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élNDlNG ‘BY \ W!
HUAB & SBNS' l
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q MSPRINGPZF .Cn‘if-M “fl
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ABSTRACT

THE SYSTEMATICS OF LYONIA (ERICACEAE)
IN NORTH AMERICA

BY
Walter S. Judd

The genus Lyonia of the family Ericaceae and
tribe Andromedeae includes approximately 35 species in
eastern North America, eastern Asia, and the Greater
Antilles. It is represented by three sections in North
America. Section Lyonia is mainly West Indian with only
hu>species on the Atlantic Coastal Plain and one in
montane areas of Mexico. The monotypic section Arseno-
% includes only Lyonia ligustrina, a widely distributed
and variable species of the Coastal Plain, Piedmont and
mDuntains in the eastern United States. Section garia
inCludes two species, both of the Coastal Plain, although
one also occurs in Cuba. Numerical taxonomic and chemo-
8ystematic investigations support the recognition of
the three sections. The phytogeography, ecology, floral
andVegetative anatomy and pollination biology were
inveStigated. The genus is distinguished from related

genera by its axillary inflorescences from the wood

fine previous seasc

r32: buds, and cons,

:5 Lie fruits.

 

 

 

 

 

(5(2) Walter S. Judd
Ch

(9 (If the previous season, two large outer scales on the
‘winter buds, and conspicuously pale, thickened sutures

of the fruits.

THE SYSTEM

 

 

n N.- ‘
1“ Parnell?

 

Depdrme'lt

THE SYSTEMATICS OF LYONIA (ERICACEAE)

IN NORTH AMERICA

BY

;\
\‘I
\

—.‘ -|.

Walter Sikhudd

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE
Department of Botany and Plant Pathology

1974

 

 

Ivish to 8X}

““f “'5‘
‘ I. V‘V
I I

.essor. Dr.
zinc-suragement (3:13

:25 and Stephen "

:n;
u.

:rzticism concert.

:' 3-50 to

Drs. Jareze
~L‘.J. Furlow for t1”.

D

 

r. Barbara I

p.

“'45-? gave the late
.\a and relate:
532:1“ - -

rm 0f chhlga:

E5 1"

2 3'5:
“{C
is:- .
y “a .
5 are listen
fittfi"

ACKNOWLEDGMENTS

I wish to express my sincere appreciation to my
xmajor professor, Dr. John H. Beaman, for his guidance
and encouragement during this study. Drs. Aureal T.
(Iross and Stephen N. Stephenson provided helpful comments
and.criticism concerning the manuscript. Special thanks
go also to Drs. James W. Hanover, Garrett E. Crow, and
John.J. Furlow for their suggestions.

Dr. Barbara Palser of Rutgers University gra-
ciously gave the late Dr. Kornelius Lem's collection
of Lyonia and related genera to the Beal-Darlington
Herbarium of Michigan State University. These specimens
were a valuable aid to this investigation.

The support of a National Science Foundation pre-
doctoral Fellowship greatly facilitated the progress of
my research, and funds for fieldwork were provided by
the Department of Botany and Plant Pathology, Michigan
State University.

I also wish to thank the curators of the herbaria
from which specimens have been used for this study. These
herbaria are listednear the beginning of the Taxonomic

Treatment .

ii

 

A special the.
:iersméing, patier.

11;;Izout the stuiy.

 

 

A special thanks goes to my wife, Beverly. Her
understanding, patience, and help have been essential

throughout the study .

iii

 

13? OF TABLES .

O
nu.- A n?!“-
..;. J? MULRES. .
fic-a-‘QFVA -
~‘.:....,‘\,‘ng‘ .

w
.

37331 CAL ACCOUNT .

u

‘-
..‘rw-

-14.. REJIIORSHIPE
EYES-SPA?“ AND E
1.322% FRUIT MOP
3:1,.
«avrescence .
. :pmgenetic

13338:.
. n

O
.a;yx ,

Ctrslla . ‘
5‘:~3ecllum .
eg'neecium. .
:riit .

'I .
~~ HON BIOLOGY
1:233.

 

TABLE OF CONTENTS

LIST OF TABLES . . . . . . . . . . . .
LIST OF FIGURES. O O O O O O O O O O 0
INTRODUCTION . . . . . . . . . . . .
HISTORICAL ACCOUNT. . . . . . . . . . .
GENERIC RELATIONSHIPS. . . . . . . . . .
PHYTOGEOGRAPHY AND EVOLUTION . . . . . . .
FLOWER AND FRUIT MORPHOLOGY. . . . . . . .
Inflorescence .
‘Morphogenetic Cycle
Flower. . . .
Calyx . .
Corolla .
‘Androeciwm

Gynoecium.
Fruit . .

POLL INAT I 0N B I OLOGY O O O C O O O O O O
EPIDERMAL APPENDAGES . . . . . . . . . .

Key to Sections and Species of Lyonia Based on
Trichome Type. . . . . . . . . . .

ECONOMIC IMPORTANCE . . . . . . . . . .
NUMERICAL TAXONOMIC STUDIES. . . . . . . .
Phenetic Studies . . . . . . . . . .

Phyletic Studies . . . . . .
Comparison of the Phyletic and Phenetic Methods

iv

Page
vi

viii

17
23
40
40
40
42
43
43
S4
55
55
59

62

64
75
77
78

130
141

 

'32"; ATIC STUD

      

lathe-is and Mater;
Results and Discus

:. ‘Sl’flC TRE‘ATYEXT

"axis Rattan.

:2; tcSections, E

":i’ v lial O - .
:geczzens Examine
lien; m.

a hut. sect

l.

LVCnia Sr

2. 1.;"05‘ a T“
x...

YRQLLTSOI

3. L‘ieria f:

\ Mitt. Sec
Ssdt. “CV.

e
' LVQni
. ‘ a ]
9.. a r
“ERRED:

Page

CHEMOSYSTEMATIC STUDIES . . . . . . . . . . 143
Methods and Materials. . . . . . . . . . 144
Results and Discussion . . . .. . . . . . 147

TAXONOMIC TREATMENT . . . . . . . . . . . l7 8
Lyonia Nuttall o o o o o 6 o o o o o o 1.78
IKey to Sections, Species and Varieties of

L onia C O O C O O O O C O O O O O 180
Spec1mens Examined. . . . . . . .. . . . 182

Lyonia Nutt. sect. Lyonia . . . . . . . . 183

 

 

 

 

 

 

 

l. L onia squamulosa Mart. & Gal. . . . . 183
2. L on1a fruficosa (Michx.) G.S. Torrey in
RoBinson . . . . . . . . . T- . 195
3. Lyonia ferruginea (Walt.) Nutt. . . . . 215
Lyonia Nutt. sect. Arsenococca (Small) Judd,
Stat. nOVO O O O O O I O O O O O O 228
4. Lyonia ligustrina (L.) DC . . . . . . 228
4a. L onia ligustrina (L.) DC. var.
Iigustrina. . . . . . . . . 231
4b. L onia’li ustrina (L.) DC. var.
o 1081 ora Michx.) Fern. . . . 258

 

‘Lyonia Nutt. sect. Maria (DC.) C. E. Wood
J. Arnold Arb. 42:48. 1961. . . . . . . . 281

5. Lyonia lucida (Lam.) K. Koch . . . . . 281

6. Lyon1a mariana (L.) D. Don. . . . . . 301
LITEMTURE CITED 0 O O O O O O O O O O O 321

 

CO‘Z-Tvarauve trq
authors ,

CCZParison of

LiSt of 01:ch 5
each ,

Characters Us

(Phenetic

HG J5? anal‘i:
loa‘chdr al

ESRGUP anal}.
lZB-Chara
3390;? anal!
“Titan
38mg? anal
flower-f
HERO?” aha?

10.
11.
12.

13.

14.

LIST OF TABLES

Comparative treatment of Lyonia by various
authors . . . . . . . . . . . .

Comparison of the sections of Lyonia . . .

List of OTU's of Lyonia, giving the code name
eaCh O O O O O O O O O O O O 0

Characters used in numerical taxonomic study
(phenetic method). . . . . . . . .

Matrix of error potentials,
set. 0 O O O O O O O

HGROUP analysis:
108-character

HGROUP analysis:
128-character

Matrix of error potentials,
set. 0 O O O O O O O

HGROUP analysis: Matrix of error potentials,
vegetative character set . . . . . .

HGROUP analysis: Matrix of error potentials,
flower-fruit characters. . . . . . .

HGROUP analysis: Matrix of error potentials,
floral characters. . . . . . . . .

HGROUP clustering of characters of Lyonia. .
Factor analysis of loo-character set . . .

Characters used in phyletic study (Wagner
me thOd) o o o o o o o o o o o 0

Character divergence values for each taxon,
used to construct Wagner tree. . . . .

Matrix of character divergence used in Whiffin-
Bierner method. . . . . . . . . .

vi

of

Page

4

13

80

81

104

105

106

107

108
119
128

131

133

138

 

I‘i‘
. .
(D

o
r
o

. -
o

. .
. '-.‘
u

I."
o

3‘

6“

 

c graphic 10C?
foliage 537%
com. ands V“

 

Rf values and i
I v ‘
m L 00.13

 

Total listing C
in each POISJ
studied

Characteristic
species of

later of cor:
of Lvon' .
. 1a 9

Chemical simii

in North A

Compounds se;
lioustrira
.oiiosn‘w

‘.

“'1 -

u agnostic c<

Comparison 0

erruc‘

.ne
comparison c

llQ“S '.
1]

Table Page

15. Geographic location of specimens from which
foliage samples for analysis of phenolic
compounds were obtained . . . . . . . 145

16. Rf values and color reactions of the phenolics
in Llonia O O O O O O O O O O 0 148

17. Total listing of phenolic compounds present
in each population of each species
studied . . . . . . . . . . . . 153

18. Characteristic phenolic compounds for each
species of Lyonia . . . . . . . . . 154

.19. Number of compounds in common among the taxa
of Lyonia studied . . . . . . . . . 167

.20. Chemical similarity between species of Lyonia
in North America . . . . . . . . . 168

21. Compounds separating L. ligustrina var.
li ustrina from L. ligustrina var.
foIiosirora. . . . . . . . . . . 170

 

 

 

 

22. Diagnostic compounds for section Lyonia. . . 171
23. Comparison of Lyonia fruticosa and L.

ferruginea . . . . . . . . . . . 210
24. Comparison of L onia li ustrina vars.

ligustrina an oliosirora. . . . . . 256

 

 

vii

 

 

 

"Grid distribu

Possible disu
lic'JStrlna
\

Drawing of flo
Drawing of flc
Drawing of fl:
Drawing of £11
Drawing of fl

Variatioh in
cease in E

VariatiOn 1
COIQr. .

variation in

LIST OF FIGURES

 

 

 

 

 

Figure
l. WOrld distribution of Lyonia . . . . .
2. Possible distribution patterns in Lyonia
ligustrina . . . . . . . . . .
3. Drawing of flower of Lyonia ligustrina. .
4. Drawing of flower of Lyonia ferruginea. .
5. Drawing of flower of Lyonia fruticosa . .
6. Drawing of flower of Lyonia mariana. . .
7. Drawing of flower of Lyonia lucida . . .
8. Variation in capsule morphology and dehis-
cence in Lyonia . . . . . . . .
9. Variation in glandular hairs in L onia;
darker shading indicates a reddish
color. . . . . . . . . . . .
10. Variation in size and shape of ferrugineous
scales of Lyonia ferruginea . . . .
11. Longitudinal views of hair types in
Section Lyonia. . . . . . . . .
12. Hair types in Sections Maria and Arseno-
cocca. . . . . . . . . . . .
.13. HGROUP phenogram of Lyonia based on 128
character set . . . . . . . . .
14.. HGROUP phenogram of Lyonia based on 108
character set . . . . . . . . .
lS. HGROUP phenogram based on vegetative char-

acters of Lyonia . . . . . . . .

viii

Page

26

34
45
47
49
51
53

57

68

70

72

74

95

97

99

 

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p -
(‘I A.
-

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L-
-

 

HGROL'P pheno;ra
acters of L?

  
  
 

3290:? phenogra
0f Lyonia

based on 10 .

N

35932"? relatio:
°n Vegetati‘

- HGRC'JP relatio

flower-frui

ThreeLd i me n 5 1C
snips anon:
data from c

F

4

:
.actor analys
based On v

based On f
LVOnia

 

 

Figure Page

16. .HGROUP phenogram based on flower-fruit char-
acters of Lyonia . . . . . . . . . 101

1]. HGROUP phenogram based on floral characters
of Lyonia . . . . . . . . . . . 103

18. HGROUP relationships between OTU's of Lyonia
based on 108 character set. . . . . . 110

13. HGROUP relationships between OTU's based
on vegetative characters of Lyonia . . . 112

ML. HGROUP relationships between OTU's based on
flower-fruit characters of Lyonia . . . 114

21. Three-dimensional model showing relation-
ships among OTU's of Lyonia, based on
data from cluster ana y81s. . . . . . 116

22. Factor analysis: Principal Factors I and II
based on vegetative characters of Lyonia . 122

23. IEactor analysis: Principal Factors I and III
based on vegetative characters of Lyonia . 124

24. Factor analysis: Principal Factors I and II
based on flower-fruit characters of
Lyonia . . . . . . . . . . . . 126

25. Index of divergence of L onia with code
letters of characters 1n specialized
State. 0 O O O O O O O O O O O 135

26. Phylogenetic trees of Lyonia produced by
Whiffin-Bierner phylet c method . . . . 140

27. :Representative chromatogram.of Lyonia
fruticosa . . . . . . . . . . . 156

23. Representative chromatogram of Lyonia
ferruginea . . . . . . . . . . . 158

29- Representative chromatogram of Lyonia
ligustrina var. ligustrina. . . . . . 160

30- Representative chromatogram of Lyonia
ligustrina var. foliosiflora . . . . . 162

ix

 

. .
:.;re

31. Representative

1:;

. Representative
mariana .
a

Phylogenetic t:

pounds dist:

 

 

 

:2. Representative
“ “‘- a u
1.. Listribution o

1“ Rexesentative

15. Dis

tribution c

8.‘ c
“it Curvatur.

co
8a and E

C. .
A Cofiearison

LVon'
Q E ,r, L

Figure Page
IL Representative chromatogram of onnia lucida . 164

32. Representative chromatogram of onnia
mariana . . . . . . . . . . . . 166

33. Phylogenetic tree showing the phenolic com-
pounds distinctive for each taxon of

Lionia O O O O O O O O O O O O 174

34. Phylogenetic trees based on chemical data,

 

 

 

 

 

 

 

 

using Whiffin-Bierner method . . . . . 176
RL Representative specimen of onnia squamulosa . 187
ML Distribution of onnia squamulosa. . . . . 189
EL. Representative specimen of onnia fruticosa . 199
38. IDistribution.of onnia fruticosa . . . . . 201
39. Leaf curvature and position in onnia 35213-

2222 and E. ferruginea . . . . . . . 209
40. .A comparison of the flowering periods of

Lzonia fruticosa and 2° ferruginea . . . 213
41. Representative specimen of onnia ferruginea . 219
42. Distribution of onnia ferruginea. . . . . 221

 

43. .Representative specimen of onnia ligustrina
var. ligustrina . . . . . . . . . 234

44. 1Distribution of onnia ligustrina var.
ligustrina . . . . . O C C O C O 236

45. .Representative specimen of onnia ligustrina
var. foliosiflora . . . . . . . . . 263

46. Distribution of onnia ligustrina var.
foliosiflora. . . . . . . . . . . 265

47. lRegions of overlap in distributions of
L onia ligustrina vars. ligustrina and
foIiosiflora. . . . . . . . . . . 279

48. Representative specimen of onnia lucida . . 286

F;;;:e
4%. Distribution 0:
it. Regresentative

Distribution 0:

 

 

_._.L -‘ ‘ .

 

 

Figure Page

49. Distribution of onnia lucida . . . . . . 288
50. Representative specimen of onnia mariana . . 305
51. Distribution of onnia mariana. . . . . . 307

xi

.__.____ am -~ ,

 

 

The Ericacea'
fillies, but infraf
iffzcult to detemi.

255 in point. Andrl
\
I:::.‘ie diverse arra

:s:e is still much

"4: should be recor

iiiinctive gene r a

«s see

ll CORtI‘OVEI 5

fine Coastal Plai
'- in some locali'
‘tI-r

75:321.
1011, The e

.. C19 species 0f

1' IQSea
sewn to the F1!”
:3 ‘i‘svxn .

3.31:4
~a in COn'JEn‘
we

INTRODUCTION

The Ericaceae are one of the more natural plant
families, but infrafamilial limits of its taxa are often
difficult to determine. The tribe Andromedeae are a
case in point. Andromeda was long used as a catch-all
for the diverse array of species in this tribe, and
there is still much confusion as to the number of genera
that should be recognized. Lyonia is one of the more
distinctive genera in the Andromedeae, but its definition
has been controversial. It is abundant in many places
on the Coastal Plain of the southeastern United States
and in some localities is a dominant component of the
VGQEtation. The extreme variability shown by several
0f the species of Lyonia has long been a source of con-
fusion and has been a stimulus for my interest in the
genus.

My research on Lyonia was initiated as a contri-
bUtion to the Flora North America data bank. Because of
the suspension of that Program, this treatment is pre-
8eI'lted in conventional form; but hopefully knowledge
°f the North American species is now more readily

ac(lessible for a computerized taxonomic information

72's. of the future.

 

use 2min Wed and e

. . . - I
anus-c in the field.
nae: ision to re:

2 1.5.. ml SPECIES.

1:21:21”: studies we:

 

:52: heavily on sta:

 

$223.“. only the sp-

‘

El"

., it is hope
’55 can eventually
“‘3 PIO"ide a has

~::‘.C‘.‘..t West India

 

system of the future. Over 4,000 herbarium specimens
were examined, and each taxon except 1;. squamulosa was
studied in the field. Fieldwork was a fundamental factor

in my decision to recognize L. fruticosa and L. ferruginea

 

as distinct species. Both the chemical and numerical
taxonomic studies were helpful in reinforcing conclusions
based heavily on standard morphological comparisons.
Although only the species native to North America are
included, it is hoped that an investigation of the entire
genus can eventually be completed. The present study
should provide a basis for interpretation of the more

difficult West Indian and Asian taxa.

 

 

‘ u -
Both Lorna 1.

Embed by Linnae‘u‘

';'V \
"qo

;..a was out m l
as glazed in Vaccii
\

:Lz'al resemblances

‘a of nisunders ta

at: :4 '
*5 «meat

ity of Lil
2:1;1'2t it was a l:

333: Pu:

sh. lBlS;

lite:
s correctly 2

LN

a. the err
x-omed
l ‘ q
«ray, lBBB; Sr:
‘35! botanists

2;: -
SPECIES ,

th

':: ‘
“‘lptiOn Of

 

HISTORICAL ACCOUNT

Both Lyonia mariana and -I_.. ligustrina were

 

described by Linnaeus in his Species Plantarum. Lyonia

 

mariana was put in the genus Andromeda but _I_._. ligustrina
was placed in Vaccinium probably because of its super-
ficial resemblances to that genus. This led to a great
deal of misunderstanding by later botanists concerning
the identity of Linnaeus' Vaccinium ligustrinum. Some
thought it was a blueberry (Marshall, 1785; Willdenow,
1799; Pursh, 1816; Nuttall, 1818; K. Koch, 1872), while
others correctly concluded that its true affinities were

with the Andromeda group (Elliott, 1817; DeCandolle, 1839;

 

A. Gray, 1886; Small, 1914; Wood, 1961), Table 1. Most
early botanists used the name Andromeda paniculata for
this species, thinking that the A. paniculata of Lin-
naeus (1753) referred to this plant. But Linnaeus'
description of A. paniculata, which is quite ambiguous,
probably refers to Leucothoé’. racemosa.

Another nomenclatural mixup concerns the identity
of Walter's (1788) Andromeda ferruginea. Pursh (1814),

Elliott (1817), and Nuttall (1818) incorrectly took his

 

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II II II II II II II mmmmmmmmmm .q
Asmooawsum
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«acoomwummun
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I Asmocaoum .m
as Ammo sou . 0cm can muaflm madam dam an chow 0H” escafiw
scans»: 4H .¢~ Iomcom . ~ loanum .<~ Ioacow . ~ Icons .K~ Ioacdm .4~ In HH .>~ I
upoaouucd svoeoupsd noofioupnd avoaounqd upoEouvcd upofiouvc¢ aswswooa> dddh¢¢flflda .A
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upmfioupqd upoaouvcd upmeoupcd moosoupcd someouvsfl mvoaounnfl II duaoad .q
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upmaonpcd mumEoupc¢ apoeoupc< momfioupcd mooEOHUQ¢ opoeouvnd noosoupad acuauufi .q
reams. .eamac .momac Assad. .mohas .mmpav .nmpas nexus
unawaam cause nuance: sosocnawz hands: Haucuunz uaosaswq
.uuonusn unowun> in «HCONA uo vacauaouu o>wuoudmeoo .H man‘s

 

 

 

 

 

 

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MAcONA MAco A case A Mano A II nnOASEdst .m
A .uoom Amocflmsuum .m AmocAmduuom .m~ Ammcwmsuumm .m~ Ammcamsuuom xm~ I
MAGONA uAGONA Mano A «Ago A Mano A snoouuaum .A
A .uomm Ammcamsuum .l Ampfimau .m Aspwmau .m “nowadu am I
macONA «AGOMA maaoxA mAcONA maaONA monumsuuou .A
I AwwHOvamoumuo .m
nwmowcoum .u .muon MHDE .u
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mAAONA «Ago A Mano A MucomA ncwuuusmwa .A
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Amvauac .M~ AspAos .A Ammomauoo .A Amumcflmume .A Ammaumc .K~ I
maumAm MACONA monuoosoA nacONA spoaouusd mqusA .A
mass: .uowm I wand: .uomm I I
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mammam mAcoxA woguooswA MACONA uuosouoqd mundane .A
loamAc lwsmac rammAc lemmas AmamAc nouns
xoom 0cm nucwm zoom .M TAHOGGMUTD Q00 .0 Hamvvfiz

 

poacwucou .A AAAAH

 

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MAGONA
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AmmooAusum .x~ .wmooAusum .x~ AmmooAusnm .x~ .mmOUAusum .xy .4 I
MAco A mEmAon MEmAon mEmAon mvmsoupnfi mmOOAusuu .A
MAco A .uoom com I I. nAAON . OJW
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msoooo MMSmomnoum
uAumz .uoom Iocomufi .uoom .mscAuu _I «HAONW .uoflm
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mAco A mEmAon msoooooconud mEuAon someonvad ucAHuuAMAA .A
MAumz .uomm MAumz .wqu I uAqun .woou
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MAco A mEmAon mAHOA 002 nAqu ooz upofioupnd usnAuda .A
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:3: description to

Er;:i:csa. Thus the:

 

-.-..- c r .2 ‘ -'
5.“: .eiricinea, a
.3321: for the true L
;2;e:s of this perio

2:21;: probably had

 

1236 not recognizin
1::ct have this pr
The most con

22;; *1.
.e early ParI

it: '.
are 14.. yAISh. A‘

113‘. 3‘ ‘
the ‘Ealrljf t}

:::3 Jva- IATKir(}~
.‘h'- -“‘-—;;
”\“a (Lam.) my..-

I.-.A'
\v‘ v A
a.e~t

1)! called

short description to mean the plant now ‘called Lyonia

fruticosa. Thus they called this plant Andromeda or

 

 

Lyonia ferruginea, and used the name Andromeda rigida
Pursh for the true 2° ferrginea. Therefore, in many

papers of this period when A. ferruginea is mentioned the

 

author probably had 1;. fruticosa in mind. Of course

 

those not recognizing these two taxa as distinct species
did not have this problem.

The most commonly used name for Lyonia lucida
until the early part of this century was Andromeda nitida
Bartr. _i_rl Marsh. Andromeda lucida Lam. was thought by
most of the early botanists to be a later homonym of A.
lucida Jacq. Andromeda lucida Jacq. is Leucothoé @-
lifolia (Lam.) Dippel, although this species is often
incorrectly called Leucothoé acuminata (Ait.) G. Don.

Until Nuttall (1818), all the species then known
of Lyonia were placed in the large polymorphic genus

Andromeda which served as a kind of catch-all. Nuttall,

 

in erecting the genus Lyonia, separated 1:. ligustrina,
g. ferruginea and 1:. fruticosa from the rest of the
species then usually lumped together in Andromeda. He
distinguished Lyonia from Andromeda on the basis of its
unusual capsule, subglobose pubescent corollas, and

awnless anthers, and stated:

‘Were not this group of species so perfectly natural
and easy to distinguish from the genuine Andromedas,

 

ll 1134‘ have bt
of that genus. 2‘
structure of the

    
  

hers: von' "”
eL 1a co...

fall victim to a dar
Luz-auntie mountai
sure of his labo

ruriana and l_.. l

5759-.
e' “Filth
Sikr‘
s‘Qlally c
\;S'
10:; Of LI
I’j‘s'. y. \
“ion of +
E‘- :1
i e varlc

it might have been retained as a subgeneric section
of that genus, notwithstanding the singular
structure of the capsule.
The name Lyonia commemorates: "the late Mr. John Lyon,
Ian indefatigable collector of North American plants, who
fell victim to a dangerous epidemic amidst those savage
Iand.romantic mountains which had so often been the

theatre of his labours" (Nuttall, 1818) . Nuttall left

12:. mariana and _I._.. lucida in Andromeda.

 

David Don (1834) first recognized the unity of
the genus as it is now circumscribed and recognized that
2. mariana and L. lucida have their closest affinities
with L. ligustrina and _L_. ferruginea. However, he did
put E. ovalifolia (an eastern Asian species) in the genus
Pieris and included the rather dissimilar species 1.2313(3-

thoé racemosa and Oxydendrum arboreum. It is interesting

 

that Q. arboreum does have slightly thickened capsule

 

valves and awnless anthers, and these similarities may
have induced Don to include it in Lyonia. Its inflor-
escence, which is a terminal panicle of racemes, is also
superficially similar to that of L. ligustrina. The
inclusion of Leucothoé racemosa may have been due to the
confusion of this taxon with Lyonia ligustrina, resulting
from the various interpretations of the Andromeda pagi-

culata of Linnaeus. Although Leucothoé racemosa has

 

awned anthers and lacks thickened sutures on its cap-
sules, it is quite similar to Lyonia ligustrina in over-

all appearance and could have been included on this basis.

 

De Candolle

9;; between Lyonia

:15 in the genus Lt-

 

52212053 and L.
E -

 
  
  

:Lu‘ed the Asian
Later treat.-.
2211872) divided

2:22: Lyonia and

SE Rug, 7 9
1“ Ln marl H I
‘ £ a!

:79“ An r0312:
:3me mi.
22*?
id‘en‘u healing
Ki: van, vie
aééspten CM”
5; h ntel‘SChe
siilhaf enhei
Se?» be). der .
61:: b“ bei Ll"
Edoenesgndel’e
Erna; “it due
~. ‘teno 51$
“ Small
’ (1914)
”grille

De Candolle (1839) recognized the close relation-
ship between Lyonia mariana and A, lucida and placed both

taxa in the genus Leucothoé. He kept A. ligustrina, A.

 

 

fruticosa and A, ferruginea in Lyonia. Like Don, he

 

 

included the Asian species of Lyonia in the genus Pieris.
Later treatments of the genus vary widely. K.

Koch (1872) divided Linnaeus' Andromeda into only two

 

 

genera: Lyonia and Andromeda (including only A, polifolia).
He put A, mariana and A. lucida in subgenus Pieris with

Zenobia speciosa, and A. ferruginea and A. ligustrina

 

 

in subgenus Eulyonia. He also pointed out an ever-present

 

danger in the determination of generic limits in the

Andromedeae:

.Am besten thut man, nur 2 Genera aus der Linne-
schen Andromeda zu bilden: Lyonia und Andromeda.
'Bei allen Lyonien kommen die Blfithen, meist zu
mehren, aus seitlichen Knospen vorjahriger Aeste.
Die Blfithenknospe bei Andromeda Polifolia ist
dagegan, wie bei den Rhodereen, gipfelstandig und die
Knospenschuppen fallen nicht ab . . . Legt man bei
des Unterscheidung der Genera allein auf die
Beschaffenheit der Bluthe einen Werth, so sieht man
sich bei der grossen Verschiedenheit im Bau der-
selben bei Lyonia gezwungen, fast aus jeder Art
ein besonderes Genus zu bilden. Und in der That
haben wir auch nach und nach fast so viele Genera
erhalten, als Arten vorhanden sind.

Thus Small (1914) was led by the diversity of the group to
recognize almost as many genera as species, putting
Lyonia mariana in Neopieris, A. lucida in Desmothamnus,

.L. ligustrina in Arsenococcus, and A. ferruginea, A.

 

fruticosa and A. squamulosa in KolismaI

 

 

Britton and
L. luciia in the ger

sutlarities in flc:

 

:1; cylindric flow.

:alyx lobes and 511':
I

:21: snail urceola -

2:325 containing r

5-“: .' ‘ l".
‘h :0 fifthglnEd 3
\

:crered sutures ,

REh-Cier's (1

Rea-e'
.....ially corres:

.L. , .
“‘5 r‘E‘VISion.

my has“ Upon t

353110;; Lv -
.JELEiSma

‘-.fnvp. .
‘ magma. is c

-'?.':e '
in dense axil

 

hm
"‘«V In

I“

S .
Cha

qu racte

.‘gste

10

Britton and Brown (1913) put both A. mariana and

A. lucida in the genus Neopieris, recognizing their many

 

similarities in floral morphology (e.g., spurred filaments,
long cylindric flowers in axillary fascicles, lanceolate
calyx lobes and similar capsules with moderately thickened
sutures containing many small seeds). Both A. ligustrina

and A. ferruginea were placed in Xolisma on the basis of

 

their small urceolate flowers, small capsules with

thickened sutures, and only very short spurred filaments.
Rehder's (1924) treatment of the genus (as Xolisma)

essentially corresponds to the classification presented

in this revision. He divided the genus into four sections,

mainly based upon the genera recognized by Small (1914).

Section Lyonia (Small's Xolisma), the type of which is

‘A. ferruginea, is characterized by having the flowers

 

borne in dense axillary fascicles (corymbs), by the
nearly unappendaged filaments and the lepidote pubescence.
The leaves are evergreen and entire, toothed or undulate;
the capsule is usually ovoid, prominently angled, with
the thickened sutures separating as a unit from the rest
10f the capsule. Its many species, 24 being enumerated
by Small (1914) , are mostly natives of the West Indies.
Section Arsenococca (Small's genus Arsenococcus)
includes the single isolated species, A. ligustrina,
which is characterized by its flowers being borne in

pseudoterminal panicles (racemes of corymbose fascicles),

Essence of ferru
:Li-shaped multicel "
;'.:'::se capsule, not
:irened sutures.

2:5. This variable

: Florida and easte-

 

 

Section Mar 1
TA

4.

...1‘3‘; ' p .4

 

56:20: .2:' .
H“ 13) and i

‘OUrtr
35‘5“.
. SPEQieS m-
‘5
>3; Ore‘ _
. sided I
I
"file .
S With 1&5
‘
.t: CS ar
. “tir
re ‘
or less

11

the absence of ferruginous scales, the presence of long
club-shaped multicellular hairs, and by the depressed
globose capsule, not prominently angled, with less
thickened sutures. The leaves are nearly always decid-
uous. This variable species is distributed from Maine
to Florida and eastern Texas.

Section Maria (Small's genera Neopieris and Des-

 

mothamnus) includes both A. mariana (the type species of

 

 

section A3333) and A. lucida. This section is character-
ized by the flowers borne in axillary clusters on the
branches of the previous season, by the noticably appen-
daged filaments, the ovoid or globose-ovoid capsule with
moderately thickened sutures, and the elongate flowers
with lanceolate calyx lobes. The leaves are entire,
evergreen or deciduous, and glabrous to quite pubescent
(usually with small, stalked glandular hairs). The two
species are native to eastern North America, primarily
on the Coastal Plain, although A. lucida is also found
in Cuba.

