong

34O6 in 1923 (MO, PH, RM);E of Alpine, Graham,

in 1965 (FPF). PARK CO.: ^^Cody,'' Weir FP

91249* in 1918 (FPF, OSC, ILL); Cody Can., 44 mi.

W of Cody, W 3258 in 1962 (COLO) ; Rattlesnake

Cr., ca. 15 air mi. WNW of Cody, Bailey in 1966

(FPF); Dry Fork, ca. 19 air mi. NW of Cody,

Stewart in 1967 (FPF); along switchbacks E of

Dead Indian Campground, ca. 22 air mi. NW of

Cody, Steivart in 1967 (FPF). YELLOWSTONE

NAT. PARK: ^'Yellowstone For. Reserve,^' Hapema?iinl906(RM).

10. ARCEUTHOBIUM DIVARICATUM

Specimens Examined: On Pinus edulis except as

noted.

UNITED STATES

ARIZONA

APACHE CO.: Carisso, Brandegee 1226 in 1875

(PH, UC); 2 mi. S of Eagar, H & L 208 in 1962

202
(FPF) ; 17 mi. NW of Sawmill on Can. de Chelly rd.,

L & Weiss 66-45 in 1966 (FPF); 3 mi. E of Lukachukai, L & Weiss 66-^7 in 1966 (FPF) ; 8 mi. SW of

Red Rock on Lukachukai rd., L <& Weiss 66-52 in

1966 (FPF). Can. de Chelly: 6 mi. SE of Chin Lee,

Ferris 10183 in 1940 (UC) and 5 mi. W of Tsalie

Dam on N Rim of Can. del Muerto, L, Ela &

Lampi 65-54 in 1965 (FPF) ; 19 mi. W of Maverick

on White River vd., H & L 904 in 1966 (FPF).

COCONINO CO.: Grand Can. Nat. Park: South

Rim: Towney in 1894 (UC), MacDougal 190 in 1898

(ARIZ, UC, US), Knowlton 295 in 1889 (US),

Spaulding FP 303 in 1909 (FPF), Hedgcock FP 4907

in 1910 (FPF), Hitchcock 25 in 1913 (US), Batram

435 in 1920 (US), Gill FP 68232 and FP 68233 in

1932 (FPF); Desert View: Cronquist in 1938 (RM),

Gill FP 68234 in 1932 (FPF), and H & Scharpf 701

in 1964 (FPF); 1 mi. SW of Bright Angel Camp,

Gill FP 68132 in 1932 (FPF) ; Bright Angel Trail,

Gill FP 68230 in 1932 (FPF) ; Grandview, Dachnoivski-Stokes in 1913 (US); El Tovar, Setschell in 1907

(UC); Duck-on-a-Rock, H 35 in 1954 (FPF) and

H & Scharpf 703 in 1964 (FPF). Flagstaff, Jones

3973 in 1884 (ARIZ, DS, POM, RM, UC); 14 mi. N

of Frazier Well, H & L 187 in 1962 (FPF) ; between

Flagstaff and Grand Can., 14 mi. SE of Rte. 64 on

Rte. 164, H 246 in 1962 (FPF); NE of Long Valley,

Ellis FP 89426 in 1939 (FPF) ; Sedona, Hedgcock FP

4917 in 1910 (FPF); North Kaibab Plateau on US

89, Gill FP 68300 in 1934 (FPF) ; 6 mi. E of Jacob

Lake on US 89, W 3185 in 1962 (COLO, FPF);

Kaibab For., Richards in 1938 (WTC); Jacob Lake

to Fredonia, Kearney & Peebles 13761 in 1937

(ARIZ, US); 30 mi. SW of Winslow on Rte. 65,

H & L 915 in 1966 (FPF); 34 mi. S of Winslow on

Chevlon rd., H & L 916 in 1966 (FPF). Walnut Can.

Nat. Mon., L 66-22 in 1966 (FPF) ; 15 mi. S of Page

on US 89, L & Weiss 69-4 in 1969 (FPF). GILA CO. :

6 mi. SE of Cedar Cr. on Rte. 73, H & L 906 in 1966

(FPF). GRAHAM CO.: San Carlos Indian Res., 3

mi. W of Eagle Cr. on Malay Gap rd., H & L III4

in 1968 (FPF). GREENLEE CO.: Red Hill near

Blue, Ellis FP 89331 in 1939 (FPF). MOHAVE

CO.: Mt. Trumbull, Hevly in 1959 (MNA); Grand

Can. Nat. Mon., 5 mi. S of Nixon Spring, H, L &

Laut 1165 in 1969 (FPF); 4 mi. E of Mt. Trumbull

P. 0., H &L 1168 in 1969 (FPF) : 10 mi. W of Wolf

Hole on Black Rock Mtn. rd., on P. monophylla,

H,Lâc Laut 1171m 1969 (FPF).

NAVAJO CO. : 4.5 mi. S of Rte. 264 on Rte. 77, ca.

12 air mi. SE of Keams Can., H & L 1084 in 1967

(FPF), 15 mi. E of Heber, Ellis FP 89337 in 1939

(FPF); 2 mi. W of Heber, L 64-19 in 1964; between

Snowflake and Show Low, Hevly in 1959 (MNA); 7

mi. W of US 60 on Mogollón Rim rd., H & L 922

in 1966 (FPF); Navajo Nat. Mon., Betatakin Can.,

Wetherill 919/3149 in 1937 (ARIZ, MNA) and

Howell 24505 in 1948 (CAS) ; Pinon, Whiting 854/

2825 in 1937 (ARIZ, MNA); 4 mi. SE of Fort

Apache on Maverick rd., H & L 905 in 1966 (FPF).

YAVAPAI CO.: Mingus Mtn., 5 mi. W of Jerome

on US 89, H & L 245 in 1962 (FPF). COUNTY

UNCERTAIN: Mahogany Range, Point Daguerre,

Palmer 2 in ISmÇUC).

CALIFORNIA

All California Collections on Pinus monophylla

INYO CO. : 8.5 mi. W of Lone Pine on Mt. Whitney

trail, Gill & Wright FP 68061 in 1931 (FPF); 10 mi.

^^above" Lone Pine, Schrieher 1017 in 1934 (UC);

SE side of Mt. Whitney, Armstrong 1024 in 1934

(UC) ; 3 mi. W of Lone Pine, Kuijt 1389 in 1957 (UC) ;

Inyo Mtns., Whippoorwill Flats on Saline Valley rd.,

W 2740 in 1960 (RSA). Death Valley Nat. Mon.:

Wildrose Can , Stewart in 1935 (US) and head of

Death Valley Can., Coville & Gilman 120 in 1931

(US). LOS ANGELES CO.: San Gabriel Mtns.:

Mescal Cr., Ewan 9941 in 1936 (MO) and Swarthout

Can., Putnam n.d., (ILL). MONO CO.: 6 mi. S of

Coleville, Wagener FP 89604 in 1936 (FPF) and

Kuijt 1413 in 1957 (UC); 0.5 mi. S of Coleville,

Peterson 63-131^ in 1963 (FPF). VENTURA CO.:

Mt. Pinos, Seymour Cr., Peirson 3251 in 1922

(POM, RSA); Mt. Pinos area, W 3217* in 1962

(COLO, FPF); E edge of Fraser Mtn., Putnam FP

98054 in 1919 (FPB); Fraser Mtn., Root in 1919

(ILL). SAN BERNARDINO CO.: San Bernardino

Mtns. : Johnson Grade, Little Bear, Gill FP 89420

in 1939 (FPF); Big Bear Ranger Sta., Gill & Wright

FP 68239 in 1931 (FPF); Dobel Mine, Bear Lake,

Gill & Wright FP 68064 in 1931 (FPF) ; Arrastre Cr.,

T. 2 N, R. 2 E, Sec. 11, Miller FP 68024 in 1962

(FPB); Arrastre Cr., 3 mi. SE of Baldwin Lake,

Jaeger in 1938 (POM); NE slope San Bernardino

Mtns., Parish & Parish I442 in 1882 (MO, UC, US) ;

N side of Sugarloaf, Mum 10771 in 1926 (POM)

1 mi. N of Baldwin Lake, H 733 in 1965 (FPF)

Bear Valley, Lieherg 3312 in 1898 (US). New York

Mtns., 5 mi. SE of Ivanpah, Gill & Wright FP 68238

in 1931 (FPF). Providence Mtns.: Brandegee in 1902

(UC); S fork of N fork of Fountain Can., Beat 660

in 1939 (JEPS) ; near Bonanza King Mine, Aiunz et

al in 1920 (POM) and Shields in 1965 (FPF).

COUNTY UNKNOWN: Bigelow in 1853-54 (US).

COLORADO

DELTA CO.: S of Delta, Hedgcock FP 9299 in 1911

(FPF). DOLORES CO.: T. 39 N, R. 17 W, Sec. 9

[ca. 4 mi. E of Cahone], Hinds 63-10 in 1963 (FPF).

GARFIELD CO.: West Elk Guard Sta., N of

Newcastle, Hinds in 1961 (FPF). LA PLATA CO.:

Without locality. Curran in 1914 (US); Durango,

Tweedy 595 in 1896 (US); N of Durango, Hunt á

Bethel FP 24428 in 1917 (FPF) ; Bayfield, Hedgcock &

203
Bethel FP 24674 in 1917 (FPF). MESA CO.:

Colorado Nat. Mon.: Wilford in 1961 (FPF); 2 mi.

S of Fruita, W 3168 in 1962 (COLO, FPF). MONTEZUMA CO.: Mancos: Baker, Earle & Tracy 94 in

1898 (CAS, MO, POM, RM, US), Tracy, Earle &

Baker 388 in 1898 (DS), and Eastwood in 1891

(COLO); 3 mi. S of Mancos, Peterson 30-61 in 1961

(FPF); 1 mi. NE of Mancos, Hinds in 1965 (FPF).

Mesa Verde Nat. Park: Erdman 120 in 1959 (COLO,

WTU), Weher 6065 in 1951 (COLO), and Bethel &

Hunt FP 29261 in 1918 (FPF); N entrance, H &

Scharpf 691* in 1964 (FPF); Spruce Tree House,

Mathias 652 in 1929 (MO). MONTROSE CO.: 13

mi. SW of Montrose, Hinds in 1960 (FPF) ; T. 48

N, R. 70 W, Sec. 9 [5 mi. W of Cimarrón], Hinds

63-14 in 1963 (FPF); 20 mi. N of Placerville,

Pesman in 1960 (CS) ; T. 46 N, R. 12 W, Sec. 7 [1 mi.

E of Ute], Hinds 63-17"" in 1963 (FPF); 11 mi. E of

Naturita on Rte. 90, H 1087 in 1967 (FPF); 8 mi.

W of Bedrock on Rte. 90, H 713 in 1965 (FPF).

OURAY CO.: 2 mi. SE of Ridgway, Payson FP

26016ml^n Ç^VF).

NEVADA

All Nevada Collections on Pinus monophylla

CHURCHILL CO.: Carroll Summit, 10 mi. E of

Eastgate on US 50; /Í cfc T^ 835 in 1966 (FPF).

CLARK CO. : Charleston Mtns. : Coville & Funston

308 in 1891 (US); Deer Cr. Campground, W 3207 in

1962 (COLO, FPF); Kyle Can., Clokey 8565 in 1939

(ARIZ, CAS, CS, MO, POM, RM, RSA, UC,

WTU) ; 2 mi. E of Nat. For. bdy. on Rte. 58, H &

Scharpf 672 in 1964 (FPF). Sheep Mtns., Spring

Can., Johnson in 1969 (FPF). DOUGLAS CO.:

Pine Nuts Hills, S of Carson City, Peterson in

1960 (FPF) ; 9 mi. S of Gardnerville, Wagener FP

89603 in 1936 (FPF) ; near Minden, McKlevey I464

in 1930 (US) ; ca. 7 mi. N of Topaz Lake on US 395,

Krehill & Nelson 467 in 1967 (FPF). ELKO CO.:

Big Springs near Cave Cr. P. 0., Mason 4748 in 1928

(UC). ESMERALDA CO.: White Mtns., Roos 1028

in 1937 (POM). LANDER CO.: 10 mi. E of Austin

on US 50, i/ * TF 833 in 1966 (FPF); Austin,TFezr

in 1916 (FPF, ILL, UC) and Jones in 1882 (POM);

Big Cr., Linsdale & Linsdale 264 in 1931 (CAS).

LINCOLN CO.: Deer Lodge, E part of co. [near

Fay], Train 2521 in 1938 (ARIZ, UC). LYON CO.:

Wassuk Range, Coal Valley, Alexander & Kellogg

5328 in 1947 (UC, WTU); 6 mi. SE of Wellington,

McMillan 197 in 1941 (RSA, UT) MINERAL CO.:

Alum Cr. S of Hawthorn, Peterson 63-327* in 1963

(FPF). NYE CO.: Grant Range, Irwin Can. near

Timber Mtn., Peterson 63-329* in 1963 (FPF); E

side of Pahute Mesa, Bostick 5237in 1967 (DS, FPF).

WHITE PINE CO. : E of Lehman Caves Nat. Mon.,

Peterson 63-338* in 1963 CFPF) and Maguire & Becraft

2535a in 1933 (UC).

NEW MEXICO

BERNALILLO CO.: Manzano Mtns., Cedro Can.,

Ellis FP 89502 in 1939 (FPF) ; Sandia Mtns., 0.5 mi.

W of Doc Long Campground, H 36 in 1956 (FPF)

and L in 1962 (FPF). CATRON CO.: Mogollón,

Hedgcock and Long FP 9936 in 1911 (FPF) ; Mogollón

Cr., Metcalfe 288 in 1902 (ARIZ, DS, FPF, MO,

POM, UC, US); 2 mi. W of Mogollón, H & L 133

in 1962 (FPF) ; Adams Diggins, 18 mi. NW of Pie

Town, L 65-5 in 1965 (FPF) ; 6.5 mi. S of Luna on

Rte. 180, H 817 in 1965 (FPF); Diamond Cr., 13

mi. S of Beaverhead on Rte. 61, L 65-41 in 1965

(FPF) ; 20 mi. N of Rte. 78 on Apache Springs rd.

L 65-44 in 1965 (FPF) ; 11 mi. SE of Reserve on new

rd. to Beaverhead, H & L 899 in 1966 (FPF) ; 1 mi.

W of Red Hill on US m, H & L 926 in 1966 (FPF) ;

11 mi. S of US 60 on Rte. 32, H & L 927 in 1966

(FPF). GRANT CO.: Telegraph Mtns., Wooton in

1902 (POM, RM, US) ; Walnut Cr., near Silver City,

Gill FP 68301 in 1934 (FPF) ; Fort Bayard, Johnson

FP 15106 in 1913 (FPF), Hedrick FP 4I and FP 286

in 1908 (FPF); Stephens Ranch, Fort Bayard

Watershed, Blumer 110 in 1905 (US); 5 mi. W of

Mimbres River on Rte. 180, H & L 125 in 1962

(FPF); 21 mi. N of Mimbres on Rte. 61, L 65-38 in

1965 (FPF). LINCOLN CO.: 1 mi. S of Nogal on

Rte. 37, H & L 101 in 1962 (FPF); Gallina Mtns.,

SW of Corona, Riffle in 1964 (FPF) and ^ cfc L

1078 in 1967 (FPF); 3 mi. SE of Ancho, L 64-30 in

1964 (FPF). MCKINLEY CO. : Continental Divide,

25 mi. E of Gallup on US 66, Ellis FP 89503 in 1939

(FPF) and L 64-12 in 1964 (FPF); 2 mi. NE of

San Mateo, L 63-29 in 1963 (FPF); 0.5 mi. S

of Old Fort Wingate, L 63-36* in 1963 (FPF).

OTERO CO.: Pinon, Wyman FP 1838 in 1914

(FPF); Box Can. near Highrolls, Viereck in 1902

(PH). RIO ARRIBA CO.: 5 mi. W of Coyote

on US 84, L 63-18 in 1963 (FPF); 15 mi. NW of

Española, L 63-136 in 1963 (FPF); 2.5 mi. W of

Canjilon, L 63-6 in 1963 (FPF); near Gobernador,

L 64-9 in 1964 (FPF); Carson Nat. For., La Jara

Cr., on Rte. 17, L 64-II in 1964 (FPF). SANDOVAL

CO. : Sandia Mtns., N side. Gill & Long FP 68137 in

1932 (FPF) and Ellis FP 89411 in 1939 (FPF); 14

mi. NW of Cuba on Rte. 44, H & Scharpf 692* in

1962 (FPF). SAN JUAN CO.: Near Cedar Hill,

Standley 7979 in 1911 (US); Navajo Res., Tunitcha

[Chuska] Mtns., Standley 7810 in 1911 (US); 5 mi.

SW of Sheep Springs, L & Weiss 66-4I in 1966

(FPF). SAN MIGUEL CO. : 3 mi. N of Pecos on Rte.

63, L 65-30 in 1965 (FPF). SANTA FE CO.: Santa

Fe, Heller & Heller 3533 in 1897 (FPF, MO, US);

Mesa NE of Santa Fe, Hedgcock & Bethel FP 24730

in 1917 (FPF, ILL); Mtns. E of Santa Fe, Fendler

312 1847 (MO); 20 mi. E of Española, L 63-16 in

1963 (FPF) ; near Rio en Media, H & Hinds 1077 in

1967 (FPF); Canoncito, Brandegee in 1879 (UC);

204
vicinity of Santa Fe, Rose & Fitch 17765 in 1913

(US) SIERRA CO.: Rhodes Pass, 20 mi. E of

Engle, Cutler 2047 in 1938 (CAS, ^10, UC) ; 2 mi. W

of Kingston on lite. 90, H&L 889 in 19G6 (FPIOSOCORRO CO.: San Mateo Mtns., 10 mi. S of Lb

60 on Rte. 52, L 65-33 in 1965 (FPF); 5 mi. W of

Magdalena on US 60, L 63-41 in 1963 (FPF) ; 25 mi.

W of Carrizozo on US 380, L 65-1 in 1965 (FPF ) ;

7 5 mi W of Claunch, L 65-20 in 1965 (FPF).

TAOS CO : Taos, Greene in 1877 (MO) ; Peñasco,

V071 Schrenk in 1907 (^10); 19 mi. S of Tres Piedras

on US 285, L 63-11 in 1963 (FPF); 11 mi. N ot

Taos on Rte. 3 at San Cristobal Jet., L 65-A5 m 196o

(FPF)- 12 mi. S of Tres Piedras on LTS 285, H 1011

in 1967 (FPF). VALENCL\ CO.: 2.3 mi. W of

Cebolleta, L 65-6 in 1965 (FPF); 3 mi. NW of

Trechado on Rte. 36, L 65-U in 1965 (FPF); 10 mi.

E of Grants on rd. to La :\Iosca Lookout, L 6o-50 m

1965 (FPF). COUNTY UNCERTAIN: Alamo Nat.

For , Long FP 19634 in 1915 (FPF) ; Embudo to Rio

Pueblo, Ó'Bryne & Long FP 18678 in 1914 (FPF).

TEXAS

JEFF DAVIS CO. : Davis Mtns. : Young 63 in 1914

ÇSlO); INIt. Livermore, Andresen & Andresen 1824*

in 1962 (FPF) ; Wood Can., Jones Ranch, Hinckley

in 1936 (ARIZ); north slope of Sawtooth yitn., on

P. cemhroides, Little & Correll 19033 in 1963 (FPF);

Mt. Livermore, Goat Can., on P. cemhroides,

Hinckley in 1935 (SRSC); H. 0. Can., on P. cemhroides, Hinckley in 1935 (SRSC); H. 0. Can., on

P. cemhroides, Hinckley 3163 in 1944 (SRSC); H. 0.

Can., 1 mi. S of summit on Rte. 166, on P. cemhroides,

H,L& Lampi 1050 in 1967 (FPF).

UTAH

CARBON CO.: Price, Floivers in 1934 (FPF, UT);

Sunnyside, Jones in 1907 (POM). DUCHESNE

CO • 11 mi. W of Duchesne on US 40, H 1052

in 1967 (FPF). GARFIELD CO.: 5 mi. SW of

Panguitch, Peterson 62-107 in 1962 (FPF); 2 mi.

W of Tropic, Buchanan 527 in 1959 (UT, WTU);

SE of Table Cliffs, Peterson 64-87 in 1964

(FPF); 21 mi. SW of Escalante on Rte. 54, H

682* in 1964 (FPF); Henry Mtns., NW slope of

Bull Mtn., Harrison III84 in 1947 (UC). IRON CO. :

Near Cedar City, Parry in 1874 (MO). JUAB CO.:

Deep Cr. Range, Stewarts Cabin, on P. monophylla,

Holmgren 3748 in 1944 (UC, WTU). KANE CO.:

0.5 mi. N of Glendale, W 3179 in 1962 (COLO,

FPF); 4 mi. SE of Mt. Carmel on US 89, H & Laut

1164 in 1969 (FPF). MILLARD CO.: House Mtns.

col? in 1938 (UT). PIUTE CO.: 3 mi. SW of

Marysvale, W 4I68 in 1967 (FPF, UT). SAN JUAN

CO.: 5 mi. N of Blanding on Rte. 47, Peterson

65-69 in 1965 (FPF) ; 8 mi. N of Blanding on Rte.

47, H 1089 in 1967 (FPF). SANPETE CO.: SE

of Ephraim, Hedgcock FP 8142 in 1912 (FPF); W of

Ephraim, Hedgcock FP 8135 and FP 8139 in 1912

(FPF) and on P. monophylla, Tidestrom 1300 in 1908

(US). WASHINGTON CO.: Zion Nat. Park, on P.

monophylla, Xelson & Nelson 2697 in 1938 (RM);

10 mi. N of Beaver Dam Summit, on P. monophylla,

W 3913 in 1966 (FPF, UT) ; 1 mi. N of Pine Park, ca.

16 air mi. W of Enterprise, H, L & Laut 1174 in 1969

(FPF). WAYNE CO.: Aquarius Plateau, Head of

Poison Cr., Rydberg & Garitón 7371 in 1905 (RM,

US); 2 mi. S of Grover, W 3171 in 1962 (COLO,

FPF). COUNTY UNCERTAIN: La Sal Altns.,

Little Springs on W slope, Rydberg & Garrett 8542

inl911(RM,US).

MEXICO

BAJA CALIFORNIA: Sierra Juárez: 21 mi. S of

La Rumorosa on rd. to Laguna Hanson, on P.

quadrifolia, L & GiU 64-34 in 1964 (FPF); 16.5 mi.

N of Laguna Hanson, on P. quadrifolia, H & Scharpf

75ein 1965 (FPF).

11. ARCEUTHOBIUM DOUGLASII

Specimens Examined: on Pseudotsuga menziesii

except as noted.

CANADA

BRITISH COLU^^IBIA: 1 mi. S of Sirdar, N of

Cî-eston, Cahier & Savile 11413 in 1953 (WTU);

Summerland Exp. Sta., Calder & Savile 11536 in

1953 (UC, US) ; Similkameen River, Macoun 79522

in 1905 (MO) ; 10 mi. SE of Kelowna, Kuijt 588 in

1954 (DAYFP, FPF); Beaver Lake rd., near Wood

Lake, Kuijt 581 in 1954 (DAYFP, FPF); 10 mi. N

of Sirdar, Kmjt 650 in 1954 (DAYFP, FPF).

MEXICO

COAHUILA: Sierra del Carmen, H, L & Muñoz

1027 in 1967 (FPF). DURANGO: 47 mi. S of

Durango on La Flor rd., H & W 514 in 1963 (FPF).

NUEYO LEON: Cerro Potosí, 11 mi. from village

"18 de Marzo" on rd. to relay tower, H & W 395*

in 1963 (FPF).

UNITED STATES

ARIZONA

APACHE CO. : 4 mi. S of Eagar, H & L 209 in 1962

(FPF); Alpine, Goodding FP 26737 in 1917 (FPF);

4.7 mi. SW of Alpine on Rte. 666, H & Scharpf 699

in 1964 (FPF); Milk Can., 8 mi. E of Nutrioso,

Parker & McClintock 7587 in 1951 (ARIZ) ; 5 mi. E

205
of Lukachukai on Red Rock rd., L & Weiss 66-48

in 1966 (FPF); 11 mi. SW of Red Rock on Lukachukai rd., L & Weiss 66-50 in 1966 (FPF) ; 2 mi. E

of Big Lake, H 820 in 1965 (FPF); 3 mi. NE of

Maverick on Fort Apache rd., H & L 902 in 1966

(FPF); 13 mi. E of McNary on Rte. 73, H & L

924 in 1966 (FPF). COCHISE CO.: Chiricahua

Mtns.: Pinery Can. rd., Gill & Ellis FP 89497 in

1939 (FPF); Pinery Can. rd., 7 mi. E of Nat. For.

bdy., H & L 152'' in 1962 (FPF); Rustler Park,

Gill FP 68056 in 1935 (FPF). Huachuca Mtns.,

Carr Can., Goodding FP 33284 in 1919 (FPF) and

Goodding 2088 in 1915 (ARIZ). COCONINO CO.:

''Kaibab For.,'^ Richards in 1938 (ARIZ, R:M,

WTU); San Francisco Peaks, Knowlton 40 in 1889

(US) ; San Francisco Peaks, Interior Basin, H & L

251 in 1962 (FPF); Grand Can., South Rim, 1 mi.

E of Grandview Point, H 39 in 1954 (FPF) and

H & L 183 in 1962 (FPF); Oak Cr. Can., Ellis

FP 89504 in 1939 (FPF); Jacob Lake, Moberg &

Gill FP 68299 in 1934 (FPF); .Alt. Eldon, near

Flagstaff, Drake FP 15122 in 1913 (FPF); San

Francisco Peaks, Spruce Cabin, Gill FP 68134 in

1932 (FPF); Flagstaff, Purpus in 1900 (UC); N

slope, Mt. Eldon, Whiting 1053/5338 in 1941

(ARIZ); Grand Can., Grandview Trail, Thornher &

Hockdoerffer 2944 in 1907 (ARIZ); San Francisco

Peaks, Goodding in 1917 (ARIZ); Baker Butte,

Andrews, Ellis & Gill FP 89229 in 1937 (FPF) ; Rim

rd., 5 mi. SE of W bdy. of Sitgreaves Nat. For.,

L 64-16 in 1964 (FPF); 3 mi. E of Long Valley, L

64-14 in 1964 (FPF); North Kaibab Plateau, 7 mi.

W of Kaibab Lodge, on Picea engelmannii^ Peterson

45-61 in 1961 (FPF); San Francisco Peaks, 7 mi.

NE of Schultz Pass on Interior Basin rd., on Ahies

lasiocarpa var. arizonica, H & L 250 in 1962 (FPF);

near Weimer Springs, W of Mormon Lake, H & L

913 in 1966 (FPF); 11 mi. S of Chevlon Ranger

Sta., // & L 918 m 1966 (FPF). GILA CO.: Sierra

Ancha, Workman Cr., H 44 in 1956 (FPF) and

L & Lampi 64-15 in 1965 (FPF) ; 26 mi. E of Payson

on Rte. 160, L 63-55 in 1963 (FPF). GRAHAM

CO.: Graham Mtns.: 1 mi. above Arcadia Guard

Sta., H & L 142 in 1962 (FPF); Hospital Flat,

H & L 226 in 1962 (FPF); Turkey Flat, Gill FP

68037 in 1936 (FPF); Treasure Park, Mielke &

Ellis FP 89694 in 1944 (FPF); Grant Cr., on Ahies

lasiocarpa var. arizonica, Stouffer FP 68239 in 1934

(FPF). GREENLEE CO.: 47 mi. S of Springerville

on US 666, Gill FP 68303 in 1934 (FPF); Blue

Summit, Gill FP 68307 in 1934 (FPF); 3 mi. S of

Hannagan Meadows on Rte. 666, Andrews in 1963

(FPF). NAVAJO CO.: 6 mi. E of Rte. 160 on

Mogollón Rim rd., H & L 920 in 1966 (FPF).

PIMA CO.: Santa Catahna Mtns.: Toumeij in 1896

(US) ; Soldier Camp, Gill FP 68127 and FP 68263

in 1932 (FPF) ; Bear Wallow Campground, H & L

166 in 1962 (FPF). YAVAPAI CO.: Spruce Altn.,

Hedgcock FP 4856 and FP 4871 in 1910 (FPF);

0.5 mi. N of Spruce Mtn., H & L 239 in 1962 (FPF)!

CALIFORNIA

PLUMAS CO.: Providence Hill, ca. 4 mi. N of

Twaiai, T. 26 N., R. 8 E., Sec. 35, Henson in 1970

(FPF). SHASTA CO.: Near Pondosa Jet., Kidjt

1367 in 1957 (UC) ; 2 mi. S W of Pondosa Jet., H & W

645 in 1964 (FPF). SISKIYOU CO.: Sisson, Hedg^

cock. Meinecke de Long FP 9669 in 1911 (FPF); Dry

Lake Lookout, Oak Knoll [4 mi. SE of Cowdry Mtn.],

Gill & Sargent FP 68179 in 1932 (FPF); N side

Cascade Gulch, :\It. Shasta, Cooke 17729 in 1947

(WTU) ; Upton, Mt. Shasta, Hall & Babcock 4078 in

1903 (UC) and Cooke 13920 in? (DS); Joe Cr., 7 mi.

S of Copper, Oregon, H & W 634 in 1964 (FPF);

West base of Black Butte, 5 mi. S of Weed on US 99,

H & W 639 in 1964 (FPF,; Siskiyou Mtns., W

fork Cottonwood Cr. [SW of Hilts], Wheeler 2783

in 1934 (MO, PO.AI, US, Z); rd. to Gumboot Lake

[ca. 12 mi. W of Dunsmuir], Smith & Bacigalupi in

1957 (UC).