The fourth section, Pieridopsis, includes only
Asian species. Those species have the flowers in axil-
lary, one-sided, usually elongated racemes and have
capsules with less strongly thickened sutures. The
leaves are entire, persistent or deciduous, and glabrous

or more or less pubescent. Species in this section can

 

:2 ions”. from Kashm;

  
  

:i central China tr

U
H“:

Wood (1961)

:‘A
5~

.Tari g ‘ ,
*aalns

' CiaCEd SPECIEC
:esrs o. the ‘1

32:51qu L. 113”:
. \¥
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' 1‘3 Cf I'
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“323:
a w to a
‘5‘ t3

12

be found from Kashmir through the Himalayas to western
and central China to central and southern Japan and
Formosa.

Wood (1961) included A. ligustrina with A. lucida

 

and A. mariana in section Maria. He states:

Section MARIA (DC.) C.E. WOOD (Xolisma sec. Maria
(DC.) Rehd.), as defined here, includes only three
isolated species which are placed together on the
basis of the filaments appendaged near the summit,
the non-lepidote pubescence, and the axillary corym-
bose (fasciculate) inflorescences.

I consider A. ligustrina to be distinct enough to warrant

 

a separate section. It differs from both A. mariana and
A. lucida in the following points: (1) its long club-
shaped glandular hairs, (2) flowers in pseudoterminal
panicles, (3) long pointed buds, (4) small urceolate
flowers, (5) small subglobose pubescent capsules, (6)
serrulate to nearly entire leaves, and (7) only small
inconspicuous spurs near the summit of the filament.
The results of numerical and chemosystematic studies
of the genus in North America also indicate the presence
of three distinct groups, corresponding to the three
sections described above (Table 2).

Rehder (1924) used the name Xolisma instead of
Lyonia. Rafinesque (1819) proposed the name Xolisma for
the Lyonia of Nuttall because Rafinesque had given the
name Lyonia to another genus, but that was only a new

name to replace Polygonella Michaux, which Rafinesque

 

 

 

 

 

 

ll I .
Inez—1:. ~.§.A¢..II.._.AU :1...»— 2.«) I...¢A._ ._ :..1£ .1.-.111. ..
I I I‘ll-'1!lll‘lllill IIIIIIII‘I Ill“ I I

I'......I:...u.....:owll.~l¢ a #3.: I...m..—.r.,v&. . .190: III.-. .I
Fu‘—~.AsfilNl lU-J‘a

 

 

.I.A...Qx.~ LO ICOAu ...l I
., Cu h... C3I-l350
. L .N ...AAESA

213

 

 

 

 

 

msosvomo mo>owA .m moosvooo no coououo>o mo>mmA .m cooumuo>o um>qu .m
was»
muMAooous «AAouou .5 vo>0 so no no oAuocAAmo oAAouou .5 oquooouo «AAouou .h
wmoAAm .00A23 mAAouoo .m msoundAm .ouAnz uo chm mAAouou .w ouooAQOA .ouAss oAAouou .w
AmvoAomwm omonfihuoo mo moAOAommM noAOAounm
mofioomuv mvoAcmm GA ocuon mno30Am .m muonfihuou cA canon muozoAm .m omonfihuoo GA ocuon uuozoAm .m
no>Am> oAaomoo mo no>A¢> Scum
mo>Am> Eoum uAcs a mu chusuumoo Spun uAcs n no ocAumHumon AAA: 0 no chvmuumon
no: mAAusms .oocoonnu moususm .v nos mAAwsms .oocoonsu moususm .v .UocoxOAnu .nouuusm .v
133.?
AAMHm ousmm .m ucocAEoum musmm .m muo> HOV ucomnd «Adam .m
cornea» sAaoAAunu
OAMAduumo cAmMME wooA .N muAucw cAdeE umoA .N no ouAuco :Amuda mooA .a
mqun AMA
rachAm commemlnsAo .mmumA AAA: nucoAm .A mqun udAsocmAm AAmEn AAA: mucuAm .A qumou ouovAom AAA! nunsAm .A
oooooocomud .uoom sAusz .uoom oAGONA .uoom
.quONA mo uaoAuoou and no sonAummeoo .~ MAAAH

:siiered incl???”I

 

:ralié nape and a s
327.13 was prOPOSEd I‘
-.~..:pi.aceae (CW-8'
Etna of Nuttall:
E.L;c:t is the olde-
snepiadaceae, whi
Lueip the next 01
l2ztall‘s Lvo '
nia.
nrw ' -
- onia Elliott

Lvon'
1a was

 

‘: Verb I
«- Jy. .h Alfierlca '

.«Owers. Barty-
till. l'sr‘;
~ “\rda Sevev
"3°31 shrubs w
4“"
“meta GeOrgl
ti 8‘“ c
r»Cla11
Y at
W". ‘1 k
‘“ Ca iec
“Slanabyetrad
V“ It’s-rent.
‘1.th " ab

C300 egetablée
mtks dwflhe ‘
C‘Jti‘ rich
5:.ILQC of

14

considered inappropriate. Therefore Lyonia Raf. is an

.invalid name and a synonym of Polygpnella. The name

 

Lyonia was proposed by Elliott (1817) for a genus in the

JAsclepidaceae (Cynanchum). This name was published before

 

Lyonia of Nuttall; thus Rehder concluded that Lyonia
IElliott.is the oldest and valid name for the genus of
.Asclepiadaceae, which made it necessary for him to
‘take up the next oldest name, which is Xolisma, for
'Nuttall's Lyonia. Lyonia Nuttall has been conserved
(yver Lyonia Elliott (Kew Bull. Misc. Info. No. 3, 1940).
Lyonia was noted very early in the exploration
<3f North America, often being taken back to Europe where
.it was grown in gardens because of its beautiful foliage

Iand flowers. Bartram (1791) refers to both A. ferruginea

 

and A. lucida several times in his Travels, indicating
that both shrubs were common elements of the vegetation
in southern Georgia and Florida even as they are today.
His description of the“ plains near the Suwannee River
.are especially interesting:

The chief trader, intending to show me some
remarkable barren plains, on our return to our
encampment; about noon we set off, when we came
‘within sight of them, I was struck with astonishment
at their dreary appearance; the view Southerly
seemed endless wastes, presenting rocky, gravelly
and sandy pine barren plains, producing scarcely
any vegetable substances, except a few shrubby,
crooked Pine trees, growing out of heaps of white
rocks, which serve only to perpetuate the perse-
cuting power and rage of fire, and to testify the
aridity of the soil; the shrubs I observed were
chiefly the following, Myrica cerifera, two or

three varieties .
ferruginae, And:
irangula, Siderc
Ilex nyrtifoliu:
Cassine, and a .
green and decrd
heartiful. . .

William Bal
rilr. Darlington‘
12 or another, t.

: 333:1“.- Anerica e

 

.. a ‘ '
..a ‘F
‘ Animator: , lE

.
‘1».-
r

...;a includes 2

...eot Eire in
By ‘ar th
.. ~ ‘ e
no : gr

H
L) r1
(0 (D
(+01
01
f

1::O"per . 1a"
Chiefi lS-he‘
it ' VI th‘

n'. .
x: ‘u '

:s'
‘ A
"VP
are .
CR“; :1
-““l

15

three varieties. . . . Prinos varieties, Andromeda
ferruginae, Andr. nitida, varieties, Rhamnus
frangula, Sideroxilon sericium, Ilex aquifolium,
Ilex myrtifolium, Empetrum, Kalmia ciliata,
Cassine, and a great variety of shrub Oaks, ever-

green and deciduous, some of them singularly
beautiful. . . .

William Baldwin in his letters to Dr. Muhlenberg
and Dr. Darlington, from 1813 to 1817, refers, at one
time or another, to every species of the genus native
'to NOrth America except A. squamulosal His description
(12 Darlington, 1843) of the Pine Barrens of southeastern
Georgia includes some interesting comments on the impor-

'tance of fire in such habitats:

By far the greatest quantity of land along the sea
board is low Pine barrens, covered with Pinus
palustris (long-leaved Pine), principally . . .
Undergrowth, Chamaerops serrulata or Saw Palmetto,
--with some fine shrubs,--as Andromedas, Bejaria,
etc. This is the most sterile offallidescfiptions
of land. . . . There is as yet, but little naked
sandy desert; but should the weather continue, a

ew years longer, as dry as it has been . . . and
fires should rage as extensively, destroying the
vegetation,--a large portion of the maritime part
of Georgia would be rendered like the deserts of
Arabia! Were I a member of the Georgia Legislature,
my most strenuous exertions would be made to pre-
vent, by law, the burning of the forests,--which
impoverishes the land, and does incalculable mis-
chief, without a single advantage resulting from

it. Yet many of the stupid people do it, to destroy
the rattlesnakes--make the grass grow--and I believe
for the fun of looking at it.

 

this judgments concerning the effect of fire on the vege-
tation are slightly exaggerated! Actually periodic
tnxrning promotes such a pine-scrub vegetation. Lyonia

:mariana and A. fruticosa both are probably fire induced

 

e‘iprosper in so
Exes (Iltis, per

userxation) .

16

and prosper in such dry sandy areas swept by periodic
fires (Iltis, personal communication, and personal

observation).

 

 

n' .

ine Fur-:1:
4.:-.-_ .i- ‘
"-5.3‘ut group.
“P5353 35 one I

15 early decal

Pun.

' " :9 gene-r

ii‘th‘b'la '7
\I he.
\

it" A .
{I an“ 501 3
\

Rafine

.... actual (
.""‘n .‘
“I; 8:101:30

“is (that

GENERIC RELATIONSHIPS

The Andromedeae have long been a taxonomically
difficult group. Most of the species were originally
treated as one genus by Linnaeus and other authors until
the early decades of the nineteenth century when many
segregate genera were recognized. Today the tribe
includes the following genera in the eastern United
States: Andromeda, Leucothoe (including Eubotrys) ,
Gaultheria, Zenobia, Lyonia, Pieris, Cassandra, Oxyden-
c_l_r_‘_1_1_m_, and Epigaea (Wood, 1961).

Rafinesque (1836) recognized the artificiality

of Andromeda as circumscribed by the botanists of his

 

day. He asserted that: ”GENERA ARE NATURAL, and that
every actual Genus that is unnatural, arbitrary, or
polymorphous, IS NOT A GENUS, but an incorrect and
artificial aggregation of aliens!"

McVaugh's (1945) first recommendation in the
recognition of segregate genera is that primary consid-
eration should be given to strong morphological char-
acters (that is, ordinarily, qualitative characters,

or those involving changes in? plant parts, or the

17

 

sauce or a‘oser
rarer than to t]
:‘tlving change
2:: of parts.

‘1'?! there a:
ifferentiating
3:5: recogniz.

:e ‘S W“ A

t—‘yh a nil

-t‘..a1 qroa
.JCSQA

J
fifth-,3
”II" V'
m, 28..
3‘
:i‘fiu
“‘31 aha
I‘.‘
‘ss *- ‘ .
\~Jls‘
10
Ana
... 5‘
' ‘tate

.‘ lZed (.“
...s and 1
“3:5
. 8017‘s;
‘2 3“. I
N‘s:

18

presence or absence of some distinctive attribute),
rather than to the weaker characters such as those
involving changes in number, shape, position or attach-
ment of parts. One of the problems in the Andromedeae
is that there are few strong morphological characters
differentiating the major groups. Thus the number of
genera recognized has varied widely and often seems to
be as much a matter of personal opinion as the result
of natural groupings of taxa. David Don who in 1834
proposed one of the first attempts at a natural arrange-
ment of the genera in the Ericaceae stated:

.As happens in other very natural families, the

Icharacters of the generic groups in the Ericaceae

are not so strongly marked as in those that are

less so; but we are not on that account to give

‘up the idea of dividing them. . . .
His treatment (1834) is quite modern, including the

following genera in the Andromedeae: Andromeda, Cassiopg,

Cassandra, Zenobia, Lyonia, Leucothoe, Pieris, Phyllodoce,

 

Bryanthum.and Daboecia. It is not within the scope of

 

this revision to consider the validity of the various
segregate genera of Andromeda.

Lyonia was one of the first segregate genera
recognized (Nuttall, 1818) . The genus is quite homo-
genous and is separated from all other genera in the
Andromedeae by a definite combination of characters,
although some of these characters appear individually

in other genera. Thus Lyonia fits McVaugh's third

 

:ezzzerdation 't
:3 supposed 9933
3.2.5 another. b

13:7."2 seems to 0

12:3 of the gene:
:arest relative
:32 can also be
\3 (Hood,
.‘Iuch conf
sateen ironic: an

‘1. ...:‘e grOlJp i

it: A.‘¥ "
N ““buOdy (192?

o
.l‘ n

«.225 between at.

':«S) and those
“=5 Spurs). 'r

z 55" v- '
-.-3.ating pl

0
\'
fires
‘ 0f fllar-e
"i:: I

19

recommendation "that the most important criterion of
any supposed genus is not the width of the gap between
it and another, but its own biological unity.” In fact
Lyonia seems to have more distinctive characters than
many of the genera recognized in the Andromedeae. Its
nearest relative is probably Pieris although relation-

ships can also be seen with Zenobia, Gaultheria and

 

 

Cassandra (Wood, 1961).

 

Much confusion has existed as to the differences
between Lyonia and Pieris with some authorities putting
the whole group into one genus. Matthews and Knox (1926)
and Anthony (1927) have doubted that any real distinction
exists between appendages on the back of the anthers
(spurs) and those at the summit of the filament (also
called spurs). This difference was used by Rehder (1924)
in separating Pieris from Lyonia. These spurs are out-
growths of filament or connective tissue and vary con-
siderably in size. In position they range from insertion
on the free portion of the filament to insertion beyond
the point of union of anther and filament, thus appearing
to be attached to the anther lobes (Matthews and Knox,
1926) . Therefore they concluded that these structures
are simply emergences, and cannot be employed as a char—
acter of generic importance. Their contention as to the
lack of a real difference in the position of the appen-

dages in these genera is only partly true. There does

 

see: to be a wide

as :cnfused beca;
:zegration tiss'

Ergir. of the as

-. ¥
‘0 U A“ P

e “31159.54
”.5?

.35 34" k
e hue tr‘i

. r

:‘F. 1
1.! I‘

t. J.

20

seem to be a wide variability in position, but in general
the spurs of Lyonia are at a lower level than those of
Pieris. The appendages of these two genera can never
be confused because those of Pieris lack the white dis-
integration tissue which is always present on the upper
margin of the appendages of Lyonia. Even when the fila-
ments of Lyonia lack the spur-like appendages near the
summit, a line of white disintegration tissue is present
on each anther-half, forming an inverted "V" at the
junction of filament and anther (Wood, 1961; Palser,
1951). The appendages of both Pieris and Lyonia should
not be confused with the awns present on the anthers of

genera such as Zenobia, Gaultheria or Leucotho'é.

 

 

Anthony (1927) also disagrees with Rehder's (1924)
use of the thickened sutures of the capsule as a unique
characteristic of the genus Lyonia. He stated that in
the description of the genus Pieris in the Genera £132.-
tarum (Bentham and Hooker, 1876) the capsule valves are
described as having margins often thickened, and there-
fore concluded that thickened capsule valves are present
in both groups. Actually nothing is farther from the
truth! Bentham and Hooker's treatment of Pieris includes
Lyonia mariana, 2° lucida (in sec. M) and I": o_vg_1_._i_-

folia (in sec. Eupieris) . All these species actually

 

belong in Lyonia, and when the transfer is made the

homogeneity of each group is greatly increased and the

:stinction be1
aims is valic
:9 also diffe:
..f.l:'.llty be ‘
I cons

inc-....bt 5:031

l
o: ‘ k . ‘

 

‘An.“
‘ v-
J:::Ju'

the t“.
1‘: .3.
h.‘ bcne We 1a

SOme b

zaler 99%”

.-
I

we :istributi

Cite»:
sh OUt e‘

21

(listinction between thickened vs. nonthickened capsule
*malves is valid. The inflorescence and leaves of Pieris
‘are also different in general aspect and can only with
difficulty be confused with those of Lyonia.

I consider Lyonia to be a natural group of species
distinct from Pieris and set off by the conspicuously
‘paler, thickened sutures of the fruit, the exclusively
maxillary inflorescences from the wood of the preceding
season, the two bracteoles at the base of the pedicels,
‘and the two large outer scales of the winter buds.

Some botanists have divided Lyonia into several
smaller genera (Britton and Brown, 1913; Small, 1914).
'The distribution of characters used in such cases is so
reticulate, when all the species are considered, that
the segregates are essentially based on single character
(tifferences. The temptation to do this is sometimes
great as the group is an ancient one and many of the
intermedate species are evidently extinct. Yet as
pointed out earlier by K. Koch (1872) , the result of
such a practice in the Andromedeae would probably be
the recognition of almost as many genera as species!

In summary, the group seems to be most natural and
clearly distinctive when separated from E23135, and

not divided into several small genera along sectional

lines.

 

Palser (l9
oray‘n on: V hr
....1. nanes .‘ue.
:‘ze: stud', reco
:fizese taxa by q

szarizing relatl

22

Palser (1951), although she used the segregate
generic names NeOpieris and Xolisma as well as Lyonia
in her study, recognized the similarity in floral anatomy
of these taxa by grouping them together in her table

.summarizing relationships within the Andromedeae.

IE2

As :-
:--:;i- '5'

D. U n

4 .e s s m a a
w. .1 u. e s e s .m
.... d n e x s a e b n l
l. r A. O .1 LL .3
¢ 5 .t k1 C U. ml 3 m ..
a S S +u n . S a . o .4.
.. . a. 3.. 3 A. .3. E . Aw ...—...; R ....u
\nu u:- ‘s: ‘9.» ... u .C. s » a: a. s s . C \
... L. p: as ... ... a . . a:
.. .. ...: .. . .... ..r. 2:. .... .. t. t... a .. 3. ....

 

 

.rr: 3.-

PHYTOGEOGRAPHY AND EVOLUTION

As treated in this study the 30 to 35 species
of Lyonia are classified in four sections. Section

Pieridopsis, comprising three or four species, occurs

 

in Asia from Kashmir to Japan, and south to Malaya. The
greatest diversity of the genus, although not the
greatest number of species, is in eastern North America
where three sections are represented. The monotypic

section Arsenococca is found only in the eastern United

 

States. Section _M_a_r_:i£ includes two species occurring
on the Coastal Plain of eastern North Merica. One of
these species, Lyonia lucida, is also native to Cuba.
Section Lyonia is the largest subgroup of the genus,
containing perhaps as many as 25 or 30 species. These
are mostly natives of the Greater Antilles, although one
is found in eastern and southern Mexico and two occur
in the extreme southeastern United States.

Most of the species in the southeastern United
States are limited to the Coastal Plain; only _I_._. £19.29...
trina is also common in the Piedmont and mountains. In

more tropical areas such as Mexico and the Greater

23

 

 

 

 

:zilles the genus
insulting isola
was areas may h

13’." 1‘

.,....-ion in secti

z.“ua 1S 5

i:::_:cr.ent of the

nu“ ‘f‘
...: ..c .Vr

«.est whiz!

Iriierr. regions .1

I I

:ZILL'Jtl’Qn pane
5:55

a Tertiary r

.....

1191’ Genera
\' Ca

"... C ‘

a: « to en
Esp-\‘Ber‘en S C
1"" :10 0L
733‘33.-n.h93 l
{ksa ACCOra
:“Ses genetic
T: ‘3!“ OUId

~15: Mia?! ‘ ‘
‘ “2': 0

law‘ a?“ g OCC ‘-

24

Antilles the genus is limited strictly to montane areas.
The resulting isolation of populations in various moun-
tainous areas may have been one factor in the extensive
speciation in section Lyonia in the West Indies.

Lyonia is a very ancient genus and was probably
a component of the Arctotertiary Geoflora, a great meso-
phytic forest which once extended around the globe in
northern regions.l Evidence for this is seen in the
distribution pattern of the genus (Figure 1). Lyonia
shows a Tertiary relic disjunction pattern between
eastern North America and eastern Asia. This pattern
was first brought to the attention of American and
European scientists by Asa Gray through a series of
papers published between 1840 and 1878 (Wood, 1972) .
Many other genera show similar patterns, a few of these

are: Calycanthus, Perideridia, Desmodium sect. Prodo-

 

carpium, Illicium, Shortia, and Gelsemium. In the

 

Ericaceae Tripetalea and Elliottia, Pieris, Epigaea,

 

 

 

1Wolfe (1966), however, believes that the concept
of the Arcto-Tertiary Geoflora is not valid. One of the
basic tenents of the Geoflora concept is that the plant
association has maintained itself with only minor changes
in composition for several epochs or periods of earth
history. According to Wolfe "it is extremely improbable
from the genetic and physiological viewpoint that many
lineages could have remained in association throughout
the Tertiary.” He states that "The floristic changes
that have occurred in the vegetation of the Pacific North-
west during the Neogene indicate strongly that the present
associations are also most probably transitory."

25

 

&
‘ .4 . ‘ V..
W .... \u . t. .\
n‘ t o g Q K's. u I N u A.
. I a 9’5 a . " ... n. u
v s . a ., . ..
n ., , . x . C . w .
u ‘ .. .

 

MHCONA mo coflusnfluumflp UHHOB

.H musmflm

 

26

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

252333355 Show
i:s::ibution patte
___a_ (Li, 19

Host of t.
555:9:3 North kte
31-? are also phy
$83165. W39“ t1"
EISiiered. men
533.9 is easily 1

:".:~ . ‘
.U- ~91.“ to faml L

% 1C activl

.ertiar' perioq
wulff (

QEMOPe

:5.“ Seem

Mary flora.

55 leave 0‘

27

(and Chiogenes show similar disjunct distributions. The
(distribution pattern of Pieris is very similar to that
of Lyonia (Li, 1952; Wood, 1972).

Most of the genera showing the eastern Asian-
eastern North American disjunction pattern are woody;
:many are also phylogenetically isolated with only a few
species. When the taxonomic status of these genera is
(considered, their relative antiquity in the phylogenetic
scale is easily seen. A large proportion of the genera
jpertain to families that are considered structurally
primitive, such as the Ranunculaceae, Santalaceae, Mag-
noliaceae, Berberidaceae, Hamamelidaceae, Theaceae, and
Ericaceae (Li, 1952). Over the years paleobotanical
findings have shown that these disjunct genera, espe-
cially the woody ones, once occurred over wide areas
from which they are now missing, and it is clear that
the extant representatives of these genera are the sur-
'vivors of the gradual climatic deterioration, volcanism,
«orogenic activity, and the glaciers that followed the
{Tertiary period (Li, 1952; Wood, 1972).

'Wulff (1943) also points out that the Ericaceae
in Europe seem to be a remnant of a more widespread
frertiary flora. He points out that the xerophytic ever-
(green.leaves of many of the species and the forced winter
dormancy in floral development in £29.51 carnea indicate

'that these species were originally adapted to less

 

agorsus clmat
Exaceae are i
gestures to HZ".
pried from st;
Bell and B; :3
find that the:

'36 i‘lnter sta:

V3

‘~
N ‘e‘aill at,"-

- z
1'“ . p

31519 for
alias“ any ‘

“tied the;
535.: t0 be
its.

"‘1 _
s 1n:
“=5

28

rigorous climatic conditions. Wulff's theory that the
Ericaceae are in "disharmony" with the low winter tem-
jperatures to which many are now exposed seems to be sup-
jported from studies of floral and shoot development

(Bell and Burchill, 1955; Lems, 1962). Bell and Burchill
found that there is a continuous developmental series in
the winter stages of the floral buds in the genera they
studied, and with unusual weather conditions, some member
of the group may be found in bloom during any one of

the 12 months of the year! Active mitosis was observed
in representatives of three genera even in the "depth of
‘winter” when the temperature was below freezing. They
found that "Vaccinium pennsylvanicum [which they studied
in detail] apparently cannot endure exposure to such
‘winter temperatures as commonly prevail in the regions
to which it is native." These facts seem to indicate
that the Ericaceae were evolved in a region where it was
possible for the plants to flower for most of the year
‘without any definite resting stage. Lems (1962) also
concluded that most species in eastern North America
.appear to be imperfectly timed to the alteration of
summer and winter. They have slow morphogenetic cycles
smith only Oxydendrum arboreum leaving no exposed flower
<1r inflorescence buds during the winter. Thus a tropical
<mrigin is indicated for the family and a majority of

‘the species still occur in tropical and subtropical

 

wazs When the
"1:185, differ‘
nr'danjru-ln de'li'e;
”....-

..L ' is
smasher .

...-

;2?‘..‘: cycle, b:

3.3 evolved co:

Les, 1962) .
Althea?

113:3: tempera

I ‘ ‘
«Canon of

I.~.

2;:o_gf1 Lyonia
“““ lament Fa

- T31.
.: fie West In
.1 3111’! three
EY‘Q
" 8 not f
3“

29

areas. When the Ericaceae first encountered summer-winter
climates, different groups evolved different responses.

Oxydendrum developed a shortened cycle, while for others

 

such as Pieris, Leucothoé and Epigaea, flower dormancy

 

 

became a means of survival. Lyonia is even less adapted
to northern regions, taking two complete seasons for one
growth cycle, but in the temperate species protective
buds evolved containing the embryonic inflorescences
(Lems, 1962).

Although we can be sure that Lyonia originated
in warm temperate or tropical conditions, the geographic
location of its origin is unknown. Lems (1962) puts it
'with the Neotropical element of the Andromedeae, a group
that has no close relatives in Asia, but which is con-
nected closely with South and Central American taxa.
.Although Lyonia is diversified in the West Indies this
development may be the result of a more recent adaptive
radiation. This view is supported by the great similarity
(of the West Indian species, all seeming to be variations
«on only three or four basic morphological types. The
«genus is not found in South America and only one species
occurs in Mexico. Lyonia also cannot be strictly Neo-
‘tropical because, as was pointed out earlier, three or
four species occur in eastern Asia. Thus the genus seems
133 have affinities with both the Neotropical and Asian-

Pumerican elements of the Andromedeae.

 

.-

mtehead, 13
Lifese stsi
States bear ]

«5-3335 Of tilt

:-.‘ p“. .
k6. Du.ger re-

[:;ZET‘.Ead (l

:I.." '
Nl‘ “In.

‘mwfinta

. m.
.z'x’“.-v ., ..'
“\m.}; V S

\
the Past

.n‘
3 :> “|
b~ “_

.iej

\a which

«“e
N\ qeglon w
1‘22“.
‘ s
Dis]
3».
“-W4
«“4 sever
n; A. ‘
‘ i‘ahta
.

a
.‘ :vc'fi,. Uri
b \L‘tta,‘ .
...:‘aY ‘10
‘ one
'7‘" ' was
‘1 17:1“. .
V 3311

30

It is evident that the geographic range of
Lyonia has not always been as it is today. There is now
considerable evidence that significant vegetational dis-
placement took place during times of glacial advance
(Frey, 1953; Whitehead and Barghoorn, 1962; Watts, 1969;
‘Whitehead, 1972).2 The full-glacial pollen assemblages
in these studies conducted in the southeastern United
States bear little resemblance to modern pollen assem-
Iblages of the same regions. Components of the modern
flora are not represented, and pollen, spores, seeds
and other remains of northern species are well represented.
Whitehead (1972) in his study of the developmental and
environmental history of the Dismal Swamp states that

Taxodium-Nyssa forests have only characterized the swamp

 

for the past 3,500 years; before that time the region
*was occupied by more northern forest types. Lyonia
lucida which today has the farthest northward extension
(of its range in the Dismal Swamp, probably migrated into
'the region with many other southern species about 3,500
years ago or slightly earlier.

Disjunct populations of E. lucida are found
around several ponds in Bartow Co., Georgia, northwest

of Atlanta. Watts (1969) found that familiar southern

 

2Braun (1947), however, felt that major changes
(If vegetation were limited to the Coastal Plain with the
floral composition of the Appalachian highlands being
only insignificantly affected.

 

:ee genera such a
:ez'lciaity of t'."
E1511: species suc‘r
51:3: appear in t?
::::l:ies that the

c

1“:u..“A§ I
“V ‘M .
,

inclui:
1:129 the Postgl.
fate had a more SI
gazlal period an
-. tie glaciers.
u‘hc \' .
~~ Jed Jersey pi
;: c-L .

we sandy pm
...:J. Lyonia r:
255 A ' -

:5 a ,
e.rly Colon;

“E? 1
L is to th
A
‘h‘
‘t I the

31

tree genera such as Liguidambar and Nyssa migrated into
the vicinity of these ponds in postglacial times. Coastal

Plain species such as Itea virginica and Psilocarya nitens

 

first appear in the fossil record in the Postglacial. He
concludes that they and probably other "Coastal Plain

disjuncts” including Lyonia lucida reached these ponds

 

during the Postglacial. It thus appears that Lyonia must
have had a more southern distribution during the full-
glacial period and has migrated northward with the retreat
of the glaciers. Thus L. mariana, which is abundant in
the New Jersey pine barrens, is also common to the north
in the sandy pine barrens of Long Island (Harshberger,
1916). Lyonia mariana is a characteristic species of
the sandhills in the Carolinas and probably was one of
the early colonizers of the sandy coastal plains when
they were uplifted after the Cretaceous inundation (Duke,
1961), Duke describes plants such as these as being
indigenous to the sandhills, but not endemic there;
rather, they are able to become established in ecologi-
cally similar habitats. Thus E. mariana may have origi-
nated in this region, migrating southward and northward
in.its colonization of the ecologically similar Coastal
‘Plain. Fernald proposed a Virginia-centered classifi-
cation of floral elements that later became distributed
throughout the sandhills area of the fall-line. Duke

(1961) noted that a northern Florida center is more

 

sizingful. Jar.
aata refugium

3223:9035; they
:fiispersal. v
resurarized 1
Tertiary uplif -
lastal Plain s
leeward across
Eli‘s 3ast soc?
Q ShO’u'S é
1152'xzct occur}

1:" v 1 ' ,
"'941 neighbe

l .

.2

32

Iueaningful. James (1961) and Woodson (1947) suggested
that a refugium existed in northern Florida during the
Cretaceous; they also favored a northern Florida center
of dispersal. When the ideas of Duke, James and Woodson
are summarized the following pattern emerges. After late
Tertiary uplift there was a dichotomous radiation of
Coastal Plain species to the middle Atlantic states and

westward across the Gulf Coastal Plain. Lyonia lucida

 

shows just such a dichotomous distribution! Lyonia
mariana shows a clear northeast pattern of migration with
disjunct occurrences in eastern Texas, Arkansas, and
several neighboring states. Possibly at one time it

'was also found on the Gulf Coastal Plain, but for some
unexplained reason became extinct in that region. As the
seeds are light and easily wind blown, long distance dis-
persal is also a distinct possibility. There are a
number of plants which show this dichotomous distribution,

a few are: Pinus taeda, Prunus angustifolia, Desmodium

 

 

strictum, Toxicodendron radicans, Leiophyllum buxifolium,

 

 

 

jBreweria pickeringii, Ruellia carolinensis, Smilax glauca

 

and Croton glandulosus. These and many more are listed

 

Iby Duke (1961) and Gillis (1971).

The distribution of L. ligustrina var. foliosi-

 

flora seems to show a similar pattern, but with addi-
'tional complications (Figure 2). It appears that radi-

ation has occurred in several directions from a point

 

Figure 2.

33

Possible distribution patterns in Lyonia

ligustrina. Note: light arrows represent

 

var. foliosiflora; dark arrows represent

 

var. ligustrina.

 

 

 

 

 

C

34

 

 

 

 

 

 

 

 

 

 

   

 

J.--
----------------o o
-—

 

 

 

 

 

 

 

 

----------o--—o---o-<b"‘

 

 

 

 

 

 

 

i.

“cocoa---------

 

'.: northern Florida.

 

3e Florida peninsu'1
vzthcoriaceous, ex‘
1) 'panicles.' :‘
L: a westward direC'
hisiana, then nor
:ai1ation of distir.
Leaves occurred. 1‘1.
35nd in a northw.