COLORADO

ARCHULETA CO.: Pagosa Springs, Hedgcock &

Bethel FP 24654 in 1917 (FPF); 3 mi. N of Piedra,

Hinds 63-6 in 1963 (FPF). CHAFFEE CO.: 6.8 mi.

S of US 285 on Bassam Park rd., H 446 in 1963

(FPF). CONEJOS CO.: 2 mi. SW of Conejos River

on Rte. 17, L 63-2 in 1963 (FPF) and Hinds 63-2 in

1963 (FPF). CUSTER CO.: 3.8 mi. NW of Isabel

Lake on Rte. 165, H 429 in 1963 (FPF) and on

Picea engelmannii, H 430 in 1963 (FPF) ; 4.7 mi. W

of Hillside, H 434 in 1963 (FPF). DOUGLAS CO.:

b.o mi. W of Rte. 105 on Jackson Cr. rd., H 206 in

1962 (FPF). EAGLE CO.: S mi. SW of Gypsum on

Cottonwood Pass rd., H 946 in 1966 (FPF). EL

PASO CO.: Palmer Lake, Bethel in 1912 (US) and in

1919 (CS); Clifton Park, Hill FP 68280 in 1932

(FPF); Palmer Lake, Hedgcock FP 15900 in 1914

(FPF, RSA, WTU); W of Palmer Lake, Hedgcock,

Thompson & dayman FP 24901 in 1917 (FPF); 5

mi. W of Monument, W 2984 in 1962 (COLO, FPF) ;

1.7 mi. above Cascade on Pikes Peak rd., H óc

Hinds 425 in 1963 (FPF); 0.3 mi. E of Bruin Inn,

N Cheyenne Can., H 538* in 1963 (FPF); Air

Force Academy, 0.5 mi. S of Cadet Area, // 54I

in 1963 (FPF); Crystal Park, Gill FP 68I4I in 1932

(FPF); Mt. .Alanitou, Gill FP 68256 in 1932 (FPF);

Halfway, Pikes Peak, Hedgcock FP 905 in 1909

(FPF); Palmer Lake Reservoir, Gill FP 68139 in

1932 (FPF). FREMONT CO.: Sangre de Cristo

Mtns., Hayden Pass rd. ca. 5 mi. SW of Coaldale,

206
Schacht in 1969 (FPF). GARFIELD CO.: 0.5 mi. S

of Douglas Pass, H 686 in 1964 (FPF) ; 14 mi. NNE

of Rifle, Hinds in 1965 (FPF). GUNNISON CO.:

Elk Mtns., Coal Cr. Can., Brandegee in 1881 (MO,

PH, UC) ; near Marble, W 3618 in 1964 (FPF, UT).

HINSDALE CO.: Little Sand Cr., 21 air mi. NW

of Pagosa Springs, Landgraf in 1965 FPF). HUÉRFANO CO.: Williams Cr., ca. 10 mi. N of Gardner,

H 432 in 1963 (FPF); N of Black Mtn., ca. 10 mi.

NNW of Gardner, Stewart in 1964 (FPF). LA

PLATA CO. : Hills N of Durango, Hunt & Bethel FP

2U29 in 1917 (FPF); W of Trimble, Hedgcock &

Bethel FP 24639 and FP 24689 in 1917 (FPF) ; Dry

Fork Cr., 8 air mi. NW of Durango, Landgraf in

1965 (FPF). LAS ANIMAS CO.: Near Stonewall

Gap, Hedgcock & Johnston FP 24945 in 1917 (FPF).

MINERAL CO. : 4.2 mi. N of Archuleta co. line on

US 160, Hinds 63-4 in 1963 (FPF). MOFFAT CO.:

Dinosaur Nat. Mon., Can. Overlook, W 4129 in 1967

(FPF, UT). MONTEZUMA CO.: W Mancos Can.,

Baker, Earle & Tracy 388 in 1898 (ARIZ, MO, RM,

POM, US); Priest Gulch, 8 mi. E of Stoner, Landgrafin 1965 (FPF) ; 14 mi. NE of Dolores on Rico rd.,

Hinds 63-12 in 1963 (FPF); Aspen Guard Sta.,

10 mi. NNE of Mancos, Hinds in 1965 (FPF);

MONTROSE CO.: Uncompahgre Plateau, Windy

Point, Peterson 51-61 in 1961 (FPF); 5.5 mi. N of

Rte. 90 on Transfer rd., Hinds 63-15 in 1963 (FPF).

OURAY CO.: Ouray, Cayman FP 26521 in 1917

(FPF) and Hedgcock FP 15915 in 1914 (FPF).

PUEBLO CO.: Beulah, Edmonston FP 295 in 1909

(FPF). SAGUACHE CO.: 5 mi. E of N Cochetopa

Pass, H 440 in 1963 (FPF) ; 2 mi. S of Poncha Pass

on US 285, H 44^ in 1963 (FPF); 4 mi. SW of Rte.

114 on Dry Gulch rd., H & L 442 in 1963 (FPF);

near Cochetopa Pass, on Picea engelmannii, L &

Schacht 63-26 in 1963 (FPF). SAN MIGUEL CO.:

7.9 mi. W of Dallas Divide on Rte. 62, Hinds in

1965 (FPF); 2 mi. W of Placerville, Payson FP

26026 in 1917 (FPF). TELLER CO.: Hotel Gulch,

1.5 mi. E of Manitou Exp. For. headquarters, H 260

in 1962 (FPF).

IDAHO

ADAMS CO.: Evergreen, Hedgcock FP 988 in 1909

(FPF) ; 13 mi. N of Council on US 95, H 977 in 1966

(FPF). BLAINE CO.: Bald Mtn. near Ketchum,

Mielke in 1957 (FPF). BONNER CO.: Hope,

Sandberg & Lieherg 4443 in 1893 (MO, POM, RM,

US); Lake Pend Oreille, Sandherg, MacDougal cfc

Heller 744 in 1892 (MO, US, Z); Priest River Weir

9995 in 1917 (ILL); Smiths Peak, Priest Lake, on

Ahies lasiocarpa, Weir 2419 in 1913 (ILL). BOUNDARY CO.: Pack River, on Ahies lasiocarpa, Weir

2415 n.d. (ILL). CAMAS CO.: Boise River, 1 mi. E

of Elmore Co. bdy., Peterson 62-50 in 1962 (FPF).

CLARK CO.: Spencer, Hurtt FP 20583 in 1912

(FPF). CUSTER CO.: ChalUs, CAll FP 68206''

in 1932 (FPF); 8 mi. W of Challis, Benloto in 1917

(ILL); Garden Cr. near Challis, on Abies lasiocarpa,

Gill FP 68207 in 1932 (FPF). FRANKLIN CO.:

Cub River near Franklin Basin, Peterson 63-364

in 1963 (FPF). IDAHO CO.: 5 mi. S of Grangeville,

Krebill 159 in 1964 (FPF), and H 968 in 1966 (FPF) ;

15 mi. N of Burgdorf on Salmon River rd., H 971 m

1966 (FPF); Salmon River at Fern Cr., ca. 23 mi.

W of Shoup, W 4í27a in 1966 (FPF, UT).

KOOTENAI CO.: Lakeside, col.? in 1889 (UC);

Canfield Mtn. [4 mi. NW of Coeur d^ Alene], Weir

2412 in 1916 (ILL). LATAH CO.: 3 mi. E of Viola,

Wicker in 1959 (WSP). LEMHI CO.: SW of Salmon

on N fork of Williams Cr., Krebill 126 in 1964

(FPF) ; 4 mi. N of Gibbonsville on US 93, H óc Hinds

1123 in 1968 (FPF) ; Wagonheimer, 3 mi. SE of North

Fork, Hedgcock FP 4457 in 1910 (FPF) ; 2.5 mi. E of

Tendoy, H & Hinds 1122 in 1968 (FPF). LEWIS

CO. : 3 mi. N of Winchester on US 95, H 967 in 1966

(FPF). MADISON CO.: Hawley Ranger Sta.,

Peterson 62-16 in 1962 (FPF). SHOSHONE CO.:

]Marble Mtn., on Ahies lasiocarpa, Weir 24I8 in 1914

(ILL). TETON CO.: Near Driggs, Peterson 62-11

in 1962 (FPF). VALLEY CO.: W of Cascade,

Thompson 13844 in 1937 (MO, UC, WTU); McCall,

Weir in 1914 (ILL) ; 3 mi. NE of McCall, H 975 in

1966 (FPF). WASHINGTON CO.: Weiser, Oinan

in 1912 (RM). COUNTY UNCERTAIN: St. Joe

Nat. For., on Abies lasiocarpa. Weir 24I6 n.d. (ILL).

MONTANA

FLATHEAD CO.: FLATHEAD LAKE: Yellow

Bay, Clemens in 1908 (DS) ; Mainland opposite Bull

Island, Cle7nens in 1908 (MO, PH); Bull Island,

Jones 9307 in 1908 (POM, US); Bigfork, Jones 9380

in 1908 (MO, POM). LINCOLN CO.: Sylvanite

[18 mi. N of Troy], on Picea engelmannii, Weir 24OO

in 1916 (FPF, ILL). MISSOULA CO.: Norman

Valley, Barkley & Barkley 3504 in 1938 (ARIZ,

COLO, MO, POM, UC, US); Pattee Cr., near

Missoula, Weir FP 91207 in 1918 (FPF, OSC);

^Taddy'^ Can., Barkley 1999 in 1937 (POM);

Pattee Can., Spaulding FP 15107 in 1913 (FPF) and

Hedgcock FP 4556 in 1910 (FPF); Missoula, Weir

9986 in 1915 (ILL) and 7426 in 1917 (ILL).

RAVALLI CO.: Blogett Cr. [W of Hamilton],

Hedgcock FP 9519 in 1911 (FPF); Spoon Cr. near

Darby, Hughes in 1917 (FPF, ILL); 10 mi. E of

Hamilton on Skalkaho Pass rd., H & Wicker 953 in

1966 (FPF).

NEVADA

WHITE PINE CO.: NE slope of Wheeler Peak,

Peterson63-337in 1963 (FPF).

207
NEW MEXICO

BERXALILLO CO.: Sandia Mtns.: Hedgcock FP

172, FP 185, FP 1538 and FP 1539 in 1908 (FPF);

Long & Seay FP 21275 in 1916 (FPF): La Luz,

Juan Tabo Trail, H 15 in 1954 (FPF); 2 mi. E of La

^Madera Ski Area, on Ahies concolor, H 56 in 1956

(FPF). CATROX CO.: 0.5 mi. W of .Alogollon,

H&L131 in 1962 (FPF) ; 6 mi. S of Luna on US 260,

H & L 213 in 1962 (FPF); Alangas Altn., H &

Scharpf697 in 1964 (FPF): Tularosa Summit, 11.5

mi. N of Rte. 78 on Apache Springs rd., L 65-43 in

1965 (FPF) ; 7 mi. E of Alogollon, on Ahies lasiocarpa

var. arizonica, Hedgcock FP 9899 in 1911 (FPF) and

H&L216 in 1962 (FPF); 2 mi. E of Bursum Campground on Rte. 78, on Ahies lasiocarpa var. arizonica,

H & L 218 in 1962 (FPF); 1.5 mi. E of Bursum

Campground on Rte. 78, on Ahies concolor, H & L

217 in 1962 (FPF) ; 2 mi. W of Rte. 78 on Bearwallow

Lookout rd., H & L 898 in 1966 (FPF). COLFAX

CO.: 8 mi. SE of Black Lake on Rte. 120, L 65-^8

in 1965 (FPF) ; 1 mi. E of Eagle Xest Lake, H FP

89958 in 1951 (FPF) and on Ahies concolor, H FP

89957 in 1951 (FPF); 9.5 mi. W of Cimarrón on US

64, L 65-21 in 1965 (FPF). GRAXT CO.: Emory

Pass, H&L120 in 1962 (FPF) ; 2 mi. X of Red stone,

H &L 127* in 1962 (FPF); 22 mi. X of Mimbres on

Rte. 61, L 65-39 in 1965 (FPF). LIXCOLX CO.:

Capitán Mtns. : 2 mi. E of Capitán Gap, H & L 106

in 1962 (FPF) and 1 mi. E, 1099 in 1968 (FPF);

West Capitán Altn. L 66-20 in 1966 (FPF); Seven

Cabins Can., Ellis, Hackleman & Gill FP 89491 in

1938 (FPF). E side of Gallinas Peak, 10 mi. W of

Corona, L & Riffle 63-43 in 1963 (FPF) smd H & L

1079 in 1967 (FPF). LOS ALAMOS CO.: 1 mi. X of

W gate to Los Alamos, L 65-31 in 1965 (FPF) ; 5 mi.

W of Los Alamos on Rte. 4, L cfc Gill 65-24 in 1965

(FPF); AIcKIXLEY CO.: Zuni Altns., Stinking

Springs rd., L 63-34 in 1963 (FPF); Chuska Altns.,

1 mi. S of Camp Ayassi [ca. 6 mi. S of Crystal],

L 64-1'' in 1964 (FPF). MORA CO.: 3 mi. XW of

Holman on Rte. 3, L 65-49 in 1965 (FPF). OTERO

CO.: Alescalero Indian Res.: Summit Can., H 26 in

1954 (FPF) ; Goat Can., H FP 89992 in 1952 (FPF) ;

Whitetail Well, H 23 in 1954 (FPF); 1 mi. E of

Apache Summit, H & Scharpf 694 in 1964 (FPF).

Silver Springs Can., 7.5 mi. XE of Cloudcroft, H óc

L 878 in 1966 (FPF). Curtis Can., 5 mi. W of State

Rte. 130, W of Alayhill, H & L 880 in 1966 (FPF);

10 mi. S of Cloudcroft on Sacramento Peak rd.,

H & L886 in 1966 (FPF). Sixteen Springs Can., 7

mi. E of Cloudcroft, H 1080 in 1967 (FPF). RIO

ARRIBA CO.: 12.5 mi. E of Canjilon on Rte. 110

L 63-7 in 1963 (FPF); 6 mi. W of Vallecitos, L63-8

in 1963 (FPF). SAXDOVAL CO.: 5 mi. SE of

Fenton Lake L 63-22 in 1963 (FPF); X end Sandia

Altns., Gill & Long FP 68136 in 1932 (FPF);

Placitas, Hedgcock FP 650 n.d., (FPF); 2 mi. X of

Sulphur Springs Jet. on Rte. 126, L cfc Gill 65-25 in

1965 (FPF). SAX JUAX CO.: 6.5 mi. SW of Sheep

Springs, L & Weiss 66-43 in 1966 (FPF) ; Tunitcha

[Chuska] Mtns., Standley 7664 in 1911 (US). SAXTA

FE CO. : Hyde State Park, L 63-39 in 1963 (FPF) ;

8 mi. E of Chimayo, L 63-24 in 1963 (FPF) ; 9 mi. E

of Santa Fe, Heller & Heller 3527 in 1897 (ARIZ, DS,

MO, US, Z). SAX AÍIGUEL CO.: 15 mi. E of

Pecos, L 63-23 in 1963 (FPF) ; 4 mi. S of Cowles on

Rte. 63, L 65-28 in 1965 (FPF). SOCORRO CO.:

Aiagdalena Altns., 3 mi. above campground in

Water Can., W 2994 in 1962 (COLO, FPF); San

Mateo Mtns.: Hughes Sawmill, Long FP 21126 in

1915 (FPF) and 11 mi. S of US 60 on Rte. 52, L

65-34 in 1965 (FPF). TAGS CO.: Peñasco, near

U.S. Hill, Long FP 19340 in 1914 (FPF); 1 mi. W of

Colfax CO. bdy. on US 64, L 65-23 in 1965 (FPF) ;

Ciénega Ranger Sta., Korstian FP 19276 in 1914

(FPF); 3 mi. E of Questa, L 65-46 in 1965 (FPF);

Santa Barbara Can., on Ahies concolor, Thome in

1963 (FPF); Taos Can., 7 mi. E of Taos on US 64,

H 876 in 1966 (FPF). TORRAXCE CO.: Manzano

Mtns., 3 mi. XW of Manzano, Riffle in 1963 (FPF).

VALEXCIA CO.: Mt. Taylor, Alarquez Can., L

63-31 in 1963 (FPF); Mt. Sedgwick, ca. 17 mi. W of

Grants, L 65-7 in 1965 (FPF). COUXTY UXCERTAIX: Front Cr. Basin, Gila Xat. For.,

Hedgcock & Long FP 9847 in 1911 (FPF).

OREGON

BAKER CO.: Lily White Guard Sta., 7 mi. XE of

Sparta, Steward & Jackman 6499 in 1953 (WTU) ; 22

mi. S of Baker on Rte. 7,H &W 584 in 1964 (FPF).

CLACKAMAS CO.: Clackamas River, T. 7 S, R.

8 E, Sec. 32, ca. 4 mi. W of the Cascade Crest, ca. 12

air mi. XE of Breitenbush, Nicholson in 1968 (FPF).

DESCHUTES CO.: 14 mi. XW of Sisters on Rte.

126, H & W 616 in 1964 (FPF). GRAXT CO.: 20

mi. X of John Day, Graham in 1964 (FPF); Prairie

City, Peebles Mill, Boyce FP 40121 in 1920 (FPF);

Dixie Butte, H 980 in 1966 (FPF) and on Ahies

lasiocarpa, H 982 in 1966 (FPF) ; 4 mi. S of Seneca

on Rte. 395, H & W 576 in 1964 (FPF) ; Skookum

Cr., 25 air mi. ESE of Seneca, H & Scharpf 1268

in 1969 (FPF). JACKSOX CO.: Rogue River Xat.

For., Yale Cr., [ca. 14 mi. SSW of Medford], Bedwell

& Childs'' in 1950 (FPF, OSC) and Childs in 1948

(OSC); 20 mi. SW of Ashland, Graham in 1963

(FPF); 10 mi. E of Butte Falls, Graham in 1964

(FPF) ; 11 mi. E of Ashland on Lake of the Woods rd.,

H & W 870 in 1966 (FPF); 20 mi. W of Ashland,

H áW636 in 1964 (FPF); Fish Lake rd., 12 mi. SE

of Butte Falls, Graham in 1965 (FPF) ; Huckleberry

Mtn., 9 mi. XE of Prospect, Graham in 1965 (FPF).

JEFFERSOX CO.: Suttle Lake, Gill FP 68192 in

1932 (FPF). JOSEPHIXE CO.: 16 mi. E of Cave

208
Jet. on Oregon Caves rd, H &W 868 in 1966 (FPF) ;

13 mi. E of O'Brien on Happy Camp rd., H & Hinds

997 in 1966 (FPF). KLAMATH CO.: Kirk, Pelican

Bay Camp, Boyce FP 40165 in 1921 (OSC) ; Pelican

Bay, Peck in 1920 (PH) ; 20 mi. W of Klamath

Falls on Rte. 66, H & W 628 in 1964 (FPF); between Lake of the Woods and Pelican Bay, Daniels

in 1914 (ILL). MORROW CO. : 35 mi. S of Heppner,

Steward à Steward 7329 in 1957 (RSA, US).

UNION CO. : 1 mi. NE of Kamela, H & W 587

in 1964 (FPF); 4 mi. W of Elgin on Rte. 204,

H & W 590 in 1964 (FPF). WALLOWA CO.: 6

air mi. SW of Jet. of Snake and Salmon Rivers,

Graham in 1964 (FPF) ; Billy Meadows [5 mi. E of

Kirkland Spring], Colville 2478a in 1907 (PH);

Kirkland area, 30 mi. N of Enterprise, Graham in

1965 (FPF). WASCO CO.: Bear Springs For.

Campground, Childs & Hansbrough FP 68261 in

1931 (FPF) and Childs & Hansbrough {Childs 69)

in 1931 (OSC); 3 mi. E of Bear Springs, Childs 57

in 1948 (OSC) ; 4 mi. W of Bear Springs, H &W 610

in 1964 (FPF). WHEELER CO.: 1 mi. N of Nat.

For. bdy., on Derr Meadows rd., 12 mi. SSE of

Mitchell, i/ 984 in 1966 (FPF); 0.25 mi. N of

Ochoco Divide on US 26, on Abies concolor, H &

Scharpf 1264 in 1969 (FPF); 1 mi. N of Ochoco

Divide, 11 air mi. W of Mitchell, H 986 in 1966

(FPF); 26 mi. S of Hardman on Rte. 207, H &

Scharpf 1272 in 1969 (FPF). COUNTY UNCERTAIN: White Pine, Starker in 1916 (FPF,

ILL).

UTAH

CACHE CO.: Logan Can., near Jardine Juniper,

Krebill 453 in 1967 (FPF). DAGGETT CO.: Big

Springs outlet, 11 mi. SW of Manila on Rte. 44, H

567 in 1964 (FPF) ; Sheep Can. SW of Manila, W

4128 in 1966 (FPF, UT); Palisades Campground,

Peterson 64-217 in 1964 (FPF). DUCHESNE CO.:

25.5 mi. SW of Duchesne on Rte. 33, H 570 in 1964

(FPF). GARFIELD CO. : S of Grover, near Wildcat

Ranger Sta., Peterson 64-92 in 1964 (FPF); 14 mi.

SE of Panguitch, Peterson 63-87 in 1963 (FPF);

Bryce Nat. Park, 3 mi. N of Far View Point, Buchanan 310 in 1957 (RSA); 23 mi. N of Boulder on

Rte. 117, H 685 in 1964 (FPF). IRON CO. : 14 mi. N

of Panguitch Lake, Peterson 62-67 in 1962 (FPF) ;

12 mi. E of Cedar City on Rte. 14, H 678 in 1964

(FPF); 13 mi. N of Panguitch Lake, on Abies

lasiocarpa, Peterson 62-28 in 1962 (FPF). KANE

Co.: Without locality, Siler in 1878 (MO); 10.4 mi.

W of Rte. 89 on Rte. 14, H 681 in 1964 (FPF);

Panguitch Lake Jet. on Rte. 14, W 4121 in 1967

(FPF, UT); 2 mi. above old Seegmiller Ranch on

lower Kanab Cr., E of Alton, W 4120 in 1967 (FPF,

UT) ; Cougar Hollow, S end Bryce Can. Nat. Park,

on Picea pungens, Peterson 62-111 in 1962 (FPF);

MILLARD CO.: Willow Cr. rd., W of Salina,

Peterson 64-63"" in 1964 (FPF) on Abies concolor,

Peterson 64-64 in 1964 (FPF). PIUTE CO. : 2 mi. W

of Nat. For. bdy. on Marysvale—^It. Belknap

rd., W 3036 in 1962 (COLO) ; Puffer Lake rd., W of

Jet., Peterson 64-83 in 1964 (FPF); Marysvale,

Tote Mine, Jones 5911c in 1894 (POAI, US);

Marysvale, Jones 5395 in 1894 (:\I0, POM US).

SALT LAKE CO.: Big Cottonwood Can., Hartley

FP 17865 in 1915 (FPF, ILL). SAN JUAN CO.:

Abajo :\Itns., near Spring Cr., Rydberg & Garrett 9864

in 1911 (RM, US); Elk Ridge, SW of Kigalia Guard

Sta., Peterson 65-76 in 1965 (FPF); N slope Abajo

Mtns., Krebill 288 in 1964 (FPF). SANPETE CO.:

Ephraim Can., SE of Ephraim, Hedgcock FP 8126

in 1912 (FPF); Great Basin Exp. Sta., Gill FP

68262 in 1932 (FPF) ; 8.7 mi. E of Ephraim on Great

Basin Exp. Sta. rd., H 574 in 1964 (FPF). SEVIER

CO.: 4 mi. NW of Joseph, Harris in 1914 (ILL);

S end of Fish Lake, H & Laut 1161 in 1969 (FPF).

SUMMIT CO.: Kamas, Hedgcock FP 93222 in 1911

(FPF). UTAH CO.: 5 mi. SW of Rte. 50 on Skyline

rd., H 572 in 1964 (FPF); Spring Lake, Parry in

1875 (MO) and on Abies concolor, Parry 86 in 1875

(MO, US). TOOELE CO.: Stansbury Mtns., end of

rd. in south Willow Can., W 3865B in 1965 (FPF,

UT). COUNTY UNCERTAIN: Ephraim Plateau,

Harris C2770B in 1927 (PH); central Utah, on

Abies concolor. Parry 10659 in 1875 (ILL, MO).

WASHINGTON

CHELAN CO. : 8 mi. NW of Leavenworth on US 2,

H, Stewart & Thompson 1136 in 1968 (FPF); Fish

Lake, Hanzlik FP 68257 in 1932 (FPF); Tip Top

Mtn., 13 mi. W of Wenatchee, Col.? in 1915 (ILL);

Chelan Lake, Weir 6162 in 1916 (ILL); Stehekin

River at north end of Chelan Lake, Graham in 1964

(FPF); 8.5 mi. E of Stevens Pass on US 2, H,

Stewart & Thompson 1137 in 1968 (FPF). CLACKA-

]MAS CO.: Clackamas River, T. 7 S., R. 8 E., Sec.

32, ca. 12 air mi. NE of Breitenbush, Nicholson in

1968 (FPF). C0LU:MBIA CO.: Wolf Fork of

Touchet River, St, John 6936 in 1925 (UC). FERRY

CO.: Sherman Cr. near Growden, Wicker in 1964

(WSP); Republic, Foster in 1912 (MO). KLICKITAT CO. : 2 mi. NW of Satus Pass, H & W 593 in

1964 (FPF). OKANOGAN CO.: Okanogan, Weir't

in 1914 (ILL); 18 air mi. E of Tonasket, Graham in

1965 (FPF) ; 5 mi. SW of Methow, Graham in 1964

(FPF) ; 9 mi. W of Republic on Rte. 30, H, Stewart &

Thoinpson 1132 in 1968 (FPF); 12 mi. W of Okanogan on Rte. 20 H, Stewart & Thompson 1134 in 1968

(FPF). YAKIMA CO.: 23 mi. E of White Pass on

Rte. 14 [2 mi. W of Nat. For. bdy.], H & W 596 in

1964 (FPF); Dog Lake, Tieton Can., Wicker in 1966

(FPF).

209
WYOMING

LINCOLN CO. : Wolf Cr. Campground on Snake

River, Targhee Nat. For., Peterson 62-6If in 1962

(FPF, RM). TETON CO. : 4 mi. E of Idaho bdy. on

Teton Pass rd., Peterson 62-12 in 1962 (FPF, RAI).

12a. ARCEUTHOBIUM GILLII subsp. GILLIl

Specimens Cited: on Pinus leiophijlla var. cliihuahuana unless as noted.

MEXICO

CHIHUAHUA: Continental Divide on Mexico NW

Railroad on ridge between Rio Chico and Rio

Caballo, on P. leiophylla, Barlow in 1911 (FPF); 1

km, S of Majalca, Andresen & Andresen 1804 in

1961 (FPF); ]Majalca, Sueur 599 in 1936 (UC); Mt.

Mohinora, Nelson 1^887 in 1898 (F, US) ; 65 mi. E of

Batopilas, Goldman 184 in 1898 (F, US); 25 mi. SW

of La Junta, H & W 292"" in 1963 (FPF) ; 34 mi.

SW of La Junta, H & W 294 in 1963 (FPF) and on

P. arizomca, H & W 293 in 1963 (FPF); 36 mi.

SW of La Junta, H & W 297 in 1963 (FPF) ; 14 mi.

E of El Vergel, H &W 307'' in 1963 (FPF); 26 mi.

SW of Col. Juárez, H & W 460* in 1963 (FPF);

12 mi. SE of Mesa Huracán on Chico rd., // & W

474 in 1963 (FPF) ; 18 mi. S of Las Varas on ^ladera

rd.,H &W 481 in 1963 (FPF) ; 48 mi. SW of Matachic on Ocampo rd., on P. lumholtzii, H & W 487

in 1963 (FPF) ; 43 mi. SW of Matachic on Ocampo

rd., H &W 484 in 1963 (FPF) and on P. ponderosa,

H & W 483 in 1963 (FPF); 29 mi. SW of Matachic

on Ocampo rd., H & W 497 in 1963 (FPF); 6 mi.

W of Matachic on Ocampo rd., H & W 499 in 1963

(FPF). SINALOA: Sierra Surutato, Municipio do

Badiraguato, on Pinus lumholtzii, Breedlove ct*

Thome 18617 in 1970 (CAS). SONORA: 'Trosty

Saddle,'^ Hartman 364 in 1890 (BM, F, ILL, K, Ny,

PH, UC, US).

UNITED STATES

ARIZONA

COCHISE CO.: Chiricahua Mtns.: Pedestal Pass,

Blumer 1515 in 1906 (ARIZ, ILL, K, :M0, R?^I,

US, Z); Pine Can., Blumer 2394 in 1907 (ARIZ,

DS, RM, Z); 6 mi. from Faraway Ranch, Portal,

Gill FP 68036 in 1936 (FPF) ; Portal, near Southwestern Research Sta., H & L 157 in 1962 (FPF);

Rustler Park, Ellis & Gill FP 89500 in 1939 (FPF) ;

Parker Can. Long FP 318 in 1909 (FPF) ; Pinery

Can., H & L 153"" in 1962 (FPF); Pinery Can. at

Jet. of N and S forks. Stone 201 in 1919 (PH).

Huachuca Mtns.: Reef, Gill & Ellis FP 89496 in

1939 (FPF). PniA CO.: Santa Catalina Mtns.:

General Hitchcock Campground, H & L 164"^ in

1962 (FPF) and H & Scharpf 705 in 1964 (FPF) ;

Oracle rd., Peterson 62-148 in 1962 (FPF) and H&L

1111 in 1968 (FPF); ]Mt. Lemmon, Peebles, Harrison

& Peebles 2565 in 1926 (ARIZ, US) : Mud Springs,

Harris C16354B in 1916 (US). Rincón :Mtns.,

Blumer 3362 in 1909 (ARIZ, :M0, UC). SANTA

CRUZ CO.: W side Huachuca .Mtns., T. 24 S, R.

18 E,H & L 161'' in 1962 (FPF) ; Santa Rita ^Itns.,

SE side of Mt. Wrightson, 1 mi. up trail from end

of Tunnel Springs rd., L 65-26 in 1965 (FPF).

COUNTY UNCERTAIN: Santa Rita Mtns.,

Pringle 13876 in 1881 (FPF).

12b. ARCEUTHOBIUM GILLII

subsp. NIGRUM

Specimens Examined:

MEXICO

DURANGO: 30 mi. E of El Salto on Rte. 40, on

P. leiophylla var. chihuahuana, H d' W 337 and 519

in 1963 (COLO, FPF), on P. lumholtzii, H ci* W 336

in 1963 (COLO, FPF), and on P. teocote, H & W 520

in 1963 (COLO, FPF) ; 32 mi. E of El Salto on Rte.

40, on P. teocote, H & W 334^ in 1963 (COLO, FPF) :

5 mi. E of El Salto on Rto. 40, on P. leiophylla, H &

W 1244 in 1969 (FPF) ; 2 mi. E of El Salto on Rto. 40,

on P. leiophylla, H & W 343'' and 525 in 1963

(COLO, FPF) ; 33 mi. S of Durango on La Flor rd.,

on P. leiophylla var. chihuahuana, H ci* W 510 in 1963

(COLO, FPF), on P. lumholtzii, H & W 509 in 1963

(COLO, FPF), and on P. teocote, H á: W 508* in

1963 (COLO, FPF); 15 mi. W of Santiago

Papasquiaro, on P. leiophylla var. chiJiuaJiuana,

H ÓC W 533 in 1963 (COLO, FPF) and on P.

lumholtzii, H (fe W 534 in 1963 (COLO, FPF) ; 18 mi.

W of Santiago Papasquiaro, on P. leiophylla, H óc W

535 in 1963 (COLO, FPF). HIDALGO: 20 mi. N of

Zimapan on Rte. 85, on P. teocote, H & W 385* in

1963 (COLO, FPF) and on P. montezumae, H & W

386 in 1963 (COLO, FPF). OAXACA: Dist. de

Teotitlán, Camino de Huaulla, on Pinus sp., Conzatti

4123 in 1921 (US). VERACRUZ: Cofre de Perote,

3 mi. SE of Perote, on P. teocote, H & W 382* in

1963 (COLO, FPF) and on Pinus sp., probably P.

teocote, W 2574 in 1959 (FPF, RSA).

13. ARCEUTHOBIUM GLOBOSUM

Specimens Examined :

MEXICO

CHIAPAS: On Pinus sp., Seler 2190 in 1896 (US);

near Siltepec, on Piîius sp., Matuda 4408 in 1941

(MO, NY); Mt. ]Male, near Porvenir, on Pinus sp..

210
Matuda 4630 in 1941 (F, NY) and on ''oak tree''

(probably in error; no host material with collection),

Matuda 4601 in 1941 (F, MO, NY, UC, US); between Cristobal de las Casas and Buenavista on rd.

to Tenejapa, on Pinus sp., hangman 3712 in 1948

(EAP, PH, US); Trail to Pokolum, near Tenejapa

Center, on Pinus sp., Breedlove 6974 in 1964 (DS,

FPF) ; on old rd. to Comitán from Mitziton, Municip. Chamula, on Pinus sp., Breedlove 8016 in 1964

(F) ; E of San Cristobal de las Casas on Rte. 90, on

P. pseudostrohus, Andresen & Steinhoff A2008 in

1962 (FPF); Valle cut SE of Las Casas, on Pinus

sp., Alexander 1199 in 1945 (NY). CHIHUAHUA:

Mesa Colorado, on Pinus sp.. Gentry 587M in 1933

(MICH); 19 mi. W of Mesa Huracán, on P. arizonica, H & W 466 in 1963 (COLO, FPF), and on

P. engelmannii, H &W ^^7* in 1963 (COLO, FPF).

DISTRICT FEDERAL: Desierto de los Leones,

on Pinus sp.. Roldan in 1919 (FPF, ILL). DURANGO: 20 mi. W of El Salto on Rte. 40, on Pinus

sp., McVaugh 11518 in 1951 (MICH); N slopes of

Cerro Huehueto, on Pinus sp., Maysilles 7260 in

1950 (MICH); San Luis del Río, 51 mi. NW of

Coyotes, on Pinus sp., Maysilles 7977 in 1955

(NY) ; 1.5 mi. E of El Salto on Rte. 40, on P. cooperi,

H &W 344* and 524 in 1963 (COLO, FPF) ; 13 mi.

W of El Salto on Rte. 40, on P. cooperi, H & W

349 in 1963 (COLO, FPT); 24 mi. W of El Salto

on Rte. 40, on P. durangensis, H & W 351 in 1963

(COLO, FPF) ; 33 mi. W of El Salto on Rte. 40, on

P. durangensis, H &W 526* in 1963 (FPF); 35 mi.

W of El Salto on Rte. 40, on P. durangensis, H &W

1246 in 1969 (FPF). JALISCO: Mamatlán, 15 mi.

SSE of Autlán by way of Chante, on Pinus sp.,

Wilhur & Wilbur 1837 and 1982 in 1949 (MICH).

MEXICO: Ojos de Agua, Nevado de Toluca, on

Pinus sp., Balls B4089 in 1938 (BM, K, UC, US);

40 km W of Toluca, on Pinus sp., Frye & Frye

2590 in 1939 (ARIZ, DS, MO, NY, RSA, UC,

WTU); Ixtacchiuatl, on Pinus sp., Purpus 1778 in

1905 (F, ILL, MO, NY, UC, US) : Amalacaxo, on

Pinus sp., Miranda 872 in 1940 (MEXU); Crucero

Agua Blanco, Temascaltepec, on Pinus sp., Hinton

8901 in 1936 (ARIZ, K, US) ; Valley of Mexico, on

Pinus sp., Borglav 1115 in 1866 (K, US); Nevado

de Toluca, on Pinus sp., N slope, (4,120 m.), Iltis,

Iltis & Flores 1701 in 1963 (WIS), on P. hartwegii,

Andresen & Steinhoff A2035 in 1962 (FPF) and on

P. rudis, H &W 369* in 1963 (COLO, FPF); 6 mi.

E of Michoacán bdy. on Rte. 15, on P. michoacana,

H & W 368 in 1963 (COLO, FPF) : 32 mi. W of

Toluca on Rte. 15, on P. montezumae, H 49 in 1956

(FPF); 3 mi. W of Rio Frío on Rte. 190, on P.

montezumae, H & W 371* m 1963 (COLO, FPF):

1.5 mi. E of Michoacán bdy. on Rte. 15, on P.

pseudostrohus, H & W 367 in 1963 (COLO, FPF).

MICHOACÁN: Cerro Tancitaro, on Pinus sp..

Leavenworth 710 in 1940 (F, MO); and on P. rudis,

Leavenicorth & Hoogstraal 1126 in 1941 (F, MO);

18 mi. S of Patzcuaro, on Pinus sp.. King & SoderStrom 5174 in 1961 (MICH, NY, UC, US); NW of

Cuidad Hidalgo, on Pinus sp., McVaugh 9952 in

1949 (MICH) ; San Alejo: on P. laivsonii. Valdivia &

Soloranzo 648 in 1963 (FPF) ; on P. rudis. Valdivia &

Soloranzo 649 in 1963 (FPF); on P. michoacana.

Valdivia & Soloranzo 650 in 1963 (FPF); on P.

montezumae. Valdivia & Soloranzo 646 in 1963

(FPF); and on P. tenuifolia. Valdivia & Soloranzo

637 in 1963 (FPF). 26 mi. E of MoreUa on Rte. 15,

on P. pseudostrohus, H 51 in 1956 (FPF) said H & W

361* in 1963 (COLO, FPF) ; 29 mi. E of MoreHa, on

P. pseudostrohus. Valdivia & Soloranzo 634 in 1963

(FPF) and on P. tenuifolia. Valdivia & Soloranzo

633 in 1963 (FPF). OAXACA: Sierra de San Felipe,

on Pinus sp., Pringle 4727 in 1894 (BM, K, MEXU,

MO, NY, PH, US, Z); 21 mi. S of Tuxtepec on rd.

to Oaxaca de Juarez, on P. rudis, Andresen & Steinhoff A1983 in 1962 (FPF); 8 mi. S of Suchixtepec,

on P. michoacana, Peterson 68-105 in 1968 (FPF).

PUEBLA: Near Rio Frío, on Pinus sp., Kenoyer in

1938 (ARIZ). SONORA: 'Tine Ridge Pass,'' on

Pinus sp., Hartman 340 in 1890 (ILL, K, NY, UC,

US). VERACRUZ: Mt. Orizaba, on Pinus sp.. Balls

B4399 in 1938 (BM, K, UC, US); Cofre de Perote,

on P. hartwegii, Beaman 2214 in 1958 (RM, UC,

US).

GUATEMALA

DEPT. ALTA VERAPAZ: Cobán, on P. tenuifolia,

von Turckheim II1815 in 1907 (ILL, MO, NY, US).

DEPT. BAJA VERAPAZ: Purulha; on Pinus sp..

Cook 7 in 1906 (FPF, ILL, US). DEPT. CHEMALTENANGO: between Tecpan and Encuentros,

Aguilar in 1955 (EAP). DEPT. HUEHUETENANGO: Sierra Cuchumatanes: Cerra Canana,

between Nucapula and Canana, on Cupressus sp.

(no host with collection), Steyermark 49044 in 1942

(F); Chemal, on Pinus sp., Johnston 1777 in 1940

(EAP, F); Km. 36, Chemal, on Pinus sp., Standley

81637 in 1940 (EAP, F, NY), 81642 in 1940 (F),

and 81683 in 1940 (F); vicinity of Touquia, on P.

rudis, Steyermark 50130 in 1942 (MO, NY, US);

between Paquix and San Miguel, on P. pseudostrohus, Molina, Burgur & Walienta 16483 in 1966

(EAP, NY); 33 km. N of Huehuetenango near

Chemal, on P. rudis, H &W 1219 in 1969 (FPF);

10 km N of Santa Eulalia on road to Barillas, on

P. rudis, H &W 1223 in 1969 (FPF). DEPT. SAN

MARCOS: Volcan Tajumulco, on Pinus sp.,

Schmidt in 1934 (F) and Steyermark 36940 (''on

roots and branches'^ in 1940 (MICH). DEPT.

ZACAPA: Sierra de las Minas, on Pinus sp., Steyermark 29788 in 1939 (MICH) and 4^524 in 1942

(MICH, NY).

211
BRITISH HONDURAS UNITED STATES

EL CAYO DIST.: Mtn. Pine Ridge, Granite Carin

rd., 6 mi. E of Augustine, ca. 30 mi. W of Stann Cr.,

Hunt 286 in 1959 (US) [Host on sheet labeled as

Pinus sp., but Hunt (1962) gives it as P. caribaea];

Altn. Pine Ridge, Raspa rd., on P. caribaea, Etheridge

BFDS 26 in 1965 (DAVFP, FPF).

14. ARCEUTHOBIUM GUATEMALENSE

GUATEMALA

DEPT. HUEHUETENANGO: Sierra Cuchumatanes: 2 km. N of Santa Eulalia on Barillas rd., on

Pinus ayacahuite, H & W 1222 in 1969 (FPF); 54

km. N of Huehuetenango on Santa Eulalia rd., on

Pinus ayacahuite, H & W 1221* in 1969 (FPF).

DEPT. TOTONICAPAN: Near km. 170 on Pan

American Highway, 15 km. W of Nahualá, on Pinus

ayacahuite, H &W 1226* in 1969 (FPF).

15. ARCEUTHOBIUM HONDURENSE

HONDURAS

DEPT. FRANCISCO MORAZAN: Near Piedra

Herrada: Rio Yeguare drainage, Lat. 14°N, Long.

87°W., on Pinus oocarpa, Williams 15964* in 1969

(BM, EAP, F); 22 km. SE of Tegucigalpa on rd. to

Escuela Agrícola Panamericana, on P. oocarpa, H,

W & Molina 1203* in 1969 (FPF); 7 km. W of

Zamorano on Tegucigalpa rd., on P. oocarpa, H &W

1209 m 1969 (FPF).

16. ARCEUTHOBIUM LARICIS

Specimens Examined: On Larix occidentalis unless

noted otherwise.

CANADA

BRITISH COLUMBIA: Burton, Kuijt 566 in 1953

DAVFP, FPF); Balfour, Kuijt 563 in 1953

(DAVFP, FPF); Slocan Lake, Kuijt in 1953

(DAVFP, FPF); Christina Cr., Kuijt 553 in 1953

(DAVFP, FPF); Grand Forks, Kuijt 556 in 1953

(DAVFP, FPF); Salmo, Kuijt 51,7* in 1953

(DAVFP, FPF); Castlegar, Kuijt 5^6 in 1953

(DAVFP, FPF); Fruitvale, Kuijt in 1953 (DAVFP,

FPF) ; Crestón, Kuijt in 1953 (DAVFP, FPF) ; St.

Mary's Lake, Kuijt 5U in 1953 (DAVFP, FPF);

3 mi. N of Moyie on Cranbrook rd., Calder & Saville

11415 in 1953 (CAS, UC, WTU); 6 mi. W of Jet.

west of Rossland, Calder & Saville 11465 in 1953

(US); Owens Lake, Weir 8362 in 1913 (ILL);

Salmon Arm, Weir 8364 in 1913 (FPF, ILL).

IDAHO

ADAMS CO.: Evergreen, Hedgcock FP 997 in 1909

(FPF). BENEWAH CO.: St. Maries, Hedgcock FP

9526 in 1911 (FPF) and Weir 659 in 1911 (ILL).

BOISE CO.: Idaho City, Weir 8371 in 1917 (ILL).

BONNER CO.: Priest River, Weir 8373 in 1917

(FPF, RM, US); Sandpoint, Wehmeyer in 1922

(US); Priest River Exp. For. Headquarters, on

Picea abies, H & Wicker 963 in 1966 (FPF); on

Pinus banksiana, Graham in 1958 (FPF) and H &

Wicker 964 in 1966 (FPF), and on Pinus resinosa,

H & Wicker 965 in 1966 (FPF); 4 mi. N of Priest

River on rd. to For. Exp. Sta., H & Wicker 961* in

1966 (FPF) and on Pinus contorta, H & Wicker 962

in 1966 (FPF). Coolin, on Pinus contorta. Weir 8385

in 1917 (ILL) and on Abies lasiocarpa. Weir 8391

in 1916 (ILL); Blue Lake, on Abies grandis. Weir

8392 in 1917 (ILL); Priest River, on Abies grandis.

Weir 3252 in 1917 (ILL). BOUNDARY CO. : Above

Upper Priest Lake, Boyce 1225 in 1924 (FPF, OSC).

IDAHO CO. : 6 mi. S of Grangeville on Snow Haven

rd., H 969 in 1966 (FPF). KOOTENAI CO.:

Clark's Fork, Coeur d' Alene Mtns., Lieberg 1672

in 1895 (MO, POM, RM, UC, US); 4th of July

Summit, W 3255* in 1962 (COLO, FPF); Granite

Sta., Sandberg, MacDougal & Heller 787 in 1892

(CAS, US); Coeur d' Alene, on Pinus ponderosa.

Weir 3271 in 1916 (ILL). LATAH CO.: Moscow

Mtn., Piper 1882 in 1894 (MO, RM). SHOSHONE

CO. : Lick Cr. Lookout [ca. 6 mi. SSW of Avery],

Gill FP 68291 in 1932 (FPF) and on P. monticola,

Gill FP 68202 in 1932 (FPF); 3 mi. W of Montana

bdy. on US 10, W 3256 in 1962 (COLO, FPF);

Monumental Butte, St. Joe Nat. For., on Pinus

monticola. Weir in 1914 (FPF, ILL) ; i^ mi. W of

Thompson Pass, on Tsuga mertensiana, H & Wicker

957 in 1966 (FPF) and on Abies lasiocarpa, H &

Wicker 958 in 1966 (FPF); 1 mi. S of Jack Waite

Mine on Woodchuck Pass rd., on Tsuga mertensiana,

H & Wicker 959 in 1966 (FPF). VALLEY CO.:

McCall, col.? in 1915 (ILL); 3 mi. NE of McCall on

shore of Payette Lake, H 973 in 1966 (FPF) and on

Abies lasiocarpa, H 974 in 1966 (FPF).

UNITED STATES

MONTANA

FLATHEAD CO.: Somers, Flathead Lake, Weir

3258 in 1916 (ILL); Big Fork, Flathead Lake, Jones

9312 in 1908 (DS, POM) and 9309 in 1908 (POM);

Columbia Falls, Williams 897 in 1892 (US).

GRANITE CO.: PhilHpsburg, Weir 8374 in 1917

(ILL). LAKE CO.: Yellow Bay, Flathead Lake,

Jones 9313 in 1908 (POM) and Butler 182 in 1908

(FPF); 4 mi. N of Dunton, Krebill 136 in 1964

212
(FPF)- E of St. Ignatius near St. Mary's Lake,

Krehill 133 in 1964 (FPF). LINCOLN CO.: Cabinet

Mtns. near Scotchmans Peak, on Larix lyalli, Weir

3250 n.d. (FPF, ILL) and on Tsuga mertensiana,

Johns in 1914 (FPF, ILL). FUNERAL CO.: 6 mi.

W of St. Jlegis on US 10, H & Wicker 956'' in 1966

(FPF) and on Pinus contorta, H & Wicker 955 in

1966 (FPF). MISSOULA CO.: Missoula, Weir FP

17048 in 1914 (FPF); Blue Cr., 35 mi. NW of

Missoula, on Picea engelrnannii, Toko & Pinney

65-34 in 1965 (FPF) ; Pattee Can. near Missoula,

on Pinus contorta, Paulson in 1916 (FPF, ILL) and

Weir 3267 in 1916 (ILL); Bitterroot Mtns. near

Lolo, on Larix lyalli, Weir 8366 in 1914 (FPF, ILL) ;

Lolo, on Pinus ponderosa, Weir 8337 in 1916 (FPF,

ILL). RAVALLI CO. : One Horse Cr. near Florence,

col.? in 1915 (ILL); Victor: on Picea engelmannii.

Weir 7196 in 1917 (FPF); on Pinus alhicaulis. Weir

8334 in 1917 (ILL) ; on Pinus contorta. Weir 8336

and 8384 in 1917 (ILL); and on Ahies lasiocarpa.

Weir 8390 in 1917 (ILL). SANDERS CO.: Thompson Falls, Hedgcock & Weir FP 944^ in 1911 (FPF) ;

and on Pinus ponderosa, Hedgcock & Weir FP 9444

in 1911 (FPF) ; 8 mi. N of Plains, on Pinus ponderosa,

Sieminski in 1964 (FPF); 2 mi. E of Noxon on US

lOA, H & Wicker 960 in 1966 (FPF).

OREGON

BAKER CO. : 6 mi. E of Medical Springs, Peterson

63-236* in 1963 (FPF) ; 12 mi. S of Baker on Rte. 7,

H & W 585* in 1964 (FPF) ; Jim Cr. Fork of Eagle

Cr., 30 mi. NE of Baker, Graham in 1965 (FPF)

and, on Pinus alhicaulis, Graham in 1965 (FPF).

CROOK CO.: Ochoco, Can. Cr., Donnelly in 1920

(FPF). GRANT CO.: 16 mi. S of John Day on Rte.

395, H & W 579 in 1964 (FPF) and on Pinus

ponderosa, H & W 578 in 1964 (FPF); 6 mi. E of

Dixie Pass on US 26, H & W 582* in 1964 (FPF);

Elkhorn Spring rd. [ca. 16 mi. E of Prairie City],

Childs 212 in 1952 (OSC); Dixie Butte, Weir 2421

1915 (FPF, ILL) and on Ahies grandis (as A. ''concolor''), Weir 2425 in 1915 (FPF, ILL); 4 mi. N of

US 26 on Dixie Butte rd., on Ahies grandis, H 981

in 1966 (FPF) ; Big Canyon, 9 air mi. NE of Seneca,

H & Scharpf 1270 in 1969 (FPF). JEFFERSON

CO.: Metolius area. Gill FP 68189* in 1932 (FPF);

Metolius River near Camp Sherman, Graham in

1964 (FPF). MORROW CO.: Alder Cr., 35 mi. S of

Heppner, Steward óc Steward 7328 in 1957 (CAS,

DS, RSA, US, WTU). UMATILLA CO.: 13 mi. E

of Kamela, H & W 588 in 1964 (FPF). UNION

CO. : 12 mi. W of Elgin, H & W 589 in 1964 (FPF) ;

15 air mi. NNE of Elgin, Orr in 1968 (FPF).

WALLOWA CO.: Minam Meadows [12 mi. SW of

Enterprise], Sampson & Pearson 168 in 1907 (FPF,

MO): Billy Meadows [30 mi. NNE of Enterprise],

Coville 2459 in 1907 (US); Wallowa Mtns., Bear Cr.,

Coville 2363 in 1907 (US) ; Wallowa INItns., Bear Cr.

sheep trail, Coville 2480 in 1907 (US). WASCO

CO. : Bear Springs Campground, I\It. Hood, Childs &

Hanshrough FP 68268 in 1931 (FPF) and Childs 27

in 1940 (OSC) ; 5.7 mi. SE of Wapinita Pass on US

26, H & W 609 in 1964 (FPF). WHEELER CO.:

3 mi S of Nat. For. bdy. on Derr JNIeadows rd., ca.

12 mi. SSE of Mitchell, H 985 in 1966 (FPF); 26

mi. S of Hardman on Rte. 207, H & Scharpf 1273

in 1969 (FPF); Ochoco Summit on US 26, H &

Scharpf 1262 in 1969 (FPF).

WASHINGTON

CHELAN CO.: Chelan Lake, Weir 8363 in 1916

(ILL); Leavenworth, Bunder n.d. (ILL). FERRY

CO.: Growden, Wicker in 1963 (FPF); Fritz Cr.

near Growden, Wicker* in 1963 (FPF); 16 mi. E of

Republic on Rte. 30, H, Stewart & Thompson 1129

in 1968 (FPF) and on Ahies lasiocarpa, H, Stewart &

Thompson 1130 in 1968 (FPF). GARFIELD CO.:

7 air mi. SSW of Peola, Orr in 1968 (FPF). KITTITAS CO.: N of Ellensburg, Brandegee 1071 m

1883 (PH, US); Wenatchee Mtns., Coville 1881 in

1901 (US); Peoh Point, ca. 3 mi. S of Cle Elum,

Will in 1968 (FPF). OKANOGAN CO.: 9 mi. W of

RepubUc on Rte. 30, on Pinus contorta. H, Steivart &

Thompson 1133 in 1968 (FPF); Loup Cr., 12 mi. W

of Okanogan on Rte. 20, H, Stewart & Thompson

1134 in 1968 (FPF). PEND OREILLE CO.:

Metaline Falls, Weir 8365 in 1916 (FPF, ILL).

STEVENS CO. : 3 mi. W of Park Rapids, Krehill 14^

in 1964 (FPF). WHITMAN CO.: N side of Kamiak

Butte, St. John 6359 in 1924 (MO, UC). YAKBIA

CO. : Havens Ranch, SE of Mt. Adams, Henderson

2535 in 1892 (WTU) ; Tieton Basin, Tweedy in 1882

(DS); ^^Yakima Region,'' Brandegee in 1882 (UC);

2.5 mi. E of White Pass on Rte. 5, H & W 597 in

1964 (FPF) and on Pinus montícola, H & W 598 in

1964 (FPF); 6.5 mi. E of White Pass, on Rte. 5,

H & W 603 in 1964 (FPF); Tieton Cr., 2 mi. SW

of Tieton Lake, on Ahies grandis, H & W 600 in

1964 (FPF); 7 mi. S of Tieton, Graham in 1966

(FPF) ; 4 mi. E of Dog Lake, Wicker in 1966 (FPF) ;

1 mi. W of Rimrock, Steivart in 1968 (FPF).

COUNTY UNCERTAIN: Cascade jNItns., Tiveedy

in 1882 (MO).

17. ARCEUTHOBWM MICROCARPUM

Specimens Examined : Host is Picea engelmannii, except as noted.

UNITED STATES

ARIZONA

APACHE CO.: INIt. Baldy, Woolsey FP 12566 in

1914 (FPF) ; 20 mi. W of Eagar on Rte. 73, H & L

207* in 1962 (FPF); 4.7 mi. SW of Alpine on US

666, on Picea pungens, H & Scharpf 700* in 1964

213
(FPF) ; 2 mi. E of Big Lake, on Picea pungens, H 819

in 1965 (FPF); 1 mi. S of Horseshoe Ciénega, 10 mi.

E of McNary, on P. pungens, H & Gilbertson 1083

in 1967 (FPF); 6 mi. NE of Maverick, on Picea

pungens, H & L 900 in 1966 (FPF) ; 5 mi. SW of

Rte. 373 on Big Lake rd., 2 air mi. SW of Greer, on

P. pungens, H & L 925 in 1966 (FPF). COCONINO

CO.: ^^Kaibab Forest,'' Richards in 1938 (RM, UC,

WTU) ; San Francisco Peaks: Flagstaff Water Camp,'

H &L 249'' in 1962 (FPF) and Interior Basin, Long

FP 21671 in 1916 (FPF). Grand Can. Nat. Park,

North Rim: Gill & Andrews FP 87301 in 1935

(FPF) ; near entrance. Gill FP 68298.m 1934 (FPF) ;

2.0 mi. E of Rte. 64, on Point Imperial rd., H 253*

in 1962 (FPF); Telephone Can., W 3183 in 1962

(COLO); 2.4 mi. E of Rte. 64 on Point Royal rd.,

on Picea pungens, Peterson 43-61 in 1961 (FPF).

GRAHAM CO. : Graham Mtns. : Kearney & Peebles

9971 in 1934 (ARIZ, US) and Ellis FP 89^30 in

1939 (FPF); Hospital Flat, Gill FP 68310 in 1934

(FPF) and Kearney & Peebles 14135 in 1938 (ARIZ) ;

Columbine, Gill FP 68290 in 1934 (FPF), Maynard

in 1963 (FPF) and on Abies lasiocarpa var. arizonica,

Stouffer & Gill FP 68291 in 1934 (FPF); High Peak

rd., 3 mi. N of main rd., H & L 224* in 1962 (FPF) ;

Soldier Cr. Campground, H & L 225* in 1962

(FPF); Webb Peak, Darrow, Phillips & Pultz 1173

in 1944 (ARIZ). GREENLEE CO.: 1 mi. S of Blue

Summit on US 666, Gill FP 68302 and FP 68305 in

1934 (FPF); US 666 N of Jet. of Gobbler Point rd.,

Peterson 62-165 in 1962 (FPF) ; 3 mi. N of Hannagan

Meadows on US 666, Andrews in 1963 (FPF) and

L 64-20 in 1964 (FPF).; Hannagan Cr. N of Hannagan Meadows on US 666, on P. pungens, H 1082

in 1967 (FPF)

NEW MEXICO

CATRON CO. : Mogollón Mtns. : On or near W fork

of Gila River, Metcalje 493 in 1903 (ARIZ, FPF,

ILL, MO, POM, UC, US); Willow Cr. Campground,

on Picea pungens, H & L 219* in 1962 (FPF) ; 4 mi.

W of Rte. 78 on Bearwallow Lookout rd., H & L

897* in 1966 (FPF).

18. ARCEUTHOBIUM OCCIDENTALE

Specimens Examined : on Pinus sabiniana except as

noted.

UNITED STATES

CALIFORNIA

ALAMEDA CO. : Berkeley, on planted P. radiata,

Offord & Scharpf in 1964 (FPF). AMADOR CO.:

6 mi. E of Jackson, Gill FP 68255 in 1932 (FPF);

1 mi. S of Jackson, W 3224 in 1962 (COLO,'

FPF); near lone, Braunton in 1904 (MO); 1 mi. S of

lone, Kuijt 1208 in 1956 (UC); Pine Grove, Hanson

718 in 1895 (MO, US). BUTTE CO.: Big Chico Cr

E of Chico, Heller III44 in 1913 (DS, MO, PH,

US); Magalia, Howell 37436 in 1962 (CAS)- near

MagaUa, W 2728 in 1960 (RSA); Chico, Griffiths in

1912 (MO); Paradise, Wall in 1935 (CAS); Bangor

Rose in 1931 (CAS). CALAVERAS CO.: 1.5 mi. E

of Copperopolis on Rte. 4, Kuijt I4IO in 1957 (UC);

rd. from Vallecito to Columbia, Quick 53-141 in

1953 (CAS). COxNTRA COSTA CO.: Mt. DiabloBethel FP 26266 in 1918 (FPF, ILL); Scharpf* in

1963 (FPF); N slope. Hicks 110 in 1917 (UC);

Mitchell Can., Jepson in 1898 (JEPS, MO) and on

P. coulteri, Bacigalupi in 1922 (DS); Little Rock

City, Kuijt 1444 in 1958 (UC); Rock City, Kuijt

1425a in 1957 (UC); Ranger Headquarters near S

entrance, H & Scharpf 856 in 1966 (FPF) and on

P. coulteri, H & Scharpf 857 in 1966 (FPF). EL

DORADO CO.: Smith Flat, Benson 47 in 1924

(DS); Placerville, W 2335* in 1962 (COLO,

FPF) ; 2-3 mi. E of Placerville on Rte. 550, Bobbins

1403 in 1943 (UC); 1.7 mi. SW of Garden Valley,

Kuijt 1257 in 1957 (UC); 3 mi. NW of Placerville

on Rte. 49, H & W 84I in 1966 (FPF); S fork of

American River, N of Placerville, Kuijt 1274 in

1957 (UC). FRESNO CO.: Auberry, Stillinger in

1920 (FPF, ILL); 10 mi. SW of Auberry on Priant

rd., H 843 in 1966 (FPF); 5 mi. N of Auberry on

North Fork rd., H 844 in 1966 (FPF). GLENN

CO.: 2.1 mi. N of Stonyford, Kuijt 1506 in 1958

(UC). KERN CO.: Walker Basin, Rothrock 429 in

1875 (MO, US); Calienti, Gill FP 68089 in 1932

(FPF); Havilah, Coville & Funston 1073 in 1891

(DS, FPF, US); Greenhorn Pass, 4 mi. W of Kernville, Gould 1010 in 1940 (DS, MO, UC, US); 1 mi.