 

..... s

w has Fll

ID the Atlartic C0

 

35

in northern Florida. One line extends southward into
the Florida peninsula and is characterized by individuals
'with.coriaceous, extremely ovate leaves, and large lax
leafy ”panicles.” A second migration probably occurred
in a westward direction along the Gulf Coastal Plain into
Louisiana, then northward into Arkansas where a secondary
radiation of distinct plants having large coriaceous
leaves occurred. A third radiation appears to have pro-
ceeded in a northwestward direction. Evidence from mor-
phological variation indicates that L. ligustrina var.

foliosiflora has migrated in a northeastward direction

 

up the Atlantic Coastal Plain as far as the Dismal Swamp
in southeastern Virginia. In the course of this migration
at least two major forms have diverged, one with glabrous
to slightly pubescent leaves and the second with thick,
very pubescent and often obovate leaves. These two forms
are most distinct in the Carolinas. Lyonia ligustrina var.

ligustrina seems to be most diverse in the southern

 

Appalachian mountains and to have migrated northeastward
from that area following the Wisconsin Ice deterioration.
The patterns of dispersal presented above show many simi-
larities with the phytogeography of the genus Asclepias

presented by Woodson (1947) .

The closely related L. ferruginea and L. fruticosa
are mainly sympatric, occurring in Florida and southeast

Georgia. Lyonia ferruginea extends slightly farther

2::212ward in Georgi
5:252: Carolina, but

5—4

c
:

 

itane areas of W“

P4

:ediately obvious
:elationships betwe-
:- squan‘ilosa? (21
rest 0‘ t1 '

. . ne section

.'..E‘.“3 are many simi

 

«matron of the eel

egerate forests c

 

saith; disjunct d

:M '
Mt: Pl ‘

has strobu
33:3

ilnlana v Far: ‘18
\

‘h: ‘
... nC

£ saccharur
:esms

. -. . Hamamelis

3173‘s
‘dsrls
\

nu smdrp' 1950 ;

36

northward in Georgia and reaches the southermnost tip of
South Carolina, but _L_. fruticosa occurs farther southward

on the Florida peninsula. Lyonia squamulosa is found in

 

montane areas of Mexico. At this point two questions are
immediately obvious: (1) What are the phytogeographic
relationships between the two Coastal Plain species and

.13 squamulosa? (2) What are their relationships with the

 

rest of the section Lyonia in the Greater Antilles?
There are many similarities between the deciduous forest
formation of the eastern United States and the montane
temperate forests of Mexico. Some species and genera
showing disjunct distributions in these two forest types
are: Pinus strobus, Liquidambar styraciflua, Carpinus
caroliniana, Fagus grandifolia and Epifsgus virginiana,

the Acer saccharum group, Nyssa sylvatica, Mitchella

 

repens, Hamamelis virginiana, Cornus florida, Menispermum

canadense, Prunus serotina, Illicium and Taxus (Miranda

 

and Sharp, 1950; Dressler, 1954; Martin and Harrell,
1957; Graham, 1964). Some authorities believe that
these species migrated to Mexico during periods of ice
advance (Dressler, 1954), others (e.g., Braun, 1955)
contend that temperate plants reached Mexico from the
United States at least by the middle of the Cenozoic.
The latter view is accepted by Martin and Harrell (1957);
they concluded that the majority of the Arcto-Tertiary

mesophytes reached the montane Mexican forest by

 

 

:i-Cenozoic at the
Pleistocene the

sotic exchange wit

Exico and Texas.
are mi"

.giated soot:

.2133} a

._ probably has

£2563: I

 

Later, W?

as large distr ib:

sax-in; I; sguamul:

ed from its

Hr“
A=KLat

=_~gEarS improbab l e

‘Wn.

‘ ene WeSt Indi.
«92*;
ed (Malfai

"Ed is
Y on ‘
9
- 80;; L.
a“1 post
keis
' OHCe 1 t
"z-ia
S‘ecldtion
J. the w
est I
: ndie
‘Crls;
M 1C af ihij
. H‘s Oce‘qr

37

mid-Cenozoic at the latest, but they agree that during
the Pleistocene there was probably additional intermittent
biotic exchange with the eastern United States along a
savannah or prairie-border woodland through northeastern

Mexico and Texas. Thus _I_.. squamulosa or its ancestors may

 

have migrated southward into Mexico in the Tertiary when
Lyonia probably had a much wider distribution then at
present. Later, when the arid western regions developed,
this large distribution was broken up into two parts,

leaving _L_. squamulosa "stranded" in Mexico and quite

 

isolated from its relatives on the Coastal Plain! It
appears improbable that 1:. squamulosa migrated into
Mexico during the Pleistocene as it is not adapted to
the dry prairie environment which it would have had to
cross. Although numerous land bridges have been postu-
lated between either Central America and North America
and the West Indies, these areas were probably never
connected (Malfait and Dinkelman, 1972) . Thus the many
species of Lyonia in the Greater Antilles must have
arrived by long distance dispersal, probably by wind,

a reasonable postulation because they have small light
seeds. Once in the Greater Antilles it appears as if
rapid speciation occurred, especially in section Lyonia.
In the West Indies Lyonia shows a definite northern
floristic affinity (Asprey and Robbins, 1953). The

genus occurs in Cuba (5 to 10 species?) , Hispaniola

 

Etc 10 species?) ,
E:e:‘:o Rico (1 spec;

is: Indies and shoe

gercus. Lyoni I

 

seed by its wind-2|

 

.::3 state that t;
33 Rants with wing;

29 “63*. Indies

 

Y" '
“70;“. 1 '
¢ lJCl

E. Eerr'jCinea l

:‘\v-
. v a
' a

 

‘I
st

he sane habitat
2:533 .
\ growing to."

Ella L _
\’ I 1
‘ % a:

fife! u
‘0 Interhen‘
5 t
hese SpecleS
s“::

~Klan
s O The
: ‘ SE") 1
«y a is
. not
”3:33,. ' a

38

(5 to 10 species?), Jamaica (2 species, Adams, 1972), and
Puerto Rico (1 species). Other genera occurring in the
West Indies and showing similar northern affinities are

Magnolia, Juglans, Fraxinus, Pinus, Phyllostylon, Salix

 

 

 

and Quercus. Lyonia probably reached each individual
island by its wind-blown seeds. Asprey and Robbins
(1953) state that there is much evidence that migration
of plants with wind-borne dissemules has occurred in
the West Indies.

Lyonia lucida, 1:. mariana, _I_._. ligustrina and

 

L. ferruginea all are very clearly demarked taxa with

 

sympatric distributions, often even growing side by side
in the same habitats. I often found 2. lucida and _I_._.

mariana growing together in the Carolinas. In southern

 

Georgia in a cut-over pine flatwoods I found 11. ligus-

trina, _Il. lucida and .I: fruticosa all growing side by

 

side! No intermediate individuals are ever observed,
thus these species must possess strong genetic isolating
mechanisms, especially since their flowering periods

overlap. The separation between _L_. ferriginea and E.

 

fruticosa is not as distinct, but in studying over 700

 

herbarium specimens only a few intermediate individuals
could be found. In the field these two taxa are very
distinct. Although they can sometimes be found growing
together, they occur in different habitats and their

flowering times are slightly different. The range of

 

. ~-, 5
:. s..anulosa doe
1: the genus, and '.
szgrificantly in m.

:sclation from the

 

 

39

£1" squamulosa does not overlap with any other species
in the genus, and this species appears to have diverged
significantly in morphological characters in its long

isolation from the rest of the taxa.

 

The flower

 

33rl’Tb-ose fascicle
-;5:ehes of COI'I'T‘L
5-355 Contracted i
sin the axii of

As in nos
mast neristern ha
c.:h

FLOWER AND FRUIT MORPHOLOGY

Inflorescence
The flowers in Lyonia are borne in axillary
corymbose fascicles, racemes, or pseudoterminal panicles
(racemes of corymbose fascicles). The racemes are some-
times contracted into axillary clusters, and each flower
is in the axil of a small leaf or bract, always also

with two lateral bracteoles at the base of the pedicel.

Morphogenetic Cycle

As in most other members of the Andromedeae, the
shoot meristem has a limited life span, and growth in
height is achieved by a succession of lateral buds rather
than by continuous growth of a terminal branch meristem.
The formation of one complete determinate branch with
leaves and flowers may be called the morphogenetic cycle.
In Lyonia (and also Zenobia and Andromeda) a vegetative
1branch.is formed in the spring and early summer which has
‘vegetative buds in the lower leaf axils and flower buds
in the upper leaf axils. Each of these contains an
embryonic raceme in which the individual flowers are as
jyet.incompletely differentiated. No pollen or ovules can

the distinguished before winter. During the second spring

40

 

he floral buds
cluster continue
3216!, with fri
5219!. The $94
:39 caPSule dur
has the morpho

In 1:. :1
1‘9 'v‘esetauve

731311011 in ti
w and
3.330!) of gro‘
Zlgflt Expect f:
‘30 SPeCies Us
:ix‘xl in With

Later than can),

he?

41

the floral buds open and the development of the flower
cluster continues. The flowers are mature by early
summer, with fruit maturation continuing during the
summer. The seeds mature in the fall and persist in
the capsule during the winter or into the third spring.
Thus the morphogenetic cycle takes two full years.

In L. mariana the floral buds are always above
the vegetative buds, but in L. lucida there is more
variation in the location of the vegetative buds. Lyonia

ferruginea and L. fruticosa exhibit less marked inter-

 

 

ruption of growth, branching almost continuously as one
might expect from plants in subtropical climates. These
two species usually have at least a few vegetative buds

mixed in with the floral buds. Lyonia ligustrina flowers

 

later than any other species in the group, probably
because of its more complex inflorescences (Lems, 1962;
and personal observations).

In L. mariana floral dormancy is often prematurely
broken and occasional plants are seen in flower in August
and September. These flowers seem to be normal morpho-
logically. Nuttall was the first botanist to record this
phenomenon, remarking in his Genera (1818): "An extremely
fine, common, and hardy species, possessing somewhat the
habit.of A, nitida when flowering again in the autumn;

its vernal flowering branches are, however, nearly

naked."

The flowe
if other genera i

e :‘etailed study

35 the tribe on (
sification the f ;
2e sepal supply

:9 Presence or a

dsition and cha:

2361 bundles, ‘
3:13‘0risin of .

rd Staminal app

I
fi together

tic SRHEr 91'0“;

w: and

...;s flirthe r poi

.I
u

‘
’n
'13}

-TQ'ZOCCa and
fines Show a c

:5‘
'b‘qnd

ilari

5‘aue .

>4
“‘1' C
0‘ awrls on

lohs.

42

Flower

The flowers of Lyonia are very similar to those
of other genera in the Andromedeae. Palser (1951) made
a detailed study of the organography and vascular anatomy
of this group in which she gives a tentative classification
of the tribe on the basis of floral anatomy. In this clas-
sification the first separation is based on the nature of
the sepal supply. Other important characters used were
the presence or absence of fibers with the septal traces,
position and character of the placenta, position of ventral
carpel bundles, the reflection of the floral parts, the
joint origin of the vascular supply to different organs,
and staminal appendages. She puts all the species of
Lyonia together in one group which she subdivides into
two smaller groups, the first containing L. lucida and

.23 mariana, and the second 2. ligustrina and L. fruticosa.

 

This further points out the distinctness of section Maria,
and the similarities in floral morphology between sections

Arsenococca and Lyonia. She points out that all these

 

species show a considerable nondivergence of vascular
strands to different organs, have horizontally "S”-
shaped filaments, and spurs which show disintegrating
‘tissue. Her statement that they all have a very short
3pair'of awns on each anther is not borne out by my

observations .

 

Tale C
eight fused s
itsed to its

Ltses can be

Lie Calyx 10':
«395, While

4575 Short tr

43

Calyx

The calyx is composed of five or rarely four to
eight fused sepals. Each lobe is nearly free, being
fused to its neighbors only at the very base. The calyx
lobes can be glabrous or densely pubescent. In section
Lyonia they are covered with peltate scales. The calyx
is usually persistent, but it falls with the leaves in
2, mariana. There is marked variation in the length of

the calyx lobes; section Maria has long and lanceolate

 

lobes, while both section Arsenococca and section Lyonia

have short triangular lobes.

Corolla
The corolla varies from cylindric to urceolate
or globose-urceolate, with five (rarely four to eight)
short recurved lobes (Figures 3, 4, 5, 6, 7). It is
'most often white, but occasionally plants with pinkish
flowers are found. It is pink or sometimes even reddish

in L. lucida. In section Arsenococca the corolla is

 

pilose, with club-shaped glandular hairs very similar

to those found on the leaves. In section Maria the
corolla is usually glabrous, although in L, mariana
there are occasionally a few nonglandular hairs, usually
following the veins. Section Lyonia is unique in that
the corolla is covered with peltate scales. The corolla

is urceolate in sections Arsenococca and Lyonia, but is

 

«cylindric to globose-urceolate in section Maria.

 

Figure 3.

44

Drawing of flower of Lyonia ligustrina.

a. Stamen, distal View, X 50; b. Stamen,

proximal view, X 50; c. Flower, X 9;

d. Longi-section of immature capsule,

X 20; e. Cut-away View of flower, X 30.

u

f

t .
O

I

 

 

Figure 4.

46

Drawing of flower of Lyonia ferruginea.

a. Stamen, proximal view, X 43; b. Stamen,
distal view, X 43; c. Flower, X 10; d.
Longi-section of immature capsule, X 20;

e. Cut-away view of flower, X 30.

 

—-

eat
(3 I.

4e \ (

V?{.‘?A

z.

 

 

Figure 5.

48

Drawing of flower of Lyonia fruticosa.

 

a. Anther and portion of filament, distal
view, X 73; b. Anther and portion of
filament, proximal view, X 73; c. Flower,

X 13; d. Longi-section of flower, X 30.

 

 

50

" 7?: ire; . y 1...“,
“Li-F 4'”, E)
f
‘ __ q‘ r:-

{ff-fiv‘
l .

,..._I

. . ,4. -
'4;."
1 . .
" ‘ '0 ‘44‘3 '.
i
2;
‘4.

I’.—— ‘le‘lfimrg‘— .'

n.1‘4‘w‘ 1:» .. a...“
\WW ‘ .- ‘

+2

 

Figure 6. Drawing of flower of Lyonia mariana. if

i
)
Ir

i-

a. Anther and portion of filament, proxi-
mal view, X 34; b. Anther and portion of
filament, distal view, X 34; c. Flower,
X 4.5; d. Longi-section of immature cap-
sule, X 13; e. Cut-away view of flower,

X 9.5.

 

 

 

 

   

                 
       

.Io MI. .0.

I. I
92.310.» «EV/7.... . a
so. us.l\l.u~...¢.~ - .. o l

   

  

51

 

n! .u-fp\0.nv.-o -

.
n L...

 

‘ fiat. \u a .. ...” HIV 0 I... u . .. . Ira ... .... . pills. a I . \
.Ho- noun! ..- a . u . ..\ ... usual .... ~..\ h ‘

... 0-. ~ . n- ... . ..ua slsn~ . .4 . v ..

f . .. I . u . . . . - . ... . o . a I . . V

.I \\ o I ’0.“ o a a a v o . a ‘nl a u
f . 06M.- . at uo~l h.» <- \\.- Vs . s I ... u 6.. I
.fl" m *9) a Ice-cu. . . 1v . h
n .n. a» s

 

Figure 7.

52

Drawing of flower of Lyonia lucida.

a. Anther and portion of filament, distal
view, X 30; b. Anther and portion of
filament, proximal view, X 30; c. Flower,

X 4.5; d. Longi-section of immature capsule,

X 17; e. Cut-away view of flower, X 11.

 

 

 

corolla and
if the Eric
$631.19 by

Peres, and

‘35.“! “90'
:1“..\v-.S ha

54

Androecium

The ten stamens are borne at the base of the
corolla and show most of the peculiar characteristics
of the Ericaceous stamen. They have inverted anthers,
opening by terminal (morphologically basal) elongate
pores, and lack apical awns. The filaments are flattened,
slightly to moderately expanded near the base, "S"-shaped,
glabrous to hairy or roughened, and with or without a
pair of short, spur-like appendages on the back near the
apex (Matthews and Knox, 1926; Anthony, 1927; Palser,
1951; WOod, 1961; and personal observations). The stamens
always have a white line of disintegration tissue on the
back of each lobe extending at least along the apex of
the filament and along the upper edge of the spurs when
present, thus forming an inverted "V" shaped pattern.
(This strange tissue is also sometimes seen on the inside
of the corolla.) The included stamens are arranged in
a tight ring with their pores facing inward and pressed
against the fusiform style (Figures 3, 4, 5, 6, and 7).

In L. ligustrina the lower portion of the filament is

 

jpubescent, and only small inconspicuous spurs are present.
CThese spurs often disappear with age.) The filaments

of _I_._. mariana are pubescent throughout their entire
length; L_. lucida has only roughened filaments, but in

both noticable spurs are found. All the members of

 

 

 

 

 

seetion Lyonia
‘- ve extremely

filaments.

The 0‘
large (especi
enlargement c
nectar secret
the sumit of
straight, and
time is trl

Pilose in 1.

...10a and L

LI. section L

55

section Lyonia native to North America lack spurs (or
have extremely small ones) and have short roughened

filaments.

Gynoecium

The ovary is superior, five locular, the placentae
large (especially in section Maria) and undivided. An
enlargement of the wall at the base usually functions in
nectar secretion. The style arises from a depression at
the summit of the ovary and is columnar to fusiform,
straight, and included (or rarely slightly exerted). The
stigma is truncate to capitate. The ovary is densely

pilose in L. ligustrina, glabrous to nearly so in L.

 

lucida and E. mariana, and densely pubescent and lepidote
in section Lyonia. The ovary is generally subglobose in
L, ligustrina and section Lyonia, but is often flattened-
ovoid in L, lucida and characteristically urn-shaped in

L. mariana.

Fruit
The fruit is a five-angled, subglobose to ovoid,
loculicidal capsule with prominent pale, thickened
sutures, which may separate from the five valves in
dehiscence (Figure 8). The placentae are persistent
at the top of the columella. The capsule of L_. mariana
is iniique, usually ovoid-triangular with a contracted

truncate apex. Sometimes the capsule of _11. lucida shows

 

 

 

 

Figure 8.

56

Variation in capsule morphology and

dehiscence in Lyonia. a.

L. squamulosa,

 

partially dehisced, X 13.5; b. E. ferruginea,

 

just beginning to dehisce, X 13.5; c.

g.

fruticosa, unopened, X 13.5; d. E. fruticosa,

 

 

completely dehisced, X 13.5; e. E. mariana,

just beginning to dehisce, X 8; f. E. lucida,

unopened, X 10; g. E. lucida, completely

 

dehisced, X 10; h. E. ligustrina var. ligus-

trina, unopened, X 8; i. E. ligustrina var.

 

foliosiflora, partially dehisced, X 8.

 

 

 

‘r‘y‘g. I

.~

'95
’i
0'...

57

'1’
'

 

 

    
   
 

   

’.'..¢O

. .“a'ohfi.

  

 

 

ha. .

41.. |

 

ararked resent

is never as dis

section Lvonia

extremely thicj

Less thickened

"‘u'

Ara usually

Marately thi

n‘ . .

*1;- mariana
\

'35 Sutures.

is a unit is n

Oi

«£53

"“R‘er Species,

58

anwrked resemblance to this unusual shape, although it
is never as distinct as in E. mariana. The fruit of
section Lyonia is always lepidote and pubescent with
extremely thickened sutures. The capsule is pilose with
less thickened sutures in section Arsenococca. Section
EEEEE usually has glabrous capsules with slightly to
moderately thickened sutures. The western populations
of E. mariana often have scattered nonglandular hairs on
the sutures. The tendency for the sutures to separate
as a unit is most strongly developed in species of
section Lyonia although it is occasionally seen in the

other species, especially E. mariana.

 

 

All ’c
elliptic flov
tissue at the
large termine

:urved f ilame

Style.
In L.
Pores open is

this Spe
Amity at
aye by the
“he 10mg ha}
teams at
‘5‘L9r

POLLINATION BIOLOGY

All the species have pendulous urceolate or
elliptic flowers, included stamens, and a rim of glandular
tissue at the base of the corolla. The anthers open by
large terminal pores which are directed inward, with the
curved filaments holding the anthers against the fusiform
style.

In E. ligustrina the anthers are mature, and the

 

pores open in buds about to Open. When the filaments
are bent outward and allowed to spring back, the pollen
is discharged. The stigma is at the same level as the
mouth of the corolla, rendering the opening so small
that bees visiting the flower cannot fail to come in
contact with it. It appears to be receptive in newly
opened flowers (Lovell and Lovell, 1935). The Lovells
also observed bumblebees sucking nectar from the flowers
of this species. The nectar accumulates in considerable
quantity at the base of the flower and is probably held
there by the densely pilose pubescence of the ovary and
the long hairs at the base of the filaments. The bee's
tongues at first passed between the curved filaments, but

later, as the bees turned half to three-fourths of the

59

 

 

 

 

 

say around the
:he filaments
exsiusively bj
The b1
bybr‘ashing t]
he dusted on ;
carried to an:
IZlCh almost I
ise Vibrathn
to be dUStEd (
Fillination CC
1'26 anthers tc
LOvel]

hash fOr two i
received sever
Mala“. and
{Bits in One
Judgir

60

vmy around the flower, their tongues passed outside of
the filaments. The flowers were pollinated almost
exclusively by bumblebees.

The bumblebees probably accomplished pollination
by brushing the curved filaments, thus causing pollen to
be dusted on its proboscis. The pollen could then be
carried to another flower and deposited on the stigma
which almost blocks the small corolla opening. Possibly
the vibration of the bee's wings also causes the pollen
to be dusted on the stigma and the bee itself. Self—
pollination could easily occur by pollen falling from
the anthers to the stigma below.

Lovell and Lovell (1935) in observing a large
bush for two hours noted that all the flowers probably
received several visits. Bombus ternarius was quite
abundant, and one speciman was seen to make sixteen
visits in one minute.

Judging from their similarity in floral morphology,
the pollination mechanisms of the other species in the
genus are probably quite similar to that of E. ligustrina.
Honeybees have been observed visiting the flowers of E.

ferruginea in northern Florida (Richard Harris, oral

 

communication).

The long spurs present on the filaments of E.
mariana and lucida possibly also function in pollination.

These spurs stick out toward the corolla and when the

 

grcboscis of a
flower it may
causing the pc
mechanism is c
sgecies of E
Late corollas,

LOVel

Y I -
\

 

The netted pa
323?. as many

61

proboscis of a bee is inserted into the bell-shaped
flower it may strike against one of these processes,
causing the powdery pollen to fall out. A very similar
mechanism is described by Knuth (1909) for several

species of Vaccinium which also have pendulous, urceo-

 

late corollas and prominent spurs.

Lovell and Lovell covered several panicles of

E. ligustrina with a fine netting to exclude all insects.

 

The netted panicles produced many seed capsules, though
not as many as from clusters of flowers which had not
been covered. Thus both self- and cross-pollination are

probably important in this Species.

 

 

 

The E:
ifications in
13.93rtance in
C'I'course, tfv
lzclasSifl’in
f‘lLCtion in t
tlll‘sate and h
filth the hate
:Etranspirat

COWar

EP I DERMAL APPENDAGES

The Ericaceae as a family show many curious mod-
ifications in trichome structure, which often are of
importance in determining relationships among species.

Of course, the trichomes are not there just to help us

in classifying! Trichomes probably have an important
function in the economy of the plant in its relation to
climate and habitat. They may be intimately concerned
with the water balance of the plant and in the regulation
of transpiration.

Cowan (1950) pointed out that the Rhododendrons
of higher altitudes, in order to survive the rigors of
winter, or periods of drought, must either severely limit
or arrest transpiration for a long period. This is
accomplished in some species by a dense covering of
hairs, but most are equipped with peltate scales. Thus
he concluded that scales when closely packed furnish the
plant with a means of restricting transpiration even more
efficiently than a covering of hairs. When these scales
are so close together that the rims of adjacent ones

overlap, a protective canopy is formed under which

62

 

 

 

transpiration
scales in Rhoc
tire function
the leaf and l
Iowan, 1950)
correlation b
:at. For exa
:zdes tend to
scales. He c
have varied 8
Efficient the

."\v
Ru

63

transpiration and gas exchange can occur. The peltate

scales in Rhododendron may also provide a water absorp-

 

tive function, by trapping the water of rain and dew on
the leaf and holding it there until it can be taken in
(Cowan, 1950). Cowan noted that there is a definite
correlation between the type of pubescence and the habi-
tat. For example, epiphytes and species from high alti—
tudes tend to be more densely covered with hairs or
scales. He concluded that "these diverse structures
have varied and contrasting functions but none is more
efficient than the well-known Rhododendron scale, a
minute organ, wonderfully made."

The Rhododendrons are not alone in their possession
of such an adaptation. Peltate ferrugineous scales quite
similar to those of Rhododendron are present in E. £25:
ruginea, E. fruticosa and E. squamulosa. In fact, the
presence of such scales is a diagnostic feature of sect.
Lyonia. Their function is probably very similar to that
of the peltate scales of Rhododendron described above.
These three species often occur in xerophytic conditions.
Eyonia fruticosa, especially, is abundant on the sandy
plains of southeast Georgia and Florida where almost
every plant shows at least some xerophytic adaptations.
Probably one of the reasons why E. fruticosa can prosper
under such harsh conditions is because of the possession

of this protective covering of scales. Lyonia ferruginea

 

 

 

 

has the additii
Leaves. Em
:‘tese plains,

restricted to
srons or the «

The c
:‘rc‘Dibly also
interest that
are generalh

NlEd1
tense of the
aClassifica
artificial.
'a‘it'n M

scales. He

64

has the additional protection provided by its revolute

leaves. Lyonia lucida, another evergreen Lyonia on

 

these plains, lacks peltate scales and is usually
restricted to more moist sites, such as slight depres-
sions or the edge of ponds or streams.

The club-shaped glandular hairs of E. ligustrina
probably also function in water regulation. It is of
interest that the mountain populations of this species
are generally quite pubescent.

Niedenzu (1890) used the leaf anatomy and pubes-
cence of the Arbutoideae and Vaccinioideae to construct
a classification for these groups. The result is rather
artificial. For example, he classified E. ferruginea
xvith Cassandra calyculata because they both have peltate
scales. He gives illustrations of the peltate scales of

Ii' ferruginea and the stalked glands of E. ligustrina.

 

I found each section of Lyonia as represented in
herth America to have a distinctive pubescence type.
Itikey is given below which brings out the distinctive
features of each group. Illustrations of each trichome

type are given in Figures 9-12.

Key to sections and species of Lyonia based on

trichome type

 

1x. Peltate scales present on leaves (Section

Lyonia) . . . . . . . . . . . . . . .

 

 

65

Papillae-like hairs abundant or moderate, pel-
tate scales usually not dark red . . . . . . C
C. Papillae-like hairs abundant (rarely
absent), scales mostly deciduous thus giving
older leaves a whitish appearance
beneath . . . . . . . . . E. fruticosa
C. Papillae-like hairs moderate, scales not"
deciduous, thus older leaves rusty colored

beneath . . . . . . . . . L. ferruginea

 

Papillae-like hairs absent (rarely present), pel-
tate scales dark red to almost transparent

. . . . . . . . . . . . . E. squamulosa

 

Peltate scales absent from leaves . . . . . . . D

D.

Leaves with multicellular, clear or white,
elongate, club-shaped hairs, sometimes also with
filiform hairs intermixed; corolla pilose

(Section Arsenococca). . . . . . E. ligustrina

 

 

Leaves with small, rounded, red-tipped, stalked
glandular hairs, glandular tip sometimes drawn out
into a definite point, filiform hairs often along
major veins; corolla nearly glabrous (Section
M)...............E
E. Glandular hairs all with small rounded tips,
reddish, usually with filiform hairs along

major veins beneath . . . . . . E. mariana

 

E. Gla
gre
elc
ha:

SU

 

 

66

Glandular hairs fewer, usually deciduous with
greater age, of various types with rounded to
elongate tips, reddish to white; filiform
hairs only on midvein of upper leaf

surface . . . . . . . . . . E. lucida

 

 

 

 

Figure 9.

67

Variation in glandular hairs in Lyonia;

darker shading indicates a reddish color.

a-b.

E. mariana, ca 160 u long;

ligustrina, ca 525 u long; d-i.

 

100-400 u long.

c. L.

E. lucida,

 

 

Figure 10.

69

Variation in size and shape of fer-

rugineous scales of Eyonia ferruginea

 

(ca 0.20 to 0.75 mm wide).

 

        

O
. . on .. .Ii .‘. .-
..u. ......r....:. ..
i . .. . .
I . .. \ ...\.
o
a u
n f“
on-aooao
I
a
. .91.
u. .
\ .\\ \o
'5‘ c
u
. .
. . o
I I .
\l.\ u
‘ O
‘ C
...alu..lu-|\

‘IO'II‘OO

u o
. u
., 0.
u.... at...
.Illv..00n..oucf t
b
O'...’ I .
0
.ID 0
l
to.
I
O ‘
u \
o o n-
o o e
o o u.
a. fell.
a
(-0
9
0
I
I

71

 

A.n CH wmcflmsuumm .m m0 mafia: Euomwafim umHfiEwm mnu muozv

 

.mcoa 2 omH mo .mmoowusum am no momwnsm mama Hmwxmnm so muwmn

 

EHOMfiHflm .o unmfln n oma mo .mmOOfiusum .m mo mamom msosowomn

 

mo uncenomuum Human mo mchEmm .0 unwound 1 00¢ mu .mmcfimsunmm
.m mo mommusm mama Hafixmnm c0 mmamom msomcwmsuumm m0 mGOHuomm

Iflmcoq .Qnm .masomq coauomm ca momma Hams mo m3mw> Hmcwosuwmcoq

 

.HH ousmflm

 

Figure 12.

73

Hair types in Sections Maria and Arseno-
cocca. a. External view of filiform

hair of Lyonia lucida, ca 200 u long;

 

b. Longi-section of filiform hair of

E. ligustrina, ca 300 u long; c. Longi-

 

section of filiform hair of E. mariana,
ca 400 u long; d. Longi-section of
glandular hair of E. lucida, ca 150 u

long.

 

 

 

Pro)
eaten. The
PUison andr
:0 kill She
Stagger-bu5
leaves on

in Old Pas
Scraping

I“
\.S\Qc

no ,
x “This“

ECONOMIC IMPORTANCE

Probably all species of Lyonia are poisonous when
eaten. The leaves of E. mariana contain the narcotic
poison andromedotoxin, C31H51010, and have been known
to kill sheep. In fact it received its common name,
stagger-bush, because of the intoxicating effect of its
leaves on sheep and cattle (Pammell, 1911; Muenscher,
1939)! In Arkansas where E. mariana is locally abundant
in old pastures, cut-over woods, sandhills, and along
road sides, farmers have tried to eradicate it by
scraping it up, etc. All such attempts have been
unsuccessful (Delzie Demaree, oral communication).

The extensive underground rhizome network produced by
E. mariana probably accounts for the difficulty in its

removal.

Lyonia ligustrina is also said to cause livestock

 

Ix:isoning, producing staggering and disorientation in
goats. Usually the major problem is in early spring
when it is one of the few things in leaf (Correll and

.Johnston, 1970; Muenscher, 1945; Delzie Demaree, oral

communication) .

75

 

 

Pannel?
species‘ of EL
these, but sir
bush probably
of L. ovalifo

- \
”ElaY Peninsu
Scats (Sarge:

An i
a“'*'910Y‘3<1 ext
diseases! in
insects (Sat

have also b
states; up
bdhdsgme f(
in amlDle c
recogniZed
with its 1
has a tent

1V0: '

Geekiful

76

Pammell (1911) states that "all three [sic]
species” of Lyonia are poisonous. He does not list
these, but since E. lucida is also often called stagger-
bush probably it, too, is poisonous. The leaves and buds

of E. ovalifolia, a small tree of the Himalayas, the

 

Malay Peninsula, China, and Japan, are poisonous to
goats (Sargent, 1893).