E of Isabella Res. on Rte. 178, Kuijt 1437 in 1957

(UC); head of Tejon Cr., Twisselmann 8832 in 1963

(CAS) ; between Glenville and Sierra Alta, 7 mi. W

of Summit Guard Sta., W 3605* in 1964 (FPF);

Kernville, Kerr in 1938 (CAS); Tehachipai Mtns.,'

Upper Cottonwood Cr., W of White Oak Lodge,

Tucker 3444 in 1938 (ARIZ); 2.5 mi. W of Wofford,

Howell 38710 in 1962 (CAS). LAKE CO.: without

locahty, Bolander 2671 in 1863 (UC, US); near Jet.

of Rtes. 53 and 20, SE of Clear Lake, Rossbach 236

in 1955 (UC); Siegler Springs, Blankinship in 1924

(CAS); Lakeside, Clear Lake, Abrams 6297 in 1916

(DS); Kelseyville, Blankinship in 1924 (RSA) and

Jussel in 1931 (CAS); Kelseyville, Rincón School,

Benson 726 in 1928 (POM); Mt. Konocti, Blankinship in 1923 (MO); 3 mi. N of Middletown on Rte.

29, Scharpf in 1965 (FPF). LASSEN CO.: 10 mi. S

of Pittville, Wagener FP 97951 in 1928 (FPB). LOS

ANGELES CO.: Liebre Mtns.: Elizabeth Lake

Can., Dudley cfe Lamb 44II in 1896 (DS); Collins

Ranch, Peirson 4302 in 1921 (RSA); Sandbergs,

Münz 4423 in 1921 (POM). MADERA CO.: Raymond, Hedgcock FP 4837 in 1910 (FPF, ILL); 9 mi.

W of Oakhurst, Kuijt 1254 in 1957 (UC); 18 mi. N

214
of Pinedale on Rte. 41, H IIU in 1968 (FPF) ; 1 mi.

W of North Fork, H 658'' in 1964 (FPF) and ii*

in 1964 (FPF) ; Hogue Ranch, 16 mi. from North

Fork, on Mammoth rd., H 847 in 1966 (FPF); 1.5

mi N of Oakhurst, Howell 41197 in 1965 (CAS);

Peckinpah Cr., Howell & Barnehij 29387 m 19o4

(CAS) :\ÍARIN CO.: Inverness Ridge, on P. muncata, Howell 19686 in 1944 (CAS, UC); ^It. Vision

rd Inverness, on P. niuricata, Schreiher 853 m 1933

(UC). :MARIP0SA CO.-. Bear Valley :Mtn., Ward

51 in 1895 (US); El Portal, Posey in 1919 (ILL);

3 mi W of Alt. BuUion, Kuijt 1253 in 1957 (UC);

near Bootjack, Howell 8135 in 1931 (CAS, US):

Mariposa, Congdon in 1903 (AIO, US). MENDOCINO CO.: Fort Bragg, on P. muricata, Mason 5639

in 1930 (UC) ; :Mouth of Gualala River, on P. muricata, Bacigalupi 1808 in 1921 (DS); Van Damme

State Park, on P. muricata, Kuijt 1214 m 19o7

(UC) ; Pine barrens near Fort Bragg, on P. muricata,

Johnson 1420 in 1929 (RSA) ; 3 mi. E of Fort Bragg,

on P. muricata, H & Scharpf 849 in 1966 (FPF) ;

Point Arena, on P. contorta subsp. holanderi, Mason

7168 in 1932 (UC); Van Damme State Park, on P.

contorta subsp. holanderi, Kuijt 1215 in 1957 (UC);

just east of airport near Albion, on P. contorta subsp.

holanderi, Peterson 65-116 in 1965 (FPF); White

Sands near Mendocino City, on P. muricata, Eastivood 18836 in 1912 (CAS); 4 mi. SE of Fort Bragg,

on P. contorta subsp. holanderi, H & Scharpf 854 m

1966 (FPF). MONTEREY CO.: Pebble Beach,

Carmel, on P. radiata, Boyce 33 in 1915 (FPF); between Pacific Grove and Carmel, on P. radiata. Gill

FP 68I45 in 1932 (FPF); Carniel, on P. radiata,

von Schrenk in 1920 (MO) ; Carmel highlands above

Yankee Point, on P. radiata. Balls 23608 in 1958

(WTU); Point Lobos State Park, on P. radiata,

Lee & Mason 9153 in 1935 (UC); Point Lobos, on

P. radiata, Kuijt 1213 in 1957 (UC); near Pomt

Pinos, on P. radiata, Dudley in 1893 (DS); Gibson

Cr., on P. radiata, Wheeler 44^2 in 1936 (POM).

Monterey: on P. radiata. Parry in 1850 (MO),

Brewer 700 in 1861 (US), Engelmann in 1880 (MO),

Meehan in 1883 (PH), Elmer 4031 in 1902 (POM),

Meinecke FP 9059 in 1910 (FPF, ILL), Bethel

FP 26279 in 1918 (FPF, ILL) and H & Scharpf 1151

in 1968 (FPF). Pacific Grove: on P. radiata, Davy

7064 in 1900 (UC), Heller 6776 in 1903 (COLO,

MO, RM, UC, US), Smith 955 in 1905 (MICH),

Condit in 1909 (UC), Beattie 5281 in 1916 (FPF),

Guppy FP 96959 in 1924 (FPB), Gill FP 68076,

FP 68093 and FP 68103 in 1932 (FPF), W 3220 in

1962 (COLO, FPF), and Peterson 63-156* in 1963

(FPF). Cypress Point: on P. radiata, Ahrams 7660

in 1920 (RM, US), and Mason 3987 in 1927 (UC).

Between Monterey and Carmel, on P. muricata.

Mason 5888 in 1931 (UC) ; Huckleberry Hill, Carmel,

on P. muricata, Mason 5515 in 1929 (UC) and

Wheeler 4428 in 1936 (PO^I); 7.2 mi. NE of Gonzales, W 3219 in 1962 (COLO, FPF). Santa Lucia

:Mtns.: San Antonio River, Brewer 582 in 1861

(US); Pine Can., Duncan 90 in 1920 (DS, PO^I);

:Miller Can., on P. coulteri, Ferris 12158 in 1929

(DS, R:M, RSA, WTU); :Miller Can., Chews Ridge,

on P. coulteri, Ray 1565 in 1949 (US); Tassajara rd.

at Chews Ridge Lookout Jet., on P. coulteri, Kuijt

1401 in 1957 (UC) and H & Scharpf 1152 in 1968

(FPF) ; ^larble Peak, 0.5 mi. NW of Anderson Peak,

on P. coulteri, Offord FP 98091 in 1963 (FPB);

Junípero Serra Peak, on P. coulteri, Howell 30162 m

1955 (CAS) ; near Cone Peak, on P. coulteri, Ferris

3649 in 1923 (DS, P0:M) and H & Scharpf 1158 m

1968 (FPF) ; :Monterey For. Reserve, on P. coidteri,

von Schrenk in 1920 (:M0) ; 5 mi. W of Jolon on

Nacimiento rd., H & Scharpf 1153 in 1968 (FPF).

NAPA CO.: Conn Valley, Jepson in 1895 (JEPS,

US) and Raven 3928 in 1951 (CAS); :Moore Cr.,

Howell :Mtn., col? in 1902 (UC); 3 mi. from Aetna

Springs on :Middletown rd., Howell 5618 m 1930

(C\S)' 14 mi. SE of :Middletown on Aetna Springs

rd., H & W 859 in 1966 (FPF). PLACER CO.:

Auburn, Loran in 1886 (UC) and Engelmann m

1880 (ILL, :\I0). SAN BENITO CO.: Pinnacles

Nat ^lon., W 3218 in 1962 (COLO, FPF) and Gill

FP 68078 in 1932 (FPF) ; 20 mi. S (E?) of Bitterwater on New Idria rd., Kuijt 1313 in 1957 (UC);

rd. to New Idria, 4.5 mi. S of Jet. N of Bitterwater,

Kuijt 1300 in 1957 (UC); near New Idria, on P.

coulteri, Quihell 1658a in 1953 (RSA) and Kuijt 1310

in 1957 (UC). SAN LUIS OBISPO CO.: Paso de

Robles, Summers 926 in 1886 (UC) ; Santa Margarita, Mason 525 in 1923 (UC) ; ?^Iariana Cr., E

side La Panza Range, Hoover 6310 in 1946 (CAS);

Cambria, on P. radiata, Gill FP 68079 and FP

68253"" in 1932 (FPF), Hoover 6448 in 1946 (CAS),

and H & Scharpf 1159 in 1968 (FPF). SANTA

BARBARA CO.: San Rafael :Mtns., Yellow Gate,

Meinecke FP 97947 (FPB) and FP 17026 (FPF,

ILL) in 1914; Pine Can., Roosevelt, Hunt FP 97958

in 1912 (FPB); Figueroa :Mtn., Pollard in 1950

(CAS) and 1956 (CAS) ; 8 mi. E of Los Olivos on

Figueroa Mtn. rd., H 935 in 1966 (FPF); 12 mi. E

of Los Olivos on Figueroa Mtn. rd., on Pinus coulteri,

H 936 in 1966 (FPF). SANTA CLARA CO.: ^It.

Hamilton Range: E slope of ^It. Hamilton, Beane

225 in 1949 (RAI); :\It. Hamilton, Chanser 6022 in

1906 (UC); Long Branch Cr., :^It. Day, Sharsmith

3848h in 1936 (DS, UC); E slope ^It. Hamilton,

Beane & James 2079 in 1949 (DS, RSA, WTU) ; 1 mi.

above Alum Rock, :Mt. Hamilton, Dutton in 1899

(DS) ; 2 mi. below summit of :Mt. Hamilton, Solbig

in 1957 (UC) ; 2 mi. E of :Mt. Hamilton on Livermore rd., H efe Scharpf 1149 in 1968 (FPF) and on

P. coidteri, H & Scharpf II48 in 1968 (FPF); near

Mt. Hamilton Observatory, on planted P. radiata,

215
H & ScharpflloO in 1968 (FPF); Summit Copernicus Peak, Sharsmith 1406a efe 6 in 1934 (UC). Mt.

ümunhum, Thomas 5519 in 1956 (DS, FPF, RSA);

Loma Prieta, on P, attenuata, Dudley in 1895 (DS)'

Stanford Arboretum, on planted P. radiata, Wight

47 in 1900 (US). SHASTA CO.: .Alorleys, Baker in

1898 (UC); 2.5 mi. W of Round Altn., Kuijt 1374

m 1957 (UC); Black Butte, Anderson in 1949 (CAS).

S0N0:MA CO.: Fort Ross, on P. muricata, Mason

4285 in 1928 (UC); 2 mi. S of Anchor Cover, on P.

muricata, Wolf 1342 in 1927 (DS). TEHAAIA CO • Mantón, Kuijt 1362 in 1957 (UC) and Root in 1919

(ILL); 12 mi. W of :\Iineral, Kuijt 1361 in 1957

(UC). TRINITY CO.: Trinity River near Swede

Cr. [ca. 6 mi. W of Big Bar], Tracy 6912 in 1924

(UC); near Weaverville, Kleeherger in 1879 (CAS)

TULARE CO. : 6 mi. E of Roads End P. 0., Hoivell

33147 in 1958 (CAS); Kern River Can., 7.6 mi. N

of Kernville, Tivisselmann n.d. (CAS). TUOLUMNE

CO.: 5 mi. E of Sonora, Wright FP 68054"" in 1931

(FPF); 2 mi. E of Sonora, W 3221 in 1962 (COLO,

FPF); E side Sonora, Gill & Wright FP 68066 in

1932 (FPF); 1 mi. W of Columbia, Peterson 63-136

in 1963 (FPF); Long Barn, Gill FP 68080 and FP

68099 in 1932 (FPF) and Wright FP 68119 in 1932

(FPF); Confidence, Boyce FP 97953 in 1915 (FPB);

2 mi. W of Rte. 49 Jet. on Rte. 120, Kuijt 1427 in

1957 (UC); 4 mi. NE of Columbia, Smith in 1956

(UC); Sonora, Bigelow 4 in 1854 (MO).

19. ARCEUTHOBWM PUSILLUM

Specimens Examined: on Picea inariana, except as

noted.

CANADA

MANITOBA

Freshford, Laut in 1968 (FPF, WINF) and on Pinus

banksiana, Laut in 1968 (FPF, WINF). Pelaire Provincial For., Laurence in 1967 (FPF, WINF);

Amana Bay, 40 mi. E of Gypsumville, on P. glauca,

Lawrence in 1967 (FPF, WINF); Sandilands For.

Reserve, Campbell in 1966 (FPF, WINF) and on

Pinus banksiana, Laut in 1967 (FPF, WINF); Glenboro, on P. glauca, Campbell in 1965 (FPF, WINF);

East Braintree, on P. glauca, Campbell in 1966 and

Lawrence in 1967 (FPF, WINF); Long Point, SE of

Grand Rapids, Shepherd in 1968 (FPF, WINF).

NEW BRUNSWICK

Grand Alann, Weatherby & Weatherby 5770 in 1927

(UC, US) and 6685 in 1934 (US); 10 mi. W of

Napadogan, Bonga* in 1964 (FPF); Acadia For.

Exp. Sta., Bonga in 1965 (FPF); Deer Island, near

Johnson Lake, on Picea sp., Malte 731/29 in 1929

(US, WTU).

216

NEWFOUNDLAND

Wild Cover, near Humbermouth, White Cliff,

Humber Dist., Rouleau 612 in 1950 (RM, UC, US);

Bay of Islands, Howe & Lang II41 in 1901 (GH);

Deer Arm, Bonne Bay; Alclver^s Cove, Fernald et al

1655 in 1929 (GH, PH), Deer Brook, Fernald et al

1656 in 1929 (GH, PH), and .Alain River, Fernald

et al. in 1929 (PH, US). Port a Port, on Picea sp.,

Fernald & St. John 10830 in 1914 (GH).

NOVA SCOTIA

Cape Breton Island: Creignish, Bonga in 1965

(FPF); 2 mi. E of Margaree For., von Schrenk in

1928 (MO, PH, POM, UC); Ingonish, on Picea sp..

Pease 20948 in 1929 (GH). Lower Argyle, Fernald

et al. 21045 in 1920 (AIO, UC, US); Gold Lake,

Birchton Branch, Fernald & Long 23788 in 1921

(GH, NY, PH) ; Hectanooga, Fernald & Pease 2IO41

in 1920 (GH, PH); Pictou, Hoive & Lang (GH,NY,

PH, RM) and on P. glauca, von Schrenk in 1929

(AIO); Tefry's Lake, Arcadia, Fernald & Pease

21043 in 1920 (GH) : East Jordan, Fernald & Pease

23878 in 1921 (GH); Alt. Uniache, Jack 3225 in

1924 (GH); Woliamakeh Beach, Port Mouton, Jack

3473 in 1924 (GH). Yarmouth Co.: Tusket, on P.

glauca. Jack 3759 in 1924 (UC) and Long & Fernald

21042 in 1920 (GH, PH); Eel Lake, on P. glauca,

Fernald et al 2IO44 in 1920 (GH, PH). Shelburne

Co. : Jack 3468 in 1924 (US).

ONTARIO

Snelgrove, White in 1918 (FPF, ILL); 7 mi. SW of

Killarney, Georgian Bay, on P. glauca, Coville in

1915 (US); near Cloche, McGregor Bay, N side

Georgian Bay, on P. glauca, Coville in 1915 (US);

Bruce Peninsula, Stokes Bay, on Larix laricina,

Hamai in 1964 (FPF); White Otter Lake, on Pinus

banksiana, McPhee & Miller, in 1955 (FPF, FPT,

WINF); Temagami For. Reserve, Watson 6762 in

1922 (WIS).

PRINCE EDWARD ISLAND

Tracadie Beach, Churchill in 1901 (MO, PH) : Tignish, Fernald et al 7327 in 1912 (NY, PH, UC, US).

QUEBEC

Richelieu, Rouleau in 1945 (UC, WIS). Lac-AuxAtocas, Pare de Mont Tremblant, Rolland-Germain

220 in 1956 (UC, US); Riv. Ste. Anne des Monts,

Fernald & Collins 218 in 1906 (US); St. Adolphe,

Comte d' Argenteuil, Riv. du Lac. St. Joseph, on

Picea sp., Rolland-Germain 7512 in 1949 (DS); Cap

Chat River, below Pirian River, Matane Co., on

Picea sp., Fernald & Pease 25031 in 1922 (GH); St.
Adolphe, Argenteuil, on P. glauca, Rolland-Germain

3061 in 1949 (R^I); Ste. Anne de la Pocatiere, on

P. glauca, Marie-Victorin 28^56 in 1928 (RAI, US,

WIS), on Picea sp., Cody et al 676 in 1947 (MO)

and Campagna 34173 in 1930 (GH); near Lake

Ouimet, Tierbonne Co., Pease 19055 in 1922 (GH);

Lac Alonroe, on ?, Rolland-Germain 361 in 1957

(US); Mont Tremblant, on Picea sp., Louis-Alphonse in 1952 (MO, US).

SASKATCHEWAN

Otosquen, Crawford in 1967 (FPF, WINF):Mi. 30

on Cumberland rd., on P. glauca, McLeod in 1968

(FPF, WINF); on Highway 123, on P. glauca at

Mi. 20, McLeod in 1968 (FPF, WINF) and jMi. 37,

McLeod in 1968 (FPF, WINF).

UNITED STATES

CONNECTICUT

LITCHFIELD CO.: North side Bear Altn., Greenman 2492 in 1910 (GH, MO) ; Bigham Pond, near

SaUsbury, Mann in 1906 (US); South Spectacle

Pond, near Kent, Gill et al FP 68284"" in 1933

(FPF) and Earger 45859 in 1909 (PH), and on Larix

laricina, Gill et al FP 68285 in 1933 (FPF) ; Norfolk,

Weatherhy 3906 in 1916 (PH, US) and Bissell in

1904 (GH).

MAINE

AROOSTOOK CO.: Fort Kent, Fernald in 1899

(UC); Monticello, Fernald & Long 13521 in 1916

(PH). CUMBERLAND CO.: Great Chebeague

Island, on Picea sp., Fernald 1741 in 1909 (GH).

FRANKLIN CO.: New Sharon, Knowlton in 1903

(GH) and Knowlton 23525 in 1905 (WIS). HANCOCK CO. : Mt. Desert Island, Great Head, on P.

glauca. Rand in 1906 (UC); Acadia Nat. Park,

Cottajn 11561 in 1938 (FPF, UT) : Gouldsboro,

Nortan in 1922 (WIS). KENNEBEC CO.: Belgrade,

Fassett 2527 in 1925 (GH, WIS). KNOX CO.: Isle

au Haut, on P. glauca, Arthur in 1900 (GH) ; Matinicus Island, on P. glauca, Norton & Chamberlain in

1916 (GH). LINCOLN CO.: Bristol, on P. glauca,

Churchill 351 in 1903 (MO) ; Squirrel Island, Boothbay Harbor, on P. glauca, Welch"" in 1958 (FPF);

Monhegan Island, on P. glauca, von Schreiik* in

1899 (MO) and Fernald in 1933 (PH, US); Linekin,

on P. glauca, von Schrenk in 1899 (MO) ; Georgetown,

on Picea sp., Morton 2063 in 1921 (WTU). Lighthouse Hill, host?, Churchill in 1921 (WIS) ; White

Island, host?, Fassett 2185 in 1925 (WIS); Ocean

Point, Fassett 3923 in 1922 (WIS). OXFORD CO.:

Norway, Bean 2241 in 1907 (GH). PENOBSCOT

CO.: Alton, Fernald 23322 in 1900 (GH, WIS).

SOMERSET CO.: Baker Brook, Headwaters St.

John River, St. John & Nichols 2281 in 1917 (US);

Pleasant Ridge, Murdoch 2010 in 1906 (F) ; Palmer

Pond, Fassett 14308 in 1931 (WIS). WASHINGTON

CO.: Indian Township, Grand Lake Stream rd.,

Gill FP 68313 in 1934 (FPF). COUNTY UNCERTAIN: :Middle Gary Pond, Spaulding & Collins in

1920 (ILL); Deer River, Weir 2394 n.d. (ILL).

MASSACHUSETTS

BERKSHIRE CO.: Becket, Hoffmann in 1904

(GH) and 1907 (AIO). ESSEX CO.: Andover, Pease

406 in 1902 (GH). HAMPDEN CO.: Springfield,

Clark & Seymour G586 in 1928 (WIS). :\IIDDLESEX CO.: Wilmington, Moore 634 in 1903 (UC);

Concord, Eaton in 1930 (RSA); Acton, on Picea sp.,

Jack in 1898 (GH). WORCHESTER CO.: Ashburnham, Murdoch 5301 in 1914 (F); Lancaster,

Fassett 2345 in 1924 (GH) ; Clinton, on P. glauca.

Jack in 1898 (F); Boylston, Jack in 1898 (MO) and

on P. glauca. Jack in 1898 (F, MO).

MICHIGAN

(Information on the A. pusillwn collections at the

University of ^Michigan (IMICH) and the Cranbrook

Institute of Science (BLH) provided by Dr. E. G.

Voss and at ]Michigan State University (]\ISC) by

Dr. J. H. Hart). ALCONA CO.: N end of Milikin

Lake, Voss 4581 in 1957 (MICH). ALGER CO.:

Chatham, on Picea sp., col? in 1901 (NY, US) and

Wheeler in 1900 (MSC) ; Rock River, Kauffman in

1927 (MICH). BENZIE CO.: Homstead Township,

North Branch of Platte River, Fröhlich 61 in 1968

(MICH). CHARLEVOIX CO.: Beaver Island: Egg

Lake, Voss 7009 in 1958 (MICH) ; Green Bay, on P.

glauca, Voss 3935 in 1957 (MICH); and Donegal

Bay, on P. glauca, Voss 6986 in 1958 (.MICH).

CHEBOYGAN CO.: Little Lake 16, near Cheboygan, Cutler in 1937 (MO, US), Voss 1539 in 1953

(MICH), and Ehlers 6127 in 1936 (MICH). ^lud

Lake bog, Gates & Gates 10682 in 1917 (R:M), Gates

14886 in 1927 (?^I0, UC) and Erlanson 484 in 1924

(MICH). Douglas Lake, Ehlers 1120 in 1920 (MICH,

WIS). Bryans Bog, Univ. of Alich. Biol. Sta., Gates

11116 in 1919 (WTU). CHIPPEWA CO.: 2.5 mi.

N of Eckermann, McVaugh 9781 in 1948 (BLH,

MICH, MO). Munuskong Bay State Park, Bessey

in 1927 (AISC). 3 mi. E of W Border of county on

Rte. M-28, Bourdo in 1961 (MSC). Whitefish township at mouth of Toquamonon River, Churchill in

1957 (MSC). Drummond Island, Warners Cove, on

P. glauca, Voss 12239 in 1966 (^NIICH). EMMET

CO.: Bliss Township, Waugoschance Point, on P.

glauca, Churchill in 1955 (MSC) and Voss IO48 in

1951 (MICH). Bliss Township, Sec. 18, on P. glauca,

Marshall 809 in 1949 (MSC). Galloway Lake, 1 mi.

N of Levering, Wood, Clover & Voss 8118 in 1953

217
(MICH) and Voss 12076 in 1966 (MICH). GOGEBIC CO.: Near Imp Lake, Gillis 3007 in 1959

(MSC) and Voss 7851^ in 1958 (MICH). Banks

Lake, Voss 12^67 in 1967 (MICH). Mud Lake,

Darlington 27^9 in 1919 (MICH). Sylvania Recreation Area, Voss 12712 in 1968 (MICH). HURON

CO.: Island in Rush Lake, Dodge in 1908 (:MICH,

MSC). IRON CO.: Basswood Pond, ca. 3 mi. SE

of Elmwood, Voss 8602 in 1959 (MICH). KEWEENAW CO.: 1.5 mi. W of Eagle Harbor, Voss 11870

in 1965 (MICH). LEELANAU CO.: Glenn Arbor

Township, Sec. 13, bog near Crystal River, Thompson L-1128 in 1948 (BLH). LUCE CO.: Barclay

Lake, 9 mi. SW of Crisp Point on Lake Superior,

Voss 3169 in 1956 (MICH). [NIACKINAC CO. : 7 mi.

E of Naubinway on US 2, on P. glauca, H 687* in

1964 (FPF). Bois Blanc Island, on P. glauca, Dodge

in 1913 (MICH, MSC) and Ehlers 51U in 1932

(iMICH). MARQUETTE CO.: Turin, Barlow 145*

in 1901 (MSC, UC). Conway Lake, Stoutarnire 3013

in 1959 (BLH). MENOMINEE CO.: Bog NE of

Wireglass Lake, ca. 5.5 mi. WSW of Carney, Voss

12696 in 1968 (MICH, MSC). MONTMORENCY

CO. : Ess Lake, 5 mi. N of Hillman, Stewart in 1966

(FPF). Lake 22 (Gaylanta Lake), Case in 1955

(MICH). SCHOOLCRAFT CO.: Long Lake, ca.

5.5 mi. NE of Meistrand, Voss 2660 in 1955 (MICH).

MINNESOTA

AITKIN CO. : 6 mi. S of Aitkin, Gill FP 68209 in

1932 (FPF) ; S of McGregor, Rosendahl U^O in 1924

(PH). BELTRAMI CO.: Waskish, Anderson* in

1965 (FPF) ; W of Upper Red Lake, T. 135 N, R.

35 E, Sec. 5, Anderson in 1967 (FPF). CASS CO.:

Cass Lake, Weir in 1916 (ILL, RM, UC) and on

Larix laricina. Weir in 1916 (FPF, ILL). CHISAGO

CO.: T. 37 N, R. 22 W, Gill FP 68208* in 1932

(FPF). CLEARWATER CO.: Itasca State Park,

Green 9 in 1947 (WIS) and on Larix laricina, French

& Tainter in 1967 (FPF) ; 2 mi. NW of Mississippi

River headwaters, Thome 19860 in 1958 (RSA);

Lake Itasca, on P. glauca, Hedgcock & Freeman FP

Jf.U5 in 1910 (FPF); Lake Itasca, Floating Bog

Bay, on P. glauca, Thome in 1962 (RSA). COOK

CO.: Superior Nat. For., T. 64 N, R. 1 W, Sec. 6,

on Rte. 12 ca. 24 air mi. NNW of Grand Marais,

Skillvng in 1964 (FPF). HUBBARD CO.: Itasca

Park, SE of Mary Lake, Grant 3197 in 1929 (GH,

MO, PH, WTU, US). ST. LOUIS CO.: 16 mi. N of

Duluth, Lakela 4503 in 1941 (MO): Floodwood,

Lakela 18527 in 1954 (US); Duluth, Normanna

township, Lakela 15555 in 1952 (WIS). COUNTY

UNCERTAIN: Gnesen Township, N of Duluth,

Lakela 7912 in 1948 (RM).

NEW HAMPSHIRE

CARROL CO.: Ossipee, on Picea sp., Grover in

1896 (GH). COOS CO.: ^'White Trail,'^ Shelburne,

on ?, von Schrenk in 1938 (^10); Randolph, Brook

Bank, Pease 3258 in 1903 (UC) ; Round Pond, Errol,

Pease 32509 in 1946 (RSA); Fernway Region, Randolph, on P. ruhens, Moore 143 in 1901 (PO^^I, UC).

CHESHIRE CO.: Nelson, on Picea sp., Batchelder

in 1931 (:M0). GRAFTON CO.: Hanover, Jesup &

Eggelston in 1893 (DS, PH, US) and on Picea sp.,

Eggelston in 1893 (PH, R]\I); Lebanon, Jesup in

1891 (MO). R0CKINGHA:M CO.: Nottingham,

on Picea sp., Eaton 443 in 1900 (GH). COUNTY

UNCERTAIN: White :Mtn. Nat. For., SE of Blue

Brook Ranger Sta., Hedgcock FP 864O* in 1913

(FPF, ILL).

NEW JERSEY

SUSSEX CO.: Pine Swamp, Lake ^Nlashipscong,

MacKenzie in 1920 (PH) and Cain & Svenson 6597

in 1935 (UC, WIS).

NEW YORK

CHENANGO CO.: Newcomb, Hanshrough &

Spaulding FP 68210 in 1932 (FPF); Plymouth,

Coville in 1887 (US); Preston, Coville in 1885 (US).

CLINTON CO.: Platsburg, Hotel Champlain, on

P. glauca, Schwartz FP 68288* in 1934 (FPF).

COLUMBIA CO.: 1.5 mi. NW of Niverville, on

Picea sp.. Wherry in 1934 (UC). CORTLAND CO.:

Labrador Swamp, Wiegland in 1893 (^10) ; Truxton,

on Picea sp., Wiegland in 1894 (UC). ESSEX CO.:

Newxomb, on P. glauca, House 7186 in 1920 (UC) ;

FRANKLIN CO. : Saranac Inn, Spaulding FP 34543

in 1909 (FPF). HA?^IILTON CO.: Raquette Lake

Village, von Schrenk in 1932 (PH, PO.AI, R:\I, UC);

Whitney Park, Sabattis, on P. ruhens, Spaulding &

Eno FP 68312 in 1934 (FPF). HERKBIER CO.:

Without locality, on Picea sp.. Höherer n.d. (UC);

Graefender Swamp, Höherer in 1881 (PH) ; Frankfort Hill, Höherer 188 in 1892 (DS); Litchfield,

Höherer in 1881 (GH). MONROE CO.: :Mendon,

Baxter in 1908 (UC) and Killip in 1917 (ARIZ,

POM). ONEIDA CO.: Frankfort Hills, Utica, col.?

in 1881 (MO). ONONDAGA CO.: Cicero, Wiegland

6344 in 1916 (GH, MO); Syracuse, Pennington n.d.

(ILL) and on Larix laricina, Pennington n.d. (ILL).

OSWEGO CO.: Oswego, Wihhe 11532 in 1878 (MO),

Killip 7816 in 1921 (US) and on P. glauca, Sheldon

439 in 1881 (US); South New Haven, Mathews 2663

in 1927 (RM); New Haven, Rowlee in 1891 (MO).

RENSSELAER CO.: Without locality, Wihhe in

1891 (ARIZ, CS, RM, UC, US, WIS). SCHENECTADY CO.: Schenectady, Wihhe in 1891 (F, US).

ST. LAWRENCE CO.: Bean Pond, Wanakena,

218
Benedict in 1918 (ILL). SULLIVAN CO.: Forestburg, Peck in 1873 (MO). THOMPKINS CO.:

Franklin Hills, Haherer in 1882 (GH). WARREN

CO.: E lake George Marsh, Brayton, Burnham in

1897 (PH) ; Warrensburgh, Millington in 1912 (MO).

WEBB CO.: Big Moose Lake, on P. ruhens, von

Schrenk in 1932 (MO, POM, UC). WESTCHESTER

CO.: Peekskill, on Picea sp., LeRoy n.d. (NY).

PENNSYLVANIA

LACKAWANA CO. : Moosic Mtn., NE of Scranton,

col? in 1886 (PH); Little Roaring Brook, Scranton,

Dudley in 1886 (DS). MONROE CO.: Tannersville,

Williamson in 1908 (PH) ; Cranberry Marsh near

Tannersville, Batram & Long in 1907 (PH); 5 mi

NE of Dresser, Haas in 1951 (PH). PIKE CO.:

Pine Lake, Greentown, Olday 240'' in 1961 (FPF);

Spruce Pond, Broion & Saunders in 1899 (PH).

SULLIVAN CO.: Shadynook, Brown in 1901 (MO,

PH); near Lopez, Williamson in 1908 (GH, PH).

WAYNE CO.: 1.5 mi. E of Hamlin, Glowenke 2468

m 1938 (WTU).

RHODE ISLAND

PROVIDENCE CO. : Bowdish Reservoir, Glochester, Collins in 1904 (PH, UC), in 1919 (US) and in

1927 (MO).

VERMONT

CHITTENDEN CO.: Near Burlington, on Larix

laricina, von Schrenk in 1899 (MO). GRAND ISLE

CO. : Alburgh, Jones in 1899 (GH). RUTLAND CO. :

Pittsford, on Picea sp., Eggelston 1561 in 1899 (US);

Mendon, on Picea sp., Eggelston 1562 in 1899 (GH);

Brandon, Button in 1911 (MO).

WISCONSIN

(Information on the A. pusillum collections at the

University of Wisconsin (WIS) provided by Dr.

H. H. Iltis, 1969). ASHLAND CO.: Mellen, Fassett

10093 in 1927 (WIS). BURNETT CO.: West

Sweden, Ninman & Thompson FP 38713 in 1922

(FPF); Gaslyn, Davis in 1911 (WIS). DOOR CO.:

Sturgeon Bay, Davis in 1913 and 1929 (WIS).

Baileys Harbor: Davis in 1918 and 1929 (WIS), and

Fassett & Sicker 14703 in 1932 (WIS) and, on P.

glauca, von Schrenk in 1934 (FPF, WIS), Pohl 1145

in 1938 (WIS), and Iltis 17558 in 1961 (WIS).

DOUGLAS CO.: Gordon, Wadmond 1271 in 1907

(GH, WIS); Solon Springs, Fassett 22395 in 1934

(WIS). FLORENCE CO.: Boot Lake, Iltis, Iltis &

Turner 22225 in 1964 (WIS) ; Long Lake, Davis in

1915 (WIS). FOREST CO.: Laona, Davis in 1915

(WIS). IRON CO.: Mercer, Davis in 1919 (WIS).

JACKSON CO.: Glacial Lake, Hartley 3991 in 1958

(WIS). LANGLADE CO.: Phlox, Davis in 1914

(WIS). LINCOLN CO.: Lost Lake, Seymour 11906

in 1950 (WIS); Gerbig Lake, Seymour 15104 in 1952

(WIS); Corning, Seymour 14645 in 1952 (WIS) and

Schlising 124 in 1952 (WIS). MANITOWOC CO.:

W of Kellnersville, Fassett, Hoffman & Staffield

18265 in 1934 (WIS). ONEIDA CO.: Rhinelander,

Anderson in 1963 (FPF) ; 9 mi. W of Crandon on

US 8, H 688 in 1964 (FPF) ; W of Woodruff on Rte.

20, Anderson"" in 1965 (FPF). PRICE CO.: Camp

Merrill, Davis in 1911 (WIS): Butternut Lake,

Davis in 1911 (WIS). RUSK CO.: Hawkins, Davis

in 1918 (FPF, ILL, WIS). SAWYER CO. : Hayward,

Davis in 1924 (WIS) and Gilbert & Davis in 1932

(WIS). TAYLOR CO.: 3.5 mi. N of Rib Lake,

Anderson 265 in 1947 (WIS). VILLAS CO.: Trout

Lake, Spaulding in 1915 (FPF), Fassett 8996 in

1929 (WIS), and Fassett 13767 in 1932 (WIS). Lake

Louise, 5 mi. E of Boulder Jet., Wilson & Wilson

3067 in 1932 (POM, WIS). WASHBURN CO.:

Spooner, Davis in 1911 (WIS).

20. ARCEUTHOBIUM RUBRUM

Specimens Examined:

MEXICO

DURANGO: 29 mi. E of El Salto on Rte. 40, on

Pinus teocote, H&W 338* and 521* in 1963 (COLO,

FPF); on P. engelmanni, H&W 339* in 1963

(COLO, FPF) ; and on P. cooperi, H & W 340 in

1963 (COLO, FPF). 24 mi. E of El Salto on Rte.

40, on P. teocote, H&W 1243 in 1969 (FPF); 2 mi.

W of El Salto on Rte. 40, on P. teocote, H&W 345*

in 1963 (COLO, FPF) and on P. durangensis, H&W

346 in 1963 (COLO, FPF). 35 mi. S of Durango on

La Flor rd., on P. teocote, H & W 511* in 1963

(COLO, FPF); on P. engelmannii, H&W 513* in

1963 (COLO, FPF) ; and on P. cooperi, H & W 512

in 1963 (COLO, FPF) ; 49 mi. W of El Salto on Rte

40, on P. herrerai, H&W 1248 in 1969 (FPF). 30

mi. NW of Santiago Papasquiaro, on P. teocote,

Straw & Forman 1777 in 1959 (RSA). SINALOA:

Without locality or host, Comisión Catastral y de

Estudio de los Recursos Naturales del Estado,

Gomales 1717 in 1920 (K, US).

21. ARCEUTHOBIUM STRICTUM

Specimens Examined: On Pinus leiophylla var.

chihuahuana except as noted.

MEXICO

DURANGO : City of Durango and vicinity, Palmer

774 in 1896 (BM, F, ILL, MO, NY, UC, US); 38

mi. E of El Salto on Rte. ^0, H & W 329* in 1963

219
(COLO, FPF); 36 mi. E of El Salto on Rte. 40,

H&W 1242 in 1969 (FPF) and on P. teocote, H&W

S18* in 1963 (COLO, FPF); 16 mi. S of Durango on

La Flor rd., H (& W 500'' in 1963 (FPF); 24 mi. S

of Durango on La Flor rd., on P. teocote, H&W

505* in 1963 (COLO, FPF) and on P. engehnannii,

H&W 506* in 1963 (COLO, FPF).

22. ARCEUTHOBIUM TSUGENSE

Specimens Examined: All on Tsuga heterophylla,

except as noted.

CANADA

BRITISH COLUMBIA : Vancouver Island: Rosen^

dahl & Brandegee 136 in 1901 (COLO, MO) ; Ucuelet,

Henry 1822 and 1881 in 1917 (RM); Pender Harbour, Spaulding FP 41633* in 1923 (FPF); Sproat

Lake, on Pinus contorta, Rosendahl & Butters 1092

in 1906 (WTU) and Rosendahl 1958 in 1907 (UC);

6 mi. E of Duncan Jet., Kuijt in 1954 (DAVFP,'

FPF); 6 mi. W of Nanaimo Lakes Jet., Kuijt in

1954 (DAVFP, FPF): Cowichan Lake, Kuijt in

1954 (DAVFP, FPF) and Smith in 1965 (DAVFP,

FPF); Robertson River Valley, Kuijt* in 1954

(DAVFP, FPF); 8 mi. W of Sooke, Kuijt in 1954

(DAVFP, FPF); Chemainus, Porter in 1954

(DAVFP, FPF) and on Pinus contorta, Kuijt in

1954 (DAVFP, FPF); Metchosin, on P. contorta,

Kuijt 602 in 1954 (DAVFP, FPF); Home Lake rd.,

on P. contorta, Kuijt in 1954 (DAVFP, FPF); Goldstream summit of Mt. Finlayson, on P. contorta.

Ziller in 1965 (DAVFP, FPF). Sechelt, Kuijt 618 in

1954 (DAVFP, FPF) and on P. contorta, Kuijt in

1954 (DAVFP, FPF). Vancouver, Stanley Park,

Hopping FP 97939 in 1928 (FPB) and Kuijt in

1954 (DAVFP, FPF); Vancouver, Univ. of British

Columbia Botanic Gardens, Kuijt in 1954 (DAVFP,

FPF) ; Agassiz, Weir 2^49, n.d. (ILL) ; Savary Island,

on Pmus contorta, Davidson in 1918 (FPF, ILL);

Daisy Lake, ca. 30 mi. N of Squamish, Boyce 2122

in 1931 (FPF).

UNITED STATES

ALASKA

Juneau, Anderson 64O6 in 1941 (PH, RM); Eagle

River, 25 mi. N of Juneau, Laurent in 1965 (FPF) ;

Chicagof Island, Basket Bay, Laurent & Labue* in

1965 (FPF) ; Kupreanof Island, 15 mi. SW of Petersburg, Graham in 1966 (FPF) ; 1 mi. S of Little Basket

Bay, on Picea sitchensis, Laurent in 1965 (FPF);

near Sitka, on T. mertensiana, Weir in 1913 (FPF,

ILL); near Ketchikan, Weir in 1915 (ILL) and

Drake in 1917 (FPF, ILL); Mt. Verstovia, Sitka,

Coville & Kearney 916 in 1899 (US); Sitka, Anderson

"Mn 1915 (US).

CALIFORNIA

ALPINE CO. : Mosquito Lake, 11 mi. NE of Alpine,

on Tsuga mertensiana, H & Scharpf 665* in 1964

(FPF) and on Pinus monticola, H & Scharpf 666 in

1964 (FPF). MENDOCINO CO. : 3 mi. E of Mendocino, H & Scharpf 850in 1966 (FPF). PLACER CO. :

Emigrant Gap, on Tsuga mertensiana, Jones in

1881 (POM). PLUMAS CO.: Mt, Elwell [6 mi. SW

of Blairsden], on Tsuga mertensiana. Lieberg 5363

in 1900 (ILL, US). SHASTA CO.: Lassen Nat. Park:

2.5 mi. N of S entrance, on Tsuga mertensiana, W

3235 in 1962 (COLO, FPF) ; 23 mi. SE of N entrance,

on main rd., on Tsuga mertensiana, H&W 652 in

1964 (FPF); 1-2 mi. S of Helen Lake, on Tsuga

mertensiana, Scharpf FP 38027 n.d. (FPB). SISKIYOU CO.: Marble Mtns.: Kidder Cr., on Tsuga

mertensiana. Gill & Sargent FP 68180* in 1932

(FPF) ; Chimney Rock Lake, on Tsuga mertensiana,

Hemphill in 1968 (FPF, UT) and on Picea breweriana, Hemphill in 1968 (FPF, UT); near Chimney

Rock, on Pinus monticola, Hemphill in 1968 (FPF,

UT). Siskiyou Mtns.; Applegate Cr. Divide, on

Tsuga mertensiana, Meinecke FP 97938 in 1913

(FPB) and on Picea breweriana. Meinecke FP 97941

in 1913 (FPB). TEHAMA CO.: Mineral, on Tsuga

mertensiana. Lory FP 97940 in 1911 (FPB)

COUNTY UNCERTAIN: Bald Mtn., Hubert in

1918 (ILL).

IDAHO

These Idaho records on Tsuga heterophylla are tentatively listed here pending confirmation (see text).

LATAH CO.: Bovill, Weir in 1918 (FPF, ILL).

COUNTY UNCERTAIN: St. Joe Nat. For., Weir

2452 in 1914 (ILL) and Weir 6838 in 1916 (FPF,

ILL).

OREGON

BENTON CO.: Mary^s Peak, on Abies procera,

Shea in 1968 (FPF). CLACKAMAS CO.: Still Cr.

Trail near Rhododendron, Gill FP 68193* in 1932

(FPF), Fault, Childs & Hansbrough FP 68266 in

1931 (FPF), and on Pinus monticola, Langley &

Goodding in 1931 (RM); 4 mi. E of Rhododendron,

Mielke FP 58275 in 1930 (OSC) ; 2 mi. W of Government Camp on US 2d, H & W 605 in 1964 (FPF)

and on Abies amabilis, H&W 606 in 1964 (FPF);

Twin Bridges [5 mi. W of Government Camp on

US 26], Thompson 9422 in 1928 (DS, MO). COOS

CO.: Siskiyou Mtns., 10 air mi. NNE of Agness, on

T. mertensiana, Graham in 1965 (FPF). Near S fork

of Coos River, 38 air mi. W of Roseburg, Stewart

in 1968 (FPF). DESCHUTES CO.: 11 mi. W of

Sisters on McKenzie Pass rd., on Tsuga mertensiana,

H & Hinds 991 in 1966 (FPF) and on Abies lasio220
carpa, H & Hinds 992 in 1966 (FPF). DOUGLAS

CO.: Myrtle Cr.—Deadman Cr. divide, ca. 11 air

mi. N of Tiller, Graham in 1965 (FPF) ; Just N of

Crater Lake Nat. Park, Diamond Lake rd., on T.

mertensiana, Graham in 1964 (FPF) ; 20 mi. NE of

Union Cr. on Rte. 230, H lip in 1968 (FPF) and

on Pinus montícola, H 1142 in 1968 (FPF). JACKSON CO.: Union Cr., on T. mertensiana, Applegate

6038 in 1929 (DS) ; Huckleberry Mtn., 10 mi. NE of

Prospect, on T. mertensiana, Graham in 1965 (FPF);

8 mi. NW of W entrance to Crater Lake Nat. Park,

Graham in 1963 (FPF). KLAMATH CO.: Crater

Lake Nat. Park: Anna Springs Ranger Sta., on T.

mertensiana. Gill FP 68184 in 1932 (FPF); Wineglass, on Pinus albicaulis, Gill FP 68182 in 1932

(FPF); 0.5 mi. S of Wineglass, on T. mertensiana.

Gill FP 68183 in 1932 (FPF) ; 5 mi. E of main rd.

on N rim of Crater Lake, on T. mertensiana, H & W

624 in 1964 (FPF) and on Pinus albicaulis, H & W

625 in 1964 (FPF); Crater Lake, Palisades, on T.

mertensiana, Coville 1478 in 1902 (US); 15 mi. SW

of Crescent Lake, Childs 65 in 1937 (OSC); 2 mi.

N of Windigo Pass, on T. mertensiana, H & W 621

in 1964 (FPF) and on Pinus montícola, H & W 622

in 1964 (FPF). LANE CO.: E Fork of S Fork of

McKenzie River, 8 mi. S of McKenzie Bridge,

Stewart & Orr in 1968 (FPF) ; near head of INIosquito

Cr., on Abies amabilis, Stewart & Orr in 1968 (FPF);

Head of Olallie Cr., on Abies amabilis. A, grandis,

and A. lasiocarpa, Stewart & Orr in 1968 (FPF). 19

mi. E of Oakridge on Rte. 58, H II40 in 1968 (FPF) ;

near Foley Springs, on Abies sp., Coville & Applegate 1090 and 1091 in 1898 (US); Brice Cr., 20 air

mi. SE of Cottage Grove, Graham in 1965 (FPF);

McKenzie Pass. On Abies lasiocarpa, Childs FP

91561 in 1947 (OSC); on Pinus albicaulis, Gill FP

68188"" in 1932 (FPF), Lachmund & Childs FP 68269

in 1930 (FPF), and Chüds FP 91560 in 1947 (OSC);

on T. mertensiana. Gill FP 68187* in 1932 (FPF)

and Childs FP 91562 in 1947 (OSC). 1 mi. W of

McKenzie Pass: on A. lasiocarpa. Boyce 2298 in

1930 (FPF) ; on Pinus albicaulis, Boyce 2299 in 1930

(FPF) ; and on T. mertensiana, Boyce 2300 in 1930

(FPF). 1.5 mi. W of McKenzie Pass: on A. lasiocarpa, W 3241 in 1962 (COLO, FPF); on P.

albicaulis, W 3243 in 1962 (COLO, FPF) and

on T. mertensiana, W 3242 in 1962 (COLO,

FPF). LINCOLN CO.: 3 mi. E of Yachats,

Graham in 1966 (FPF); Dicks Fork Cr., 4 mi. S of

Waldport, H & Scharpf 1252 in 1969 (FPF) ; T 15 S,

R 11 W, Sec. 7, ca. 3 air mi. SE of Yachats on Picea

sitchensis, Ferguson in 1969 (FPF). LINN CO.:

Minto Mtn., on T. mertensiana, Mielke FP 40694

in 1931 (OSC); Marion Lake, on T. mertensiana,

Coville & Applegate 1156, in 1898 (US); 12 mi.

W of Santiam Pass on US 20, H & Scharpf 1254 in

1969 (FPF), on Abies amabilis, H & Scharpf 1255

in 1969 (FPF), and on Picea engelmannii, H &

Scharpf 1256 in 1969 (FPF). TILLAMOOK CO.:

Near Nehalem, Childs FP 40545 in 1930 (OSC).

WASCO CO. : 5.7 mi. SE of Government Camp on

US 26, H & W 607 in 1964 (FPF) and on A. amabilis,

H & W 608 m 1964 (FPF).

WASHINGTON

CHELAN CO. : Lyman Glacier W of Chelan Lake,

on T. mertensiana. Weir 2456 in 1916 (ILL); Chelan

Lake, Weir n.d. (FPF, ILL) : N of Chelan Lake, on

P. albicaulis. Weir 3242 in 1916 (FPF, ILL).

CLALLA]\1 CO.: Lake Crescent, on Tsuga sp.,

Jones 3439 n.d. (WTU) ; Piedmont, Crescent Lake,

Boyce 608'' in 1920 (FPF) and FP 40065 in 1920

(OSC) ; near Blyn, W 3240 in 1962 (COLO). GRAYS

HARBOR CO.: Wesport, Gill FP 68195 in 1932

(FPF); 3.5 mi. NW of Montesano, H 1250 in 1969

(FPF). KING CO.: Seattle, Piper in 1883 (PH),

and on T. mertensiana'!. Piper 663 in 1892 (DU, MO,

US, WTU); Berlin [5 mi. WNW of Skykomish],

Hedgcock FP 4765 in 1910 (FPF); Ronald Bog, N of

Seattle, Günther in 1930 (WTU); Corea, Otis 1036

in 1920 (US); 4 mi. W of Stevens Pass on US 2,

H, Stewart & Thompson 1138 in 1968 (FPF) and on

Abies amabilis, 1139 in 1968 (FPF). KITSAP CO.:

Mountaineer's Cabin, on Tsuga sp., Jones 4299 in

1931 (WTU). KITTITAS CO.: Near Ellensburg,

Brandegee 1070 in 1883 (PH, US) ; E side Snoqualmie

Pass, W 3252 in 1962 (COLO) and on Abies amabilis,

W 3251 in 1962 (COLO). LEWIS CO.: 5 mi. N of

Cayuse Pass on Rte. 143, on Abies grandis, H & W

602 in 1964 (FPF), 4 mi. W of White Pass on Rte.

14, Wicker in 1966 (FPF) and on Abies amabilis

Wicker in 1966 (FPF). PACIFIC CO.: 3 mi. N of

Nemah on Rte. 101, H & W 6O4 in 1964 (FPF).

PIERCE CO.: Mt. Rainier Nat. Park: Longmire:

Gill FP 68198 in 1932 (FPF), Gill & Warren FP

68199 in 1932 (FPF), Flett in 1915 (ILL), Bookman

FP 68265 in 1931 (FPF), and on A. amabilis, Flett

in 1916 (ILL) and Gill & Warren FP 68197 in 1932

(FPF) ; 5 mi. E of Longmire, W 3248 in 1962 (COLO,

FPF) and on A. amabilis, W 3249 in 1962 (COLO,

FPF); White River entrance, Gill FP 68196* in

1932 (FPF). Ashford, Flett in 1915 (FPF, ILL);

Huckleberry Cr., 8 mi. S of Greenwater, Steivart in

1968 (FPF) and on A. amabilis, Stewart in 1968

(FPF); Silver Springs, 32.5 mi. SE of Enumclaw

on US 410, H &W 601 in 1964 (FPF). SAN JUAN

CO.: Oreas Island, Mt. Constitution, Graham &

Thompson in 1965 (FPF) and on Pinus contorta:

Foster in 1907 (WTU), Cowles 372 in 1907 (US),

Rigg & Frye in 1908 (WTU), Zeller & Zeller 1194

in 1907 (MO), Wright in 1914 (US), Beattie 5801 in

1921 (FPF), Barker 254 in 1926 (DS) and Graham &

Thompson in 1965 (FPF). SKAGIT CO.: Near

221
Power Dam on Skagit River, col.? n.d. (WTU).

SKAMANIA CO.: Government Soda Springs,

Boyce 596 in 1920 (FPF) and Boyce FP 40057 in

1920 (OSC); 24 mi. SE of Rändle on Trout Lake

rd., W 3247 in 1962 (COLO, FPF); Tillicum Cr.,

18 air mi. ESE of Spirit Lake, Graham in 1965

(FPF) and on Abies amahilis, Graham in 1965

(FPF); Dry Cr., 20 air mi. N of Stevenson, Graham

in 1965 (FPF). SNOKOMISH CO.: 15 mi. N of

Seattle, Witirock 5097 in 1927 (WTU). THURSTON

CO.: Black Hills, Meyer 1685in 1939 (MO). WHATCOM CO.: Twin Sister Range, on T. mertensiana,

Muenscher 10431 in 1939 (DS); Bellingham to Mt.

Baker, on T. mertensiana, Weir in 1915 (ILL).

COUNTY UNCERTAIN: Upper Nisqually Valley,

on T. mertensiana, Allen 303 in 1896 (DS, MO, UC).

23a. ARCEUTHOBIUM VAGINATUM

subsp. VAGINATUM

Specimens Examined :

MEXICO

CHIHUAHUA: Sierra Madre, on Pinus sp., Nelson

6077 in 1899 (K, US) ; Sierra Madre near Guachochi,

on Pinus sp., Goldman 177 in 1898 (US); 36 mi. SW

of La Junta, on P. ponderosa var. scopulorum, H &W

295 in 1963 (COLO, FPF); 39 mi. SW of La Junta,

on P. ponderosa var. scopulorum, H & W 298"^ in

1963 (COLO, FPF); 6 mi. E of El Vergel, on P.

ponderosa var. scopulorum, H & W 310 in 1963

(COLO, FPF) ; 1 mi. W of El Vergel, on P. ponderosa var. arizonica, H & W 311 in 1963 (COLO,

FPF) ; 10 mi. W of El Vergel, on P. ponderosa var.

arizonica, H & W 313* in 1963 (COLO, FPF) : 10

mi. E of El Vergel, on P. ponderosa var. scopulorum,

H &W 308 in 1963 (COLO, FPF); on Ocampo rd.

SW of Matachic, on P. ponderosa var. arizonica, at

48 mi., H &W 486 in 1963 (COLO, FPF); 63 mi.

H &W 494'' in 1963 (COLO, FPF); and at 69 mi.

H & W 491 in 1963 (COLO, FPF). COAHUILA:

General Cepeda, Pinus sp.. Nelson 6730 in 1890

(US). DISTRICTO FEDERAL: Desierto de los

Leones, on Pinus sp., Alexander & Hernandez 2333

in 1945 (BM). DURANGO: Ciénega de Ibarra, on

Pinus sp., Martinez in 1940 (F); 13 mi. W of El

Salto on Rte. 40, on P. cooperi, H &W 348"* in 1963

(COLO, FPF): 15 mi. W of El Salto on Rte. 40, on

P. cooperi, H & W 531* in 1963 (COLO, FPF); 18

mi. W of El Salto on Rte. 40, on P. durangensis,

H &W 1245 in 1969 (FPF) ; 26 mi. S of Durango on

La Flor rd., on P. engelmannii, H & W 507* in

1963 (COLO, FPF); 47 mi. S of Durango on La

Flor rd., on P. cooperi, H & W 515 in 1963 (COLO,

FPF). HIDALGO: Real del Monte, on Pinus sp..

Coulter 17, n.d. (GH, K). JALISCO: Nevado de

Colima, on Pinus sp., Jones 487 in 1892 (ILL, MO,

POM, US); Pringle 4368 in 1893 (BM, ILL, K,

MEXU, MO, NY, PH, UC, Z); Reiche in 1913

(MEXU); McVaugh 10071 in 1949 (MICH) and

12916 in 1952 (MICH). MEXICO: Nevado de

Toluca, Ojos de Agua, on Pinus sp.. Balls B4O86 in

1938 (BM, K, UC, US); Telapón Peak N of Río

Frío, on Pinus sp., Miranda II4 in 1940 (MEXU)

and on P. hartwegii, Beaman 244I in 1958 (UC);

Ixtacchiuatl, on Pinus sp., Purpus 3836 in 1905

(POM); Río Frío, on Pinus sp.. Hartley FP 89795

in 1947 (FPF); 6 mi. W of Río Frío on Rte. 190, on

P. montezumae, H &W 370* in 1963 (COLO, FPF).

NAYARIT: Sierra Madre, near Santa Teresa, on

Pinus sp.. Rose 3460 in 1897 (US). NUEVO LEON:

Cerro Linadero, on Pinus sp., Meyer & Rogers 2932

in 1948 (MO); 15 mi. SW of Galeana, on Pinus sp.,

Mueller & Mueller 1013 in 1934 (F, MEXU, MICH) ;

Galeana, Hacienda Pablilio, on Pinus sp., Taylor

182 in 1936 (ARIZ, F, MEXU, MICH, RSA);

Pablilio, 15 mi. S of Galeana, on Pinus ponderosa

var. arizonica, Griffiths in 1966 (FPF). Cerro Potosí:

on P. rudis, Andresen & Steinhoff A2050 in 1963

(FPF); 2 mi. from village ''18 de Marzo^^ on rd. to

Relay Tower, on P. ponderosa var. arizonica, H & W

391* in 1963 (COLO, FPF); 10 mi. from village

''18 de Marzo'' on rd. to Relay Tower, on P. ponderosa var. arizonica, H & W 393 in 1963 (COLO,

FPF) ; near Relay Tower, on P. rudis, H & W 397*

in 1963 (COLO, FPF) and on P. culminicola, H &W

398 in 1963 (COLO, FPF). OAXACA: 1 mi. N of

Suchixtepec, on P. lawsonii, Peterson 68-106 in 1968

(FPF). SINALOA: Sierra Surutato, Municipio de

Badiraguato, 3 mi. SE of Los Ornos, on Pinus

herrerai, Breedlove & Thome 18535 in 1970 (CAS).

TAMAULIPAS: Rte. to Pena Nevada out of

Hermosa, on Pinus sp.. Standard, Tauber & Taylor

2495 in 1949 (DS, UC, WTU). PUEBLA: 1.7

mi. E of Paso de Cortez, between Popocatepetl and

Ixtacchiuatl, on P. hartwegii, H & W 374* in 1963

(COLO, FPF). VERACRUZ: Cofre de Perote, 6

mi. SE of Perote, on P. montezumae, H & W 378*

in 1963 (COLO, FPF) and 5 mi. SE of Perote, on

P. montezumae, H & W 380 in 1963 (COLO, FPF).

ZACATECAS: Sierra Madre, on Pinus sp.. Rose

3535 m 1897 (US).