An infusion of the leaves of E. ovalifolia is
employed externally in the treatment of cutaneous
diseases, and the young leaves are used to destroy
insects (Sargent, 1893). Several species of Lyonia
have also been used as ornamental shrubs. Sargent
states: ”Most of the species of Andromeda produce
handsome foliage and beautiful flowers often arranged
in ample clusters, and their value as garden plants is
recognized in all temperate regions." Lyonia mariana
‘with its large white flowers is especially beautiful but
has a tendency to have a rather straggling growth.
JLyonia lucida and E. ferruginea also would make very
Jbeautiful ornamentals, and more study should be directed

toward their growth and culture.

NUMERICAL TAXONOMIC STUDIES

The use of electronic computers in the analysis
of large amounts of comparative data has become a powerful
and popular technique in plant systematics. The field
of numerical taxonomy began in the 1950's and has gained
‘widespread attention since the publication of Sokal and
Sneath's Principles of Numerical Taxonomy in 1963. It is
still somewhat controversial with various people advocating
one or another of the many methods that have been proposed
since the 1950's. Two major viewpoints exist: the 2E3:
23523 school and the phyletic school. The followers of
the phenetic school believe that numerically generated
classifications should reflect relationships based on
‘as many characters as possible and should not take phylo-
geny into account (Hsiao, 1973; Sokal, 1963; Sokal and
Sneath, 1963; Sokal and Crovello, 1970). In contrast,
the followers of the phyletic school believe that quanti-
'tative evolutionary schemes are the primary goal (Kluge
,and.Farris, 1969; Cain and Harrison, 1960; Farris, 1970;

Whiffin and Bierner, 1972; Wagner, 1962, 1969; Mayr, 1965).

77

In t]
approaches w

Ilith the 0th

Clu:
malfléd ve
3330313 is a
Ii.--

78

In this study both the phenetic and the phyletic
approaches were tried. The results of each are compared

with the other and with previous classifications.

Phenetic Studies

Cluster analyses were performed using a slightly
modified version of program HGROUP of Veldman (1967).
HGROUP is a hierarchical grouping program which utilizes
a generalized distance function based on the concept of
error sum of squares or within-group variance (i.e., the
sum of squared deviations from group means). The program
uses the total within-groups variation as the function to

be minimallyincreased at each step in the grouping pro-

 

cess. Principal components analyses were made using
program FACTOR of Veldman (1967). This program reduces
a set of variables that were used to gather data (e.g.,
leaf length, corolla color) to a smaller set of new,
uncorrelated variables which are defined solely in terms
of the original dimensions, and which retain the most
”important" information contained in the original data.
'The process can be thought of as the construction of a
:neW'space which is maximally representative of the space
«defined by the original variables (Veldman, 1967).

All programs were run on the CDC 6500 computer
at Michigan State University.

Taxa for numerical analysis were first selected

‘by conventional herbarium methods (i.e., species were

 

 

 

sorted “to 1
within and d
variable E.
three 9609“
an Operatic
northern va
western POE
an indepenc
considered
seven taxa
A
from the I
examinatiI
acters th
specimens
measured.
character
such as 1
presence
be invar
fliese We
“3 (Tab

Chained

m'
"&

tation

-L

79

sorted into sets showing continuous patterns of variation
within and discontinuous patterns among sets). The widely

variable E. ligustrina var. foliosiflora was divided into

 

three geographical regions, each of which was treated as
an Operational Taxonomic Unit (OTU). The more homogeneous
northern variety was treated as a single OTU; the disjunct
western populations of E. mariana were also considered

an independent OTU. In all other cases each species was
considered to be an OTU. In all, ten OTU's representing
seven taxa were selected (Table 3).

A tentative list of characters was abstracted
from the literature, and many others were added after
examination of specimens from all taxa. The 144 char-
acters thus obtained were measured on each of 10 to 84
specimens of each OTU. In all, 366 specimens were
:measured. The measurements were continuous in such
characters as length or width, multistate in features
such as color or degree, and dichotomous in cases of
;presence or absence. Several characters were found to
Ibe invariant, or were difficult to measure with accuracy;
these were eliminated, leaving 128 characters for analy-
sis (Table 4). The specimens from which the data were
obtained are indicated by an asterisk (*) in the specimen
citations of the taxonomic treatment.

Cluster analyses of the OTU's were carried out

using all 128 characters and also the 79 vegetative

 

 

LIGM

LIGE

LIG]

MAR

LUQ

80

TABLE 3. List of OTU's of Lyonia, giving the code name of

each.

 

Code Name

Taxon

 

FRUT

FERR

SQUA

LIGW

LIGM

LIGE

LIGL

MARW

LUCI

onia fruticosa

)3

. ferruginea

 

k* lb

. squamulosa

 

L. ligustrigg var. foliosiflora
Tbklahoma, Texas, Afkansas,
Louisiana)

 

 

L. ligustrina var. foliosiflora
YTennessee, Mississippi, Alabama)

 

 

L. ligustrina var. foliosiflora
TFlorida, and northward on
coastal plain)

 

 

E. ligustrina var. ligustrina

 

 

L. mariana
Twestern disjunct populations)

L. mariana
Teastern Coastal Plain)

E. lucida

 

*2an 4. en
(E

%

C}

x

1. VEGETAT

1. Hab
2. Nun
one
[rei—
{Characters
"4. 3. Lu:
4 Lo
5. De
la
me
6. Le
ir
7, 0‘
m
i]
8' DI
d1
h.
(
9. L.
i:

 

81

TABLE 4. Characters used in numerical taxonomic study
(phenetic method).

 

Character Character States

 

I. VEGETATIVE CHARACTERS

1. Habit 1. Small shrub (1' to 4')
2. Large shrub (5' to 10')
3. Small tree (over 10')
2. Number of nodes in *
one year's growth As actual measurement

(Characters 3 through 17 refer to twig characters)

3. Luster l. Dull
2. Somewhat shiny
3. Very shiny
4. Longitudinal ridges 1. Absent
2. Slightly angular
3. Very angular
5. Density of nonglandu- 1. Glabrous
lar hairs at inter- 2. Sparse
nodes (mature) 3. Moderate
4. Dense
6. Length of hairs at 1. Short
internodes (mature) 2. Medium
3. Long
7. Density of ferrugi- 1. No scales
neous scales at 2. Sparse
internodes (mature) 3. Moderate
4. Dense
8. Density of glandular l. Glabrous
and nonglandular 2. Sparse
hairs at internodes 3. Moderate
(immature) 4. Dense
9. Length of hairs at 1. Short
internodes (immature) 2. Medium
3. Long

TABLE 4. Cc

Cl

“

10. Den:
neo
int
(in

11- Col
int

12. Ra

13- Si

14. c

15,

lg.

17

I
[3.

3“: at

is.

82

 

 

TABLE 4. Continued
Character Character States
10. Density of ferrugi- 1. No scales
neous scales at 2. Sparse
internodes 3. Moderate
(immature) 4. Dense
11. Color of scales at 1. All or most dark orange
internodes (mature) 2. All brown or black
3. Half orange, half white
or clear
4. All white or clear
12. Rato of small to 1. All small
large scales 2. Some large, mostly small
(mature) 3. Half and half
4. Some small, mostly large
5. A11 large
13. Shape of scales 1. Small and regular
(mature) 2. Large and regular
3. Small and irregular
4. Large and irregular
14. Color of scales at 1. All dark orange
internodes (immature) 2. All brown or black
3. Half and half
4. All white or clear
15. Shape of scales 1. Small regular
(immature) 2. Large regular
3. Small irregular
4. Large irregular
l6. Persistence of 1. Persistent
scales 2. Deciduous
17. Texture of older 1. Smooth
twigs 2. Somewhat rough
3. Shredding bark

(Characters 18 through 25 concern

18.

Shape

l.
2.

the winter buds)

Round
Elongate

 

it

ll
22
2

 
 

83

 

 

TABLE 4. Continued
Character Character States
19. Apex 1. Rounded
2. Sub-acute
3. Acute
20. Length *Measured in mm
21. Diameter at widest *Measured in mm

22.

23.

24.

25.

26.

(Characters 27 through 76 refer to

27.

28.
29.

point

Rato of length to
diameter

Density of hairs on
bud

Density of scales

Color of scales
Variance in leaf

size

Base

Angle made by base

Serration type
(distal portion)

*Calculated from 20 and 21

PI sawed .boawrd obbJNFH
O O O O O 0 O O O O I 0

LAN
0

I'U'IthNH
ooooo

Glabrous
Sparse
Moderate
Dense

None
Sparse
Moderate
Dense

Dark orange
Light orange
Clear

Leaves becoming very
reduced

Moderately reduced
Not reduced at top of
branch

the leaves)

Attenuate
Cuneate
Obtuse
Truncate
Cordate

Measured in degrees

1.
2.
3.

Entire
Serrulate
Serrate

 

TABLE 4.

Ce

%

Ch

K

30.

31.

32.

33.

34.

35.

35,

37

38.

39

40.

41
42.

Serr
(prc

Revc
(prc

Revc
(die

Per:

Nunu
per
(prc

per
(dis

Len:
(prc

Len:
(di:

Hei.

Thi.

84

 

 

TABLE 4. Continued
Character Character States
30. Serration type 1. Entire
(proximal portion) 2. Serrulate
3. Serrate
31. Revolute margin 1. Not revolute
(proximal) 2. Slightly revolute
3. Very revolute
32. Revolute margin 1. Not revolute
(distal) 2. Slightly revolute
3. Very revolute
33. Persistence 1. Deciduous
2. Evergreen
34. Number of serrations *Actual number counted
per unit length
(proximal)
*
35. Number of serrations Actual number counted
per unit length
(distal)
*
36. Length of serrations Measured in mm
(proximal)
37. Length of serrations *Measured in mm
(distal)
38. Height of teeth 1. Very short
2. Medium
3. Long
39. Thickness of blade 1. Thin
2. Moderately coriaceous
3. Very coriaceous
40. Apex 1. Obtuse
2. Mucronate
3. Acuminate
4. Acute
41. Angle made by apex *Measured in degrees
*
42. Length Measured in mm

«)6.

47.

48,

49,

so,

51.

Rat

Rat
ler

Co

Lu

85

 

 

TABLE 4. Continued
Character Character States
43. Length from base to *Measured in mm

44.
45.

46.

47.

48.

49.

50.

51,

widest point
Width at widest point

Ratio of length to
width

Ratio of length to
length at widest
point

Color above

Luster above

Density of non-
glandular hair on
veins below

Density of non-
glandular hairs on
veins above
(mature)

Length of non-
glandular hairs on
veins below (mature)

* .
Measured in mm

*Calculated from 42 and 44

*Calculated from 42 and 43

WM
000 o

WNH WNH

Light green
Medium green
Dark green

Dull
Moderately shiny
Very shiny

No pubescence

Only hairs on mid vein
Slight pubescence on
mid vein and lateral
veins

Moderate or quite
pubescent on mid vein
and lateral veins

Mid vein glabrous but
pubescent on lateral
veins

No pubescence

Slight pubescence on
mid vein

Moderate pubescence on
mid vein

Very pubescent on mid
vein with some pubes-
cence on laterals

Short
Medium
Long

TABLE 4.

52.

S3.

54.

 

55.

se.

59.

86

 

 

TABLE 4. Continued
Character Character States
52. Length of nonglandular 1. Short
hairs on veins above 2. Medium
(mature) 3. Long
53. Density of glandular l. Glabrous
and nonglandular hairs 2. Sparse
on blade below 3. Moderate
(mature) 4. Dense
54. Length of glandular 1. Short
and nonglandular 2. Medium
hairs on blade below 3. Long
(mature)
55. Density of scales on 1. None
blade above (mature) 2. Sparse
3. Moderate
4. Dense
56. Density of scales on 1. None
blade below 2. Sparse
3. Moderate
4. Dense
57. Color of scales on 1. All brown or black
blade below 2. All or most dark orange
3. Half orange half white
4. All white or clear
58. Ratio of small to 1. All small
large scales on 2. Some large, mostly small
blade below (mature) 3. Half and half
4. Some small, mostly large
5. All large
59. Ratio of small to 1. All small
large scales 2. Some large, mostly
(immature) small
3. Half and half
4. Some small, mostly
large
5. All large

87

 

 

TABLE 4. Continued
Character Character States
60. Density of scales on 1. None
leaf blade above 2. Sparse
(immature) 3. Moderate
4. Dense
61. Shape of scales 1. None
(mature) 2. Regular
3. Irregular
4. Both
62. Shape of scales 1. None
(immature) 2. Regular
3. .Irregular
4. Both
63. Persistence of small 1. Persistent
scales on lower sur- 2. Partly deciduous
face 3. Deciduous
64. Persistence of large 1. Persistent
scales on lower sur- 2. Partly deciduous
face 3. Deciduous
65. Glandular hairs on 1. Yes
blade above 2. No
66. Average size of *Measured in mm
scales
67. Prominence of major 1. Not prominent
veins below 2. Moderately so
3. Very prominent
68. Curvature of blade 1. Curved upward
2. Flat
3. Curved downward
69. Prominence of major I. Not prominent
veins above 2. Moderately so
3. Very prominent
70. Depression of veins 1. Depressed
above 2. Level with surface
3. Raised

 

TABLE 4. C

71. Ma]

72. Le:

73. Ra

74. De

75. Le

76- De

77. Pc

88

 

 

TABLE 4. Continued
Character Character States
71. Marginal vein present 1. Yes
2. No
72. Length of petiole *Measured in mm
73. Ratio of length of *Calculated
blade to length of
petiole
74. Density of non- l. Glabrous
glandular hairs 2. Sparse
on petiole 3. Moderate
4. Dense
75. Length of hairs on 1. Short
petiole 2. Medium
3. Long
76. Density of scales on 1. Absent
petiole 2. Sparse
3. Moderate
4. Dense

(Miscellaneous vegetative characters)

77.

78.

79.

II.

80.

Position of branches

Water relations

Location

Inflorescence type

1.
2.
3.
4.

1.
2.
3.

Rigidly ascending
Arching

Only somewhat arching
No particular orien-
tation

Growing in standing
water

Moist soil

Dry soil

Coastal Plain, Pied-
mont or mountains
Coastal Plain
Tropical mountains

FLOWER AND INFLORESCENCE CHARACTERS

Reduced raceme (fascicle)
Raceme
Panicle

 

TABLE 4 .

31. Nu
ir
32. Re
ta
83. r;-
m
m
84. N
h
e
85. 1
1
86, 5
87, <
88, 1
89.
90.
91.
92.
91

89

 

 

TABLE 4. Continued
Character Character States
81. Number of flowers per *Actual number counted
inflorescence
82. Relationship to vege- 1. Flower buds always
tative growth above
2. Some mixing
3. Flower buds below
vegetative buds
83. Number of panicles 1. Does not apply
making up pseudoter- 2. Actual number counted
minal inflorescence
84. Number of leafy *Actual number counted
bracts per inflor-
escence
*
85. Total length of Measured in mm
floral bracts
86. Shape of corolla l. Urceolate
2. Elongate
3. Ovoid-elongate
87. Color of corolla 1. White
2. Pink
3. Red
88. Pubescence of 1a Glabrous
corolla 2. Sparse hairs
3. Dense hairs
4. Scales only
5. Scales and hairs
39. Length of corolla *Measured in mm
*
9C’. Width of corolla Measured in mm
91. Ratio of length to *Calculated
width
92. Length from base to *Measured in mm
widest point
93. Length of calyx *Measured in mm

lobes

T‘LE 4. 4

%

Cl

‘——_

94. Ra

95. Ni

96. Ra

 

97. PU

98, Pt

99- Pe

no. L.
101.

102,

103.

104

 

90

TABLE 4. Continued

 

Character Character States

 

94. Ratio of length of *Calculated
corolla to length
of calyx lobes

 

95. Width of calyx *Measured in mm
lobes
*
96. Ratio of length to Calculated
width of calyx .
lobes __
97. Pubescence of calyx l. Glabrous
(abaxial) 2. Sparse hairs
3. Dense hairs
4. Scales only
5. Scales and hairs
98. Pubescence of calyx 1. Glabrous
(adaxial) 2. Sparse hairs
3. Dense hairs
4. Scales only ,
5. Scales and hairs
99. Persistence of l. Deciduous
calyx 2. Partly deciduous
3. Persistent
100. Length of filaments *Measured in mm
101. Curvature of fila- 1. Straight
ments 2. S-shaped
3. Several curves
102. Pubescence of fila- 1. Smooth
ments 2. Spines
3. Spines and long hairs
on lower portion
4. Long hairs covering
entire filament
103. Length of anther 1"Measured in mm
104. Spur length 1. Spurs absent

2. Short spurs
3. Long spurs

 

 

NUS 4. C

107. 5‘:

 

112. T

or, PM

114. ‘

115

115

91

TABLE 4. Continued

 

Character Character States

 

105. Length of style *Measured in mm

 

106. Pubescence of ovary l. Glabrous

2. Sparse hairs

3. Dense hairs

4. Scales only

5. Scales and hairs
107. Shape of ovary l. Subglobose

2. Ovoid

3. Elongate-ovoid ‘
108. Length of pedicel *Measured in mm

I * O
109. Length of bractioles Measured in mm

110. Pubescence of pedicel l. Glabrous
2. Sparse hairs
3. Dense hairs
4. Scales only
5. Hairs and scales
111. Width of bractioles *Measured in mm
112. Time of flowering l. March-April
2. May-June
3. July-August
4. September-October
5. November-February
III. FRUIT AND SEED CHARACTERS
* .
J~13. Length of capsule Measured in mm
114. Width of capsule *Measured in mm

 

115. Ratio of length and *Calculated
width

1. Oval to elongate-oval

2. Subglobose

3. Ovoid with a flattened
top

4. Urn-shaped

116. Shape of capsule

 

 

TABLE 4.

 

119.

120.

126,

127.

 

 

92

 

 

TABLE 4. Continued
Character Character States
117. Thickening of sutures 1. Slightly thickened
2. Moderately thickened
3. Very thickened
118. Pubescence of capsule l. Glabrous
2. Very slight pubescence
3. Moderate pubescence
4. Very pubescent (hairs)
5. Scales only--sparse
6. Scales only--dense
7. Scales and hairs
119. Color of capsule l. Yellow-tan
2. Light brown-orange (or
greenish)
3. Dark brown-brown
120. Separation of 1. Not separating
sutures from valves 2. Partially separating
of capsule 3. Sutures completely
separate
121. Number of capsules *Actual number counted
per inflorescence
122. Number of seeds per *Actual number counted
capsule
123. Length of seeds *Measured in mm
124. Width of seeds *Measured in mm
125. Ratio of length to 1”Calculated
width
126. Color of seed 1. Yellowish tan
2. Brown
3. Dark brown
127. Depression of apex 1. Slightly depressed
of capsule 2. Moderately depressed
3. Very depressed
*
128 Length of pedicel Measured in mm

\

(on mature capsule)

 

 

 

 

‘I'l

 

characters
33 floral .
characters
loo-charac
acters we:
Vere found
Here elim;
was also

Finjures 1
tea OTU‘ s
the 108-c
9139110ng
tive, fr]
Table 5 .
based up
respondi
Tables E
Pornon

Show“ is

93

characters, the 49 fruit and floral characters, and the
33 floral characters. A cluster analysis of the 128
characters themselves and a factor analysis of a similar
loo-character set were also made to determine which char-
acters were most closely correlated. Several characters
were found to be highly linked, a good portion of these
were eliminated, resulting in a 108-character set, which
was also analyzed by cluster analysis. The phenograms in
Figures 13 and 14 represent the cluster analysis of the
ten OTU's for the complete (128-character) set, and for
the 108-character set. Figures 15, 16 and 17 show
Phenograms produced in cluster analyses of the vegeta-
tive, fruit and floral, and floral data sets, respectively.
Table 5 gives the distance between each pair of OTU's
based upon analysis of the 108-character set. Cor-
responding values for the other data sets are found in
Tables 6, 7, 8, and 9. A graphic representation of a
Portion of the information presented in these tables is
Shovm in Figures 18, 19, 20, and 21.

The phenograms resulting from the 128 and 108
chali‘acter sets are very similar. The phenogram based on
vegetative characters is also quite close to these two.
All the phenograms indicate that the genus consists of
three distinct groups. Lyonia ferruginea, E. squamulosa
and E. fruticosa are closely clustered in the first

group (section Lyonia), E. ligustrina is by itself in

 

 

 

 

 

 

 

 

 

Figure 13.

94

HGROUP phenogram of Lyonia based on 128
character set. Units on axis indicate
the sum of squared deviations from group

means or error index.

‘ 39

 

95

FERR SQUA FRUT LIGM LIGE

 

L__J

LIGW LIGL MARE MARW

 

 

 

 

 

 

100

200

’_.___-._——

i 300

360

480

540

 

—————-—-—’—-——-—'—-—-——

 

E—JL

, —.__.___._._ ...—_—

 

_-—_—

 

 

 

 

 

LUCI

 

 

 

 

 

96

Figure 14. HGROUP phenogram of onnia based on 108

character set.

I \n
..Ié'j

1330

 

‘Q‘

“ 100

I: 200

 

 

400

460

FERRSQJAFRUT

 

 

 

if

 

97

MARW LUCI
L194 LIGE LIGW LIGL MARE

 

 

 

 

 

 

 

 

 

 

 

 

98

i
A.

Figure 15. HGROUP phenogram based on vegetative

characters of Lzonia.

 

99

FERR SQUA FRUT LIGW LIGE LIGW LIGL MARE MARW LUCI

 

 

 

 

 

50

 

 

 

 

 

 

 

 

 

 

100

 

 

 

 

140

 

 

 

380

1000

 

 

 

 

 

100

Figure 16. HGROUP phenogram based on flower-

fruit characters of onnia.

 

 

101

LIGL LIGW LIGM LIGE FRUT SQUA FEPR MARW MARE LUCI

o L.__J LT—J 1 __J |__1__j

L___

 

 

 

 

 

50

 

 

 

100

 

 

 

150

200

 

 

 

 

 

250 ——

 

 

102

 

Figure 17. HGROUP phenogram based on floral char-

acters of Lzonia.

 

 

103

FERR SQUA FRUT LIGM LIGE LIGW LIGL MARE MARW LUCI

° L4 Le L___. LJ

 

 

 

 

20

 

40

 

 

 

60

 

 

 

80

 

 

 

 

1130 I T J

 

104

 

 

 

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109

 

Figure 18. HGROUP relationships between OTU's of
onnia based on 108 character set; solid
lines indicate closest related OTU.
dotted lines indicate second closest

OTU.

 

 

110

 

 

lll

 

Figure 19. HGROUP relationships between OTU's

based on vegetative characters of

onnia.

112

 

 

Figure 20.

113

HGROUP relationships between OTU's

based on flower-fruit characters of

onnia.

 

114

46.4

 

  

 

   

 

 

115

 

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116

 

 

 

 

the

(sec

IESI

117

 

the second group (section Arsenococca), and E° lucida and
g, mariana are loosely clustered in the third group
(section Earls). Using just flower and fruit characters
results in a much closer clustering of E. mariana with

E, lucida, indicating the many similarities of these

two taxa in reproductive structures. The grouping of l
E. mariana and g. lucida is much less strong when only "

vegetative characters are used. The three species, 1 ‘

 

 

£° fruticosa, £0 ferruginea and 2° squamulosa all seem ‘
to be equidistant from each other. gyonia squamu1osa

and E, fruticosa seem to be slightly closer in flower-

 

 

fruit characters, but E. ferruginea and E. squamulosa

are closer when only vegetative characters are considered.
When a.line is drawn at a error index level of 240 on

the phenogram based on 128 characters, the genus is
delimited into three groups, each corresponding to one

of the sections. Six groups are present when a line

is drawn at the 60 error index level. This lower

error level indicates that much more homogenous

groups are involved. These six groups correspond to

each of the species in the genus native to North America.
SeCtion Maria is clustered first with section Arsenococca
When Vagetative characters are used, indicating the
Vegetative similarities between these two groups. On

the other hand, section Arsenococca is much more similar

t o .
o seat-1°11 Lyonia in floral morPhOJ-OQY: and clusters

 

 

th
se
Va
in
la

 

118

first with this group when just flower-fruit characters
are considered. Over all, section Arsenococca seems to
be most closely related to section Maria. Lyonia

ligustrina var. ligustrina is most closely clustered

 

with the western part of the range of E. ligustrina var.

foliosiflora, while both the middle and eastern parts

 

of the distribution of var. foliosiflora cluster very
closely. This indicates the similarities between the
Arkansas populations of var. foliosiflora and the
typical variety. It is sometimes hard to identify
these Arkansas populations, especially late in the
season, but they clearly belong with the southern
variety, having large coriaceous leaves, rather lax

inflorescences and reddish twigs. The western popu-

 

lations of E. mariana also appear to be rather distinct.
They have been recognized as forma vestita by Rehder
(1924) and tend to be more pubescent than the eastern
populations.

The clustering of the 128 characters themselves
also indicated a division of the genus into three major
groups; Table 10 lists the characters in each group.
Group I includes characters distinctive of section
Elfiflfiav'groups II and III.indicate characters important

In sections Arsenococca and Maria, respectively. This

 

clUSterj-ng also indicated that all the characters

1 .
nv°lV1n9 ferrugineous scales are highly correlated.

 

 

1Jl9

TABLE 10. HGROUP clustering of characters of Lygnia.

 

 

GROUP I GROUP II GROUP III
7 Density of ferr. scales 18 Shape of bud 86 Shape of corolla

at internodes, mature

10 Density of ferr. scales 20 Length of bud 89 Length of corolla
at internodes, immature

11 Color of ferr. scales 22 L/w of bud 90 Width of corolla
at internodes, mature

12 Ratio of small to large 29 Serration type 92 Length of corolla
ferr. scales, mature from base to widest

pt.

13 Shape of ferr. scales, 30 Serration type 93 Length of calyx lobes
mature

14 Color of ferr. scales 34 No. of serrations/ 95 Width of calyx lobes
at internodes, immature unit distance

15 Shape of ferr. scales, 35 No. of serrations/ 96 L/w of corolla
immature unit distance

16 Persistence of small 36 Length between 100 Length of filaments
scales serrations

24 Density of scales on bud 38 Length of teeth 103 Length of anthers

25 Color of scales on bud 54 Length of hairs on 105 Length of style

lower leaf surface

55 Density of scales on 80 Inflorescence type 107 Shape of ovary
upper leaf surface,
mature

56 Density of scales on 81 No. of flowers/infl. 114 Width of capsule
lower leaf surface,
mature

57 Color of scales on lower 83 No. of panicles 122 No. of seeds/capsule
leaf surface, mature

58 Ratio of small to large 84 No. of bracts/infl. 123 Length of seeds
scales on lower 1f.
surface

60 Density of scales on 85 Total length of bracts
upper 1f. surface,
immature

61 Shape of scales on leaf 121 No. of capsules/fruit
mature cluster

64 Persistence of large
scales

66 Size of scales

76 Density of scales on

petiole

82 Relation of flowers 1:0
veg. growth

 

97 Pubescence of calyx
1
10 Pubescence of pedicle

11
7 Thickening of sutures

 

 

 

120

This probably indicates that all these characters are
controlled by one complex of closely linked genes. Other
characters such as length of corolla and length of fila-
ments, or number of flowers per inflorescence and inflor-
escence type seem to be logically correlated.

The results of factor analyses based on vegetative
characters and flower-fruit characters appear in Figures
22, 23 and 24. It is easily seen that this study also
indicates a clear separation of the genus Lyonia into
three groups. Figure 22 points out the distinctness of
£3 mariana and L. lucida in vegetative morphology, while
Figure 20 shows their similarities in flower and fruit
characters. All three sections seem to be equally distant
from each other, with L. lucida showing a slight tendency
to group with section Lyonia (note Figure 22). Figure 23

and to a lesser extent Figure 22 illustrate the distinct-

 

ness of E. ligustrina var. ligustrina from E. ligustrina

 

var. foliosiflora. Principle factor III seems only to

 

separate L. lucida from all the other species.

Factor analysis of a loo-character set provided
further evidence of the three strong subgroups within
Lyonia. Characters highly correlated with Principal
Factor I are important in distinguishing section Lyonia;
Principal Factor II clearly represents section §£§227
ococca, while characters most closely associated with

Principal Factor III are those important for section

 

 

 

 

 

121

 

 

Figure 22. Factor analysis: Principal Factors I

and II based on vegetative characters

of Lyonia.

 

   

122

 

 

II
11.0
. MARE
‘ O MARW
1 0.5
d
'1 -o.s Q‘ LUCI 0.5 1.0
FERR ‘ 0.10 I I f T I I g
SQUA .LIGL
O FRUT LIGW
. LIGM
4 LIGE
d-0.5
.
«-100

 

 

 

 

 

 

 

 

Figure 23.

123

Factor analysis: Principal Factors I

and III based on vegetative characters

of Lyonia.

 

~1.o

FERR

r

.FRUT

LUCI

124

 

‘ 0.5

 

$1.0

LIGM

.MARE LI

 

LIGL

 

 

 

 

 

 

 

Figure 24.

125

Factor analysis: Principal Factors I

and II based on flower-fruit characters

of Lyonia.

 

126

 

 

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4-1.o

 

 

 

 

127

$2533. (A list of the characters most highly correlated
with the first four principal factors is given in Table 11.)
Principal Factor IV seems to be closely correlated with
characters that are important in distinguishing E. gaggif

cosa from L. ferruginea. This points out one of the

 

advantages of a numerical taxonomic investigation. Not
only are major groups clearly indicated, but also char-

acters important in delimiting these groups are indicated!

 

For example, from the intercorrelation analysis of these
100 characters it is seen that the amount of curvature
of the leaf blade is highly correlated with: (1) the
length-width ratio of the leaves, (2) the depression of
the veins above, (3) the growth form, (4) the amount of
revolution of the leaf margins, (5) the type of leaf
apex, (6) the time of flowering, and (7) the prominence
of the major veins on the lower leaf surface. This
combination of characters makes little sense until it

is realized that it is precisely these characters which

separate L. fruticosa from L. ferruginea. They are cor-

 

 

related because they occur in definite combinations in

these species. Lyonia ferruginea has extremely curved

 

leaves with depressed veins vs. the flattened, reticulate-

 

veined leaves of L. fruticosa. It is a large shrub or
tree, while 2, fruticosa is a small shrub, often not
over knee high. Lyonia ferruginea flowers in the early

spring, but 2. fruticosa does not usually flower until

 

 

TABLE 11.

128

Factor analysis of loo-character set. Charac-
ters most highly correlated with first four
principal factors.

 

Principal Factor

Correlated Characters

 

II

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0000
o

11.

12.
13.

14.
15.
16.
17.
18.
19.
20.
21.

10.

11.