23h. ARCEUTHOBIUM VAGINATUM

subsp. CRYPTOPODUM

Specimens Examined: on Pinus ponderosa var.

scopulorum except as noted.

MEXICO

CHIHUAHUA: 5 mi. S of Garcia on El Largo rd.,

on P. ponderosa var. arizonica, H & W 461* in 1963

(COLO, FPF); 17 mi. N of El Largo on Garcia rd.,

H & W 462 in 1963 (COLO, FPF); 9 mi. N of El

Largo on Garcia rd., on P. engelmannii, H & W 464

in 1963 (COLO, FPF); 16 mi. W of Mesa Huracán,

on Chico rd., on P. engelmannii, H & W 471 in

222
1963 (COLO, FPF) : 13 mi. SE of Mesa Huracán on

Chico rd., H &W 476 in 1963 (COLO, FPF) : 5 mi.

W of Las Varas on Mesa Huracán rd., on P. engelmannii, H & W 479 in 1963 (COLO, FPF) ; 18 mi.

5 of Las Varas on :Madera rd., H & W 480"" in 1963

(COLO, FPF) ; 45 mi. SW of Matachic on Ocampo

rd., on P. ponderosa var. arizonica, H & W 4^5 in

1963 (COLO, FPF); 66 mi. SW of .Matachic on

Ocampo rd., on P. ponderosa var. arizonica^ H & W

492 in 1963 (COLO, FPF). COAHUILA: Sierra del

Carmen: 1 mi. W of Ocampo, H,L& Munoz 1023*,

on P. ponderosa var. arizonica in 1967 (FPF) and

1024 on P. arizonica var. stonniae in 1967 (FPF);

6 mi. W of Ocampo, on P. ponderosa var. arizonica,

H, L& Muñoz 1026 in 1967 (FPF). Sierra del Pino,

on P. ponderosa var. arizonica, Johnston & Mueller

591 in 1940 (GH). SONORA: 2.7 mi. N of La Mesa,

on P. engelmannii, Tucker 2563 in 1952 (ARIZ, UC).

UNITED STATES

ARIZONA

APACHE CO. : 6 mi. W of Window Rock, H & L

188 in 1962 (FPF); 4 mi. S of Eagar, H & L 211 in

1962 (FPF); 2 mi. E of Big Lake, H 821 in 1965

(FPF); Lukachuikai Mtns., Goodman & Payson

2927 in 1936 (WTU); Fort Apache Res., 5 mi. NE

of Maverick, H &L901 in 1966 (FPF); Fort Apache

Res., 14 mi. W of Maverick on Fort Apache rd.,

H & L 903 in 1966 (FPF); 8 mi. E of McNary on

Rte. 73, H & L 923 in 1966 (FPF). Navajo Res.,

6 mi. NW of Sawmill, L & Weiss 66-44 in 1966

(FPF); 6 mi. E of Lukachuikai on Red Rock rd.,

L & Weiss 66-49 in 1966 (FPF) ; 11 mi. SW of Red

Rock on Lukachukai rd., L cfc Weiss 66-51 in 1966

(FPF). COCHISE CO.: Chiricahua Mtns., Burrall

55 in 1908 (FPF) and Raue 50 in 1956 (ARIZ):

Rustler Park, Gill FP 68047 in 1935 (FPF); 1 mi.

S of Rustler Park, H 808 in 1965 (FPF); Onion

Saddle, on P. ponderosa var. arizonica, H & L 154

in 1962 (FPF) and on P. engelmannii, H & L 155

in 1962 (FPF) ; Barfoot Peak on P. ponderosa var.

arizonica, Bluyner 1452 in 1906 (ARIZ, FPF, ILL,

RM, US); Rustler, on P. ponderosa var. arizonica,

Ferris 9953 in 1940 (DS, RM, RSA, UC, WTU);

Gulching rd., on P. engelmannii. Blumer 1557 in

1907 (ARIZ, FPF, ILL, RM, US, K); 10 mi. W of

Portal, on P. engelmannii, Gill FP 68038 in 1936

(FPF). Huachuca Mtns.: Carr Can., Gill, Ellis &

Sowell FP 89498 in 1939 (FPF) and Goodding FP

33285"" in 1919 (FPF); head of Carr Can., on P.

ponderosa var. arizonica. Gill, Ellis & Sowell FP

89499 in 1939 (FPF); Miller Can., Goodding in 1909

(ARIZ); head of Ramsey Can., Doherty FP 15240

in 1914 (FPF, ILL); 1.5 mi. W of Reef, on P.

ponderosa var. arizonica, H & L 233 in 1962 (FPF).

COCONINO CO.: Jacob Lake, Reed & Barkley

4381 in 1939 (POM); Flagstaff, MacDougal 23 in

1898 (ARIZ, RM, UC); 15 mi. S of Williams, Gill

FP 68252 in 1932 (FPF); Williams, Hedgcock &

Bethel FP 244ÍS in 1917 (FPF); 2.5 mi. E of Fort

Valley Exp. For., Gill FP 68249 in 1932 (FPF);

Fort Valley Exp. For., Long FP 19724 m 1915

(FPF); Grand Can., S Rim, Grandview Point, H &

L 185 in 1962 (FPF); 3 mi. N of Weimer Springs,

W of Mormon Lake, H & L 912 in 1966 (FPF);

13 mi. NE of Glints Well on Rte. Qo, H & L 914

in 1966 (FPF); 3 mi. S of Chevlon Ranger Sta., 43

mi. S of Winslow, H & L 917 in 1966 (FPF); Jacob

Lake, Gill FP 68296 in 1934 (FPF); Dry Park, NW

of Kaibab Lodge, Peterson 64-72* in 1964 (FPF);

Kaibab Plateau, Richards in 1938 (UC) ; 4 mi. E of

Grandview, Grand Can., Newlon 856 in 1922

(JEPS) ; Museum of Northern Arizona, N of Flagstaff, Heiser 848 in 1944 (WTU); Grand Can.,

Toumey 8 in 1893 (UC); 5 mi. E of Woods Can.

Lake, Mason, Mason & Phillips 2422 in 2422a in

1964 (ARIZ); Oak Gr., Goodding 17 in 1916 (ARIZ);

Observatory Hill, Flagstaff, Thornher in 1930

(ARIZ); Hualapai Indian Res., ca. 5 mi. NW of

Frazier's Well, Truesdell in 1965 (FPF); Coconino

Rim rd., 1 mi. SE of Sitgreaves Nat. For. bdy.,

L 64-15 in 1964 (FPF) and 0.5 mi. N of Jet. with

Payson-Heber rd., L 64-17 in 1964 (FPF); 16 mi.

S of Happy Jack, L 64-13 in 1964 (FPF). GILA

CO.: Final Mtns.: Ice House Can., Kirhy & Long

FP 18735 in 1914 (FPF) and Kirhy FP 12673 in

1914 (FPF); Bob Tail Ridge, Gill FP 68295 in 1934

(FPF); 5 mi. above Sulphide del Rey, Gill & L

63-51 in 1963 (FPF). 21 mi. E of Payson (3 mi. E

of Indian Gardens), L 63-54 in 1963 (FPF); 2 mi.

W of Pine, W 2705 in 1960 (RSA) ; 17 mi. NE of

Young, L & Lampi 65-16 in 1965 (FPF). Sierra

Ancha: Aztec Peak, Johnson n.d. (ARIZ); Sierra

Ancha Summit, between Roosevelt and Pleasant

Valley, Harrison, Kearney & Peebles 5979 in 1929

(ARIZ, US); Baker Mtn., Harrison 7868 in 1931

(ARIZ); Sierra Ancha, Peebles & Smith 13264 in

1937 (ARIZ, US); Sierra Ancha, Workman Gr.,

L & Lampi 65-14 in 1965 (FPF). GRAHAM CO.:

Graham Mtns.: Pine Crest, Maguire 12138 in 1935

(RM); 1 mi. above Arcadia Guard Sta., H & L

140b in 1962 (FPF) and on P. ponderosa var. arizonica, H & L 140a in 1962 (FPF); Tripp Can.,

8 mi. S of Nat. For. bdy., H &LI46 in 1962 (FPF) ;

Hospital Flat, H & L 227 in 1962 (FPF) and Gill

FP 68309 in 1934 (FPF) ; Columbine, Shreve 5239

in 1917 (DS); Swift Trail, Peebles 12981 in 1936

(ARIZ, US). San Carlos Apache Res., Malay Gap,

H & L 1115 in 1968 (FPF). GREENLEE CO.: 3

mi. N of Nat. For. bdy. on US 666 (17 mi. N of

Clifton), H & L 136 in 1962 (FPF); 47 mi. S of

Springerville on US 666, Gill FP 68304 in 1934

(FPF); 10 mi. S of Mogollón Rim on US 666, L

64-22 in 1964 (FPF). MOHAVE CO.: Hualapai

223
Mtn. Park, Gill & Andreios in 1959 (FPF), W 3029

in 1962 (COLO), and H 715 in 1965 (FPF) ; Hualapai

Mtn., Kearney & Peebles 12721^ in 1935 (ARIZ,

US). NAVAJO CO.: 1 mi. W of Show Low on Rte.

60, L 63-45 in 1963 (FPF); 4 mi. E of Rte. 160 on

Mogollón Rim rd., H & L 919 in 1966 (FPF); near

Deer Springs, 26 mi. E of Rte. 160 on Mogollón

Rim rd., H & L 921 in 1966 (FPF). PIMA CO.:

Santa Catalina Mtns.: Loran in 1881 (UC) and

Thornher in 1930 (ARIZ); Soldier Camp, Gill FP

68129 and FP 68251 in 1932 (FPF), and on P.

ponderosa var. arizonica, Hedgcock & Long FP 9759

in 1911 (FPF); Summerhaven, Loomis, Peebles &

Harrison 2179 in 1926 (ARIZ) and H & L 168 in

1962 (FPF); Mt. Lemmon, Nichol in 1925 (ARIZ);

Palisade Ranger Sta., H & L 172 in 1962 (FPF)

and on P. ponderosa var. arizonica, H & L 171 in

1962 (FPF); Mt. Lemmon Ski Lodge, H & L 169

in 1962 (FPF); Summit of Marshall Can., Gill FP

68131 in 1932 (FPF); 1 mi. S of Rose Lake Jet.,

on P. ponderosa var. arizonica, H & L 165 in 1962

(FPF); Sabino Can., on P. ponderosa var. arizonica,

Hedgcock & Long FP 9781 in 1911 (FPF, ILL).

Rincón Mtns., Mt. Oeboa, on P. ponderosa var.

arizonica, Blumer 3563 in 1909 (ARIZ, DS, FPF,

ILL); Mica Mt., on P. ponderosa var. arizonica, Ela

in 1968 (FPF). SANTA CRUZ CO.: Santa Rita

Mtns., Mt. Wrightson trail: 4 mi. from end of rd.,

H 795 in 1965 (FPF); 1.5 mi. from end of rd., on

P. ponderosa var. arizonica, H 797 in 1965 (FPF).

YAVAPAI CO.: Groom Cr., H & L 177 in 1962

(FPF) and Taylor FP 186^1 in 1914 (FPF); Mingus

Mtn., ca. 2 mi. S of US 89A, H & L 243 in 1962

(FPF); Beaver Cr. Ranger Sta. Drake FP 15119 in

1914 (FPF); Crown King, Ancona & Long 12829 in

1914 (FPF); Horsethief Basin, 7 mi. S of Crown

King, H & Laut 1180 in 1969 (FPF); Prescott,

Hedgcock FP 4856 in 1910 (FPF, ILL); near Prescott, Hedgcock FP 4854 and FP 4855 in 1910 (FPF) ;

Copper Basin, Tourney 292 in 1892 (ARIZ); Santa

Maria Mtns., 5 air mi. WNW of Camp Wood, H &

Laut 1179 in 1969 (FPF).

COLORADO

ARCHULETA CO. : Pagosa Springs, Bethel in 1897

(CS) and Smith in 1894 (PH); 20 mi. S of Pagosa

Springs on US M, H & L 1051 in 1967 (FPF) ; S of

Pagosa Springs, Hedgcock & Bethel FP 24656 in 1917

(FPF); 10 mi. W of Pagosa Springs, Hinds 63-5 in

1963 (FPF). BOULDER CO.: Sugarloaf Mtn.,

Ramaley & Bobbins 1784 in 1906 (COLO, RM);

between Sunset and Ward, Tweedy 4973 in 1902

(RM); Mt. Alto, Ramaley 824 m 1901 (COLO,

RM) ; S Boulder Peak, Ewan 12721 in 1941 (COLO) ;

Tucker's Meadow, Wolcott n.d. (COLO); Lyons,

Thomson FP 26558 in 1917 (FPF); SW of Boulder,

Hedgcock FP 22571 in 1916 (FPF, UC); 1 mi. E of

Rte. 160 on Left Hand Can. rd., H 203 in 1962

(FPF) and on P. contorta subsp. latifolia, H 204 in

1962 (FPF); Boulder, Hedgcock, Thompson & Gayman FP 24755 in 1917 (FPF) ; near Boulder, W 2802

in 1961 (COLO); between Nederland and Rollinsville, Jones 23161 in 1962 (ILL); 5 mi. N of Nederland on Rte. 160, on P. flexilis, H 202 in 1958

(FPF); Ridge S of Left Hand Can., 2 mi. SE of

Ward, on Pinus flexilis, Sprakling & Plumb in 1966

(FPF). CLEAR CREEK CO.: Mill Cr., Brookvale,

Churchill in 1918 (RM); 0.5 mi. S of divide between

Idaho Springs and Central City, H & Staley 280 in

1962 (FPF); Empire, Weir 6163 in 1918 (ILL);

Hamlin Gulch rd., 1.4 mi. from Fall River rd., 7 air

mi. NW of Idaho Springs, H 1056 in 1967 (FPF).

CONEJOS CO.: River Springs Ranger Sta., ca. 15

mi. W of Antonito, Hinds 63-3 in 1963 (FPF); 3.1

mi. W of Nat. For. bdy. on Rte. 17, L 63-1 in 1963

(FPF). COSTILLA CO.: Costilla Estate at New

Mexico bdy., Edmonston FP 1511 in 1908 (FPF).

CUSTER CO.: Greenwood, Edmonston FP 561 in

1909 (FPF); Alvarado Campground, 9 mi. SW of

Westcliffe, H 433 in 1963 (FPF) ; 4.5 mi. W of Hillside, H 435 in 1963 (FPF); 3.5 mi. W of Hillside on

Spruce Cr. rd., on P. contorta subsp. latifolia, H 436

in 1963 (FPF); Westcliffe, Clay Ranger Sta., Hedgcock, Thompson & Johnson FP 24828 in 1917 (FPF).

DENVER CO.: Denver, Case Golf Course, H &

Stewart 832 in 1965 (FPF). DOLORES CO.: Doe

Springs rd., T. 40 N, R. 17 W, Sec. 25, [12 mi. ESE

of Dove Cr.], Hinds 63-11 in 1963 (FPF). DOUGLAS CO. : 1.5 mi. E of Franktown on Rte. m, H 544

in 1963 (FPF); Missouri Ridge rd.. Manitou Exp.

For., H 257 in 1962 (FPF). ELBERT CO.: 5.5 mi.

S of Elbert on Rte. 157, H 420 in 1963 (FPF) ; 6 mi.

W of Kiowa on Rte. 86, H 543 in 1963 (FPF). EL

PASO CO.: Cheyenne Can., Christ 1864 in 1935

(CS); Colorado Springs, Jones 801 in 1878 (ARIZ,

POM, RM); 14.5 mi. E of Monument, H 419 in

1963 (FPF); 1.5 mi. E of Monument, H 542 in

1963 (FPF), 1.7 mi. from Cascade on Pikes Peak

rd., H & Hinds 426 in 1963 (FPF); Bruin Inn, N

Cheyenne Can., H 539"" in 1963 (FPF); Colorado

Springs, Saunders in 1893 (RM) ; Pikes Peak, Halfway, Hedgcock FP 903 in 1909 (FPF) ; Mt. Manitou,

Gill FP 68247 in 1932 (FPF) ; Palmer Park, Austin

Bluff, Gill FP 68248 in 1932 (FPF) ; NE of Manitou,

Hedgcock FP 22710 and FP 22711 in 1916 (FPF);

5 mi. W of Monument, W 2983 in 1962 (COLO,

FPF). Monument; Hedgcock FP 670 in 1909 (FPF),

Hartley FP 1613 in 1909 (FPF), Hedgcock <Sc Pierce

FP 15951 in 1914 (FPF, ILL, RSA, UC, WTU),

Hedgcock FP 22550 in 1916 (FPF, UC), and Hedgcock FP 24909 in 1917 (FPF). Fremont Exp. For.

[2 mi. W of Manitou Springs], Hedgcock FP 9174

and FP 9175 in 1911 (FPF) and FP 22704 and FP

22705 in 1916 (FPF); Palmer Lake, Hedgcock FP

224
15898 in 1914 (FPF, UC), Hedgcock, Thompson &

Gayman FP 2J^902 in 1917 (FPF), and Gill FP

682U in 1932 (FPF). FREMONT CO.: '^Canyon

City/' Sierra Majada, Brandegee 11295 in 1877

(FPF, ILL). GILPIN CO.: Rollinsville, Hedgcock

FP 22599 in 1916 (FPF). HINSDALE CO.: Piedra

River, T. 37 N, R. 3 W, Sec. 14, ca. 15 air mi. NW

of Pagosa Springs, Peterson 19-61 in 1961 (FPF).

HUÉRFANO CO. : 10 mi. N of Gardner on Ophir

Cr. rd., H 431 in 1963 (FPF); La Veta, Hedgcock &

Thompson FP 24809 in 1917 (FPF). JEFFERSON

CO.: Gunbarrel Cr., Hill FP 68278 in 1932 (FPF);

10 mi. W of Deckers, H 421 in 1963 (FPF) ; 8 mi.

SW of Morrison on US 285, H 453 in 1963 (FPF);

Buffalo Cr., Hill FP 68277 in 1932 (FPF); near

Golden, Weir 8958 in 1918 (ILL). LA PLATA CO.:

Tacoma Power Plant, on Pinus flexilis, Loughridge

273 in 1934 (CS) ; W of Trimble, Hedgcock & Bethel

FP 24688 in 1917 (FPF, UC) ; 10 mi. W of Durango

on US 160, H & Scharpf 690 in 1964 (FPF). LARIMER CO. : Estes Park, Cooper 243 in 1904 (ARIZ,

RM); Prospect Mtn. near Estes Park, Maize in

1966 (FPF); ^Tt. Collins,^' Buffum 5690 in 1896

(RM); ''Mountains of Larimer Co.,'' Crandal in

1889 (RM); 4 mi. SW of Log Cabin, H 200 in 1960

(FPF); Rist Can. Picnic Ground, H 201 in 1961

(FPF) ; near Cherokee Park (lat. 40° 53'N-, northern

limit?), H 262 in 1962 (FPF); Poudre River 4 mi.

E of Rustic, H 410 in 1963 (FPF) ; Stove Prairie

Hill, col.? Plants of Colo. 2274 in 1896 (RM); Skyland Ranch, Rocky Mtn. Nat. Park, Nelson 3205

in 1938 (RM); Rist Can., Cameron, PL Colo. 2279

in 1897 (CS); Brinwood, Stove Prairie, col? PI.

Colo. 2277 in 1898 (FPF) ; 1 mi. SW of Glen Haven,

H & Staley 287 in 1962 (FPF) ; Assoc. Camp, Rocky

Mtn. Park, Case FP 38702 in 1922 (FPF) ; Rattlesnake (Pinewood) Park, H 87 in 1961 (FPF); Rist

Can., 7.2 mi. W of La Porte, H 552 in 1964 (FPF);

Rist Can., near Picnic Ground, on P. contorta subsp.

latifolia, H 199 in 1961 (FPF); Stove Prairie Hill,

Pennock, on P. contorta subsp. latifolia, PI. Colo. 899

in 1892 (CS) ; 1 mi. E of Pennock Pass, on P. contorta subsp. latifolia, H, Hinds & Staley 281 in 1962

(FPF) and H 560 in 1964 (FPF); N of Stratton

Park, 13 mi. WNW of Fort ColHns, on P. contorta

subsp. latifolia, H 709 in 1962 (FPF) ; Bear Gulch,

5 mi. W of Masonville, on P. contorta subsp. latifolia,

H 710 in 1964 (FPF). LAS ANIMAS CO.: Mesa de

Maya, Green Can. Rogers 6089 in 1948 (COLO,

US) ; Stonewall Gap, Hedgcock & Johnson FP 24944

in 1917 (FPF); Monument Park, H 1096 in 1968

(FPF). MESA CO.: Sawmill Mesa rd., 20 mi. SW

of Delta, Hinds 63-19 in 1963 (FPF). MONTEZUMA CO.: Water Can., T. 38 N, R. 13 W, Sec.

30, ca. 10 mi. N of Mancos, Hinds 63-8 in 1963

(FPF). MONTROSE CO.: 10 mi. W of Paradox,

Harrington 4392 in 1949 (CS); Hanks Valley rd.,

T. 46 N, R. 11 W, Sec. 18, ca. 10 mi. NE of Norwood,

Hinds 63-16 in 1963 (FPF). PARK CO.: 1.5 mi.

NW of Bailey, H 282 in 1962 (FPF); 1.7 mi. S of

11 Mi. Reservoir rd. on Blue ]\Itn. rd., near Lake

George, H 4^4* in 1963 (FPF). PUEBLO CO. : 5 mi.

W of Rye on Rte. 165, H 427 in 1963 (FPF).

SAGUACHE CO.: Spanish Cr., near N Cochetopa

Pass, H 438 in 1963 (FPF) ; E of North Cochetopa

Pass, on P. contorta subsp. latifolia, H 437 in 1963

(FPF) and on P. aristata, H 439 in 1963 (FPF).

SAN MIGUEL CO.: Iron Springs Mesa [7 mi. NW

of Placerville], Payson FP 26022 in 1917 (FPF).

TELLER CO.: Florissant, Ramaley 1333 in 1905

(RM); 6.2 mi. S of Divide on Rte. 67, H & Hinds

423 in 1963 (FPF) and on P. aristata, H & Hinds

422 in 1963 (FPF); Elklum (Elkton?), Hedgcock &

Johnston FP 26406 in 1917 (FPF); Woodland Park,

Gill FP 68138 and FP 68246 in 1932 (FPF).

COUNTY UNCERTAIN: Uncompahgre Divide,

Payson 396 in 1914 (COLO, RM) ; Trinchera Estate,

E side San Luis Valley, Edmonston FP 175 in 1908

(FPF); Rocky Ford, Pool FP 20552 in 1912 (FPF).

NEW MEXICO

BERNALILLO CO.: Manzano Mtns.: 12 mi. S of

US 66 on Rte. 10, H 21 in 1954 (FPF). Sandia INItns.,

Doc Long Campground, Gill & Long FP 68250 and

FP 68258 in 1932 (FPF) and Muenscher & Muenscher 14562 in 1939 (MO, UC, WTU). CATRON

CO.: Datil :Mtns., White House Can., 10.5 mi. NW

of Datil on US QO, H & L 1116 in 1968 (FPF) ; 1 mi.

E of Mogollón, H & L 132 in 1962 (FPF); 7 mi. E

of Mogollón, H & L 221 in 1962 (FPF) ; Mogollón

Cr., Woodroio & Long FP 18399 in 1914 (FPF);

Mangas Mtn., H & Scharpf 698 in 1964 (FPF);

Tularosa Summit, 11.5 mi. N of Rte. 78 on Apache

Springs rd., L 65-42 in 1965 (FPF); 5 mi. E of

Beaverhead Ranger Sta. on Rte. 59, H & L 893 in

1966 (FPF) ; Can. Cr. Mtns., 12 air mi. W of Beaverhead Ranger Sta., H & L 895 in 1966 (FPF); 1.5

mi. E of Rte. 78 on Loco Mtn. rd., H & L 896 in

1966 (FPF); 1 mi. E of Fox Mtn., 20 mi. N of

Apache Cr., H & L 930 in 1966 (FPF). COLFAX

CO. : 1 mi. E of Taos Co. line on US 64, L 65-22

in 1965 (FPF). DONA ANA CO.: Organ Mtns.,

Fillmore Can., Wooton in 1903 (FPF). EDDY CO.:

Guadalupe Mtns. : Johnson FP 18638 in 1914 (FPF) ;

Dark Can., H & L 113* in 1962 (FPF). GRANT

CO. : W side Emory Pass, H & L119 in 1962 (FPF) ;

1 mi. NW of Redstone, H & L 126 in 1962 (FPF);

Silver City, Gaetz FP 182 in 1908 (FPF) ; N of Pinos

Altos, Hedgcock FP 813 in 1909 (FPF); 20 mi. N of

Mimbres on Rte. 61, L 65-37 in 1965 (FPF); Burro

Mtns., near summit of Jack's Peak, H & L 1103 in

1968 (FPF); Redstone Cr., on P. ponderosa var.

arizonica. Gill FP 68308 in 1934 (FPF) ; Snow cabin

225
rd., 4 mi. W of Rte. 25, on P. ponderosa var. arizonica, H & L 890 in 1966 (FPF). LINCOLN

CO.: Ruidoso, Fisher & Steyermark 38183 in 1938

(KM); Capitán Mtns., Capitán Gap, H & L 104*

and 108 in 1962 (FPF); Capitán Mtns., west

Capitán Peak, L 66-21 in 1966 (FPF); Capitán

Mtns., T. 7 S, R. 16 E, Parks & Long FP 18383 in

1914 (FPF) and Parks FP 18385 in 1914 (FPF);

Capitán Mtns., E slope of Capitán Peak at Pine

Lodge, Martin 883 in 1945 (WTU) ; Gallinas Peak,

10 mi. W of Corona, L & Riffle 63-^2 in 1963 (FPF) ;

White Mtns., Valley of Carrizo, Wickens 2403 in

1932 (PH); S side of Capitán Mtns., 15 mi. E of

Lincoln, H & Scharpf 695 in 1964 (FPF) ; Jicarilla

Mtns., 7 mi. SE of Ancho on White Oaks rd., L

64-31 in 1964 (FPF) ; Tucson Mtn., NW of Capitán,

L 64-32 in 1964 (FPF). LOS ALAMOS CO.: Bandelier Nat. Mon., L 65-55 in 1965 (FPF). MCKINLEY CO. : 2 mi. W of McGaffey, L 63-37 in 1963

(FPF). MORA CO. : 17 mi. N of Mora on Rte. 38, L

65-53 in 1965 (FPF). OTERO CO. : Avis, Wyman FP

12815 in 1914 (FPF) ; Sierra Blanca Peak, Wolf 2880

in 1928 (DS, RSA); Sacramento Mtns., James Can.,

Wickens 2199 in 1932 (PH) ; Cloudcroft, H & W 454

in 1963 (FPF). Mescalero Apache Res. : Carrizo, H &

L 103 in 1962 (FPF); PauFs Can., H 24 in 1954

(FPF) Goat Can., H FP 98511 in 1952 (FPF) ; above

Ruidoso, Hinckley in 1963 (ARIZ) ; Whitetail area,

on P. strohiformis, L 64-2 in 1964 (FPF). 13 mi. SE

of Cloudcroft on Rte. 23, H & L 879 in 1966 (FPF),

Tall Pines Camp, 2 mi. NW of Weed, H & L 884

in 1966 (FPF), Upper Peñasco Can., 13 mi. S of

Cloudcroft, H &L885m 1966 (FPF). RIO ARRIBA

CO.: Chama, Hedgcock & Bethel FP 29454 in 1917

(FPF); 12 mi. NW of Jet. of Lagunitas rd. and Rte.

285, L 63-13a in 1963 (FPF); 24 mi. E of Canjilon,

L 63-9 in 1963 (FPF); 2 mi. S of Tierra Amarilla,

L 63-4 in 1963 (FPF) ; 3 mi. SW of Tres Piedras,

L 63-25 in 1963 (FPF) ; 2.3 mi. N of Gallina, L 63-21

in 1963 (FPF) ; 2 mi. W of Tres Piedras, W 2986

in 1962 (COLO, FPF) ; 1 mi. N of S Jicarilla Res.

bdy. on Rte. 95, L 64-7 in 1964 (FPF) ; 18 mi. E of

CO. bdy. and 12 mi. S of Rte. 17, L 64-8 in 1964

(FPF). SAN JUAN CO.: Chuska Mtns., 6 mi. SW

of Sheep Springs, L & Weiss 66-42 in 1966 (FPF).

SAN MIGUEL CO. : Windsor's Ranch, Pecos Nat.

For. Standley 4173 in 1908 (FPF); Glorieta Mesa,

3.5 mi. S of Rowe, H 1097 in 1968 (FPF). SANTA

FE CO.: Sante Fe, Hedgcock & Bethel FP 24712 in

1917 (FPF, ILL, UC); Borrega area E of Española,

L 63-27 in 1963 (FPF); Cañoncito, Brandegee in

1879 (UC); Can. 4 mi. E of Santa Fe, Heller 3608

in 1897 (ARIZ, Z) ; near Santa Fe, W 2988 in 1962

(COLO). SIERRA CO.: Kingston, Metcalfe 964 in

1904 (FPF, POM, UC) ; near Boiler Peak on Rte.