Density of scales on twigs
(mature)

Density of scales on twigs
(immature)

Texture of bark

Apex of bud

Density of scales on bud
Length of leaves

Length from base of leaf to
widest point

Width of leaves

Color of leaves

Density of scales on upper leaf
surface

Density of scales on lower leaf
surface

Density of scales on petiole
Relation of vegetative to
floral buds

Pubescence of corolla
Pubescence of calyx

Spur length

Pubescence of pedicel

Shape of capsule

Pubescence of capsule

Width of seeds

Thickness of sutures of fruit

Shape of bud

Length of bud

Length to width ratio of bud
Serration type (distal)
Serration type (proximal)
Number of serrations per unit
length (distal)

Number of serrations per unit
length (proximal)

Length of pubescence
Inflorescence type

Number of flowers per inflor-
escence

Number of bracts per inflor-
escence

 

 

 

129

 

 

TABLE 11. Continued
Principal Factor Correlated Characters
II (cont.) 12. Total length of bracts per
inflorescence
13. Length of peduncle
14. Length of bracteoles
15. Width of bracteoles
16. Length of capsule
17. Width of capsule
18. Shape of corolla
III 1. Density of nonglandular hairs
on bud
2. Density of nonglandular hairs
on veins of upper leaf surface
3. Length of hairs on veins of
lower leaf surface
4. Presence of marginal vein
5. Density of nonglandular hairs
on petiole
6. Color of corolla
7. Curvature of filaments
8. Thickness of blade
9. Longitudinal ridges on stem
10. Presence of revolute margin
11. Angle made by apex of leaf
12. Luster of leaves
13. Presence of glandular hairs on
lower leaf surface
14. Position or growth form of
branches
IV 1. Curvature of leaf blade
2. Depression of veins on upper
leaf surface
3. Habit (tree or shrub)
4. Type of leaf apex
5. Prominence of major veins on
upper leaf surface
6. Prominence of major veins on
lower leaf surface
7. Length of petiole
8. Length-width ratio of leaf

 

130

late spring or early summer. Many other distinctions are
listed and are discussed in the taxonomic treatment. Thus
the knowledge of one good distinguishing character can
lead to the discovery of many more! Of course, the human
mind is the best "computer" and perceptive botanists have
long recognized the distinctive characters separating

these two taxa.

Phyletic Studies

Twenty-four characters were selected and assigned
generalized (primitive) and Specialized (advanced) states
using the method of Wagner (1962, 1969). Table 12 lists
these characters. Then each taxon was scored Q'if gen-
eralized and l’if specialized for each of the characters.
All the taxa are listed in Table 13 with the character
divergence values for each. The total divergence index
of each taxon was determined by adding the individual
divergence values of each character together. Then
mutual groupings of characters between taxa were deter-
mined, and the taxa were arranged in sequence according
to these groupings. They were plotted on a concentric
graph (Figure 25), the radii being determined by the
mutual groupings of characters and the distance being
determined by the divergence of each taxon.

Several instances of parallel derivation of

Characters are evident. These include the deciduous

 

 

V

131

 

 

TABLE 12. Characters used in phyletic study (Wagner
method).
Code Generalized Specialized
Letter State State
A Leaves evergreen Leaves deciduous
Bl Flowers in racemes Flowers in corymbose
fascicles
BZ Flowers in racemes Flowers in panicles
C Flowers white Flowers pink or red
D Corolla cylindric Corolla strongly urceo-
late
E Calyx persistent Calyx deciduous
F Plant not lepidote Plant lepidote
G Small spurs or none on Long spurs on filament
filament
H Capsule ovoid or sub- Capsule urn-shaped
globose
I Sutures of capsule less Sutures of capsule
thickened strongly thickened
J Thickened sutures not Thickened sutures separ-
separating from valves ating from valves as a
unit
K Tall shrub or small tree Small shrub
L Floral buds not always Floral buds usually
above vegetative buds always above vegetative
buds
M Leaves coriaceous Leaves thinner
N Branches terete or Branches round
angled
O Calyx lobes broad and Calyx lobes elongate

short

132

 

 

TABLE 12. Continued
Code Generalized Specialized
Letter State State

P Ovary with massive p1a~ Placentae less massive--

T1

T2

centae--many small
ovules

Buds globose, usually
small

Buds with many outer
scales

Inflorescence with
leafy bracts

Filaments not pubescent

Filaments not pubescent

Filaments elongate

Capsule almost com-
pletely glabrous

Lower leaf surface
without dense covering
of hairs

Glandular hairs small
or absent

fewer ovules
Buds elongate, larger
Buds with only two outer

scales

Inflorescence lacking
leafy bracts

Filaments pubescent on
lower half only

Filaments pubescent
for their entire length

Filaments short

Capsule pubescent, at
least at base

Lower leaf surface with
a uniform covering of
densely packed hairs

Glandular hairs large,
club-shaped

 

133

Character divergence values for each taxon,

used to construct Wagner tree.

TABLE 13.

 

FRUT SQUA LUCI MARI LIGL LIGF

FERR

Character

 

1* 1010000000111011101* 1101..

1* 1010000000111011111* 1101

11* 00101111111110010* 10000

01* 10001000010010010000000

01* 01010011000001010001100

01* 01010011100001010001110

01* 01010011000001010001110

*
lad * 112
AnbnuCnuqununuIwuxaanmuonrnuRnamlTHUvaua

13 12

13

10

Total

 

Character does not apply.

Character R was not included in the calculations.

**

134

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136

habit and round branches in Lyonia mariana and in a.

ligustrina; and small urceolate flowers with short fila-

 

ments and pubescent capsules in E. ligustrina and in the

 

E. ferruginea group. Lyonia mariana and the E. ferruginea

 

 

group show parallel development of the reduced racemose

inflorescences and strongly thickened sutures of the cap-

 

sule. Lyonia fruticosa and E. mariana have both evolved
as dwarf rhizomatous shrubs of sandy soils.

Lyonia lucida is the most primitive species

 

studied; E. mariana and E. ligustrina are the most
advanced. The E. ferruginea group (section Lyonia) is
moderately advanced with E3 squamulosa occupying the
most primitive position and E. fruticosa the most
advanced. The results also strongly indicate that the
genus is divided into three major parts, corresponding
to the three sections recognized in the taxonomic treat-
ment. Both E. mariana and E. lucida are grouped in
section flagia in spite of their obvious vegetative dif-
ferences. The evolutionary development of the characters
in each section can easily be traced in Figure 25.

The Wagner method requires numerous assumptions
regarding which represents the primitive and which repre-
sents the advanced state of each character. These
decisions, although made after an extensive study of
the genus and a thorough overview of related genera, are

somewhat subjective and introduce a major source of error.

137

Thus a second method was used for comparison which does
not require that any assumptions be made as to which state
of each character is primitive and which is advanced.

This method (Whiffin and Bierner, 1972) requires only

that one taxon be chosen as representing the most primi-
tive (ancestral) taxon.

Twenty-three characters (essentially similar to
those used in the Wagner method above) were used and arbi-
trarily one state was designated as primitive (O) and the
other as advanced (1). Then a table of designated char-
acter states of each character for each taxon was con-
structed. From this a matrix (Table 14) of differences
was formed which records the number of characters for
which each pair of taxa exhibit different states. By
using this matrix and following the steps outlined by
Whiffin and Bierner (1972) a phyletic tree can be con-
structed. The resulting trees will be different if a
different taxon is selected as the most primitive, although
I found that in every case the major grouping of species
was similar (Figure 26). These phyletic trees clearly
indicate that the genus is composed of three distinct
groups. Again these three groups correspond to the
three sections of the genus present in North America.

In every case E. lucida and E. mariana are clearly linked,
with E, mariana as the more advanced, and E. lucida pro-

viding a connection to the other members of the genus.

138

 

 

 

TABLE 14. Matrix of character divergence used in Whiffin-
Bierner method.
LUCI MARI FERR FRUT SQUA LI GL LIGF
LUCI 2 3
MARI 15 23
FERR ll 7 23
FRUT lO 8 22 23
SQUA 12 8 20 19 23
LIGL 8 10 10 9 ll 23
LIGF 9 11 ll 10 12 22 23
Note: Numbers indicate number of characters in common.

Symbols for taxa are the same as given in Table 3
except that MARI indicates E. mariana and LIGF
indicates E. ligustrina var. foliosiflora.

 

 

139

 

Figure 26. Phyloqenetic trees of Lyonia produced by
Whiffin—Bierner phyletic method. A. Tree
assuming that E. lucida is most primitive.

B. Tree assuming that E. squamulosa is

 

most primitive.

 

140

LIGL
LIGF
MARI

FRUT

SQUA

 

 

 

 

? LUCI

 

   

 

A
FRUT
FERR
LIGF LUCI
’ SQUA

 

 

 

 

 

141

Lyonia ligustrina var. ligustrina is always more advanced

 

than var. foliosiflora, which links this group to either

 

section Maria or section Lyonia depending upon which
species is taken as the most primitive. Lyonia ferruginea,

E. fruticosa and E. squamulosa are always closely con—

 

 

nected with E. squamulosa in the most primitive position

 

unless either E. fruticosa or E. ferruginea is specifi-

 

 

cally designated as primitive in constructing the network.
A high degree of similarity is evident between
the methods of Wagner (1962, 1969) and the "quick method"
of Whiffin and Bierner (1972). Both indicate that Lyonia
is divided into three clearly demarked groups. Both also

indicate that E. mariana is more advanced than E. lucida

 

 

and that E. ferruginea and E. fruticosa are probably more

advanced than E. squamulosa. The relationships of the

 

three sections to each other are not as clear. The Wagner
method seems to indicate a closer tie between Arsenococca
and Maria, while the modified method indicates that

Arsenococca is closer to Lyonia.

 

Comparison of the Phyletic and
Phenetic Methods

Both the phyletic and phenetic methods lead to
the same conclusions. They both show that Lyonia is
divided into three clearly demarked groups which I have
recognized as the sections Lyonia, Arsenococca, and

.Maria. It is interesting that these three subgroups were

142

very early recognized by botanists using only "classical"
taxonomic methods. These studies also indicate that
section Maria has the greatest amount of intrasectional
variation, especially when only vegetative characters

are considered. They also indicate which characters

are important in differentiating each section from the
others. A more detailed discussion of these characters
can be found in the Systematic Treatment. Both studies
indicate the distinctness of each species recognized in

the taxonomic treatment.

CHEMOSYSTEMATIC STUDIES

With the advent of modern methods of analysis,
comparative studies of the chemistry of plants have
become much more widespread. Such progress has been par-
ticularly valuable to the systematist because it has
opened up a new dimension for study and has provided
evidence confirming interpretations established by more
traditional methods (Brehm, 1966). Sometimes the com-
pounds themselves are identified and the pathways of
their synthesis determined (Belzer and Ownbey, 1971);
in other studies such as those of Alston and Turner
(1959) or Furlow (1974), the chromatographic patterns
are compared without determining the actual chemicals
involved. The compounds are treated as ordinary taxo-
nomic characters and, as with any taxonomic characters,
mmltiple correlations, rather than single differences,
are considered the most useful in determining relation-
ships.

Giannasi and Rogers (1970) list three reasons for
using chemical data in taxonomic studies: (1) to test
classifications based on morphology, (2) to determine

species specific patterns, and (3) to detect possible

143

144

phyletic relationships. Just like any other taxonomic
character, chemical data can give us insights into intra-
specific variation and speciation (Turner, 1970).

In this study phenolic chemical characters were
used to provide a comparison with the classifications
based upon morphology and to help clarify taxonomic
relationships which could not be perceived from morphology
alone.

As pointed out by Bate-Smith (1963) and Heywood
(1966) the results of chemosystematic studies must be used
along with other kinds of data, not by themselves. Care
must also be taken in evaluating the results! Numerical
analysis of unidentified compounds can sometimes be mis-
leading especially if the sample size is small, as is

often the case in such studies (Weimarch, 1972).

Methods and Materials

Samples of leaves of five species of Lyonia were
obtained from widely separated parts of the geographical
range of each taxon (Table 15). These were collected
from early to late summer from one- or two-year-old
branches and were dried for approximately 96 hours in
plant presses (natural drying). Two to four populations
of each species were sampled. Within each population
specimens were taken from five individuals. The indi-

‘Vidual samples from each population were combined. A

145

 

 

 

 

 

 

 

 

 

 

 

 

TABLE 15. Geographic location of specimens from which
foliage samples for analysis of phenolic com-
pounds were obtained.

Collection .
N ler Taxon Location

18, 19, 20, E. mariana Near Wading River, Bur-

21, 22 lington Co., N.J.

319 E. mariana Near Pritchardville,
Beaufort Co., S.C.

277 E. mariana Near Morven, Brooks Co.,
Ga.

62, 65, 66, E. lucida Near New Bern, Craven

67, 70 Co., N.C.

100, 101, 102, ‘E. lucida Near Jamestown, Berkeley

103, 105 Co., S.C.

326 E. lucida Along 0.8. 82 near
Liberty-Long Co. line,
Long Co., Ga.
267 E. fruticosa North of Sparks, Cook
Co., Ga.
314 L. fruticosa Amelia Island, Nassau
— Co., Fla.
264 L. ferruginea Southwest of Tifton,
— Tift Co., Ga.
300 E. ferruginea Jekyll Island, Glynn
Co., Ga.

33, 34, 36, E. ligustrina North of Pocomoke City,

38, 40 var. ligustrina Worchester Co., Md.

239 E. ligustrina Near Highlands, Macon
var.IIgustrina Co., N.C.

42, 43, 44, E. ligustrina Near South Mills, Camden

50, 52 var. foliosiflora Co., N.C.

252 L. ligustrina Northeast of Ashburn,

 

var. fdlioSIflora

 

Turner Co., Ga.

 

146

total of 15 samples was analyzed. Voucher specimens are
depOsited in the Beal-Darlington Herbarium of Michigan
State University (MSC).

The phenolic compounds were extracted from dried
leaves following, in general, the method of Hanover and
Hoff (1966), Hanover and Wilkinson (1970) and Furlow
(1974). Four-tenths of a gram of leaf material from each
of the five collections at a particular site were combined
into a single sample weighing 2.0 grams. Phenolic sub-
stances were extracted by homogenizing the samples in a
blender with 100 m1. of boiling water for 2 min., filter-
ing, washing with 50 ml. of boiling water and then repeat-
ing the entire process once more except with two 50 m1.
boiling water washes. This method was found by Hanover
and Hoff (1966) to effectively eliminate such interfering
substances as tannins and chlorophyll from the extract.

The water extract was washed five times with
50 ml. portions of ethyl ether in a separatory funnel
to remove waxes and fats, followed by 50 ml. portions
of normal butyl alcohol.

Compounds were separated by two-dimensional
descending paper chromatography. Fifty microliters of
extract were applied with a micropipet to a point on
the upper surface of 46 by 57 cm. Whatman 3MM chromato-
graphic filter paper 8 cm. from each edge in the upper

left-hand corner. The papers were folded on a line

147

5.5 cm. from the upper edge and placed, in groups of
eight, in a chromatographic chamber, which was equil-
ibrated for 2 hrs. with the lower portion of a mixture

of n-butanol, acetic acid, and water (4:1:5). The sheets
were then irrigated with the upper portion of the solvent
and allowed to develop until the solvent front neared the
bottom of the sheets (about 16 hrs.). After drying, the
chromatograms were irrigated in the second direction with
a mixture of acetic acid and water (3:17), and allowed to
develop until the solvent again had nearly reached the
bottom of the sheets (about 5 hrs.). They were then
thoroughly dried at room temperature.

Completed chromatograms were examined without
chemical treatment and in daylight and under both long-
and short-wave ultraviolet illumination after a 30 min.
exposure to fumes of ammonium hydroxide. Spots were out-

lined, their R values calculated, and their color

f
reactions to each treatment noted (Table 16). Spots
produced by the same compound on various sheets were

identified, numbered, and tallied, but no attempt was

made to determine the chemical identity.

Results and Discussion
Seventy-four compounds were found in the butanol
fractions of the five species studied; of these, 8
(numbers 1, 2, 12, 17, 18, 21, 22, and 28) occurred on

every sheet and four (numbers 3, 16, 20, and 23) were

TABLE 1 6 .

148

Rf values and color reactions of the phenolics
in L onia. Color abbreviations: L = light;

B1 = blue; R = red; G = green; Y = yellow;

P = purple; Br = brown; V = violet; O = orange.

 

Rf n-Butanol,

Rf Acetic

 

 

Com— acetic acid, acid, Ordinary UV Light
pggnd water water Light Untreated NH
(4:1:5) (3:17) 3
1. 0.00 0.06 0 LBr BrR
2. 0.88 0.00 LY V V
3. 0.37 0.06 —— LY Y
4. 0.33 0.12 -- LB Y
5. 0.42 0.14 -- P B1P
6. 0.26 0.08 -- P LG
7. 0.28 0.14 -- LP P
8. 0.64 0.13 -- P PBl
9. 0.56 0.01 -- LBl LP
10. 0.44 0.30 -- LBr Br
11. 0.67 0.39 Y BrR CY
12. 0.55 0.44 Y BrR Y
13. 0.35 0.46 LY -- LPR
14. 0.30 0.43 -- -- L0
15. 0.23 0.44 -- -- L0
16. 0.31 0.56 -- LBrR LR
17. 0.47 0.58 LY BrR Y
18. 0.67 0.58 YO BrR BrRO
19. 0.52 0.57 LO Br BrR
20. 0.77 0.63 -- P Bl

149

TABLE 16. Continued

 

Rf n-Butanol, Rf Acetic

 

 

Com- acetic acid, acid, Ordinary UV Light
pggnd water water Light Untreated NH
(4:1:5) (3:17) 3
21. 0.55 0.73 -- P 81G
22. 0.60 0.86 -- P PBl
23. 0.49 0.85 -- P P
24. 0.51 0.81 -- LGBl G
25. 0.35 0.82 -- -- G
26. 0.34 0.82 -- -- PBl
28. 0.98 0.90 LY LG LP
29. 0.56 0.26 -- -- G
30. 0.19 0.29 -- -- LB
31. 0.16 0.39 -- -- L0
32. 0.51 0.15 -- -- LB
33. 0.58 0.33 LY LBr Y
34. 0.54 0.50 YO Br CY
35. 0.67 0.67 -- Bl G
36. 0.64 0.64 LY RBr O
37. 0.40 0.68 -- P PBl
38. 0.38 0.78 LY LOBr Br
39. 0.29 0.79 -- LBrP LBrP
40. 0.62 0.59 -- -- Y
41. 0.24 0.19 -- -- G
42. 0.27 0.07 -- -- CY
43. 0.56 0.09 -- -- P

150

TABLE 16. Continued

 

Rf n-Butanol, Rf Acetic

 

 

Com- acetic acid, acid, Ordinary UV Light
P§:?d water water Light Untreated NH
(4:1:5) (3:17) 3
44. 0.21 0.44 -- -- LY
45. 0.36 0.32 -- LBr LY
46. 0.30 0.62 -- -- LY
47. 0.30 0.50 -- LBr LR
48. 0.41 0.42 -- GBl GBl
49. 0.46 0.38 -- -- L0
50. 0.80 0.70 -- G B1
51. 0.61 0.79 -- RBr PBl
52. 0.75 0.88 -- B1 B1
53. 0.59 0.90 LY RBr RBr
55. 0.81 0.51 -- B1 B1
56. 0.43 0.79 -- BlG G
58. 0.43 0.71 -- -- B1
59. 0.76 0.81 -- LBlY LBl
60. 0.75 0.42 -- LBl LBl
61. 0.98 0.44 -- L81 L31
62. 0.70 0.90 LY RBr PBl
63. 0.55 0.66 -- P BlP
64. 0.51 0.90 -- LY YG
65. 0.66 0.71 -- LBl --
66. 0.48 0.76 LY O O
67. 0.40 0.85 LY LO 0

151

TABLE 16. Continued

 

Rf n-Butanol, Rf Acetic

 

 

Com- acetic acid, acid, Ordinary UV Light
p§:?d water water Light Untreated NH
(4:1:5) (3:17) 3
68. 0.48 0.76 -- L81 B1
69. 0.67 0.76 -- LBl B1
70. 0.67 0.85 -- LP P
71. 0.57 0.64 -- -- G
75. 0.53 0.68 LY Br RBr
76. 0.49 0.86 -— Br Br
77. 0.65 0.81 -- LBl --
78. 0.63 0.90 -- LBr LBr
79. 0.59 0.89 -- LBr LBr

 

152

present in every species but not on every sheet. The
number of spots on individual sheets ranged from 18 to
27; from 25 to 35 compounds were found in the species
studied (see Table 17). At least one unique compound
appeared in every taxon (Table 18). A representative
chromatogram for each species is given in Figures 27
through 32.

Chemical similarities among the taxa were calcu-
lated using the paired affinity index of Ellison, Alston
and Turner (1962); the resulting values are shown in
Table 20 (see also Table 19).

The results of this study indicate that each
taxon has a definite pattern of phenolic compounds and
can be recognized readily on this basis. Some taxa such
as E. mariana and E. lucida seem to be uniform chemically.
The population of E. mariana sampled in Burlington Co.,
N.J. is extremely similar to the population sampled in
Beaufort Co., S.C.! Similarly, the population of E.
lucida sampled in Craven Co., N.C. is quite similar to
the population in Long Co., Ga. Thus the chemical
characters seem to compare well with the morphological

characters which are also uniform in these two species.

On the other hand, E. ligustrina, a very polymorphic

 

species, had the most different phenolic compounds of
the Species studied, and showed the greatest intra-

specific variation. The northern E. ligustrina var.

 

153

TABLE 17. Total listing of phenolic compounds present in each population of each
species studied.

 

—— -::=_:9 may? Arm—1:."—

Spot MARI MARI MARI FRUT FRUT FERR FERR LUCI LUCI LUCI LIGF LIGF LIGL LIGL
No. 18 319 277 314 267 300 264 326 62 101 252 42 33 239

 

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><><><><><><><><><

xxxxxxxx
xxxxxxxx

XXXXXXXXXXXXX

><><
XX

 

Note: Numbers skipped were rejected as not representing distinct compounds, in
most cases they were included with another spot.

154

TABLE 18. Characteristic phenolic compounds for each
Species of Lyonia.

 

Compound Number Frequency (in percent of sheets)

 

I. E. mariana

5 100
7 33
8 66
14 66
25 33
26 33
63 66

II. E. lucida

64 100
65 100
66 A 100
67 100
68 100
69 100
70 100
71 100
75 50
76 50
77 50
78 100
79 100

III. E. fruticosa

 

49 50

IV. E. ferruginea

 

58 50
59 50
60 50
61 50
62 100

V. E. ligustrina

 

30 75
31 25
32 25
37 75
38 25
39 50
40 25
41 25
42 25
43 25
44 25
45 25

46 25

 

155

 

.mmoofiusum macomq mo Emumoumeouso m>flumucmmmummm

 

.nm mummflm

156

 

 

 

 

l-BUIAKL ACHIC ACID, HATER (4:1:5)

 

A
'

 

.7”

 

—; mncmn.mm (3:17)

 

 

157

 

.mmcflmsuumm choNq mo Emumopmeouno m>Humucmmmuomm

 

.mm ousmflm

158

 

 

I-BUMNOL, ACETIC ACID. HATER (4:1:5)

 

j:
3

 

v

N"

 

 

#: ACETIC ACID. HTER (3.17)

 

159

 

.mcflnuwsmfla

 

.um> mcfluumsmfla macomq mo Emumoumfiouco m>wunucmmmummm

 

.mm musmflm

 

160

 

 

 

 

mm

 

 

 

34.3 5:: 53¢ :53 £153.:

 

 

Inca

N'ETK AUE, IATEP (3.175

A

 

 

161

 

.muoHMAmOMHom

 

.um> mcfluumsmfia macomq mo Emumoumfiouso m>flumusmmmumom

 

F

.om wusmflm

162

 

 

 

 

I

 

8..
:3
E
i
55
3
S
i
i

 

 

A KEIIC ACID. UTE! (3:17)

 

163

 

.mowosa MHCQNM mo Emumoumeouno m>flumucmmmumwm

 

.Hm mesons

 

164

 

 

 

C
HATER (4:1:5)

 

ACETIC ACID.

l-BUTANOL,

 

 

\

 

 

% ACETIC ACID. HATER (3:17)

'

 

 

 

165

 

.mcmwuna second mo EmumoumEonso m>wumucmmmummm

 

.Nm musoflm

166

 

 

 

 

 

 

 

33.: a; .23 ...—E g

 

 

ACETIC ACID. HATER (3:17)

'

 

 

167

 

TABLE 19. Number of compounds in common among the taxa
of Lyonia studied. I

 

 

MARI FRUT FERR LUCI LIGF LIGL L_fi
MARI 28
FRUT 15 29
FERR 12 25 29
LUCI 16 15 13 31
LIGF 16 17 16 14 30

 

LIGL 16 18 16 15 20 25

 

 

 

168

 

 

TABLE 20. Chemical similarity between species of L onia
in North America. (Numbers represent pEIEEd—
affinity values.)

MARI FRUT FERR LUCI LI GF LI GL

MARI 100

FRUT 35.7 100

FERR 26.7 75.8 100

LUCI 37.1 33.3 27.8 100

LIGF 38.1 40.5 37.2 29.8 100

LIGL 43.2 50.0 44.4 36.6 57.1 100

 

 

 

 

169

ligustrina is more homogenous in phenolic compounds than

 

the southern var. foliosiflora. Again the phenolics seem
to accurately reflect the total variability of each taxon,
in that the southern variety is the more diverse morpho-
logically. The two widely separated populations of E.

ligustrina var. ligustrina both have very similar chemical

 

 

profiles, while a greater difference is shown between the
geographically closer populations sampled of var. Egiigr
siflora.

Definite differences in chromatographic patterns
occur between E. ligustrina var. ligustrina and E. Eigggf

trina var. foliosiflora. The phenolic compounds separating

 

these two varieties are listed in Table 21 below.

Lyonia and E. ferruginea are both moderately
variable and quite distinct in their phenolic compounds.
Lyonia fruticosa possesses the following compounds not

found in E. ferruginea: ll (50% of sheets of L. fruti-

 

cosa), 24 (100%), 49 (50%), 51 (100%), 58 (50%), 59 (50%),
60 (50%), 61 (50%), and 62 (100%). Thus the evidence from
phenolic compounds further supports the recognition of
'Uhese two taxa as distinct species. The p0pulation of

E. fruticosa from.Nassau Co., Fla. which showed some

 

morphological similarity to E. ferruginea is much more

similar to E. fruticosa in its phenolic pattern than it

 

is to»E. ferruginea, and it does not appear to be a hybrid

 

between these two species.

 

 

170

TABLE 21. Compounds separating L. ligustrina var. ligus-
trina from E. ligustrIna var. foIIOSiflora.

 

 

 

 

 

 

Frequency (in percent

Compound Number of sheets)

 

 

 

 

 

I. E. ligustrina var. ligustrina f
9 100
24 50 _
31 50
32 50
38 50

II. L. ligustrina var.
foliosiflora

4 100
6 100
39 100
40 50
41 50
42 50
43 50
44 50
45 50

46 50

 

 

171

From Tables 18, 19 and 20 it is seen that E.

 

 

fruticosa and E. ferruginea are probably very closely

related and quite distant from the other taxa studied.

 

The most closely related from outside section Lyonia

seems to be E. ligustrina. Eyonia ferruginea and E. fru-

 

 

ticosa show the following group of compounds which are

absent from all other taxa.

 

TABLE 22. Diagnostic compounds for section Lyonia.

 

Frequency (in percent

Number of Compound of sheets)

 

52 100
53 100
56 100
35 75
47 75
48 50
50 50
55 50
62 50
49 25
58 25
59 25
60 25
61 25

 

.A close relationship is also indicated for E. ligustrina

‘var. ligustria and E. ligustrina var. foliosiflora. These

 

 

 

tw0>taxa have a number of compounds in common and form a
<1istinct group separate from E. mariana, E. lucida or the

IL. ferruginea group. Lyonia lucida does not seem to be

 

closely related to any other taxon. Its closest ties

seem to be with E. mariana in that they both possess

 

172

compounds 10 and 15 which occur in no other taxon. Lyonia

mariana seems to show relationships to both E. lucida and

E. ligustrina. It shares compound 19 with E. ligustrina.

 

 

The above information is summarized in the phylogenetic
tree showing the compounds held in common by each succes-
sive group of taxa (Figure 33). This tree gives a clear
total picture of the possible interrelationships within
the genus as a whole and seems to support the division
into the same three sections as recognized on the basis
of morphological characters alone., The only difference
is that the chemical data seem to indicate that section

Arsenococca is slightly closer to section Lyonia whereas

 

the total morphological classification puts it closer to
section EEEEE. In floral morphology section Arsenococca
and section Lyonia do show several similarities (see also
Figure 34).

The variability of phenolic compounds in higher
plants has been discussed by Alston (1966), Alston and
Turner (1963), Hanover and Hoff (1966), Taylor (1971)
and others, who point out that chemical composition can
vary with season, age, tissue, growth site, individual,
etc. These factors were controlled as much as possible
in this study. Fresh material of E. lucida (no. 326)
was chromatographed to compare with the phenolic pattern

obtained from dried material from the same population.

173

 

 

Figure 33. Phylogenetic tree showing the phenolic

compounds distinctive for each taxon of

Lyonia.

 

 

 

 

 

Figure 34.

175

Phylogenetic trees based on chemical data
(Table 20), using Whiffin-Bierner method.

a. Tree with Lyonia ferruginea taken as

 

the most primitive. b. Tree with E.

lucida taken as the most primitive. c.

Tree with E. fruticosa taken as the most

 

primitive.

176

MARI
1|) LUCI

 

 

 

 

LIGL
FRUT
LIGF * 9
(JD FERR
A
FERR
FRUT
LIGL
T‘P MARI
LIGF
d' LUCI
B
MARI
LUCI
q. FERR
LIGL

 

LIGF .1 \ FRUT

 

 

 

177

Although some differences were noted, the resulting
patterns were quite similar. Thus extracts prepared
from dried material provide a reasonable indication of

the phenolics present in the living plant.

A
1. i

 

TAXONOMIC TREATMENT

Lyonia Nuttall

Lyonia Nutt. Gen. N. Am. Pl. 1: 266. 1818, nom.

cons., non Eyonia Elliott. Sketch Bot. S.C. 1: 316. 1817

 

(= Cynanchum L., Asclepiadaceae) nec Lyonia Rafinesque.

 

Med. Repos. N.Y. ser 2, 5: 353. 1803 (= Polygonella Michx.,

Polygonaceae).

Xolisma Raf. Amer. Monthly Mag. & Crit. Rev.

4: 193. 1819.
Neopieris Britton and Brown. Ill. Fl. ed 2, 2: 690.

 

1913.
Desmothamnus Small, Shrubs Fla. 96. 1913.

Arsenococcus Small E3 Small and Carter. F1. Lan-

 

caster Co. 218. 1913.

Evergreen or deciduous shrubs, rarely small trees,
'usually from an underground rhizome, with round to terete
or angled branches. Leaves alternate, short-petioled,
entire or shallowly toothed or serrulate, nearly glabrous,
or vwith hairs or peltate scales; winter buds ovoid, with
two outer scales. Inflorescence axillary (sometimes
appearing terminal but falsely so), corymbose, racemose or

paniculate, the racemes sometimes contracted into

178

 

179

axillary fascicles; each flower in the axil of a small

bract and with two lateral bracteoles at the base of the

pedicel, these sometimes quickly deciduous. Calyx 5-,

rarely 4- to 8-lobed, the lobes valvate or reduplicate

in bud, persistent or deciduous in fruit. Corolla

cylindric-campanulate to urceolate or globose-urceolate, 1
with 5 (4 - 8) short lobes, white to red, glabrous to
pilose or lepidote. Stamens 10, rarely 8 - 16; filaments
flattened, S-shaped, expanded near base, glabrous to hairy
or roughened, with or without a pair of short spurlike
appendages in the back near the apex; anthers obtuse, the
lobes parallel, lacking apical awns, dehiscent by large
terminal pores, always with a white line of disintegration
tissue on the back of each lobe extending at least along
the apex of the filament and along the upper edge of the
appendages when present; pollen tricolporate, surface
psilate, indistinctly tectate, in tetrads without viscin
strands. Stigma truncate to capitate; style columnar to
fusiform, straight, not exerted; ovary superior, five-
locular, the placentae large, undivided; disc an enlarge-
ment of the ovary wall, variously develOped. Capsule
subglobose to ovoid, five-angled, loculicidal, with paler,
thickened sutures which may separate from the five valves
in dehiscence; placentae persistent at the top of the

columella; seeds scobiform, with a loose thin testa.

180

Asia (Kashmir to Japan, south to Malaya), Greater Antilles,
and North America (eastern United States and Mexico).

Lectotype species: Lyonia ferruginea (Walt.)

 

Nutt.: see I.C.B.N. 261. 1956, and Rickett and Stafleu,

Taxon 9: 75. 1960.

Key to Sections, Species and Varieties
ofIEyonia

 

 

A. Lower surface of leaves, pedicels, and calyx lepidote
(Section Lyonia) . . . . . . . . . . . . B.
B. Capsule subglobose, 2.5-4.0 mm. long; leaves

only slightly pubescent or glabrous beneath. .
. . . . . . . . . . l. E. squamulosa
B. Capsule ovoid, oblong-ovoid or oval, 3.0-
6.0 mm. long; leaves usually moderately
pubescent beneath . . . . . . . . . . C.
C. Leaves conspicuously reduced toward the
ends of the branches, major veins not
depressed, margin not revolute or only
slightly so; ultimate branchlets

rigidly ascending . . . 2. E. fruticosa

 

C. Leaves not conspicuously reduced toward
the branches, major veins usually
depressed, margin usually revolute;
ultimate branchlets not rigidly

ascending. . . . . . 3. E. ferruginea

A

 

181

A. Lower surface of leaves, pedicels, and calyx glabrous

or pubescent but not lepidote . . . . . . . . D.

D.

Inflorescence paniculate; corolla urceolate,
4 mm. long or less; calyx lobes less than 2 mm.

long; leaves pilose (Section Arsenococca, E.

 

ligustrina the only species) . . . . . . . E.

 

E. Inflorescence naked or with only a few
leafy bracts (chiefly mountains and Pied-
mont south of Va. and mountains, Piedmont
and Coastal Plain north of Va.). . . . .

. . . . . . . . .4a. var. ligustrina

 

E. Inflorescence conspicuously leafy bracted
or at least the lower panicles with large
bracts (chiefly Coastal Plain from SE Va.
to W to Tex. and Ark.) .4b. var. foliosiflora
Inflorescence corymbose (fasciculate); corolla
elongate, longer than 4 mm; calyx lobes 2 mm
or longer; leaves glabrous or with slight to
moderate pubescence on veins, sometimes glandu-
lar dotted (Section EEEEE) . . . . . . . F.
F. Leaves evergreen, rigidly coriaceous, with
intramarginal vein; corolla less than 5 mm

wide, usually pink; flowers in leaf

axils . . . . . . 5. L. lucida

 

 

182

F. Leaves thinner, deciduous, lacking intra-
marginal vein; corolla more than 5 mm
wide, usually white; flowers borne on

leafless branches . . 6. E. mariana

Specimens Examined
In the citation of herbarium specimens, abbrevi-
ations of institutions follow the fifth edition of Index

Herbariorum (Lanjouw and Stafleu, 1964) and Taxon 15:

 

335, 1966. The herbaria and their abbreviations are:
Arnold Arboretum (A); Duke University (DUKE); Escuela
Nacional de Ciencias Biolégicas, Mexico (ENCB); Florida
State University (FSU); University of Georgia (GA); Gray
Herbarium (GH); Mississippi State College (MISSA); Uni-
versity of Michigan (MICH); Missouri Botanical Garden
(MO); Michigan State University (MSC); University of
North Carolina, Chapel Hill (NCU); New York Botanical
Garden (NY); University of Texas (TEX); National Museum,
Smithsonian Institution (US); Vanderbilt University
(VDB); University of Wisconsin (WIS).

The specimens from which data were obtained for
the numerical taxonomic study are indicated by an asterisk
(*). Ordinarily one specimen per county has been cited

for species in the United States.

A: I
|

 

183

Lyonia Nutt. sect. Lyonia
Andromeda L. sect. Lyonia Gray, Syn. F1. N. Am,
2(1): 32. 1878, in part. Xolisma Raf. sect. Lyonia Rehd.,
J. Arnold Arb. 5: 54. 1924.
Lyonia Nutt. sect. Eulyonia K. Koch, Dendr. 2(1):

119. 1872, in part.

Evergreen shrubs or small trees with terete or
angled branches. Leaves entire or shallowly toothed and
lepidote. Flowers in dense axillary fascicles; corolla
urceolate; filaments unappendaged or with only very small
spurs. Capsule elpidote with pale much thickened sutures
which separate as a unit from the five valves in dehiscence.
Coastal Plain from Florida to South Carolina, montane

areas of eastern and southern Mexico, and the Greater

Antilles.

l. Lyonia squamulosa Mart. & Gal.

Lyonia sqgamulosa Mart. & Gal., Bull. Acad. Brux.
9: 542. 1842. Xolisma squamulosa (Mart. & Gal.) Small,
F1. N. Am. 29(1): 66. 1914. Type: MEXICO: "NE. du
coffre du Perote, surout pres de la Venta del Esquilon,

a 4,500 pieds de hauteur absolute." E. Galeotti 1797

(BR?, not seen).

A widely branched shrub usually under 3 m
(occasionally to 5 m). Young branches covered with

ndJnIte ferrugineous scales, becoming glabrous or with

184

only sparse covering of scales with greater age, also
often with short nonglandular hairs. Buds lepidote, ovoid,
1.0-2.0 mm long. Leaves evergreen, moderately coriaceous,
alternate, short petioled, entire or rarely shallowly
toothed, sometimes with revolute margins, elliptic, oval,
ovate, or obovate-elliptic, 1.0-6.0 cm long, 0.5-3.0 cm
wide, with tips acute, acuminate or rounded and bases
attenuate or cuneate, adaxial surface lusterous, glabrous
to slightly lepidote, with slight pubescence on mid-vein,
abaxial surface densely lepidote, rarely with nonglandular
hairs intermixed. Flowers in axillary corymbose fascicles;
pedicels lepidote, each with a pair of bracteoles at the
base. Calyx S-lobed, persistent, lobes triangular, 1.0-
2.0 mm long, 0.5-2.0 mm wide, abaxial surface lepidote

and sometimes also nonglandular pubescent, adaxial sur-
face sparsely to moderately nonglandular pubescent.
Corolla urceolate, with 5 short lobes, white, lepidote,
2.5-4.0 mm long, 2.5-3.5 mm wide. Stamens 10, inserted

at the base of the corolla; filaments flattened, expanded
:near the base, S-shaped, roughened, unappendaged or with
'very short spur-like appendages on the back near the apex;
anthers obtuse, the lobes parallel, lacking apical awns,
dehiscent by large terminal pores, always with a white line
«of disintegration tissue in the back of each lobe extend-
ing at least along the apex of the filament and into the

short appendages if present. Style columnar or fusiform,

 

185

not exerted; ovary superior, 5-1ocular, tomentose and
lepidote. Capsule subglobose, S-angled, 2.5-4.0 mm long,
2.5-4.0 mm wide, loculicidal, with paler thickened sutures
which separate as a unit from the five valves in dehis-
cence, lepidote and often nonglandular pubescent, espe-
cially at the base. Seeds ca 80 per capsule, very

narrow, averaging 1.5 mm long (Figure 35).

Distribution: Mexico, in temperate montane
regions along the eastern Sierra Madre from Nuevo Leon
southeast to Veracruz, Puebla, and Oaxaca, Chiapas and
Guerrero, mostly between 1000 and 2400 meters (Figure 36).

Flowering: throughout the year but especially
March-June.

Representative specimens: MEXICO. Chiapas.

4 miles north of Jitotol on rd. to Pueblo Nuevo Solista-
huacan, municipio of Jitotol, 5500 ft, Breedlove £222
(ENCB, US); Lago de Monte Bello, 25 miles east of La
inrinitaria, Municipio of La Trinitatia, 5200 ft, Breedlove
714959 (FSU*); near the Lago de Monte Bello, southeast of
Comitan, Miranda 2__6_§_4_ (US); 5 miles south of Pueblo Nuevo
Solistahuacan along rd. to Jitotol, Municipio of Jitotol,
5300 ft, 39133 & Breedlove E2233 (ENCB*); 3 kilometers
northwest of Pueblo Nuevo Solistahuacan, Municipio of
Pueblo Nuevo, 5400 ft, Zuill 126 (ENCB). Guerrero.

10 kilometers southwest of Amojileca, Municipio of

Chilpancingo, 2050 meters, Rzedowski 23567 (ENCB, use”.

 

v

 

Figure 35.

186

Representative specimen of Lyonia squamu-

losa (Ventura 9205, ENCB).

 

 

187

Fan.
10c

I‘Jtodc

' nt

  

HMMMIU us LA -‘UIIA MACDOIAL as M“ ”MARIA!

[WI W‘NK‘O IAOIOIAL “UFO. I) '

PLANT“ WAS

rochn

AH

 

188

 

 

.mmoHsEdmwm nanomm mo coausnwuumwo

. mm 85m:

 

189

 

 

 

 

 

 

 

 

 

no. 94.
30.55.3404 (a: 4(30- n .288
g.0>30.v >00. 000.
«...-l... I.- in IL I a
CI '1 . D
n.4x
‘
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x f
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n
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a a

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....

.‘fl

 

 

 

190

Hidalgo. Jacala, Kenoyer 312'! 16 November 1937 (M0);
El Estribo, Carretera Tulancingo-Tenango, 44 kilometers,

2000 meters, Leyva 536 (ENCB, TEX*); Tenango de Doria,

 

2100 meters, Puig 2100 (ENCB); alrededores de Zacualtipan,

Quintero 808 (ENCB MSC*); 6 kilometers southeast of

 

Zacualtipan, 2150 meters, Rzedowski & Madrigal 29457
»(ENCB*); Cerro de las Cruces, Aqua Blanca, Vela 586

(ENCB*). Nuevo Leon. Cerro del Viejo, 15 miles west of

 

Dulces Nombres, Municipo of Zaragoza, Meyers & Rogers £323!
20 August 1948 (M0); Dulces Nombres, 24N, 99, 5-100.5° W,
Meyers & Rogers EZEE (MO); Potrero Redondo west to Puerto.
a Laguna Sanchez, municipio of Villa Santiago, Mueller
.EEEZ (TEX*). Oaxaca. Cumbre de Huehuetlan near Teotitlan,

Conzatti 4111 (US); without definite locality, 6000 ft,

 

Galeotti 1849 (US); near Cerro Zempoaltepetl, southwest

 

slopes of mountain near Tlahuitoltepec, 2400 meters,

Hallberg 953 (ENCB*, US); Patio de Arena, ca 9500 ft,

 

Ihigh point of trail from Zacatepec to Tamazulapan,
:Leonard 3931 (NCU*); Villa Alta, Eg£§y_32§ (ENCB).
:Puebla. Serras Decoration archeveché, near Puebla,
Arsene 1221!: (MO, US); Chinantla, Liebmann 8629 (MSC*);
£1350 meters, Municipio of Pahuatlan, Egig_§22§ (ENCB);
near Honey Station, 5800 ft, Pringle E993 (US); Zaca-
poaxtla, Salazar _s_.3_1_., 15 April 1913 (US). Egg Eu_i_s_
Potosi. 4 kilometers northeast of Guadalcazar, 1800

meters, Dias 3739 (ENCB); Sierra de San Miguelito, near

 

 

191

the Cueva del Mezquite, 2200 meters, Rzedowski 5451

 

(ENCB*); Cerro Grande, ca 6 kilometers northwest of

Guadalcazar, 2000 meters, Rzedowski 5956 (ENCB); Mesita

 

de Gallos, ca 20 kilometers southeast of Zaragoza, 2050

meters, Rzedowski 6100 (ENCB*). Tamaulipas. highest part

 

 

of the Sierra de Tamaulipas, 900-1500 meters, Rd. from
Rancho Las Yucas to Santa Maria de los Nogales, Martinez
& Luyando F-l917 (TEX*). Veracruz. Santa Ana, Xalacingo,

2150 meters, Vela & Martinez 1189 (ENCB); Agua Cruz,

 

municipio of Jalacingo, 1650 meters, Ventura EQEQ (ENCB);
Paso Enriquez, municipio of Yecuautla, 1600 meters,
Ventura EZEZ (ENCB*, WIS); Esquilon, municipio of
Jilotepec, 1310 meters, Ventura 3331 (ENCB); La Florida,
municipio of Atzalan, 1750 meters, Ventura §§§§_(ENCB);
Tezchiquiapa, municipio de Jalacingo, Ventura 222§_(ENCB).
Martens and Galeotti (1842) reported both E.

squamulosa and E. ferruginea from Mexico. Sargent (1893)

 

recognized only E. ferruginea as occurring in Mexico,

 

while Standley (1924) listed only E. squamulosa, but
stated: "The Mexican plant differs only in its slightly
shorter capsules and it is doubtful whether itrepresents
a distinct species."

Small (1914) included all Mexican plants in E.

gquamulosa, separating this taxon from E. fruticosa and

 

E. ferruginea by its subglobose capsules and the width

 

of the calyx. Rehder (1924) implied that several members

 

 

192

of the genus occur in Mexico, and Wood (1961) stated that

E. ferruginea occurs in Mexico about the eastern Sierra

 

Madre from San Luis Potosi south to Oaxaca, but leaves
the question somewhat open by saying that "one or two"
species occur in "eastern and southern Mexico." Lyonia

§quamulosa is a variable species, thus Martens and Galeotti

 

were only describing two extremes from the total pattern
of variation.
This variation is interesting in that it parallels

that between E. fruticosa and E. ferruginea on the Coastal

 

Plain in the southeastern United States. Some plants

tend to have large elliptic leaves with a dense covering
of dark ferrugineous scales on the abaxial surface, while
others are small shrubs with ascending branches bearing
small obovate leaves with acute or blunt tips and light
gold or whitish scales! The former are superficially like

3E. ferruginea; the type specimen may be of this group.

 

.Martens and Galeotti state that it blooms in May and
compare it with the West Indian E. jgmaicensis. The
latter taxon shows some similarities to E. fruticosa.
Dhartens and Galeotti may have called such plants E.

ferruginea, especially if they were following the nomen-

 

<:1ature of Nuttall (1818). The flowering time is given
.as June. I found it impossible to satisfactorily separate
two groups of Mexican plants as there is a continuous

range of variation between them and in many respects the

 

 

193

characters seemed to show a reticulate pattern within the

taxon as a whole. Lyonia squamulosa is clearly separated

 

from E. fruticosa by the glabrous and densely lepidote

 

abaxial leaf surface, its subglobose angular capsules, its
larger size, the more elliptic leaves, and its usual lack
of very reduced leaves along the upper portions of the

branches. It can be differentiated from E. ferruginea by

 

the general lack of nonglandular pubescence on the abaxial
leaf surface, the unrevolute or only slightly revolute
leaf margins, the lack of noticeable depression of the
major veins of the leaf, and by its subglobose angular

capsules. Lyonia squamulosa also has a tendency to have

 

darker orange or brown peltate scales, and more elliptic
and ovate leaves. Lumping this taxon with either E.

ferruginea or E. fruticosa would completely wreck the

 

 

consistency of the characters upon which these taxa are
based.

A specimen collected by Rzedowski (no. EEEQ) in
San Luis Potosi is very unusual in that it has a fine
nonglandular pubescence on the lower leaf surfaces.
Extremely revolute leaves are present on Rzedowski 5451,
also collected in San Luis Potosi. A specimen from
Veracruz (Ventura 5655) showing small obovate leaves
with rounded tips is also unusual. More collecting is
needed to determine if these characters are limited to

these areas. There is still insufficient material to

194

reach a complete understanding of the variation of this
taxon but it seems to be quite polymorphic, showing

relationships with both E. fruticosa and E. ferruginea.

 

Possibly this variability is due to its widely scattered
distribution in temperate montane areas.

The vegetation of the montane areas on the eastern
escarpment of the Anahuac Plateau or Central Mesa (begin-
ning about Xilitla, San Luis Potosi and extending through
Hildago, Puebla, and part of Veracruz) and certain high-
lands in Oaxaca and Chiapas were discussed by Miranda

and Sharp (1950). Lyonia squamulosa is listed as occurring

 

in the understory of the mixed oak forest with Gaultheria

hirtiflora and Heberdenia penduliflora. Important trees

 

 

are Quercus, Liquidambar, Symplocos, Prunus and Phoebe.
It is also listed as an understory component of the pine
forest where it occurs with the following shrubs: 922:
ostegia icosandra, Zinowiewia integerrima, Ascyrum hyper-

icoides, Vernonia patens, and Eyrica pringlei; trees in

 

the overstory include Eiggg and Quercus. Lyonia squamulosa
is also important in disturbed communities which follow
after the original pine or mixed oak forest has been
destroyed by timber cutting, agriculture or fire. It
sometimes invades in almost pure stands! In such com-
munities it commonly occurs with Myrica pginglei, Bac-
charis conferta, Rubus spp. and Pteridium aquilinum var.

feei (Miranda and Sharp, 1950).

195

Although Eyonia squamulosa has not yet been
reported from Guatemala (Standley and Williams, 1966)
it should be searched for there. It has been collected
in Chiapas, and Steyermark (1950) suggests that the vege-

tation of the Cuchumatanes and Sierra de las Minas in

 

Guatemala may have much in common with the montane regions

of Mexico.

2. Lyonia fruticosa (Michx.) G.S. Torrey E2 Robinson

 

Lyonia fruticosa (Michx.) G.S. Torrey E3 Robinson,

 

Prox. Am. Acad. Arts 51: 527. 1916. Andromeda ferruginea

Walt. var. fruticosa Michx. Fl. Bor. Am. 1: 252. 1803.

 

Xolisma fruticosa (Michx.) Nash, Bull. Torrey Bot. Club

 

22: 153. 1895. Lyonia ferruginea (Walt.) Nutt. var. fru-
ticosa (Michx.) Rehder Ea Bailey, Cyclop. Am. Hort. 3:

960. 1900. Type: "in Florida and Georgia" Michaux (P?,

not.seen).

A small shrub, often colonial, usually rigidly
ascending, up to 1.5 m tall (rarely 3 m). Young branches
tfliin, rigid, coated with minute ferrugineous scales and
often densely nonglandular pubescent, becoming glabrous
or lightly pubescent with age. Buds ovoid, lepidote,
(1.5-2.0 mm long. Leaves evergreen, moderately coriaceous,
alternate, short—petioled, entire or rarely obscurely
toothed, only rarely with revolute margins, obovate to

oblanceolate, sometimes oval or elliptic, 0.5 to 5.5 cm

 

196

long, 0.3 to 2.8 cm wide, with tips acute, abruptly
acuminate or obtuse, and bases cuneate, obtuse or atten-
uate, adaxial surface glabrous to slightly lepidote with

a slight nonglandular pubescence on mid-vein, abaxial

 

surface lepidote and usually nonglandular pubescent,
scales easily falling off thus mainly absent on older
leaves, leaving their lower surface with a whitish color;

leaves much reduced toward the end of the branches, flat

I...

or with a slight upward curve, with prominently reticulate,
not depressed veins. Flowers in axillary corymbose fas-
cicles, often densely clustered in the upper portions of
the branches; pedicels lepidote and nonglandular pubescent,
each with a pair of bracteoles at the base. Calyx 5-1obed,
persistent, lobes triangular, 1.0-1.5 mm long, 0.5-1.0 mm
'wide, with abaxial surface lepidote, and adaxial surface
'with.sparse to moderate nonglandular pubescence. Corolla
'urceolate, with 5 short lobes, white, lepidote, 2.5-4.0 mm
Long, 2.5-4.0 mm.wide. Stamens 10, inserted at the base
<3f the corolla; filaments flattened, expanded near the
base, S-shaped, roughened, unappendaged or with very small
spur-like appendages on the back near the apex; anthers
cflrtuse, the lobes parallel, lacking apical awns, dehiscent
by large terminal pores, always with a white line of dis-
ithegration tissue on the back of each lobe extending
along the apex of the filament and into the short spurs

if present. Style columnar or fusiform, not exerted;

197

ovary superior, 5-locular, tomentose and lepidote. Cap-
sule ovoid, 5-angled, 3.0-5.0 mm long, 2.5-4.5 mm wide,
loculicidal, with paler thickened sutures which separate
as a unit from the five valves in dehiscence, lepidote
and moderately to densely nonglandular pubescent, espe-
cially at the base. Seeds ca 140 per capsule, very narrow,
averaging 2.2 mm long (Figure 37).

Distribution: Coastal Plain from Florida to
Georgia (Figure 38).

Flowering: throughout the year but mainly in
May and June.

Representative specimens: UNITED STATES. Florida.
Alachua Co.: 9 miles south of Gainsville, 92525 s.n.,
28 December 1935 (NCU). Baker Co.: 5 miles east of
(Jlustee, Weigand & Manning 2389 (GH). Bradford Co.:
locality unknown, Murrill £32}! (MO). Brevard Co.: 10

Iniles south of Melborne, Deam 57588 (NY). Broward Co.:

 

 

4 sales west of Deerfield, Eggg_§2§§§ (NY). Charlotte Co.:
(Saloosa Forest, Parrott El (DUKE). Clay Co.: 3 miles
west of Green Cove, Murrill 357 (US). Collier Co.: Marco
IIsland, Lakela 22222 (FSU, GH). Columbia Co.: 24.5 miles
ruxrth of Lake City, Godfrey E3112 (FSU*). Dade Co.:

Buena Vista, Moldenke 326a (NY*, MO); Miami, Tracy 9210
(GB, MSC, TEX, NY, US). DeSoto Co.: south of Frostproof,
Small E DeWinkeler 9588 (NY*). Duval Co.: vicinity of

Mayport, Godfrey E Reinert 61200 (FSU*) . Gilchrist Co.:

 

 

198

 

Figure 37. Representative specimen of Lyonia

fruticosa (Nash 700, MSC).

 

199

   

bani: mm (Michx.)form
m‘. [.10- m

PLANTS 0‘ CENTRAL PENINSULAR FLORIDA
.‘ouzcuo m vu..~nv ()0 comm . nu coulnv
"v I): - y ~A‘u “1.: f. I.“

700. Xolisma fruhcou .\l . ...

bar-nu ...c. 3...".... .4,” ”n.4,”,

 

200

Figure 38. Distribution of Lyonia fruticosa.

 

201

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

a H r
.
... . I
v

p
4 I': V

b 4

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1g}, .
. J?
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m ,1 ‘ . -
nonun- ‘ x '

-_-- t ' i x - ' \
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I I. n- ' ' ' ,.--/ , \
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202

3.8 miles west of Gilchrist-Alachua Co. line, about 6
miles east of Trenton, EE£§_EE.21° £223 (FSU, GH, NCU, US).
Glades Co.: Indian Mound near Citrus Center, Small 9915
(GH). Highlands Co.: 3.5 miles south of Baias Den, 33y
EE‘EE. 2E2; (NCU*). Jefferson Co.: 4 miles east of

Drifton, Kral 6422 (FSU*, GA, GH). Lake Co.: vicinity

 

of Eustis, Nash 700 (A, MSC, NY, US). Lee Co.: Pine

 

Island, Tracy 7263 (GH, MSC, NY, US). Leon Co.: junction

 

 

of US 20 and Fla. 267, Musselman 4123 (NCU). Levy Co.:

 

4 miles north of Cedar Key, Godfrey 56618 (DUKE, FSU, GH).
Liberty Co.: between Telogia and Sumatra, Godfrey 62634
(FSU, NY, TEX). Madison Co.: 8 miles north of Greenville,

Kral 3758 (FSU). Manatee Co.: near Bradentown, £353

 

Churchill E32}! 2 April 1923 (A, GH). Marion Co.: on

 

Rt. 40, 2.3 miles west of junction with Rt. 19, Egg§.§,g.,
16 July 1959 (MSC). Martin Co.: north of Port Salerno,
Lakela 32231 (GH*). Nassau Co.: Amelia Island, Judd 314
(MSC*). Osceola Co.: marsh near Kissimmee, Hunnewell 8708
(GH). Palm Beach Co.: 3 miles east of Sunshine Parkway
toward Boca Raton, McDaniel 9136 (DUKE). Pasco Co.: 1
mile east of Gowers Corner, Bay. 22 El: 2231 (GH*).
Pinellas Co.: 5 miles south of Port Richey, 33y. EE.E£°
22§2_(FSU, NCU*). Polk Co.: near Lake Wales, Demaree
ggggg (FSU*). Putman Co.: Welaka, University of Florida
Conservation Reserve, EEE§.§rE-r 17, 18 July 1959 (FSU,

MSC*, US). St. Johns Co.: without definite locality,

203

Reynolds 274 (US). St. Lucie Co.: 3 miles southwest of

 

Fort Pierce, McCart 10485-A (NCU). Sarasota Co.: near
Phillippe Creek, about 3 miles south of Sarasota, Ward 3135
(GH, NCU). Taylor Co.: at Steinhatchee, Godfrey 55953

(FSU*, GH, NCU); vicinity of Keatons Beach, Godfrey 61656

 

(FSU*, US). Volusia Co.: northeast edge of DeLand Air-

port, Prichard 648 (NCU*). Wakulla Co.: Apalachicola

 

National Forest, along Forest Rd. 309 between Helen and

 

Lost Creeks, Clewell 1662 (FSU*). Georgia. Appling Co.:

4 miles northwest of Baxley, £222§.l§lgi (DUKE). Bacon
Co.: 1.8 miles east of Alma, Duncan EEEEE (GA). Berrien
Co.: 2 miles north-northwest of Ray City, Faircloth 1947
(NCU). Brantley Co.: just east of Brantley-Ware Co. line,

Judd 291 (MSC*). Brooks Co.: 6 miles south of Morven,

 

Bozeman 2211 (NCU*); 3.7 miles west of Lowndes-Brooks Co.
line, Faircloth 3083 (NCU*). Camden Co.: along Crooked
River, at the Crooked River State Park, Bozeman 22§§_(NCU*).
Charlton Co.: 1.7 miles north of St. George, Bozeman 3312
(NCU). Clinch Co.: at Argyle, Clewell g§§l_(FSU*). Coffee
Co.: 7.6 miles east of Douglas, Duncan EEEEE (GA). Col—
quitt Co.: Indian Creek, 3.3 miles northwest of Berlin,
Faircloth 2664 (NCU). Cook Co.: about 7.5 miles north of
Sparks, 2293.321 (MSC*). Echols Co.: Tom Creek Swamp,

0.7 mile north of Fla. state line, Bozeman EEEQ (NCU*).
Effingham Co.: 4 miles north of Stillwell, Duncan EEEEE

(GA). Glynn Co.: 0.3 mile east of junction of US 84 & l7,

204

Bozeman 6440 (NCU). along Rt. 17, north of Fancy Bluff

Creek, Mellinggr 232°! 11 June 1960 (GH*). Lanier Co.:

 

along US 84, just east of the Alapaha River, Judd 281

 

(MSC*). Liberty Co.: 3 miles southwest of Hinesville,
Wiegand E Manning 2387 (GH). Long Co.: along US 82, about

0.1 mile west of Liberty-Long Co. line, Judd 325 (MSC*).

 

Lowndes Co.: 1.5 miles west of Dasher, Faircloth 1653

 

(NCU). McIntosh Co.: 1.3 miles south of north tip of
Sapelo Island, Duncan EQEEE (DUKE, GH, NCU, US). Pierce
Co.: about 0.5 mile south of Patterson, 22123.222l3.(NCU)'
Tattnall Co.: 3.7 miles 24° southwest of Glenville,
Padgett 111 (GA). Tift Co.: near Tifton, Haltern 292:!
August 1926 (GA). Ware Co.: near Waycross, Godfrey §Q§Q§_
(FSU, GH, TEX); 0.3 miles west of Ware-Brantley Co. line on

US 84, Judd 290 (MSC*). Wayne Co.: 5 miles east of Broad-

 

hurst, Bozeman 6186 (NCU).
This taxon is quite distinct from the closely

related and sympatric Lyonia ferruginea; there is a long

 

tradition beginning with Michaux (1803) in support of this
view. Recently there has been a tendency to lump E.

fruticosa with E. ferruginea (eg., Radford, Ahles, and

 

Bell, 1968); this I consider to be a serious error in
that the two taxa are separated by several good morpho-

logical characters and very few plants appearing to be

 

intermediate have been discovered. Perhaps Redford EE.El°

 

205

confused these two taxa because only L. ferruginea occurs

 

in the range of the Flora 2E the Carolinas.

 

Michaux (1803) lists E. fruticosa as a variety of

 

E. ferruginea, giving as its distinguishing characters

 

the growth form, the prominent reticulations of the veins
of the leaf, the nonrevolute margins and the difference

in flowering time (E.g., June for E. fruticosa and April

 

for E. ferruginea). Asa Gray also considered these two

 

taxa as distinct, pointing out that E. ferrgginea has

 

narrower (sic) less reticulated leaves while E. fruticosa

 

has mostly cuneate-obovate or rhomboidal leaves that are
conspicuously reticulated. Elliott (1817) and Pursh

(1814) called this taxon Andromeda ferruginea, and Nuttall

 

(1818) called it Lyonia ferruginea, giving the name

 

Andromeda (or Lyonia) rigida to the plant known as E.

 

 

ferruginea today! This situation deve10ped from the

 

ambiguous original description of Andromeda ferruginea
given by Walter (1788). Pursh, Elliott and Nuttall took

the description to refer to E. fruticosa, when in fact,

 

it actually describes E. ferrgginea! In Walter's descrip-

 

tion "follis rotundatis margine revolutis, subtus fer-

rugineis” can only refer to E. ferruginea as it has

 

markedly revolute leaves with a dense covering of fer—

rugineous scales beneath. In contrast, E. fruticosa

 

usually has nonrevolute leaves and tends to loose its

ferrugineous scales as the leaves age. This confusion

 

206

does not alter Nuttall's transfer of Walter's Andromeda

 

ferruginea to the genus Lyonia in 1818. Thus Lyonia

 

ferruginea (Walt.) Nuttall is the correct name for E.

 

ferruginea, although Nuttall would have been thinking of

 

what is now E. fruticosa when he used this name! G. Don

 

(1834) and DeCandolle (1839) in following Nuttall, made _ f

the same error. I
Small (1914, 1933), Wood (1961), Long and Lakela

(1971) and Lems (1962, and unpublished notes) all con-

sidered E. fruticosa to be specifically distinct from

 

E. ferruginea. Part of the difficulty involving the

 

identity of this species is that the characters which

separate it from E. ferruginea are best seen in the field.

 

Nash (1895) reports:

 

[L onia fruticosa] is specifically distinct from
XoIisma ferruginea, which grows from 6-12 ft. high,
Has Iight green leaves which are very much rolled
in on the margins, and flowers early in March.

X. fruticosa never attains a height of more than

6 feet, has a much stricter habit, very dark leaves
which are not revolute, and does not flower until
two months later. The difference is quite marked
in herbarium specimens, but in the field there can
be no mistaking it.

 

 

How the leaves are held on the stem, the amount of curva-
ture of the leaf, the appearance and depression of the
major veins, the density of ferrugineous scales on the
abaxial leaf surface, and the growth form or appearance
of the plant are all good characters in differentiating

L. fruticosa from E. ferruginea. The importance of these

 

 

vegetative characters was noted quite early, for example,

 

207

Elliott (1817) states that the leaves of E. ferruginea are

 

"entire" and "convex" with "revolute" margins in contrast

to the "very entire, generally flat" leaves of E. fruticosa

 

(Figure 39).

The table on page 210 summarizes the characters
that Wood (1961), Long and Lakela (1971), Small (1914,
1933), Lems (unpublished notes) and I have found to be

most helpful in differentiating E. fruticosa from E. fer-

 

ruginea.
Small (1914, 1933) used the corolla length as a

distinguishing character, giving the values of 2.5-3 mm

for E. ferruginea as compared to 3.5—5 mm for E. fruticosa.

 

Long and Lakela (1971) also used the corolla length in
differentiating these two taxa, giving a value of 2-3 mm

for E. ferruginea and 4-5 mm for E. fruticosa. Actually

 

 

this is not a good character for separating the two species
as the range of overlap is too great. K. Lems (unpublished

notes) suggested also that E. fruticosa has more densely

 

tomentose leaves, larger flower clusters, wider and shorter

leaves and shoots which continue to grow after flowering.

 

Lyonia fruticosa does tend to have shorter leaves and thus
leaves that are wider in proportion to their length than

those of E. ferruginea, but there is a wide range of

 

overlap between the two taxa in this character. Lyonia

fruticosa does have a slight tendency to be more pubescent,

 

but this character, too, breaks down frequently. I could

208

 

Figure 39. Leaf curvature and position in Lyonia

fruticosa and E. ferruginea. a. E. fru-

 

ticosa. b. E. ferruginea.