59, 15 mi. E of Beaverhead Ranger Sta., H & L 894

in 1966 (FPF). SOCORRO CO.: Magdalena Mtns.,

Mouth of Hop Can., Diehl 878 in 1903 (POM);

San Mateo Mtns. : N side of San Juan Peak, L 65-32

in 1965 (FPF) and 11.5 mi. S of US 60 on Rte. 52,

L 65-35 in 1965 (FPF). TAGS CO.: 4 mi. E of

Questa on Rte. 38, L 65-47 in 1965 (FPF) ; 10 mi.

S of Rancho de Taos on Rte. 3, H 877 in 1966

(FPF). TORRANCE CO.: 5 mi. N of Tajique, L &

Riffle 63-56 in 1963 (FPF). VALENCIA CO.: Ojo

Redondo, T. 11 N, R. 12 W, L 63-38 in 1963 (FPF) ;

S of San Mateo, Mt. Taylor, L 63-32 in 1963 (FPF) ;

14 mi. E of Grants on rd. to La Mosca Lookout,

L 65-51 in 1965 (FPF). COUNTY UNCERTAIN:

Front Cr. Basin, Gila Nat. For., Hedgcock & Long

FP 9835 in 1911 (FPF); Gila Nat. For., Munro FP

15114ml^l4: (FPF).

TEXAS

CULBERSON CO.: Guadalupe Mtns.: Johnston

3185 in 1958 (SRSC) ; Ridge above McKittrick Can.,

Moore & Steyermark 3470 in 1931 (DS, PH, UC,

US) ; the Bowl, Warnock & McVaugh 5452 in 1947

(SRSC); above Frijole, McVaugh 8156 in 1947 (DS,

SRSC). JEFF DAVIS CO.: Davis Mtns.: Upper

Madera Can., Palmer 33429 in 1928 (PH); Mt.

Livermore, Hinckley in 1937 (ARIZ), SR), Hinckley

2650 in 1943 (SRSC) and Warnock in 1936 (SRSC) ;

N slope of ^It. Livermore, H, L & Lampi 1046"^

in 1967 (FPF).

UTAH

GARFIELD CO.: Bryce Can. Nat. Park: Bryce

Point, W 3035 in 1962 (COLO, FPF) and :Mistletoe

Ridge, Buchanan 99 in 1956 (ARIZ). Ruby's Inn,

Harris C2912813 in 1929 (UC); Aquarius Plateau,

Vickery 616 in 1960 (RSA, UT) ; 20 mi. N of Boulder

on Rte. 117, H 684 in 1964 (FPF). KANE CO.:

N of Navajo Lake, Peterson 49-61 in 1961 (FPF);

8 mi. S of Navajo Lake on Zion rd., W 4123 in 1966

(FPF, UT). SAN JUAN CO. : Elk Mtns. : Rydberg &

Garrett 9328 in 1911 (RM, US); Elk Ridge, 2 mi. E

of Kigalia Guard Sta., Peterson 65-74 in 1965 (FPF).

La Sal Mtns., Pine Ridge near Rte. 46, Peterson

65-83 in 1965 (FPF). Abajo Mtns., S side of Abajo

Mtn., Krebill 235 in 1964 (FPF). SEVIER CO.:

Wildcat Knolls, W of Emery, Peterson 64-96* in

1964 (FPF); Emery, Williams in 1918 (FPF, ILL).

WAYNE CO.: Near Grover, Peterson 64-93* in

1964 (FPF) ; Wayne Wonderland, Milner 7250a in

1935 (UT). COUNTY UNCERTAIN: La Sal Mtns.,

Stithem in 1960 (FPF); Muddy Watershed, Manti

Nat. For., Korstian in 1918 (FPF, ILL, WTU).

226
23c. ARCEUTHOBIUM VAGINATUM

subsp. DURANGENSE

Specimens Examined :

MEXICO

DURANGO: 36 mi. W of El Salto on Rte. 40, on

Pinus durangensis, H & W 353* and 527* in 1963

(COLO, FPF) ; 45 mi. W of El Salto on Rte. 40, on

P. montezumae, H &W 354* in 1963 (COLO, FPF)

SLudH & W 1237 in 1969 (FPF). Km. 1148, 50 mi.

E of Rte. 15 on Rte. 40, on P. herrerai, H &W 1249

in 1969 (FPF). SINALOA: Km. 1179, 46 mi. E of

Rte. 15 on Rte. 40, on P. montezumae, H & W 1234

in 1969 (FPF).

KASHMIR

Nanga Parbat, near Rama, Wehster & Nasir 6471

in 1955 (K, US); Baltal, Steivart 7549 in 1922 (K,

PH); Pahalagon, Singh in 1965 (FPF); Sonamarg,

Stewart in 1921 (MO) ; Baltal to Sonamarg, Stewart

21297 in 1940 (US) ; Kishenganga Valley, W. Gurais,

col? in 1892 (BM, K); Gurais Valley, Duthie 14182

in 1893 (BM); Sonamarg in Sind Valley, Duthie

I4I8I in 1892 (K); Kali River, Nabi, Duthie 24933

in 1900 (K) ; Guohai Valley, Astor District, Duthie

12292 in 1892 (K); 0.5 mi. S of Sonamarg, 59 mi.

NE of Srinigar, W &W 4S58 in 1968 (FPF, UT);

0.5 mi. above Tangmarg, 24 mi. W of Srinigar,

W & W 4359 in 1968 (FPF, UT); above Sumblili,

Duthie 11204 in 1892 (K).

24. ARCEUTHOBIUM VERTICILLIFLORUM

Specimens Examined:

MEXICO

DURANGO: 36 mi. E of El Salto on Rte. 40, on

P. engelma,nnii, H & W 331* and 517* in 1963

(COLO, FPF) and I24I in 1969 (FPF); 24 mi. E of

El Salto on Rte. 40, on P. engelmannii, H & W 34I*

in 1963 (COLO, FPF) ; 7 mi. E of El Salto on Rte.

40, on P. cooperi, H & W 342* in 1962 (COLO,

FPF).

Old World Taxa

25. ARCEUTHOBIUM CHÍNENSE

Specimens Examined :

CHINA

YUNNAN: W of von Fumin, vicinity of Yanggai

and Dsolin-ho, on Keteleeria davidiana, HandelMazzetti 4^99 in 1914 (US); around Hoji between

Hoching and Sunygueh, on Keteleeria sp., Schneider

2951 in 1914 (K, US); West flank of Tali Range,

Lat. 25° 40' N, on ^^conifers,'' Forrest 15557 in 1917

(K). SZECHWAN: vicinity of Huei-li Hsien, on

Keteleeria sp., Yü 1558 in 1932 (GH).

26. ARCEUTHOBIUM MINUTISSIMUM

Specimens Examined : All on Pinus griffithii.

WEST PAKISTAN

SWAT: 25 mi. above Bahrein, E of Kalam, Rodin

5695 in 1952 (K, UC, US) ; 4 mi. W of Utrot, Kalam

Agency, Thatcher in 1962 (FPF); above Utrot,

Stewart & Rahman 25213 in 1953 (BM).

INDIA

HIMACHAL PRADESH: Sangla-Rupin rd., Parker

in 1928 (UC). UTTAR PRADESH: Gangotri,

Dudgeon & Kenoyer 132-2-1 in 1920 (PH). PRADESH UNCERTAIN: Gangrinini, Dudgeon &

Kenoyer 561 in 1920 (MO).

NEPAL

Nampa Gadh, Western Nepal, Duthie 5947 in 1886

(BM, K) ; Kali Valley near Kangua, Western Nepal,

Duthie 3359 in 1886 (BM, K) ; Near Tinkar, Parker

2085 m 1923 (K).

27. ARCEUTHOBIUM OXYCEDRI

No specimens are cited for this taxon because (1)

it is poorly represented in North American herbaria

and (2) its distribution is fairly well known (see p.

169).

28. ARCEUTHOBIUM PINI

Specimens Examined :

CHINA

YUNNAN: E flank of Lichiang Range, Lat. 27°

30' N, on Pinus, Forrest 6672 in 1900 (K) ; MekongSalwin divide; Lat. 28° 12' N, on Pinus, Forrest

14194 in 1917 (K). SZECHWAN: Between Molien

and Tialou beyond des Yalung, on Pinus tabulaeformis, Handel-Mazzetti 2629 in 1914 (US). TIBET.

SE Tibet: Kongbo Prov. Charko-Lilung, Valley of

Lilung Chu, Lat. 29° 08' N, Long. 93° 54' E, on

Pinus sp., Ludlow, Sherriff and Taylor 44^5 in 1938

(BM); Tongyuk, on Pinus sp., Kingdon-Ward

12089 in 1935 (BM).

227
EXSICCATAE. INDEX TO NUMBERED COLLECTIONS

The first number is the collector's number for the

specimen followed by the taxon number (1 through

28) in parentheses indicating our determination of it.

Abrams, 2818(8), 6297(18), 7660(18), 8677(8), 12130(lâ),

12752(8), 12753(S).

Alexander 1199(13).

Alexander & Hernandez 2333(2Ssi).

Alexander & Kellogg 5328(10).

Allard 17432(5).

AWen 303(22).

Ancona & Long FP 12829(2dh).

Anderson 232(22), 265(19), 6406(22).

Andresen & Andresen 1804(12si), 1824(10).

Andresen & Cochrane 1729(9).

Andresen & Miles 631(4).

Andresen & Simpson 1787(4).

Andresen & Steinhoff A 1983(13), A 2008(13), A 2035(13),

A 2047(Q), A 2050(23SL).

Andrews, Ellis & Gill FP 89229(11).

Applegate 2478(3), 6038(22).

Archer 6396(8).

Armstrong 1024(10).

Ashton 66(9).

Bacigalupi 1808(18).

Baker 209(8), 1006(8).

Baker, Earle & Tracy 94(10), 388(11).

Balls B 4086(23Q), B 4089(13), B 4228(2), B 4399(13),

23608(18).

BsLrhe 314(8).

Bsirker 254(22).

Barkley 1999(11).

Barkley & Barkley 3504(11), 3505(3).

Barlow 145(19).

Batram 455(10).

BQQ\ 660(10).

Beaman ^^^4(13), ^44i(23a).

Bean ^^4^(19).

Beane ^^5(18).

Beane & James 2079(18).

Beattie 5281(18), 5801(22).

Beàv^oil FP 91035(8).

Bedwell & Childs FP 91033(8), FP 91034(8).

Benson 47(18), 726(18), 4907(3), 5705(8).

Bethel 317(3), FP 24449(8), FP 26266(18), FP 26279(18),

FP 27111(3).

Bethel & Hunt FP 29261(10).

Bethel, Schmoll & Clokey 4314(9), 4315(9).

Bigelow 4(18).

Blakely 57S'5(la).

Blankinship 436(9).

Blumer 110(10), 1452(23h), 1515(12Q), 1557(23h), 2394(12a)

3362(12Q), 3563(23h), 5177(3).

Bolander 2671(18), 5095(3), 6380(8).

Bookman FP 68265(22).

Borglav 1115(13).

Bostick 5^57(10).

Boyce 30(lh), 31(7), 32(7), 33(18), 40(3), 596(22), 608(22),

703(3), 1225(16), 1905(lsi), 1906(ls¿), 1907(8), 1908(8),

2121(8), 2122(22), 2298(22), 2299(22), 2300(22), FP

Boyce (continued)

15981(l2i), FP 15982(lh), FP 15983(8), FP 40057(22),

FP 40065(22), FP 40120(3), FP 40121(11), FP 40165(11),

FP 40179(8), FP 40198(3), FP 97944(S), FP 97953(18),

FP 97955(8).

Brandegee 1069(3), 1070(22), 1071(16), 1226(10), 11295(23h),

12201(3).

Breedlove 1109(12.), 6974(13), 8016(13).

Breedlove & Thorne 18535 (23a), 18617 (12íí).

Breitung 5586(3).

Brewer 582(18), 700(18).

Brown 580(8), 675(U), 794(3), 795(3), 941(8).

Buchanan 99(23h), 310(11), 5^7(10).

Buñnm5690(23h).

Burrall55(23b).

Butler 182(16), 270(1^), 272(9), 273(8).

Cain & Svenson 6597(19).

Calder & Savile 11374(3), 11413(11), 11415(16), 11465(16),

11536(11).

Calder, Parmelee & Taylor 16644('à), 19159(3).

Calder, Savile & Ferguson 14363(3), 15273(3).

Cameron PI. Colo. 2279 (23b).

Campagna 54^75(19).

Cantielow 1659(8).

Csise FP 38702(23h).

Chsinser 6022(18).

Childs 10(8), 14(^), 22(8), 27(16), 28(lsi), 52(8), 53(8), 57(11),

61(8), 65(22), 67(3), 68(8), 212(16), 245(3), FP 40545(22),

FP 68272(8), FP 91560(22), FP 91561(22), FP 91562(22),

FP 91575(8), FP 91615(8).

Childs & Hansbrough FP 68261(11), FP 68268(16).

Christ Í504('3D), 1877(9).

Churchill 55i(19).

Clark & Seymour G 586(19).

Clausen i777(8).

Clokey 2684(3), 5429(9), 5430(9), 5431(8), 5432(8), 5433(U),

5434(1^), 5492(9), 8565(10).

Clokey & Anderson 6574(8).

Cody et al. 676(19).

Conzatti 4123(12h).

Cook 7(13).

Cooke 11576(9), 11593(8), 13574(1Q), 13920(11), 16034(3),

16132(1Q), 16212(9), 17722(10), 17729(11).

Cooper 23(3), 65(9), 1064(3), 243(23b), 1547(8).

Cooper & Silva 923(7).

Copeland 4^4(la), 1425(8).

Cotia^m 11561(19).

Coulter ^7(23a), 7655(3).

Coville 1478(22), 1881(16), 2363(16), 2459(16), 2478a(ll),

2480(16), 2487(3), 2488(3), 2489(3), 2490(3), 2491(3).

Coville & Applegate 1090(22), 1091(22), 1156(22).

Coville & Funston 308(10), 311(9), 1073(18), 1460(8), 1482(8),

1596(3).

Coville & Oilman ^^0(10).

Coville & Kearney 916(22).

Cowles 57^(22).

Cox FP 68228(3).

Cronquist 1613(3), 2829(3).

Cusick 2324(8), 2691(3), 2701(8).

Cutler 2047(10).

Darlington ^7^^(19).

228
Darrow, Phillips & Pultz 1173(17).

Davidson & Gill FP 89991 (S).

Davy 7064{IS).

Detting 4^55(8).

Diehl575(23b).

Doherty FP 15240(2Sh).

Drake FP 15119{2Sh), FP 15122{n).

Dudgeon & Kenoyer 561{2Q), 132-2-1(26).

Dudley 755(8), 1395(lh), 1978(S).

Dudley & Lamb 44^(18).

Duncan 90(1S).

Duthie 3359(26), 59^7(26), 11204(26), 12292(26), 14181(26),

14182(26), 24933(26).

Eastwood 12138(S), 18836(18).

Eaton 44S(19).

Edmonston FP 51(3), FP 175(2Sh), FP 188(3), FP 199(3),

FP 295(11), FP 561(23h), FP 1511(23h).

Eggelston 1561(19), 1562(19), 7459(3), 9718(8), 10599(3).

Ehlers 1120(19), 5144(19), 6127(19).

Ekman 1313(5), 12024(o).

Ellis FP 89331(10), FP 89337(10), FP 89411(10), FP

89426(10), FP 89430(17), FP 89431 (^), FP 89502(10)

FP 89503(10), FP 89504(11).

Ellis & Gill FP 89500(12si).

Ellis, Hackleman & Gill FP 89491(11).

Elmer 1246(S), 4031(18).

Englevth FP 91031 (Isi).

Erdman 1^0(10).

Erlanson 484(1^)-

Ethevidge BFDS 26(13).

Evans FP 17832(3), FP 17843(3).

Ewan 3564(7), 9924(8), 9941(10), 10110(7), 10111(8),

12721(23h), 15154(9).

Fassett 2185(19), 2345(19), 2527(19), 3923(19), 8996(19),

10093(19), 13767(19), 14308(19), 22395(19).

Fassett & Sieker 14703(19).

Fassett, Hoffman & Staffeld 18265(19).

Faull, Childs & Hansbrough FP 68266(22).

Fendler 283(23b), 312(10).

Fernald 1741(19), 23322(19).

Fernald & Collins 218(19).

Fernald & Long 13521(19), 23788(19).

Fernald & Pease 21041(19), 21043(19), 23878(19), 25031(19).

Fernald & St. John 10830(19).

Fernald et al. 1655(19), 1656(19), 75^7(19), 21044(19),

21045(19).

Ferris 3649(18), 7828(lsi), 9953(23h), 10183(10), 12158(18).

Ferris & Duthie 4^^(8), 467(3), 857(8).

Yiker 695(8).

Fisher & Steyermark 38183(23h).

Forrest 6672(28), 10169(28), 14194(28), 15557(25).

Frolichöi(19).

Frye & Frye 2590(13).

Fuertes 1923(5).

Gaetz FP 182(23h).

GalhresiihFP 1691(3).

Garrett FP 15833(9), FP 38106(9), FP 38107(9), FP 38174(9).

Gates 11116(19), 14886(19).

Gates & Gates 10682(19).

Gayman FP 26295(3), FP 26521(11).

Gentry 587 M(13).

Geyer 577(8).

Gilbert 112(17).

Gill FP 68036(122.), FP 68037(11), FP 68038(23h), FP

68046(3), FP 68047(23h), FP 68056(11), FP 68060(1^),

FP 68063(3), FP 68076(18), FP 68078(18), FP 68079(18),

FP 68080(18), FP 68087(8), FP 68089(18), FP 68093(18),

FP 68097(8), FP 68099(18), FP 68103(18), FP 68127(11),

FP 68128(4), FP 68129(23h), FP 68130(4), FP 68131 (23h),

FP 68132(10), FP 68134(11), FP 68138(23h), FP 68139(11),

FP 68140(3), FP 68141(11), FP 68143(9), FP 68144(9),

FP 68145(18), FP 68182(22), FP 68183(22), FP 68184(22),

FP 68185(3), FP 68186(12.), FP 68187(22), FP 68188(22),

FP 68189(16), FP 68190(3), FP 68191(8), FP 68192(11),

FP 68193(22), FP 68195(22), FP 68196(22), FP 68198(22),

FP 68200(3), FP 68202(16), FP 68203(8), FP 68205(3),

FP 68206(11), FP 68207(11), FP 68208(19), FP 68209(19),

FP 68223(lh), FP 68224(1^), FP 68227(3), FP 68229(3),

FP 68230(10), FP 68232(10), FP 68233(10), FP 68234(10),

FP 68235(9), FP 68237(8), FP 68243(8), FP 68244(23h),

FP 68246(23h), FP 68247(23h), FP 68248(23b), FP

68249(23h), FP 68251(23h), FP 68252(23h), FP 68253(18),

FP 68255(18), FP 68256(11), FP 68262(11), FP 68263(11),

FP 68290(17), FP 68291(16), FP 68295(23h), FP

68296(23b), FP 68297(U), FP 68298(17), FP 68300(10),

FP 68301(10), FP 68302(17), FP 68303(11), FP 68304(23h),

FP 68305(17), FP 68306(4), FP 68307(11), FP 68308(23h),

FP 68309(23h), FP 68310(17), FP 68313(19), FP 89313(4),

FP 89420(10), FP 89984(3).

Gill & Andrews FP 87301(17).

Gill & Ellis FP 89496(122), FP 89497(11).

Gill, Ellis & Hackleman FP 89492(4), FP 89490(4), FP

89492(4).

Gill, Ellis & Sowell FP 89498(23h), FP 89499(23b).

Gill & ELinds FP 89981(3).

Gill &Lightle(55-5i (23b).

Gill & Long FP 68136(11), FP 68137(10), FP 68250(23h),

FP68258(23h).

Gill & Sargent FP 68179(11), FP 68180(22), FP 68181(lh).

Gill & Wagener FP 68039(8), FP 68040(8), FP 68041(7).

Gill & Warren FP 68197(22), FP 68199(22).

Gill & Wright FP 68061(10), FP 68064(10), FP 68066(18),

FP 68072(8), FP 68160(7), FP 68161 (U), FP 68162(8),

FP 68225(12), FP 68238(10), FP 68239(10), FP 68240(8).

Gill et al. FP 68284(19), FP 68285(19).

Gillis 5007(19).

Glowenke 2468(19).

Goldman Í77(23a), 184(122).

Gonzales 1717(20).

Goodding 17(23h), 309(6), 2088(11), FP 26737(11), FP

33284(11), FP 33285(23h).

Goodman & Payson 2927 (23h).

Gorm2.n 576(3), 1753(9).

Gould 1010(18).

Grant 1113(8), 3197(19).

Green 9(19).

Greene & Richter 112(3), 113(3), 114(3), 115(3), 116(3),

118(3), 119(3), 121(3), 122(3), 123(3), 124(3), 125(3),

126(3).

Greeman & Greeman 4650(3).

Greenma^n 2492(19).

Griffith & Morris 505(8).

Guppy FP 96959(18).

Haberer 188(19).

ÍÍ2II 6642(8).

Hall &Babcock-^075(11).

229
Hall & Chandler J^26{'i).

Hall & Harbour 574(3).

Handel-Mazzettii ^^,^5^(28), 4999(2ö).

Hansbrough FP 68270&).

Hansbrough & Spaulding FP 68210(19).

Hansbrough, Mielke & Joy FP 68273{8).

Hanson 718{1S).

Ranzlik FP 68257(11).

Ravger 45859(19).

Harrington 4392(2Sh).

Harris C 2770B(11), C 16354B(12íí), C 2912813(23h), FP

Harrison 7868(2Sh), 11184(10).

Harrison & Larson 7859(2>).

Harrison, Kearney & Fulton 5979(22>h).

Hartley 3991(19), FP 1613(2Zh), FP 1774(9), FP 17865(11),

FP89795(23si).

Hartman 340(13), 364(12si).

Harvey FP 4159(8).

Hawksworth 9(7), 15(11), 21(23h), 23(11), 24(2Sh), 26(11),

30(U), 35(10), 36(10), 39(11), 44(11), 49(13), 51(13),

56(11), 81(3), 82(9), 84(9), 85(9), 87(23h), 89(3), 189(3),

190(3), 191(3), 192(3), 193(3), 196(9), 199(23b), 200(23h),

201(23h), 203(23h), 204(23h), 205(3), 206(11), 246(10),

252(U), 253(17), 254(1^), 255(9), 256(9), 257(23h), 258(3),

260(11), 261(3), 262(23h), 263(3), 264(3), 265(3), 266(3),

267(3), 268(3), 269(3), 270(3), 271(3), 272(3), 273(3),

274(3), 275(3), 276(3), 277(9), 278(3), 282(23h), 284(9),

285(3), 286(3), 406(3), 407(3), 408(3), 409(3), 410(23h),

411(3), 412(3), 413(3), 414(S), 415(3), 416(3), 417(9),

418(3), 419(23h), 420(23h), 421 (23h), 424(23h), 427(23b),

429(11), 430(11), 431(23h), 432(11), 433(23h), 434(11),

435(23h), 436(23b), 437(23b), 438(23b), 439(23b), 440(11),

444(11), 445(3), 446(11), 447(3), 448(3), 449(3), 450(3),

451(3), 452(3), 453(23b), 538(11), 539(23b), 540(9), 541(11),

542(23b), 543(23b), 544(23b), 545(3), 546(3), 547(3), 548(3),

549(3), 550(3), 551(3), 552(23b), 553(3), 554(3), 555(3),

559(3), 561(3), 562{^)), 563(3), 566(3), 567(11), 568{:3),

569(3), 570(11), 571(9), 572(11), 574(11), 658(\8), 659(8),

660(7), 678(11), 679(9), 680(9), 681(11), 682(10), 684(23b),

685(11), 686(11), 687(19), 688(19), 689(9), ?6'^(23b),

710(23b), 711(3), 712(3), 713(10), 715(23b), 728(8), 730(7),

731(8), 733(10), 734(8), 735(8), 736(8), 738(8), 739(8),

7W(7), 742(8), 745(8), 751(8), 752(8), 795(23b), 796(0),

797(23b), 808(23b), 817(10), 819(17), 820(11), 821(23b),

827(9), 828(3), 829(9), 830(3), 843(18), 844(18), 847(18),

848(8), 876(11), 877(23b), 934(3), 935(18), 936(18), 937(U),

938(lb), 939(3), 945(3), 946(11), 947(3), 967(11), 968(11),

969(10), 970(8), 971(11), 972(3), 973(10), 974(10), 975(11),

976(3), 977(11), 978(8), 979(3), 980(11), 981(10), 982(11),

983(8), 984(11), 985(10), 986(11), 988(3), 989(8), 1011(10),

1052(10), 1055(9), 1056(23b), 1057(9), 1058(9), 1080(11),

1082(17), 1087(10), 1089(10), 1096(23b), 1097(23b),

1140(22), 1141(22), 1142(22), 1143(3), 1144(18), 1188(3),

1250(22), FP 98857(11), FP 89958(11), FP 89992(11)

FP 98511 (23b).

Hawksworth & Bailey 1008(9), 1009(3), 1010(3).

Hawksworth & Baranyay 90(3), 91(3), 92(3).

Hawksworth & Gilbertson 1083(17).

Hawksworth & Gill 194(9), 195(9), 198(9).

Hawksworth & Hawksworth 1146(7), 1147(7).

230

Hawksworth & Hinds 197(9), 422(23b), 423(23h), 425(11),

426(23b), 990(la), 991(22), 992(22), 993(3), 994(7), 996(8),

995(8), 997(11), 998(la), 999(la), 1000(lb), 1001(8),

1002(12.), 1004(3), 1005(3), 1007(3), 1077(10), 1118(3),

1119(3), 1120(3), 1121(3), 1122(11), 1123(11), 1124(3).

Hawksworth & Landgraf 279(9).

Hawksworth & Laut ií(5i(11), 1164(10), 1179(23b), 1180(23b).

Hawksworth & Lightle 101(10), 103(23b), 104(23b), 105(a),

107(4), 109(4), 106(11), 108(23b), 113(23b), 119(23b).

120(11), 121(4), 125(10), 126(23b), 127(11), 131(11),

132(23b), 133(10), 136(23b), 140a(23b), 140b(23b), 142(11),

146(23b), 152(11), 153(122.), 154(23b), 155(23b), 156(4),

157(122i), 159(123), 161(122i), 164(12a), 165(23b), 166(11),

167(4), 168(23b), 169(23b), 171(23b), 172(23b), 177(23b),

183(11), 184(U), 185(23b), 187(10), 188(23h, 207(17),

208(10), 209(11), 210(4), 211(23b), 212(4), 213(11), 216(11),

217(11), 218(11), 219(17), 220(4), 221(23b), 224(17),

225(17), 226(11), 227(23b), 228(4), 233(23b), 235(0),

236(122.), 239(11), 243(23b), 245(10), 247(9), 249(17),

250(11), 251(11), 442(11), 878(11), 879(23b), 880(11),

884(23b), 885(23b), 886(11), 889(10), 890(23b),

891(4), 893(23b), 894(23b), 895(23b), 896(23b), 897(17),

898(11), 899(10), 900(17), 901 (23b), 902(11), 903(23b),

904(10), 905(10), 906(10), 912(23b), 913(11), 914(23b),

915(10), 916(10), 917(23b), 918(11), 919(23b), 920(11),

921 (23b), 922(10), 923(23b), 924(11), 925(17), 926(10),

927(10), 929(4), 930(23b), 1051(23b), 1078(10), 1079(11),

1084(10), 1098(4), 1099(11), 1102(4), 1103(23b), 1105(4),

1107(4), 1109(0), 1111(12^), 1114(10), 1115(23b), 1116

(23b), 1117(4), 1168(10).

Hawksworth & Scharpf 661(8), 662(12.), 663(lb), 664(3),

665(22), 666(22), 667(lsi), 668(8), 670(3), 671(9), 672(10),

674(12.), 675(8), 676(12.), 690(23b), 691(10), 692(10),

694(11), 695(23b), 696(4), 697(11), 698(23b), 699(11),

700(17), 701(10), 702(12.), 703(10), 705(12a), 765(8), 767(8),

783(8), 786(10), 789(8), 849(18), 850(22), 854(18), 855(12.),

856(18), 857(18), 1148(18), 1149(18), 1150(18), 1151(18),

1152(18), 1153(18), 1158(18), 1159(18), 1252(22), 1255(22),

1256(22), 1257(Is), 1261(3), 1262(10), 1264(U), 1265(8),

1268(11), 1269(3), 1270(10), 1271(8), 1272(11), 1273(10).

Hawksworth & Staley 280(23b), 287(23b), 288(9).

Hawksworth & Stewart 832(23b).

Hawksworth & Wicker 949(3), 950(3), 951(3), 952(3), 953(11),

954(3), 955(10), 956(10), 957(10), 958(10), 959(10), 960(10),

961(10), 962(10), 963(10), 964(10), 965(10), 966(8), 1126(3),

1127(3).