 

 

 

TABLE 23.

210

Comparison of Lyonia fruticosa and E. ferruginea.

 

 

 

E. fruticosa

 

E. ferruginea

 

 

10.

11.

Leaves much reduced 1.
toward the ends of the
branches

Margins of leaves 2.
usually not revolute

Leaves tending to be 3.
obovate or oblanceolate

Leaves generally small 4.

Small shrub with rigidly 5.
ascending branches

Leaves flat or arched 6.
upward

Leaf veins not depressed, 7.
prominently reticulate

Ferrugineous scales 8.
usually deciduous-~thus
absent from older leaves

Peak of flowering in 9.
March-April

Slightly longer flowers, 10.
2.5 to 5.0 mm

(average 3.3 mm)

Very abundant in cut 11.

over or disturbed areas

Leaves only slightly
reduced toward the end
of the branches

Margins revolute

Leaves elliptic, ovate
or obovate

Leaves very small to
quite large

Shrub or small tree,
branches not rigidly
ascending

Leaves with convex
curvature

Leaf veins depressed,
not reticulate

Ferrugineous scales
usually not deciduous--
thus present on older
leaves

Peak of flowering in
May-June

shorter
2.0 to 4.0 mm
2.9 mm)

Slightly
flowers,
(average

Not as common in dis-
turbed regions

 

211

not find any significant difference in the number of
flowers per inflorescence.

A difference in flowering time was observed by
Michaux (1803), Pursh (1814) and Nash (1895), but Elliott
(1817) states "the different periods of flowering, as they
were determined by Michaux by cultivation, mark a dif-
ference of habit; yet in the woods this difference is'
not obvious, but there appears to be a constant succession
of flowers from April to June." Which one of these views
is correct? In examining herbarium specimens I found
that both views are correct, in part (Figure 40). In
Georgia there is a clear difference in flowering periods

with E. ferruginea blooming in March and April, and E.

 

fruticosa blooming in June and July! Farther south this

 

sharp distinction breaks down and both can be found

flowering at once, but even in Florida E. ferruginea blooms

significantly earlier than E. fruticosa. In plants grown

 

under uniform greenhouse conditions at East Lansing,

Michigan, E. ferruginea began to flower early in March,

 

while E. fruticosa did not begin to flower until late in

 

April, not coming into full bloom until the end of June!
Most of the specimens observed from the vicinity

of Tampa, Florida, are completely lacking in nonglandular

pubescence on the abaxial leaf surface. This interesting

variation seems to be concentrated in just that region,

 

 

 

212

 

A.mmcflmsuumm

.m n I “mmmmmmmmw .q u o "muozv .mmoflmsuumm am now

 

 

mmooflusum mflcomq mo moofluwm mcflum3oam may mo comHHnmEoo d

 

.os musmHm

213

 

5;

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00000

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>42 an? i mum in

Irp-Ip-r.IL .P- I»
¢¢¢¢¢¢¢¢¢I¢u0
06¢¢¢¢¢¢ .QO

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I

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On

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214

although scattered specimens with only a very slight

pubescence have been observed from throughout the range

of the taxon.

Lyonia fruticosa occurs in hammocks and pinelands

 

in sandy acid soils on the Coastal Plain. It does not
extend as far north as E. ferruginea; I have seen no
collections from South Carolina, and have not seen it
growing in either Jasper or Beaufort counties in South
Carolina despite a careful search for it.

Eyonia fruticosa commonly occurs with other

 

ericaceous shrubs such as L. ligustrina var. foliosiflora,
L. mariana, L. lucida, Leucothoe racemosa, Gaylussacia

spp., Vaccinium spp. and Befaria racemosa. Other

 

associated shrubs include Clethra alnifolia, Ilex glabra,
Cyrilla racemiflora, Cliftonia monophylla and Myrica

cerifera.

 

Harshberger in his Vegetation of South Florida

(1914) included E. fruticosa in his Sand-Pine (Rosmary

 

Scrub), Slash—Pine and Pond—Pine formations. The pond-
pine formation occurs on the ancient dunes, related in
origin to the coastal dunes, which have covered an older,
:flat land of limestone, which is seen in the valleys, or
swales between the ridges. The dominant tree is Pinus
calausa, the understory includes several species of Quercus,
aIui Serenoa repens, Ceratiola ericoides, Vaccinium myrsin-

:Ltes and Lyonia fruticosa. The vegetation of these

 

 

£1_;‘

 

215

ancient dunes is essentially xerophytic because the
water, which falls as rain, rapidly percolates through
the sand. The coriaceous leaves and covering of scales
may be xerophytic adaptations, and indeed I observed in

collecting that E. fruticosa could be kept for several

 

hours without wilting! The slash-pine (Pinus caribaea)

 

formation occurs on the geologically older flatlands.
This forest consists of tall slash-pines of even age
with an open crown admitting light to the forest floor

beneath. Common understory species include Serenoa repens,

 

Myrica cerifera, Quercus spp., Geobalanus oblongifolius,

 

Vaccinium myrsinites, Gaylussacia dumosa, Lyonia fruticosa,

Befaria racemosa, and Crossopetalum floridanum. During

 

 

the dry season the plants are essentially under xeric con-
ditions. The water which falls as rain is quickly lost
by percolation through the sand and by limestone holes

to underground channels (Harshberger, 1914).

3. Lyonia ferruginea (Walt.) Nutt.

 

Lyonia ferruginea (Walt.) Nutt., Gen. N. Am. Pl.

 

l: 266. 1818. Andromeda ferruginea Walt. Fl. Car. 138.

 

1788. Cassandra ferruginea (Walt.) Ndz., Bot. Jahrb. Syst.
11: 146, 148, 224. 1889. Xolisma ferruginea (Walt.) Heller,
Cat. N. Amer. Pl. 6. 1898. Type: (BM?, not seen).
Andromeda ferruginea Walt. var. arborescens Michx.,
Fl. Bor. Am. 1: 252. 1803. Eyonia ferruginea (Walt.) Nutt.

var. arborescens (Michaux.) Rehd. $3 Bailey, Cyclop. Am.

 

H

216

Hort. 3: 960. 1900. Type: "in Florida and Georgia"
Michaux, (P?, not seen).
Andromeda rigida Pursh, Fl. Am. Sept. 292. 1814.

Eyonia rigida (Pursh) Nutt., Gen. N. Am. Pl. 1: 266. 1818.

 

Type: "in barren sandy woods: Carolina to Florida" (not

seen). .I

A widely branching shrub or small tree, often

 

colonial, to 6 m tall. Young branches thin, rigid, some-
what angular, divergent, lepidote and nonglandular
pubescent, becoming glabrous or only slightly pubescent
with greater age; bark of trunk divided into long narrow
ridges by shallow longitudinal furrows, reddish brown,
separating into short thick plates. Buds ovoid, lepidote,
1.0-2.5 mm long. Leaves evergreen, moderately coriaceous,
alternate, short petioled, entire, usually with revolute
margins, elliptic to ovate or obovate, 1.0-7.5 cm long,
0.5-3.0 cm wide, with tips acute or short acuminate, and
bases attenuate or cuneate, adaxial surface glabrous to
slightly lepidote with slight nonglandular pubescence

on midvein, abaxial surface lepidote and usually non-
glandular pubescent; scales persistent, thus usually
present on older leaves; leaves 222 greatly reduced in
size toward the end of the branches, often with a down-
ward curvature; veins obscure, quite depressed. Flowers
in axillary corymbose fascicles, pedicels lepidote, some-

times also nonglandular pubescent, each with a pair of

217

bracteoles at the base. Calyx S-lobed, persistent, lobes
triangular, 1.0-2.0 mm long, 0.5-1.0 mm wide, with abaxial
surface lepidote, and adaxial surface glabrous or with
sparse nonglandular pubescence. Corolla urceolate, with
5 short lobes, white, lepidote, 2.0-4.0 mm long, 2.0-4.0 mm
wide. Stamens 10, inserted at the base of the corolla;
filaments flattened, expanded near the base, S-shaped,
roughened, unappendaged or with very short spur-like
appendages on the back near the apex; anthers obtuse, the
lobes parallel, lacking apical awns, dehiscent by large
terminal pores, always with a white line of disintegration
tissue on the back of each lobe extending along the apex
of the filament and into the short spurs if present.
Style columnar or fusiform, not exerted; ovary superior,
5-locular, tomentose and lepidote. Capsule ovoid, S-angled,
3.0-6.0 mm long, 3.0-4.0 mm wide, loculicidal, with paler
thickened sutures which separate as a unit from the 5
valves in dehiscence, lepidote and moderately to densely
nonglandular pubescent, especially at the base. Seeds
ca 90 per capsule, very narrow, averaging 2.3 mm long,
Figure 41.

Distribution: Coastal Plain from Florida to
South Carolina (Figure 42).

Flowering: mainly in March, April and May.

Representative specimens: Florida. Bay Co.:

5 miles north of Lynn Haven, Lazor 5009 (FSU*); St. Andrews

 

 

218

 

Figure 41. Representative specimen of Lyonia fer-

 

ruginea (Griscom 21596, GH).

 

 

219

   

Lu‘( ‘\

"BIA U} \URTNWL‘TTJL\ FLUuID\
L. “A... Man i m \n

LI nun. “In "I

 

 

220

Figure 42. Distribution of Lyonia ferruginea.

 

221

 

222

State Park, Wooten 208 (FSU*). Brevard Co.: north of

 

Coca, Rhodes 8305 (A). Calhoun Co.: 3 miles west of

 

Blountstown, Coker 3.3., 26 February 1947 (NCU). Clay

 

Co.: sandy bank of Kingsley Lake drain, Camp Blanding,
Totten §.E., 2 April 1943 (NCU, NY*). Columbia Co.:
on bluff of Santa Fe River, northeast of High Springs,

Godfrey 52847 (DUKE, FSU, GH, MISSA, NCU, NY). Dixie

 

Co.: 0.8 mile west of Suwannee River, about 13.9 miles

north of Suwannee, Traverse 736 (GH*). Duval Co.:

 

vicinity of Mayport, Godfrey E Reinert 61199 (FSU*).

 

Franklin Co.: near St. Theresa, Godfrey 53100 (DUKE,

 

FSU, NCU, NY*); St. Theresa, Griscom 21596 (GH). Gulf

 

Co.: near Wewahitchka, Demaree E9332 (NCU). Hernando
Co.: Weeki-Wachee, Cooley §11§_(FSU, NY*). Highlands
Co.: near Lake Placid, Demaree E22§2_(FSU*, NCU). Hills-
boro Co.: sand barrens Hillsboro River, 2.5. Churchill
.§\E°' 9 April 1923 (GH). Jefferson Co.: without definite

locality, Lighthipe 3.3., 15 March 1891 (NY). Lake Co.:

 

near Astor, Clewell 2108 (FSU*); vicinity of Eustis, Nash
El (A, MSC, NY, US). Lafayette Co.: 4.5 miles north of
Day, Duncan 18240 (GA). Leon Co.: near Lake Bradford,

Godfrey 57997 (FSU*). Levy Co.: Cedar Key, Godfrey 56607

 

 

(DUKE, FSU, GH). Liberty Co.: near Hosford, E.g.

Palmer 38526 (A, US). Marion Co.: Ocala National

Forest, Perry 1643 (DUKE). Orange Co.: near Orlando,

 

§.g. Palmer 38306 (A, US). Osceola Co.: north of Lake

 

223

Tohopekaliga, Lemaire 362 (FSU). Pasco Co.: near Jessa-

 

mine, Barnhart 2540 (NY). Pinellas Co.: near Dunedin,

 

Tracy 6648 (GH, MSC). Polk Co.: about 2 miles northeast

 

of Lakeland, Ford EE.El° 2440 (NCU). Putnam Co.: Welaka,

University of Florida Conservation Reserve, Lems 3.3.,

 

17, 18 July 1959 (MSC). St. Johns Co.: about 2.2 miles

south of Crescent Beach, Wood 9329 (NCU*). Seminole Co.:

 

 

6 miles southwest of Sanford, Duncan 4607 (GA). Sumter

 

Co.: west of Bushnell, Kral 7870 (GA, US). Taylor Co.: I

 

between Buckville and Perry, Godfrey 52855 (DUKE, FSU, GH,

 

MISSA, NY*). Volusia Co.: sand dunes opposite Daytona,

Small E DeWinkeler 9423 (NY). Wakulla Co.: St. Marks

 

Wildlife Refuge, Godfrey 71410 (FSU*); between Panacea

 

and Sopshoppy, St. Marks Wildlife Refuge, Trott 181
(FSU*). Walton Co.: without definite locality, Ashe
3.3., 20 March 1927. Georgia. Appling Co.: without

definite locality, Duncan 16290 (US). Bacon Co.: 7.5

 

miles east of Alma, Bozeman 4471 (NCU). Berrien Co.:
6.1 miles north-northeast of Nashville, Faircloth 2142
(NCU). Brantley Co.: on Ga. 259, ca 5.0 miles south of

junction with US 84, Bozeman 4014 (NCU). Brooks Co.:

 

5.5 miles north-northwest of Quitman, Faircloth 1982
(NCU). Bryan Co.: 2 miles west of the Ogeechee River
on Ga. 204, Bozeman £332 (NCU). Camden Co.: Little
<Cumberland Island, Butler 3.3., 20 April 1968 (NCU).

Charlton Co.: 0.3 mile west of Bell Bridge, Bozeman 3033

224

(NCU). Chatham Co.: 5.5 miles south of Victory Drive,
Savannah, Coker 2.n., 27 March 1948 (NCU*). Clinch Co.:
near Homerville, 3333‘3.3., October 1923 (NCU). Coffee
Co.: near Gaskins Spring, Hammock of Seventeen Mile

Creek, Harpgr 2047 (A, GH, NY, US). Colquitt Co.: 1.8

 

miles north of Berlin, Faircloth 3278 (NCU). Cook Co.:

 

6.6 miles west of Adel, Faircloth 2359 (NCU). Echols

 

Co.: 6.9 miles east of Lake Park, Faircloth 2308 (NCU).

 

Effingham Co.: about 3 miles south of Stillwell, Bozeman
'EEZE (NCU). Evans Co.: 3.8 miles northwest of Claxton,
Duncan EEEEZ (NCU*, TEX). Glynn Co.: Jekyl Island,
93£f£!.§r2°' 21 March 1890 (A*); near Brunswick, Sargent
.§\E°' 31 March 1914 (A*). Liberty Co.: just east of
city limits of Hinesville on US. 82, £333_EEE (MSC*).

Long Co.: near Moody's Camp, Bozeman 2503 (NCU*).

 

Lowndes Co.: 12.6 miles north-northeast of Valdosta,

Faircloth 3705 (NCU). McIntosh Co.: about 0.6 miles

 

southeast of Fort Barrington, Bozeman 434 (NCU*). Mont-

 

gomery Co.: along county rd. S-1125, 0.4 mile east of
Oconee River, Bozeman EZEE_(NCU). Pierce Co.: west of
Little Satilla River on Ga. 32, Bozeman E2323 (FSU, GH,
NCU, NY, TEX). Telfair Co.: 4.7 miles west of McRae,
2£E§§.§E.El°.lgl (GA). Thomas Co.: 9.3 miles northeast

of Thomasville, Faircloth 2491 (NCU*). Tift Co.: ca

 

0.1 mile northeast of Little River near US 319, Judd 264

(MSC*). Turner Co.: 4.1 miles east-northeast of

225

Ashburn, Faircloth 4402 (NCU*). Ware Co.: near Suwannee

 

Lake, Harper El (NY). South Carolina. Beaufort Co.:

 

near Bluffton, Mellichamp_E.3., 1873 (NY*); on Rt. 46

 

just 2 miles south of city limits of Bluffton, Judd 320

 

(MSC*). Jasper Co.: about 6 miles south of Hardeeville,
31312519333 (NCU*).

The differences between this species and E. EEEEEf
3333 have been discussed above. It is important to repeat
that Pursh (1814) and Nuttall (1818), because they mis-
understood Walter's original description (1788), used the

name E. ferruginea to refer to E. fruticosa. This does

 

 

not effect Nuttall's transfer of Walter's Andromeda fer-

 

ruginea to Lyonia.

Lyonia ferruginea is a variable species, showing a

 

wide range of leaf shape and size, amount and nature of
pubescence, amount of curvature of the leaf margins and
depression of the major veins of the leaf. Toward the

extreme southern edge of the range (about the latitude

of Highlands County, Florida) forms with very small and
extremely revolute leaves occur.

Lyonia ferruginea extends farther westward on the

 

Florida panhandle and farther northward into Georgia than

does E. fruticosa. Its farthest northern extent is in

 

JaSper and Beaufort counties in South Carolina where it
is quite rare. I discovered a small colony near Blufton

(Beaufort Co.) where it occurs as an understory shrub

226

in an oak-pine forest. Other understory elements are

palmettos, Ilex spp. and Vaccinium spp. Only this one

 

clump could be located in one day's search of the area.
Lyonia fruticosa was not found and the open pine-palmetto
flatwoods where it is abundant were absent. The woods
which shelters this small colony is near developed areas
and thus is probably in danger of being destroyed in the
near future.

Lyonia ferruginea occurs in hammocks with acid

 

humus, sandhills, pine flatwoods, and pine—evergreen
oak scrub. From my limited collecting experience it
seems to be more common in shaded areas, or areas where
oaks are common, or where oaks and pines are mixed. In

contrast, E. fruticosa abounds in the sunny Open pine-

 

palmetto flatlands, or in scrub vegetation (mainly pal-

mettos, ericads, Ilex, Myrica and other small shrubs)

 

dominating areas where the pines have been cut. Lyonia

figrruginea seems to be less abundant in southeastern

 

Georgia than E. fruticosa, possibly due to the activities

 

of man which have tended to increase the plant communities

favorable to E. fruticosa at the expense of those favoring

 

E. ferruginea.

I found both species growing side by side along
[1.5. 84 in a pine-oak-palmetto forest just east of the
ICLapaha River in Lanier Co., Georgia. Lyonia fruticosa
was the more common and no plants showing intermediate

Characters could be located.

227

Most of the plants appearing intermediate between

E. ferruginea and E. fruticosa can usually be determined

 

 

fairly easily as one or the other taxon after more careful
observations have been made. I believe that this inter-
mediacy is more the result of environmental conditions
than genetic difference. In moist shaded situations

where E. fruticosa is being crowded by surrounding vege-

 

tation its branches are often not rigidly ascending as
is typical. A typical individual when transplanted from
a cut over pine-palmetto flatwoods and grown in a green-
house was found to develop lax, nonascending branches!

Rapidly growing young shoots of E. ferruginea tend to be

 

rigidly ascending and superficially resemble E. fruticosa.

 

In the field Myrica cerifera is similar in overall

 

appearance, especially from a distance.

Lyonia ferruginea and E. fruticosa are both often

 

 

infected by the fungus Exobasidium vaccinii which causes

 

strange shaped galls due to tissue hypertrophy on all

or part of a leaf, a flower bud or an individual flower.
The mycelium is intercellular with branched haustoria
entering the host cells; the basidia extend above the
layer of epidermal cells. The galls are white or pink,
soft and succulent when young, becoming brown and hard
with greater age. This disease is seldom serious except
in shaded areas in wet seasons. Exobasidium vaccinii

also occurs on a wide variety of other ericaceous shrubs

228

such as Vaccinium, Rhododendron, Arctostaphylos, Leu-

cothoe, Gaylusaccia, Cassandra and Andromeda (Westcott,

 

1960; Stevens, 1925).

 

Lyonia Nutt. sect. Arsenococca (Small) Judd, stat. nov.

Arsenococcus Small E3 Small and Carter, Fl.

 

Lancaster Co. 218. 1913, Xolisma Raf. sect. Arsenococcus

(Small) Rehder, J. Arnold Arb. 5:54. 1924.

Deciduous shrubs with round or only slightly
ridged branches. Leaves serrulate to nearly entire,
pilose-glandular, not lepidote. Flowers in axillary or
pseudoterminal inflorescences which are small panicles
(racemes of corymbose fascicles) aggregated near the
end of the growth of the preceding year; calyx lobes
triangular; corolla urceolate, pilose; filaments
pubescent with small spurs. Capsule glabrous to pilose
(glandular pubescent), with pale moderately thickened
sutures, usually not separating as a unit from the five
valves in dehiscence. Coastal Plain, Piedmont and
mountains from Maine south to Florida and west to
eastern Texas, Arkansas and Oklahoma.

Type species: Lyonia ligustrina (L.) DC.

4. Lyonia ligustrina (L.) DC

Rhizomatous widely branching shrub to 4 m. Young

branches slender, glabrous or minutely pubescent, often

229

somewhat flattened at the nodes, yellowish-brown to ashy
gray or reddish. Buds oblong-ovoid, pointed, reddish,
glabrous, 1.5-4.0 mm long. Leaves deciduous, sometimes
partly evergreen in the southern portions of the range,
thin to slightly coriaceous, alternate, short petioled,
serrulate to nearly entire, sometimes with slightly
revolute margins, elliptic to obovate, oblanceolate, or
ovate, 1.5-8.0 cm long, 0.5-4.5 cm wide, with tips acute
to acuminate or mucronate, and bases cuneate or obtuse,
adaxial surface nearly glabrous to densely pilose (glan-
dular pubescent), abaxial surface pilose, often with non-
glandular hairs along the veins. Flowers in axillary or
pseudoterminal inflorescences which are small panicles
(racemes of corymbose fascicles) aggregated near the end
of the growth of the preceding year, naked, with small
bracts or with conspicuous leafy bracts; pedicels pilose
to nearly glabrous, each with a pair of quickly deciduous
bracteoles at the base. Calyx 5-lobed, persistent, lobes
triangular, 0.5-1.5 mm long, 0.5-1.0 mm wide, with abaxial
surface pilose, occasionally also nonglandular pubescent,
adaxial surface glabrous. Corolla urceolate or globose,
with 5 short lobes, white, pilose, 2.0-4.0 mm long, 2.0-
4.5 mm wide. Stamens 10, inserted at the base of the
corolla; filaments flattened, markedly expanded near

the base, S-shaped, the lower portion pubescent, the

upper part strongly roughened, with short spurs on the

230

back near the apex; anthers obtuse, the lobes parallel,
lacking apical awns, always with a white line of disin-
tegration tissue on the back of each lobe extending along
the apex of the filament and into the short spurs. Style
columnar to fusiform, not exerted; ovary superior, 5-
locular, densely pilose. Capsule globose or subglobose,
slightly 5-angled, 2.0-3.0 mm long, 2.0-4.0 mm wide,
loculicidal, with pale thickened sutures, usually not
separating from the 5 valves in dehiscence, sparsely to
densely pilose-glandular. Seeds ca 70 per capsule, very
narrow, averaging 1.4 mm long.

Distribution: Maine southward to Florida, then
west to Arkansas, east Texas and Oklahoma.

Flowering: late April - early July.

This species is extremely variable in stature,
pubescence, leaf size and shape, and inflorescence
structure. Michaux (1803) knowing it from New England
to Florida, described it as ”Magnitudine et figura
admodum variant.” Numerous intergrading varieties have
been described, most of which have few significant genetic
differences. Lyonia ligustrina can be found anywhere
from the grassy balds in the Appalachians to the low
Coastal Plain cypress swamps, and with enough morpho-

logical variations to match the extremes in habitat!

231

This extreme variation has led to many taxonomic and
nomenclatural difficulties which will be discussed with

each variety.

Lyonia ligustrina goes by several common names,

 

some of which are Male-berry, Male—blueberry, He-huckle-
berry, and Seedy-buckberry. All probably result from

the fact that vegetatively it looks like a blueberry but
it is a disappointment to would-be berry-pickers because

of its hard dry capsules--of course inedible!

4a. Eyonia ligustrina (L.) DC. var. ligustrina

 

 

Lyonia ligustrina (L.) DC., Prodr. 7: 599. 1839.

 

Vaccinium ligustrinum L., Sp. Pl. 351. 1753. Andromeda

 

ligustrina (L.) Muhl., Cat. Pl. Amer. sept. 44. 1813.

 

Xolisma ligustrina (L.) Britt., Mem. Torrey Bot. Club 4:

 

135. 1894. Arsenococcus ligustrinus (L.) Small E3 Small
and Carter, Fl. Lancaster Co. 218. 1913. Lyonia ligustrina
var. typica Fernald, Rhodora 43: 625. 1941. Type: "in
Pennsylvania." 52$E.ILINNI probably sheet no. 497.11,
microfiche photo seen).

Andromeda parabolica Duhamel, Arb. ed. nov.
(Nouv. Duh.) l: 191. 1801. Lyonia parabolica (Duhamel)
K. Koch, Dendr. 2: 199. 1872. Type: (unknown).

Andromeda paniculata L. var. nudiflora Michx.,
Fl. Bor. Am. 1: 255. 1803. Type: "in frigidioribus,

per Etats-Unis." (P?, not seen).

232

Leaves elliptic to obovate or oblanceolate, 1.5-
8.0 cm long, 1.0-4.5 cm wide, deciduous, usually not
coriaceous, serrulate or nearly entire, usually not
lustrous above, with nonrevolute margins, adaxial surface
Sparsely to densely pilose, abaxial surface moderately
to densely pilose, with slight to dense nonglandular
pubescence on the major veins. Flowers in axillary or
pseudoterminal, naked or nearly naked panicles aggregated
near the end of the growth of the preceding year. Calyx
with abaxial surface pilose-glandular, occasionally also
nonglandular pubescent. Capsule globose to subglobose,
usually moderately to densely pilose, often also non-
glandular pubescent (Figure 43).

Distribution: South Carolina, northern Georgia
and Alabama to Maine: mountains, Piedmont and Coastal
Plain (Figure 44).

Flowering: May - early July.

Representative specimens: Alabama. Calhoun Co.:

9.1 miles north-northeast of Jacksonville, Clark 16039

 

(NCU). Cherokee Co.: Little River Canyon Park, Kral 33351

 

(FSU, VDB*). Cullman Co.: without definite locality,

Eggert s.n., 19 June 1897 (NY). DeKalb Co.: DeSoto State

Park, near Fort Payne, Kral 36502 (VDB*). Russell Co.:

 

slopes on side of high ridge about 2 miles west of Phenix

City, Duncan 9636 (GA, MO). Connecticut. Fairfield Co.:

 

near New Canaan, Harger 7684 (GH). Hartford Co.:

 

233

 

Figure 43. Representative specimen of Lyonia ligus-

 

trina var. ligustrina (J. A. Churchill s.n.,

 

 

4 July 1935, MSC).

234

 

AM ERICAN FLOR \

nynnin linustrina

Pannsvlvanin, *hWette 20., ‘hiw .y].

rocky gloves of the ?wu'hiorhany iiver mi
43.1113.

b )uly 1°35

J1)hl1 \. (.hllrvlfill. \I.I|.

235

 

Figure 44. Distribution of Lyonia ligustrina var.

 

ligustrina.

 

 

 

 

6
3
2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

IUIIVI-

 

237

Southington, Andrews 3.3., 26 August 1903 (GH*). Middle-

sex Co.: near Essex, Eneguist 451 (NY). New London Co.:

 

near Stonington, Harger 6310 (NCU). Delaware. New

 

Castle Co.: Highland Woods off Darley Rd., Reed 51105

(MSC*). Sussex Co.: 4 miles west of Rehoboth Beach,

near Love Creek, McVaugh 6545 (GH, NY). District of

 

Columbia. Takoma Park, Painter 304 (DUKE). Georgia.

 

Bartow Co.: 4.5 miles southeast of Adairsville, near

Pond no. 10, Greear 64411_(GA, NCU). Carroll Co.: 8.5

 

miles north of Carrolton, Carnes 66 (GA). Cherokee Co.:
about 6.2 miles south 50° west of Canton, Duncan 8360
(GA). Clarke Co.: near Athens, Harper s.n., 2 June 1897
(NY). Cobb Co.: Camp Burt Adams, Duncan 3919 (GA).

Dade Co.: summit of Lookout Mountain, McVaugh 9043
(NCU*, TEX). Dawson Co.: Dawson-Pickins Co. line, about
0.7 mile south of Oglethorpe Mt., Duncan 1188 (GA).
Elbert Co.: along Deep Creek, 8.9 miles west 2° north

of Elberton, Duncan 11696 (GH*, MO). Forsyth Co.:

 

Suwannee Mt., Duncan 3609 (GA, US). Fulton Co.: ridge

 

just southeast of Campbelltown, Duncan 3657 (GA).
Habersham Co.: at Tallulah Gorge, Duncan 13592 (GA).
Hall Co.: Hunt tract, bank of Chattacooche River,
Bishop s.n., June 1940 (GA). Hart Co.: about 10 miles
north of Hartwell near Chandler Ferry, Duncan 7812 (GA).
(Lumpkin Co.: 2.9 miles southeast of Woody Gap, Duncan

116389 (GA). Madison Co.: about 6 miles from Ila,

238

Montgomery 445 (GA). McDuffie Co.: vicinity of Thomson,

 

Bartlett 1653 (NCU, VDB). Murray Co.: near spring at

 

gap west-southwest of Bald Mountain, Duncan 7255 (GA).

 

Rabun Co.: Tallulah Falls, Perry and Straham 976 (NY*).

 

Stephens Co.: along stream just above Toccoa Falls,

Duncan 10034 (CA). Towns Co.: Brasstown Bald, Duncan

 

1460 (GA). Union Co.: North Georgia Experiment Station,
3 miles south of Blairsville, McKay s.n., 15 July 1931

(GA, GH, TEX). Walker Co.: Lookout Mt., Ruth 440 (NY).

 

White Co.: ridge of Tray Mt., Duncan 1811.5 (NY*). Whit-

 

field Co.: Taylors Ridge, Wilson 152 (NY). Kentucky.

 

 

Bell Co.: near Middlesboro, Gleason 8828 (NY*). Casey

 

Co.: southeast corner of county, Braun 2676 (A, US).

 

Laurel Co.: near Corbin, Horsey 2265 (GH). McCreary Co.:

 

south fork of Cumberland River, Braun 1016 (US). Powell

 

Co.: Stanton, McFarland 2398 (US). Maryland. Anne

 

 

Arundel Co.: 0.8 mile north of junction of Md. 175 & 3

in Millerville, Ahles 53582 (NCU*). Baltimore Co.: along

 

Tramble Rd. 0.2 mile east of Pulaski Hwy., Baltars 1827

 

(DUKE, FSU). Cecil Co.: Gray's Hill, 2 miles east of

Elkton, Pennell 13276 (NY). Charles Co.: Wolfden Creek,

 

Cedarville State Forest, Simpson s.n., 5 July 1952 (US).

Garrett Co.: Mt. Lake Park, Shreve 540 (US). Hartford

 

Co.: 0.5 mile south-southwest of Havre de Grace, Shull

157 (NY). Prince Georges Co.: Lanham, Bartlett 2170

 

(NCU, VDB*). St. Marys Co.: near Christmas Hill,

 

239

Walker 39292 (US). Talbot Co.: Miles Creek, 2 miles

 

northeast of Trappe, Earle 1693 (GH). Worchester Co.:

 

along edge of Corker's Creek, 4 miles south of Snow Hill,

 

Judd 33 (MSC). Maine. Cumberland Co.: near Standish,

Fernald 5 Long 14313 (A). Franklin Co.: Horseback Pond,

 

near Chesterville, Rossbach 6376 (NCU*). Kennebic Co.: .4

 

north shore of Great Pond, Rome, Rossbach 5403 (NCU).

 

Knox Co.: south shore of Newbert Pond, Appleton, Ross-

bach 1557 (NCU). Lincoln Co.: Murray Hill, Boothbay,

 

Gilbert s.n., 14 July 1897 (GH). Oxford Co.: Lovell,

Johnson 111 (NY*); Albany, Songo Pond, Wheeler 555703

 

 

(VDB*). Sagadahoc Co.: Five Islands, Rehder 1023 (A).