Hawksworth & Wiens 283(3), 292(122.), 293(122.), 294(122.),

295(232.), 296(0), 297(122.), 298(222.), 307(12Q), 308(232.),

310(232), 311(232.), 313(232.), 314(0), 329(21), 331(24),

334(12b), 336(12b), 337(12b), 338(20), 339(20), 340(20),

341(24), 342(24), 343(12b), 345(20), 346(20), 348(232.),

349(13), 350(0), 351(13), 353(23^), 354(23^), 361(13),

367(13), 368(13), 369(13), 370(232.), 371(13), 374(23^),

378(232.), 380(232.), 382(12b), 385(12b), 386(12b), 391 (23a),

392(0), 393(232.), 394(2), 395(11), 396(2), 5^7(23a),

398(23^), 454(23b), 460(122i), 461 (23b), 462(23b), 464(23b),

466(13), 471 (23b), 474(123), 475(0), 476(23b), 479(23b),

480(23b), 481(122.), 484(122.), 483(122.), 485(23b), 486(232.),

487(12a), 491(232.), 492(23b), 493(0), 494(23a), 497(12^),

499(122), 500(21), 505(21), 506(21), 507(232.), 508(12b),

509(12b), 510(12b), 511(20), 512(20), 513(20), 514(11),

515(232), 516(0), 517(24), 518(21), 519(12b), 520(12b),

521(20), 524(13), 525(12b), 526(13), 527(23c), 530(0),
Hawksworth & Wiens (continued)

55í(23a), 533il2h), 534{l2h), 535{12h), 536(6), 565(9),

575(9), 576(11), 577(S), 578(16), 579(16), 580(8), 581(S),

582(16), 583(8), 584(11), 585(16), 586(3), 587(11), 588(16),

589(16), 590(11), 591(3), 592(lsi), 593(11), 594(8), 595(8),

596(11), 597(16), 598(16), 599(3), 600(16), 601(22), 602(22),

603(16), 604(22), 605(22), 606(22), 607(22), 608(22),

609(16), 610(11), 611(8), 612(3), 613(U), 614(3), 615(la),

616(11), 617(8), 618(3), 619(8), 620(3), 621(22), 622(22),

623(3), 624(22), 625(22), 626(l2i), 627(8), 628(11), 629(la),

631(lsi), 632(lsi), 633(8), 634(11), 635(U), 636(11), 637(lix),

639(11), 641(lsi), 642(9), 643(U), 644(7), 645(11), 646(8),

648(U), 649(8), 650(lh), 651(3), 652(22), 654(7), 655(8),

656(8), 657(3), 833(10), 835(10), 836(8), 838(3), 839(3),

840(8), 841(18), 842(8), 858(8), 859(18), 860(8), 861(8),

862(7), 863(8), 865(8), 866(8), 867(8), 868(11), 869(8),

870(11), 871(3), 872(8), 873(8), 57^(la), 57^^(8), 1193(b),

1199(b), 1209(15), 1219(13), 1221(U), 1222(14), 1223(13),

1226(14), 1229(2), 1234(23c), 1237(23c), 1242(21), 1244

(12b), 1245(232.), 1246(13), 1248(20), 3339(2), 3359(2),

3404(12h), 3414(13), 3465(21), 3490(20), 3507(23c).

Hawksworth, Andrews & Buchanan 80(3).

Hawksworth, Hinds & Staley 281(23h).

Hawksworth, Lightle & Gilbertson 1110(4).

Hawksworth, Lightle & Lampi 1046(23h), 1050(10).

Hawksworth, Lightle & Laut 1165(10), 1171(10), 1174(10).

Hawksworth, Lightle & Muñoz 1023(23b), 1024(23h),

1026(23b), 1027(11), 1032(4).

Hawksworth, Stewart & Thompson 1128(3), 1129(16),

1130(16), 1131(3), 1132(11), 1133(16), 1134(16), 1135(11),

1136(11), 1137(11), 1138(22), 1139(22).

Hawksworth, Wiens & Molina 1203(15).

Hedgcock FP 172(11), FP 185(11), FP 624(3), FP 635(9),

FP 650(11), FP 670(23h), FP 813(23h), FP 879(3), FP

888(3), FP 903(23h), FP 905(11), FP 988(11), FP 997(16),

FP 998(8), FP 1538(11), FP 1539(11), FP 1614(9), FP

1649(3), FP 1858(lâ), FP 1873(8), FP 1897(8), FP 1908(3),

FP 1909(8), FP 4S38(3), FP 44^7(11), FP 4556(11), FP

4765(22), FP 4835(8), FP 4837(18), FP 4854(23h), FP

4855(23h), FP 4871(11), FP 4907(10), FP 4917(10), FP

8126(11), FP 8135(10), FP 8139(10), FP 8142(10), FP

8640(19), FP 9174(23h), FP 9175(23b), FP 9299(10),

FP 9339(3), FP 9519(11), FP 9526(16), FP 9782(4), FP

9899(11), FP 11147(9), FP 15834(3), FP 15848(3), FP

15849(3), FP 15898(23h), FP 15900(11), FP 15909(9),

FP 15915(11), FP 15965(3), FP 19185(9), FP 22549(3),

FP 22550(23h), FP 22571(23h), FP 22592(3), FP 22598(3),

FP 22599(23h), FP 22704(23b), FP 22705(23h), FP

22706(9), FP 22707(9), FP 22710(23h), FP 22711(23h),

FP 22718(3), FP 22719(3), FP 24909(23h), FP 47157(3),

FP 47529(3), FP 47756(3), FP 48627(8), FP 49442(8), FP

49544(3), FP 54116(3), FP 54990(8), FP 54991(8), FP

59245(3), FP 67206(3), FP 68059(8), FP 93222(11).

Hedgcock & Bethel FP 2441S(23h), FP 24639(11), FP

24654(11), FP 24656(23h), FP 24688(23h), FP 24689(11),

FP 24712(23h), FP 24730(10), FP 29454(23h).

Hedgcock & Freeman FP 4145(19).

Hedgcock & Johnston FP 24944(23h), FP 24945(11), FP

26406(23h), FP 26425(3), FP 26068(3), FP 26433(3).

Hedgcock & Long FP 9759(23h), FP 9781 (23b), FP 9835(23h),

FP 9847(11), FP 9936(10).

Hedgcock & Meinecke FP 4788(8), FP 4789(8), FP 4822(7),

FP 4823(lsi), FP 4827(7), FP 4833(7), FP 4834(lsi), FP

4836(8), FP 4838(7).

Hedgcock & Pierce FP 15951 (23b).

Hedgcock & Thompson FP 24809(23b).

Hedgcock & Weir FP 9442(16), FP 9443(3), FP 9444(16),

FP 9489(3).

Hedgcock, Meinecke & Long FP 9668(8), FP 9669(11).

Hedgcock, Payson & Johnston FP 24852(9).

Hedgcock, Thompson & Gayman FP 24755(23h), FP

24901(11), FP 24902(23h).

Hedgcock, Thompson & Johnson FP 24828(23b).

Hedrick FP 4H10), FP 286(10).

Heiser 848(23h).

Heller 925(8), 3608(23b), 6776(18), 10660(8), 11060(lsi),

11144(IS).

Heller & Heller 3527(11), 3533(10).

Henderson 5^7(8), 2535(16), 2539(3), 3581(3), 11501(3),

14130(8).

Hendrix 616(9).

Henry 1822(22), 1881(22).

Hicks 110(18).

Hill FP 68236(3), FP 68276(9), FP 68277(23h), FP

68278(23h), FP 68279(3), FP 68280(11).

Hinckley 2650(23h), 3163(10).

Hinds 63-2(11), 63-3(23h), 63-4(11), 63-5(23h), 63-6(11),

63-8(23h), 63-10(10), 63-ll{23h), 63-12(11), 63-13(3),

63-14(10), 63-15(11), 63-16(23h), 63-17(10), 63-19(23h),

63-21(3).

mnton 3278(2), 8901(13).

Witchcock 25(10), 559(l3i).

Hitchcock & Martin 7^^5(3).

Holdrige 1328(5).

Holmgren 3748(10).

Hoover 3610(18), 6448(18).

Hopkins 426(3).

Hopping 391(8), FP 97939(22).

House 7^5^(19).

Howard & Howard 8129(5), 9125(5).

Howe &Langíí4^(19).

Howell 386A(8), 929(8), 2459(U), 5618(18), 8135(18),

15674(U), 19686(18), 22902(lh), 24505(10), 27038(3),

30162(18), 33147(18), 34093(8), 34204(8), 37436(18),

38710(18), 38805(7), 38824(8), 38844(lh), 41197(18).

Howell & Barneby 29387(18).

Howell & Eastwood 7204(9).

Hubert FP 91292(8).

Hunt 286(13), FP 25071(8), FP 97958(18).

Hunt & Bethel FP 24428(10), FP 24429(11).

-Hura FP 20583(11).

Iltis 17558(19).

Iltis, Iltis & Flores 1701(13).

Iltis, Iltis & Turner ^^^^5(19).

Jack 3225(19), 3468(19), 3473(19), 3759(19).

Jaenicke FP 21285(lsi).

Jiminez 1069(5).

Johnson 1420(18), FP 15106(10), FP 18638(23h).

Johnston 1688(7), 1720(8), 1777(13), 3185(23h).

Johnston & Mueller 591 (23h).

Johnston & Thompson FP 24881(3), FP 24883(9).

Jones 4^7(23a), 801(23h), 3439(22), 3973(10), 4299(22),

5076(3), 5372(3), 5395(11), 5911c(ll), 9307(11), 9309(16),

9312(16), 9313(16), 9380(11), 23161(23h).

231
Kearney & Peebles 9971(17), 12724{2Sh), 13761(10),

14135(17).

Kiener 5686(3).

Kildare 2634(lsi), 8785(8), 10484(8).

Killip 7816(19).

King & Soderstrom 5174(1S).

Kirby FP l£673(2Sh).

Kirby & Long FP 18735(23h).

Knowlton 40(11), 295(10), 23525(19).

Kors,údin FP 19276(11).

Krebill i^^(ll), 127(3), 133(16), 136(16), 147(16), 154(8),

159(11), 235(2Sh), 244(S), 247(7), 453(11), 288(11), 453(11),

454(9), 538(3).

Krebill & Nelson 467(10), 468(3).

Kuijt 528(3), 535(3), 536(3), 544(16), 546(16), 547(16),

553(16), 556(16), 563(16), 566(16), 581(11), 588(11), 589(3),

591(3), 592(3), 598(3), 602(22), 618(22), 650(11), 1208(18),

1213(18), 1214(18), 1215(18), 1216(1Q), 1253(18), 1254(18),

1257(18), 1272(12.), 1274(18), 1277(8), 1300(18), 1310(18),

1313(18), 1330(3), 1331(3), 1332(3), 1333(8), 1335(3),

1337(8), 1339(8), 1340(1Q), 1341 (ISL), 1343(3), 1344(3),

1345(8), 1346(8), 1347(8), 1348(8), 1350(3), 1352(3),

1354(3), 1355(3), 1356(8), 1357(8), 1358(3), 1361(18),

1362(18), 1365(13.), 1366(19), 1367(11), 1369(12.), 1374(18),

1389(10), 1390(3), 1393(1^), 1394(7), 1401(18), 1407(8),

1410(18), 14îl(^), 1412(3), 1413(10), 1415(9), 1423(8),

1425a(18), 1427(18), 1428(8), 1431(lsi), 1432(8), 1437(18),

1444(IS), 1471(8), 1497(8), 1501(3), 1502(7), 1503(lsi),

1506(18), 1513(3).

Lachmund & Childs FP 68269(22).

LsikelsL 4503(19), 7912(19), 18527(19).

Langman 3713(13).

Leavenworth 710(13).

Leavenworth & Hoogstraal 1126(13).

Lee & Mason 9153(18).

Leiberg 520(3), 5884(9).

Leonard 4690(5).

Lieberg 1672(16), 3312(10), 5363(22).

Lightle 63-1 (23h), 63-2(11), 63-4(23h), 63-6(10), 63-7(11),

63-8(11), 63-9(23h), 63-11(10), 63-13a(23h), 63-13b(10),

63-16(10), 63-18(10), 63-21 (23h), 63-22(11), 63-23(11),

63-24(11), 63-25(23h), 63-27(23h), 63-29(10), 63-31(11),

63-32(23h), 63-34(11), 63-36(10), 63-37(23h), 63-38(23b),

63-39(11), 63-41(10), 63-45(23h), 63-54(23h), 63-55(11),

^4-1(11), ^4-2(23b), 64-7(23h) 64-8(23h), 64-9(10),

64-11(10), 64-12(10), 64-13(23h), 64-14(1^), 64-15(23h),

64-16(11), 64-17(23b), 64-19(10), 64-20(17), 64-21(4),

64:-22(23h), 64-30(10), 64-31(23h), 64-32(23h), 65-1(10),

65-4(10), 65-5(10), 65-6(10), 65-7(11), 65-20(10),

65-21(11), 65-22(23h), 05-23(11), 65-26(12a), 65-28(11),

65-30(10), 65-31(11), ^5-32(23b) 65-33(10), 65-34(11),

65-35(23h), 65-36(4), 65-37(23h), 65-38(10), 65-39(11),

65-40(4.), 65-41(10), 65-42(23h), 65-43(11), 65-44(10),

65-45(10), 65-46(11), 65-47(23h) 65-48(11), 65-49(11),

65-50(10), 65-51(23h), 65-53(23h), 65-55(23h), 66-19(4),

66-20(11), 66-21(23h), 66-22(10).

Lightle & Gill 64-34(10), 64-35(8), 65-24(11), 65-25(11).

Lightle & Lampi 64-15(11), 65-14(23h), 65-16(23h).

Lightle & Riffle 63-42(23h), 63-43(11), 63-56(23h).

Lightle & Schacht 63-26(11).

Lightle & Weiss 66-41(10), 66-42(23h), 66-43(11), 66-44(23h),

66-45(10), 66-47(10), 66-48(11), 66-49(23b), 66-50(11),

66-51 (23b), 66-52(10), 69-4(10).

Lightle, Ela & Lampi 65-54(10).

Linsdale & Linsdale 264(10).

Little & Correll 19033(10).

Loll FP 11832(9).

Loomis, Peebles & Harrison 2179(23h).

Long FP 318(122), FP 19340(11), FP 19634(10), FP

19724(23h), FP 21126(11), FP 21128(4), FP 21671(17).

Long & Fernald 21042(19).

Long and Seay FP 21275(11).

Lory FP 97940(22).

Loughridge 273(23h).

Love & Love 5740(3).

Ludlow, Sherriff & Taylor 4465(28).

Macbride ^(577(9).

Macbride & Payson 3270(3), 3654(3).

MacDougal ^5(23b), 190(10).

McKelvey 1464(10).

Mackenzie 78(3).

McMillan 197(10).

Macoun 79522(11).

McVaugh 8156(23h), 9781(19), 9952(13), 10071(232),

11518(13), 12916(232).

McVaugh & Wilber 10130(2).

Maguire 12138(23h).

Maguire & Becraft 2535a(10).

Maguire & Maguire 20377(9), 20378(9).

Malte 731/29(19).

Marie-Victorin 28456(19).

Marshall 809(19).

Martin 883(23h), 902(4).

Mason 525(18), 1079(3), 2590(8), 3987(18), 4285(18),

4748(10), 5515(18), 5639(18), 5888(18), 7168(18).

Mason, Mason & Phillips 2422(23b), 2422a(23h).

Mathias ^5^(10).

Matthews ^^^5(19).

Matuda 4408(13), 4601(13), 4630(13).

Maysilles 7260(13), 7977(13).

Mearns 4513(3).

Meinecke FP 9059(18), FP 10754(U), FP 17026(18), FP

17053(3), FP 17062(7), FP 17100(8), FP 20151(7), FP

20152(8), FP 97934(3), FP 97938(22), FP 97941(22),

FP 97947(18), FP 97948(8).

Meinecke & Boyce FP 17193(12), FP 17194(12), FP 17196(8).

Metcalfe 288(10), 493(17), 964{23h), 1173(4).

Meyer 688(8), 1685(22).

Meyer & Rogers 2932(232).

Mielke FP 40694(22), FP 58275(22).

Mielke & Ellis FP 89694(11).

Mielke & Zentemyer FP 89238(9).

Milburge 253(8), 1328(8).

Miles FP 15108(8).

Miller FP 68024(10), FP 97956(lh), FP 98023(9), FP

98034(7), FP 98112{lh).

Milner 7250a(23h).

Miranda 114(232), 872(13).

Mitchell FP 68212(8).

Moberg & Gill FP 68299(11).

Molina, Burgur & Wallenta 16483(13).

Moore 143(19), 634(19).

Moore & Steyermark 3470(23h).

Mori 64-86(3).

Morion 2063(19).

Mueller & Mueller 1013(23SL).

232
Muensáier 10431(22).

Muenscher & Muenscher 14562i2Sh).

Muir 64-327(S).

Munns FP 9440(S).

Munro FP 15114{2Sh).

Munz 4^^5(18), 6260(S), 8350(8), 10566(8), 10771(10),

15220(2»), 21347(8).

Munz & Johnston 8576(lsi), 8705(8).

Murdoch 2010(1^), 5301(1^).

Nelson 845(?>), 866(8), 3205(2'¿h), 3320(9), 4887(12SL), 4959(9),

6077(2dsi), 6730(233^), 8247(9).

Nelson & Nelson ^^57(10), 5515(3), 6833(3).

Newcomb 156(lsi), 165(8).

^ewlon856(23b).

Ninman & Thompson FP 38713(19).

O'Bryne & Long FP 18678(10).

Oñorá FP 98091(18).

Olday 240(19).

Oleson 328(3).

Osterhout 2523(3).

Otis 1011(8), 1021(8), 1036(22).

Overbay FP 68281(8), FP 68282(3).

Owenhy 875(3).

Owenby & Brown 2428(8).

Owenbey & Owenbey 2032(3).

Palmer ^(10), 774(21), 33429(23h).

Parish & Parish 966(8), 144^(10).

Parker 2085(26).

Parker & McClintock 7557(11).

Parks FP 18385(23h).

Parks & Long FP 18383(23b).

Parks & Parks 24063(8).

Parks & Tracy 11532(lh).

Parry 86(11), 574(3), 10659(11).

Patterson ^57(3).

Payson 396(23b), FP 26016(10), FP 26022(23h), FP

26026(11).

Payson & Armstrong 3406(9).

Pease 406(19), 3258(19), 19055(19), 20948(19), 32509(19).

Feck9525(lh), 9594(3), 14322(3), 21639(3).

Feeh\esl2981(23h).

Peebles & Smith 13264(23h).

Peebles, Harrison & Peebles 2565(12si).

Peirson 77^0(3), 1721(8), 2322(8), 2818(7), 3251(10), 4302(18).

10136(3).

Peterson 19-61 (23h), 30-61(10), 37-61 (U), 39-61 (la),

43-61(17), 45-61(11), 49-61(23h), 51-61(11), 52-61(9),

126-61(3), 62-4(3), 62-5(3), 62-10(3), 62-11(11), 62-12(11),

62-14(3), 62-16(11), 62-17(3), 62-18(3), 62-28(11), 62-47(3),

62-49(8), 62-50(11), 62-51(3), 62-53(9), 62-60(3), 62-64(11),

62-67(11), 62-75(9), 62-107(10), 62-111(11), 62-145(4),

62-148(120), 62-165(17), 63-87(11), 63-129(3), 63-131(10),

63-134(3), 63-136(18), 63-139(lsi), 63-149(3), 63-156(18),

63-166(8), 63-224(8), 63-225(8), 63-236(16), 63-295(5),

63-315(8), 63-327(10), 63-329(10), 63-337(11), 63-338(10),

63-361(3), 63-364(11), 64-63(11), 64-64(11), 64-72(23b),

64-83(11), 64-87(10), 64-92(11), 64-93(23b), 64-96(23b),

64-100(8), 64-144(8), 64-217(11), 65-69(10), 65-74(23b),

65-76(11), 65-83(23b), 65-88(3), 65-89(3), 65-90(3),

65-116(18), 65-339(9), 65-370(9), 65-395(8), 65-403(8),

68-91(2), 68-105(13), 68-106(23a).

Petty 64-568(3).

Fickle 57-15(8).

Piper 663(22), 1882(10), 3701(3).

Pohl 1145(19).

Pond FP 193(8).

Pool FP 20552(23b).

Porter 8756(3).

Pringle 4368(23^), 4727(13), 13876(12^).

Proctor 10771(5), 10772(5).

Purpus 1778(13), 3836(23si), 6491(2).

Futnsim FP 98054(10).

Quibell 280(8), 1658a(18).

quibeW & QnibeW 2641(3).

Quick 53-27(U), 53-32(7), 53-141(18), 54-87(lb), 60-21(U).

Ramaley 824(23b), 1333(23b), 3754(3).

Ramaley & Robbins 1784(23b).

Raue 50(23b).

Raven 3678(3), 3928(18), 8349(3), 10456(3), 16874(8).

Ray 1565(18).

Reed 1831(3)

Reed & Barkley 4^5^ (23b).

Riley 47-208(3).

Robbins 1403(18).

Rodin 877(3), 5695(20).

Rogers 6089(23b).

Rolland-Germain ^^0(19), 361(19), 3061(19), 7512(19).

Roos ^0^5(10), 1240(8).

Ross 1917(1SL).

Root & Goodding FP 68274(lb), FP 68275(9).

Rose 3460(23a), 3535(230), 60113(8).

Rose & Fitch 17765(10).

Rosendahl <5^ö(22), 1958(22), 4450(19).

Rosendahl & Brandegee 136(22).

Rosendahl & Butters 1092(22).

Rossback 236(18), 237(8).

Rothrock 69(11), 213(8), 4^9(18).

Rouleau Öi^(19).

Rydberg & Bessey 3938(3).

Rydberg & Garitón 7371(10).

Rydberg & Garrett 8542(10), 9328(23b), 9864(11).

St. John 3051(8), 6359(10), 6936(11).

St. John & Nichols 2281(19).

Sandberg, MacDougal & Heller 744(11), 757(16), 892(3).

Sampson & Pearson 21(3), 168(10).

Sandberg 925(8), 9358(3).

Sandberg & Leiberg 593(8), 4443(11).

Scharpf FP 38026(8), FP 38027(22), FP 98107(lb), FP

98114(8).

Schlising 124(19).

Schmoll 498(3).

^ámeiáQv 2951(25).

Schoenfeldt 3600(8).

Schreiber 776(3), 853(18), 891(1Q), 1017(10), 1948(3).

Schwartz FP 68288(19).

Seay FP 33403(3).

Seemann 2138(24).

^Q\QX 2190(13).

Seymour 11906(19), 14645(19), 15104(19).

Sharsmith 1406a & 6(18), 3848h(18).

Shear 57^4(3).

Shear & Bessey 55^7(3).

Sheldon 439(19), 8845(8).

ShreYe5239(23b).

SlosinFP 97943(8).

Smith 955(18).

233
Snell 1047(9).

Solbrig ^4^0(3).

Spaulding FP 303(10), FP 15107(11), FP 34543(19), FP

41633(22).

Spaulding & Eno FP 68312(19).

Spenser 1018 (S).

Standley 4173(2Sh), 7664(11), 7810(10), 7979(10), 16335(S),

81637(13), 81642(13), 81683(13).

Stanford 0^.9(3).

Stanford, Tauber & Taylor 2495(2Ssi).

Steward 6605(3).

Steward & Jackman 6499(11).

Steward & Sowder 6803(8).

Steward & Steward 7328(1Q), 7471(3).

Stewart 7549(26), 21297(26).

Stewart & Rahman 25213(26).

Stewart & Stewart 7329(11).

Steyermark 29788(13), 36940(13), 42524(13), 49044(13),

50130(13).

Stone 20l(12si).

Stouñer FP 68239(11).

Stouffer & Gill FP 68291(17), FP 68292(4), FP 68293(4).

Stoutsunire 3013(19).

Straw & Forman 1777(20).

Sueur 599(123i).

Suksdorf 1364(8), 2246(1SL).

Summers 926(18).

Taylor 182(230,), FP 18641 (23h).

Thomas 5519(18), 10262(8).

Thompson L-1128(19), 9422(22), 13844(11), 15105(3).

Thomson FP 26558(23h).

Thornber & Hockdoerffer 2944(1^)-

Thorne 19860(19).

Tidestromi500(10).

Toko & Pinnay 65-34(16).

Toumey 5(23b), ^S'^(23b).

Tracy 1060(1Q), 2355(8), 6912(18), 8950(lh), 1057l(lh),

16464(S).

Tracy, Earle & Baker 388(10).

Train 2150(9), 2154(1Q), 2521(10).

Tucker 2563(23h), 3444(1^)-

Tweedy 449(3), 595(10), 4973(23h).

Twisselmann 8681(8), 8832(18).

Umbach55>4(3).

Valdivia & Soloranzo 633(13), 634(13), 637(13), 646(13),

648(13), 649(13), 650(13).

Vickery (51(5 (23b).

von Turckheim II 1815(13), 3241(5).

Voss 1048(19), 1539(19), 2660(19), 3935(19), 4^81(19),

6986(19), 7009(19), 7854(19), 8602(19), 11870(19),

12076(19), 12239(19), 12467(19), 12696(19), 12712(19).

Wsiámond 1271(19).

Wagener FP 89603(10), FP 89604(10), FP 97951(18).

Walpole 234(3), 323(3), 417(8).

Ward 51(18), 595(8).

Warnock & McVaugh 5452(23h).

Waters 242(3).

Watson 6762(19).

Weatherby 1667(7), 1674(lh), 3906(19).

Weatherby & Weatherby 5770(19), 6685(19).

Weber 1957(3), 6065(10), 6093(3), 6725(3).

Webster & Nasir 6471(26).

Weir 659(16), 2374(3), 2376(3), 2379(3), 2382(3), 2384(3),

2387(3), 2394(19), 2400(11), 2412(11), 2415(11), 2416(11),

Weir(continued)

2418(11), 2419(11), 2421(16), 2423(lsi), 2425(16), 2426(lh),

2427(lh), 2429(12.), 2449(22), 2452(22), 2456(22), 3185(8),

3187(8), 3189(8), 3191(7), 3193(8), 3194(8), 3200(8),

3202(8), 3206(8), 3216(9), 3218(9), 3239(7), 3240(7),

3242(22), 3246(9), 3250(16), 3252(16), 3258(16), 3267(16),

3271(16), 3288(7), 6162(11), 6163(23h), 6838(22), 7196(16),

7426(11), 7473(3), 8243(8), 8252(8), 8254(9), 8334(16),

8336(16), 8337(16), 8345(9), 8362(16), 8363(16), 8364(16),

8365(16), 8366(16), 8371(16), 8373(16), 8374(16), 8384(16),

8385(16), 8389(3), 8390(16), 8391(16), 8392(16), 8440(3),

8442(3), 8444(3), 8456(3), 8475(8), 8958(23h), 9780(3),

9829(3), 9868(8), 9872(9), 9874(3), 9875(3), 9986(11),

9995(11), FP 17047(8), FP 17048(16), FP 29660(8), FP

91207(11), FP 91249(9).

WetheriW 919/3149(10).

Wheeler ^^^(8), 330(3), 1115(8), 2783(11), 3005(1Q), 4428(18),

4452(18).

WhitGà 3171(3).

Whiting 854/2825(10), 1053/5338(11).

Whitney 1703(1^), 1717(3).

Wibbe 11532(19).

Wickens 2199(23h), 2403(23h).

Wiegland 0544(19).

Wiens 2335(18), 2444(^), 2574(12h), 2705(23h), 2728(18),

2740(10), 2741(8), 2802(23h), 2808(7), 2926(9), 2966(3),

2983(23h), 2984(11), 2986 (23b), 2988(23h), 2994(11),

3029(23b), 3032(8), 3033(lâ), 3034(9), 3035(23h), 3036(11),

3168(10), 3171(10), 3178(9), 3179(10), 3180(12.), 3183(17),

3185(10), 3204(12), 3205(8), 3207(10), 3210(12), 3211(8),

3212(8), 3217(10), 3218(18), 3219(18), 3220(18), 3221(18),

3222(8), 3223(12), 3224(18), 3226(3), 3227(8), 3228(12),

3229(8), 3230(7), 3231(8), 3232(12), 3233(12), 3235(22),

3236(8), 3238(7), 3239(8), 3240(22), 3241(22), 3242(22),

3243(22), 3244(12), 3245(8), 3246(8), 3247(22), 3248(22),

3249(22), 3251(22), 3252(22), 3253(8), 3254(8), 3255(16),

8256(16), 3257(9), 3258(9), 3260(3), 3605(18), 3606(8),

W07(8), 3609(7), 3610(12), 3611(7), 3612(8), 3613(12),

^614(3), 3615(lb), 3616(8), 3617(9), 3618(11), 3865(9),

"^3658(11), 3913(10), 41^0(11), 4121(11), 4122(12),

4123(23b), 4124(9), 4125(9), 4127(9), 4127a(ll), 4127b(8),

4128(11), 4129(11), 4168(10).

Wiens & Wiens 4358(26), 4359(26).

Wiggins 2106(8), 2725(8), 2817(8), 9322(8).

Wiggins & Demoree 5018(8).

Wight 47(18).

Wilbur & Wilbur 1837(13), 1982(13).

Williams 897(16), 2402(3), 15964(10).

Wilson & Wilson 50^7(19).

Wittrock 5057(22).

Wolf 1342(18), 2880(23b).

Wolf & Rothrock 70(3).

Wood, Clover & Voss 8118(19).

Woodrow & Long FP 18399(23b).

Woolsey FP 12566(17).

Wootson 23(3).

Wright FP 68048(3), 68050(lb), FP 68052(12), FP 68054(18),

FP 68107(3), FP 68109(8), FP 68111(8), FP 68114(3),

FP 68115(12.), FP 68116(12), FP 68117(8), FP 68119(18),

FP 68123(lb), FP 91563(8).

Wyman FP 1838(10), FP 12815(23b).

Young (55(10).

Yu 1558(25).

Zeller & Zeller 1194(22).

234 sur U.S. GOVERNMENT PRINTING OFFICE: 1972 O—412-429