Somerset Co.: near Pittsfield, Knight 1840 (GH). border

 

of Lake Megunticook, Friesner 14056 (NY). York Co.:

 

0.75 mile north of Bauneg Beg Pond, True 5147 (GH).

 

Massachusetts. Barnstable Co.: near Orleans, Palmer

 

44807 (GH). Berkshire Co.: Mount Washington, E. 2' &

H. N. Moldenke 11651 (NY). Dukes Co.: Chilmark,

 

Martha's Vineyard, MacKeever MV-437 (NY). Essex Co.:

 

near Rockport, Cushman 3354 (TEX*). Franklin Co.: near

 

East Hawley, Livingston s.n., 4 August 1960 (NCU).

 

Hampden Co.: Pole Ridge Swamp, Wilbraham, Clark 2 Sey-

mour G759 (DUKE*). Hampshire Co.: 0.5 mile south of

 

Easthampton, Manning s.n., 24 June 1930 (VDB). Middle-
sex Co.: near Long Bridge, Newton, Gilbert s.n.,

19 June 1894 (GH). Nantucket Co.: Polpis, Nantucket

240

Island, MacKeever N532 (NY*). Norfolk Co.: Randolph

 

Ave., Blue Hills, Seymour s.n., 4 September 1908 (DUKE).
Plymouth Co.: Marion, ZEE£.§\E°I September 1888 (NY).
Suffolk Co.: Boston, 292E.§'E°' 1816 (US). Worchester
Co.: Pine Hill, Lancaster, BrownԤ.g., 20 August 1942

(TEX). New Hampshire. Carroll Co.: bog near Chocorua,

 

Lems s.n., 22 June 1960 (MSC). Cheshire Co.: near East

 

Jeffrey, B. J. Palmer 43033 (A). Grafton Co.: East

 

Hebron, Nelson, s.n., 10 July 1917 (NY*). Hillsboro Co.:

Peterboro, Batchelder s.n., 11 July 1913 (NY). Sullivan

 

Co.:' East Washington, Parker s.n., 23 August 1868 (NY).

New Jersey. Atlantic Co.: Newtonville, Bassett E Long

 

§.g., 12 August 1923 (GH*). Bergen Co.: Franklin Lakes,

Moldenke 10745 (NY). Burlington Co.: Wading River,

 

Long 12167 (GH). Camden Co.: Merchantville, Long 21373

 

(GH, NY*). Cape May Co.: 2 miles southeast of Goshen,

Lems s.n., June 1962 (MSC). Cumberland Co.: 2 miles

 

north of Dividing Creek, Adams 1136 (A*). Gloucester
Co.: Riders Bog, near Nialaga, Bassett s.n., 23 August
1923 (GH). Hunterdon Co.: near Peters Creek, 1 mile

southwest of Linvale, Benner 7379 (GH). Middlesex Co.:

 

about 2 miles northwest of Cranbury, Long 51877 (GH).

 

Monmouth Co.: near Locust, Williams 1 (GH*). Morris

 

Co.: Chatham Twp., Tanglewood Lane, Members, Summit
Nature Club (§.§.) 165 (NY). Ocean Co.: 2 miles east

of Cedar Grove, Fogg 4849 (GH*). Passaic Co.: near

 

 

I-—

241

Little Falls, Barnhart 1440 (NY*). Somerset Co.: near

 

Watchung, Moldenke 1312 (DUKE, NY). Sussex Co.: 1 to 2

 

miles north of Sparta, Adams 3 Wherry 1541 (A). Warren

 

Co.: Marble Hill, near Phillipsburg, Porter s.n., 11

June 1886 (NY). New York. Dutchess Co.: near Clove,

 

Standley g Bollman 11980 (US). Orange Co.: Black Rock

 

 

Forest, Raup 7805 (GH). Putnam Co.: Cold Spring, Barn-

 

hart 1005 (NY). Rensselaer Co.: Sandlake, House 5181

 

 

(NY). Richmond Co.: Bradley Ave., Staten Island, Dowell

3842 (GH). Rockland Co.: Clarkstown Twp., Lehr 370

 

(NY*). Suffolk Co.: Hauppauge Bog, Cold Spring Harbor,

Cain 484 (NY*). Tompkins Co.: South Hill, Ithaca,

 

Eames E Thomas 4770 (GH*). Ulster Co.: Rockland,

 

Camp_3049 (NY). Warren Co.: 2 miles south of Luzerne,

 

F2gg_19329 (NY*). Washington Co.: northwest of Tripoli,
Burnham s.n., 30 June 1896 (GA). Westchester Co.: Ken-
sico, Stabler s.n., 25 June 1886 (GH). North Carolina.

Alamance Co.: 18.7 miles west of Chapel Hill, Radford g

Rabb 714 (NCU). Alexander Co.: 3.25 miles northwest

 

of Vashti, Radford 13764 (NCU). Alleghany Co.: off Blue

 

Ridge Parkway, about 4.5 miles south of its junction

with NC. 89, Fox 2 Godfrey 2030 (FSU, GH, NY*). Anson

 

Co.: Burnsville, g,£.§. g.§., 20 July 1917 (NCU). Ashe

Co.: 4 miles northeast of Jefferson, Radford 6233 (NCU).

 

Avery Co.: Linville, Hunnewell 9277 (GH*). Bertie Co.:

 

southwest of Lewiston, Ahles 46137 (NCU*). Bladen Co.:

 

 

242

Rockfish Creek, Radford & Radford 2077 (NCU). Buncombe

 

Co.: near Biltmore, Biltmore Herbarium 24 (GH, NCU, NY).
Burke Co.: exposed north side of Table Rock, Stewart
1§44 (NCU, NY). Caldwell Co.: east of Blowing Rock,
Heller 1295 (GH, NY). Catawba Co.: 1.8 miles north of
Long Island, McNeely 1248 (NCU). Chatham Co.: 4 miles

north of Sanford, Radford 4 Radford 2001 (NCU). Cherokee

 

Co.: 1 mile southeast of Unaka, Radford 11221 (NCU).
Chowan Co.: 0.1 mile north of junction of NC. 32 & 37, ‘
Ahles 47846 (NCU). Clay Co.: just north of Brasstown,
Hardin 1411_(FSU, GH). Cleveland Co.: 3.5 miles south

of Boiling Springs, Fox 3945 (GH, NY). Columbus Co.:

 

3.8 miles southeast of Cherry Grove, 8311_1132§_(NCU).
Davidson Co.: near Yadkin River, near Yadkin College,
Totten s.n., 18 June 1931 (NCU). Davie Co.: 10 miles
northwest of Mocksville, Radford 14444 (NCU). Duplin

Co.: 5.5 miles north of Beulaville, 12135 34Q§4 (NCU).
Durham Co.: 2.7 miles north of Weaver, Bradley 2241 (NCU).
Edgecomb Co.: 0.3 mile northeast of Gethsemane, Radford

32057 (NCU). Franklin Co.: 3.2 miles west of New Hope,

 

Ahles 16562 (NCU). Gaston Co.: near summit of Kings

 

Mt., Costing 1gg1_(DUKE). Graham Co.: 10 miles west

of Robbinsville, Radford 14222 (NCU). Granville Co.:

0.7 mile west and 1.8 miles south of Culbreth, Ah1gs‘11211
(NCU*). Guilford Co.: in Pleasant Garden, Melvin s.n.,

2 June 1953 (NCU). Halifax Co.: 1 mile north of Enfield,

243

Fox 4 Godfrey 2747 (GH). Harnett Co.: just south of

 

Olivia, Wilbur 9134 (DUKE). Haywood Co.: near summit

 

of Cold Mt., Ramseur 515 (NY). Henderson Co.: about 5

 

miles northeast of Brevard, Wilbur 7107 (DUKE*). Hert-

 

ford Co.: Chowan River Swamp off the Loop Rd. of the
Bigwoods, Allen s.n., 16 July 1957 (DUKE*). Iredell

Co.: 4 miles east-northeast of Harmony, Ahles 45222

 

(NCU). Jackson Co.: Whiteside Mt., Hardin 1111 (GH).
Johnston Co.: 4.3 miles east of Smithfield, Radford 4
Stewart 1914 (NCU, TEX). Lee Co.: 0.5 mile west of
Patterson's Creek and 0.5 mile south of Deep River,

Kessler 294 (NCU). Lenoir Co.: 0.5 mile south of Deep

 

Run, Radford 22231 (NCU). Lincoln Co.: near Clark

 

Creek, north of Lincolnton, Bell 15283 (NCU). Macon

 

Co.: north of Highlands, Godfrey 51465 (FSU*); banks

 

of Cane Creek, Godfrey 60008 (FSU*). Madison Co.: 4

 

miles south of Marshall, Ahles 42405 (NCU*). McDowell

 

Co.: 6 miles southeast of Marion, Fox 4805 (NY*).

Mitchell Co.: Mount Mitchell, Schallert 5561 (DUKE).

 

Montgomery Co.: 5 miles south of Troy, Radford 14241.
(NCU). Moore Co.: Horseshoe Bend, Kessler £11 (NCU).
Nash Co.: 1.1 miles east of Middlesex, 42133.14431 (NCU).
Northampton Co.: 2.1 miles west-northwest of Garysburg,
Ah1gs_4§§gg (NCU). Orange Co.: Duke Forest, Costing
figgg (DUKE). Perquimans Co.: Hertford, Bartley g Pontius

473 (NY). Person Co.: by NC. 57, 0.3 mile southwest of

244

US. 501, Bell 12298 (NCU). Polk Co.: Lake Lanier, Tryon,

 

Smith §,g., 25 June 1933 (NCU). Randolph Co.: 8.2 miles
northwest of Coleridge, Bell 14248 (NCU). Richmond Co.:
along Drowning Creek, on US. 1 just south of Moore Co.

line, Fox 3671 (GH*). Robeson Co.: southwest of Tolars-

 

ville, Ahles 28964 (NCU). Rockingham Co.: 1 mile north

 

of Intelligence, Radford 13711 (NCU). Rowan Co.: 1.2

 

miles southeast of Gold Hill, Horton 1187 (NCU). Sampson

 

Co.: 1.1 miles west-southwest of Newton Grove, Ahles
29964 (NCU). Standley Co.: north of Big Lick, Radford 4

Haesloop 7051 (NCU). Stokes Co.: Hanging Rock State

 

Park, Fox 4 Godfrey 2004 (GH). Surry Co.: Pilot Mt.,

 

Fox 861 (GH*, NY). Swain Co.: about 3 miles northwest

of Bryson City, Bell 3277 (NCU). Transylvania Co.:

 

Toxaway Falls, Hardin 2127 (FSU). Tyrrell Co.: 2.8
miles southeast of Cross Landing, Radford §1§31_(NCU).
Union Co.: 2 miles west-southwest of Marshville, Ah1g§
21522_(NCU). Wake Co.: Lake Johnson, Godfrey 49421 (GH).

Warren Co.: Poplar Mt., Seaman 2927 (NCU). Watauga Co.:

 

Blowing Rock Mt., §m§11_4 Heller 114 (NY, US). Wilkes
Co.: about 1 mile west of Ball's Mill, Stewart 4.4.,

15 June 1939 (NCU). Wilson Co.: 3 miles south-southwest
of Black Creek, Radford 14924 (NCU). Yancey Co.: south-

east of Micaville, Ahles 46921 (NCU*). Ohio. Ross Co.:

 

Liberty Twp., near Jones' Crossing, Bartley 4 Pontius 21_

(NY). Pennsylvania. Adams Co.: 5.75 miles west-southwest

 

 

245

of Wenksville, Tanger 4381 (DUKE). Bucks Co.: 2.5 miles

 

northwest of Neshaminy, McDowell 395 (GH). Carbon Co.:

 

2.75 miles northeast of Albrightsville, Adams 3463 (A).

Centre Co.: 1 mile west of Hublersburg, Wahl 520 (GH*).

 

Chester Co.: Bradford Hills, E. B. Bartram 1262 (GH*).

 

Clinton Co.: 5.5 miles northwest of Beech Creek. Fogg

17832 (NY). Cumberland Co.: near Bridgeton, Byhouwer 4

 

Kobuski 103 (A). Delaware Co.: 1.5 miles east-southeast

 

of Newtown Square, Travis 41 (A). Fayette Co.: Ohi0py1e,

along the Youghiogheny River, Bright 16920 (TEX*); slopes

 

of the Youghiogheny River at falls, 1, 4, Churchill §.g.,

 

4 July 1935 (MSC). Juniata Co.: Runs Gap, Tuscarora Mt.,

3.5 miles south of Port Royal, Adams 5237 (A). Lackawanna

 

Co.: 4 miles south of Scranton, Glowenke s.n., 16 May

 

1936 (NY*). Lebanon Co.: South Mt., Penryn, Heller 615
(GH, NY*, US). Lehigh Co.: 5/8 mile south-southeast of

Crackersport crossroads, Pretz 10765 (VDB). Luzerne Co.:

 

along the Susquehanna River above Nescopec, Heller 14215
(A). Lycoming Co.: Trout Run, Greene s.n., 19 June 1898

(A). Northampton Co.: 1 mile north of Bangor, Schaeffer

 

19743 (DUKE). Northumberland Co.: 0.5 mile north of
Montandon, Manning s.n., 12 July 1964 (GH). Perry Co.:
Hominy Ridge, 1.5 miles north-northwest of Markelville,

J. W. §_§, 1, Adams 3013 (A). Philadelphia Co.: at

 

Tinicum, MacElwee 576 (GH*, NY). Pike Co.: near Tafton,

 

E. g. Palmer 43580 (NY). Wayne Co.: without definite

 

 

246

locality, Moyer §,g., 1867 (US). Westmorland Co.:

 

without definite locality, Jennings 1451 (GH). Rhode

 

Island. Newport Co.: Newport, Mearns 611 (NY). Provi-

 

dence Co.: near Harrisville, E. g. Palmer 44967 (A*).
Washington Co.: Wickford, Kennedy §,g,, 17 June 1908

(GH). South Carolina. Aiken Co.: 1.3 miles west of

 

junction of US. 1-76 and SC. 254, Ahles 55471 (NCU).

 

Anderson Co.: Beaverdam Creek, near Tugaloo River,

Ahles 13455 (NCU). Calhoun Co.: west-southwest of

 

Elloree, Ahles 30372 (NCU*). Chesterfield Co.: Little

 

Black Creek near SC. 265, Radford 12354 (NCU*). Clarendon
Co.: 3 miles southwest of Manning, Godfrey 4_Tyron 906a
(GH, NY). Darlington Co.: east side of Kilgore's Branch,

Smith 939 (NCU). Florence Co.: 1.8 miles west-southwest

 

of Hyman, B311'Z§§§_(NCU). Greenville Co.: above River
Falls, Rodgers 4 Mullens 61111 (NCU). Lee Co.: 1 mile
south of Lynchburg, Radford £4114 (NCU). Lexington Co.:
6 miles southeast of Lexington, Radford 122§§_(NCU).
Oconee Co.: 7 miles northwest of Jocassee, Radford 1111i
(NCU). Pickens Co.: 3 miles north of Rocky Bottom, near

US. 178, Radford 16729 (NCU*). Saluda Co.: 1.3 miles

 

west of Ward, Radford 23023 (NCU*). Spartanburg Co.:

 

below Rainbow Lake Dam, north of Spartanburg, Bell 8341
(NCU*). Williamsburg Co.: 5.5 miles north-northeast of

Mouzon, Radford 24812 (NCU). Tennessee. Blount Co.:

 

top of Gregory's Bald, Wilson 1783 (GH*). Campbell Co.:

 

247

near Pioneer, Sharp 35'31. 3828 (NCU). Carter Co.: near
Roan Mt., Rehder 5.3., 20 June 1900 (A). Coffee Co.:

just on the outskirts of Manchester, Godfrey 69807 (FSU*,

 

NCU). Cumberland Co.: just west of Crossville, Kral
40544 (VDB*). Fentress Co.: near Clarkrange, Sharp 4

Underwood 2884 (NCU). Green Co.: Greystone and mountains

 

beyond, Anderson 1068 (GH). Grundy Co.: just south of

 

Beersheba Springs, Sharp 23,31. 43580 (NY, VDB). Hamilton

Co.: Walden Ridge, above Soddy, Underwood 4 Sharp 2357

 

(NCU). Johnson Co.: Shady Valley, Sharp 4 Underwood 1542
(FSU, GH*, VDB). Loudon Co.: 0.75 mile north of Lenoir

City, Wilson 4274 (GH). Rhea Co.: near Rockwood, Coffman

 

§.p., 13 June 1899 (US). Scott Co.: without definite
locality, Braun §.p. (US). Sevier Co.: Greenbrier,

Pinnacle Mt., Sharp & Wilson 1302 (GH). Unicoi Co.:

 

about 3 miles southeast of Erwin, Zobel 5.2., 16 July
1965 (DUKE). Union Co.: Rhodelia unit, Norris Lake

Forest, Morrison 5.3., 15 June 1937 (FSU). Van Buren

 

Co.: near Piney Creek Falls, Shanks 22.31.‘45Q5 (NCU).
Vermont. Bennington Co.: Pownal, Eggleston 1424 (NY*).
Rutland Co.: Tadmire Hill, Pawlet, Dutton 1115 (A).

Windham Co.: near Westminster Co.: Blanchard 55 (NY*).

Virginia. Accomac Co.: Accomac, Gleason 8562 (NY*).

 

 

Albemarle Co.: at Albemarle-Nelson Co. line on Va. 6,

James 12136 (NCU). Allegheny Co.: edge of Potts on

 

Potts Mt., Harvill 14178 (NCU). Amelia Co.: without

 

248

definite locality, Lewis 5.5., 3 June 1936 (NCU). Amherst

 

Co.: on US. 60 along Huffs Creek, James 12230 (NCU).

 

Appomattox Co.: northeast of Appomattox, James 7321 (NCU).

 

Arlington Co.: Lyon Village, 0.5 mile north of Clarendon,
Hermann 10002 (GH, NY*). Augusta Co.: Magnolia Swamp,
Freer pp 31. 12110 (NCU). Bath Co.: 6.2 miles northeast

of Cow Pasture River on Va. 42, James 8074 (NCU). Bedford

 

Co.: SharptOp, Peaks of Otter, Eggleston 18645 (NY, US).

 

 

Botetourt Co.: 10 miles northwest of Roanoke, Totten 5.3.,
20 June 1936 (NCU). Brunswick Co.: 10.5 miles west-
northwest of Emporia, K£g1_15555 (FSU). Buckingham Co.:

1 mile northeast of county line on Co. Rd. 605, Ramsey 23
31. 1515 (NCU). Campbell Co.: southwest of Concord,
53m25‘1515 (NCU*). Charlotte Co.: 1.7 miles west of
Little Roanoke Creek on Va. 40, ggmgs_1511_(NCU*). Chesa-

peake: 5 miles west of Hwy. 168 on Hwy. 624, James 1769

 

(NCU*). Chesterfield Co.: just east of junction of Co.

Rd. 619 & US. 95, Jame§_5107 (NCU). Craig Co.: along

 

Craig Creek, just east on Newcastle, Eggg_15115_(GH).
Culpeper Co.: 1.6 miles south of junction of US. 211 &

Va. 229, 12mg§.1515 (NCU). Dinwiddie Co.: about 10 miles
west of Stony Creek, §£2l.£l§£1 (NCU). Essex Co.: 0.8
mile northwest of junction of Co. Rd. 617 & US. 17, 12225
5455 (NCU). Fairfax Co.: near Vienna, Sargent 1551 (FSU).
Fauquier Co.: 3 miles below Thorofare Gap, Pond Mt.,

Allard 11329 (GH*). Franklin Co.: 2.8 miles northwest

249

of junction of Co. Rd. 890 & Co. Rd. 646, James 10604
(NCU). Frederick Co.: 4.1 miles southwest of junction

of US. 50 & Co. Rd. 704, James 10895 (NCU). Giles Co.:

 

Mountain Lake Biological Station, Salt Pond Mt., Thorne

11221 (FSU*). Grayson Co.: Independence, Gleason 8758

 

(NY*). Greenville Co.: about 10 miles south of Emporia,

Kral 13756 (VDB). Halifax Co.: Childrey Creek on Co. Rd.

 

632, James 2547 (NCU). Hampton: Buckroe, Robinson 441

 

 

(GH). Henry Co.: 0.6 mile south of junction of Co. Rd.

682 & Co. Rd. 683, James 10448 (NCU). Highland Co.:

 

southeast of Monterey, 55mgs_1515_(NCU). Isle of Wight
Co.: near Franklin, Heller 5.5., 7-28 June 1893 (A).
James City Co.: 1.5 miles northwest of Williamsburg,
Grimes 5555 (NY). King William Co.: banks of Mattaponi

River near King William Landing, Fernald 5_Long 11601

 

(GH, NY). Lancaster Co.: about 0.2 mile north of junction

of Va. 3 & Va. 200, James 6039 (NCU). Lousia Co.: 0.3

 

mile northeast of junction of US. 33 & Co. Rd. 631, 55555
15555 (NCU). Lunenburg Co.: south of Kenbridge, 55255
5555 (NCU). Madison Co.: Shenandoah National Park,
Walker 1455 (US). Mecklenburg Co.: east of Boydton,
Gleason 5555 (NY). Middlesex Co.: 5 miles northwest of
junction of US. 17 & Co. Rd. 610, 55555.5555 (NCU).
Montgomery Co.: 1 mile northwest of McCoy, Cooperrider
5111 (NCU). Nansemond Co.: east of Cox Landing and south

of South Quay, Fernald 5 Moore 15132 (GH, NY*). New Kent

 

250

Co.: without definite locality, James 5387 (NCU). North-

 

ampton Co.: near Seaview, Hunnewell 16896 (GH). North-

 

umberland Co.: 0.3 mile south of junction of Co. Rd.

679 & Va. 200, James 5864 (NCU). Orange Co.: 1.8 miles

 

northeast of junction of Co. Rd. 690 & Co. Rd. 643, James
13660 (NCU). Page Co.: 1.1 mile northwest of junction

Of CO. Rd. 658 & CO. Rd. 674, James 11266 (NCU). Patrick

 

Co.: near Ararat, Harvill 16495 (NCU*). Pittsylvania

 

Co.: northeast of Kentuck, James 3192 (NCU). Prince
Edward Co.: 5 miles south of Farmville, Fosberg 15546

(GH). Pulaski Co.: northwest of Pulaski, James 8509

 

(NCU). Rappahannock Co.: near Meadow Spring shelter,

Walker 2302 (US). Roanoke Co.: about 4 miles northwest

 

of Dixie Caverns, Wood 2652 (GH*). Rockbridge Co.: near

 

Lexington, 5. 5. Churchill 5.5., 23 August 1924 (GH).

 

Rockingham Co.: above Hone Quarry Camp, Allard 2114 (US).
Scott Co.: 2.5 miles east of junction of US. 58 & Co. Rd.
701, James 9660 (NCU). Shenandoah Co.: near Orkney

Springs, Steele 105 (FSU*, TEX). Smyth Co.: White Rock

 

Mt., Britton g£_§1. 5.5., 22 June 1892 (NY). Spotsylvania
80.: 1.5 miles southwest of Fredericksburg, 5. 5.‘5

E.Ԥ. Randolph 211 (GH). Stafford Co.: near Stafford,

 

Wiegand 4_Manning 2390 (GH). Surry Co.: 3 miles west

of Bacons Castle, Kral 11010 (NCU*). Sussex Co.:

 

Coppahaunk Swamp, south of Waverly, Fernald 5_Long 10367

 

(GH). Tazewell Co.: near Mountain Gate, Harvill 12138

 

'x

251

(NCU). Virginia Beach: Virginia Beach Golf Grounds,
Meredith 5.5., 17 June 1923 (A). Warren Co.: at Page—

Warren Co. line on US. 340, James 11529 (NCU). West

 

Virginia. Cabell Co.: Rickett's Place, Gilbert 556

 

(FSU, DUKE*, GH, NY, US). Greenbrier Co.: White Sulphur

Springs, Hunnewell 6747 (GH). Hampshire Co.: Capon ;

 

Springs, Hunnewell 17984 (GH). Hardy Co.: east of Trout

 

River, near Sugar Knob Shelter, Allard 9393 (US). Mineral

 

 

Co.: at Sand Spring on Rt. 42 near Elk Garden, Brown s,p.,

 

September 1966 (NCU). Monon Co.: near Dellslow, Allamong

 

5.5., 5 July 1964 (NCU*). Pendleton Co.: Shenandoah Mt.,

near US. 33, Radford 45912 (NCU*). Pocohontas Co.: near

 

Durbin, Moore 2406 (A, GH). Raleigh Co.: 1.5 miles from

 

Beckley, Tosh 923 (US). Upshur Co.: Buckhannon, Mills-

 

paugh 411 (NY).

 

Some confusion has existed as to the identity of

the Linnaean name Andromeda paniculata. Aiton (1799),

 

Willdenow (1799) and Michaux (1803) understood the

description of Linnaeus to refer to Lyonia ligustrina.

 

Pursh (1814) used Andromeda paniculata to refer to 5.

 

ligustrina var. ligustrina; Nuttall's understanding of

 

the name was similar to that of Pursh, although he trans-
ferred it to Lyonia. G. Don (1834) followed Nuttall

in using Lyonia paniculata. Others such as K. Koch

 

(1872) and Asa Gray (1878) indicated that the Andromeda

paniculata of Linnaeus was a different plant. Elliott

 

252

(1817) used Andromeda ligustrina for Lyonia ligustrina

 

 

var. ligustrina but included Andromeda paniculata in his

 

 

flora with the following remarks:

This is the description of the original 4. paniculata
of Linnaeus; which I have inserted, althougH’con-
sidered a Virginian plant, for the purpose of point-
ing out the obscurity that still hangs over this
genus. Willdenow, in his celebrated edition of this
great work, has certainly, on the authority of the
Hort. Kewensis, substituted some other plant, I

should suppose the modern A. paniculata, if the leaves
of the latter were ever ovate. Yet the figure of Pluk-
enet and the Herbarium Linnaeus ought, where they are
accessible, to remove all doubt.

 

 

In the discussion under A. ligustrina he adds:

 

To the accurate and extensive researches of Dr.
Muhlenberg, American Botanists are indebted for

the real history of this plant. It appears to have
been the original Vaccinium ligustrinum of Linnaeus;
it certainly agrees with the-Linnaeandescription of
that plant as far as it extends. Modern botanists
have viewed it as the Andromeda paniculata of
Linnaeus, but whoever attends to the description

of that plant, in the early editions of the Species
Plantarum, will readily perceive that its great
author must have had a very different one in view.

 

 

 

The views expressed above by Elliott seem to be correct.

The Linnaean discription of Andromeda paniculata is quite

 

ambiguous in that some of the characters given refer to

5. ligpstrina while others seem to be describing a com-

 

pletely different p1ant--possib1y a species of Leucothoé.

 

The description seems to fit either Leucothoé racemosa

 

(L.) A. Gray or 5. recurva (Buckley ) A. Gray in the
following respects: (1) "racemes secundis," (2) ”cor-
ollis sub-cylindricis," (3) "foliis . . . crenulatis,"
(4) "antheris aristatis." Leucothoé racemosa has

racemose inflorescences from axillary buds on wood of

 

253

the preceding season, flowers with cylindric or ovoid
corollas, serrulate or crenulate leaves, and awned

anthers! In contrast, Lyonia ligustrina has paniculate

 

inflorescences, flowers with urceolate corollas, serrulate
to nearly entire leaves, and unawned anthers. Yet
Linnaeus' description does state that the racemes are
naked and panicled, characters which could be interpreted

as referring to 5. ligustrina. Koch (1872) lists Andromeda

 

 

paniculata L. as a synonym of Leucothoé racemosa, which

 

 

he calls 5yonia racemosa. There is a specimen in the

 

Linnaean Herbarium (as seen from microfiche) which is

labeled paniculata and has the letter "K" near the base

 

of the specimen indicating that it was collected by P.
Kalm. The sheet contains a fruiting specimen of 5.

ligustrina var. ligustrina and a flowering specimen of

 

 

Leucothoé racemosa! This possibly accounts for the con-

 

fusing description given for 5. paniculata in the Species

 

Plantarum.

 

The description given by Linnaeus for Vaccinium

ligustrinum is also ambiguous and reads as follows:

 

"racemis nudis, caule fruticosa, foliis crenulatis,
oblongis.” The habitat is given as Pennsylvania and
Kalm is indicated as the collector. A specimen was
located in the Linnaean Herbarium (studied on microfiche)
which is labeled "ligustrinum"; the letter "K" is also

present indicating that it was collected by Kalm. This

254

specimen is clearly 5. ligustrina var. ligustrina and may

 

 

be the type of the name Vaccinium ligustrinum. This view

 

is strengthened by the fact that Linnaeus cites no other

sources after his description of 1. ligustrinum, and lists

 

the collector as Kalm. Thus it appears that 5. ligustrina
is the prOper name for this species.3

Andromeda racemosa L. was misapplied by Lamarck

 

(1873). Linnaeus' name actually is the basis of Leu-

cothoé racemosa.

 

Michaux (1803) was the first person to recognize
infraspecific variation in 5. 11gustrina. He recognized
two primary varieties, each of them with two subvarieties:

the first, "var. 1. nudiflora: racemes nudis.” and the

 

second, "var. 2. foliosiflora: racemes foliosis.” He

 

gives the habitat of var. nudiflora as 'frigidiorbus,

 

per Etats-Unis." and of Var. foliosiflora as "in sylvis

 

Carolinae inferioris" and "in stagnosis." Michaux is
correct in giving the naked inflorescences as the most
distinguishing character of var. ligustrina and in
recognizing the distinctness of the leafy-bracted plants

of the Coastal Plain.

 

3Evidently this information was unavailable to
K. Koch (1872) for he states: "Vaccinium ligustrinum
L. sp. p1. I, 351 ist eine Pflanze, die ohen ein gutes
Original zur hand zu haben, durch die Beschreibung kaum
noch entziffert werden kann, aber gewiss keine Andromeda.
Man kann ohnmaglich von Linné ebenso von Michaux, denken,
dass sie den unteren Fruchtknoten ihrer Vaccinium ligus-
trinum nicht bemerkt haben sollten.”

l ‘L

I“.

255

The true inflorescence of 5. ligustrina is an

 

axillary panicle or raceme of corymbose fascicles aggre-
gated near the end of the growth of the preceding year,
often forming a prominent pseudoterminal inflorescence.
The number of axillary panicles in the terminal cluster
varies from as high as 16 to as low as one in var. 11355:

trina; in var. foliosiflora there may be as many as 251

 

This difference appears to be mainly environmental. A
single plant can have a widely varying number of panicles
clustered at the ends of its branches.

Nuttall (1818) differentiated this variety from

the more southern var. foliosiflora, giving as its major

 

characters: "leaves obovate-lanceolate, nearly entire,
somewhat acuminated at both extremities, membranaceous;
panicle terminal, nearly naked . . . "

The table on the following page summarizes the
most consistent differences between the two varieties.

Populations in the southern Appalachians often
show a tendency toward obovate or oblanceolate leaves
with acuminate tips. The mountain pOpulations also are
often more pubescent. Plants with scattered large bracts
in the inflorescence occur in the mountains of western
North Carolina, the extreme northwest portion of South
Carolina and northeast Georgia.

Although the inflorescence of this variety is

referred to as naked, usually a few small bracts are

 

256

present. Thus plants of this variety have often been

misidentified as var. foliosiflora, even when collected

 

from as far north as Maine!

TABLE 24. Comparison of 5yonia ligustrina vars. ligustrina
and foliosiflora.

 

 

 

 

 

 

 

var. ligustrina var. foliosiflora
1. Panicles naked or with l. Panicles conspicuously
only a few bracts leafy bracted

2. Leaves usually membranous 2. Leaves often coriaceous,
especially late in

season
3. Leaves usually dull 3. Leaves often lustrous
4. Margins of leaves not 4. Margins often revolute
revolute or only slightly
so

5. Twigs usually not reddish 5. Twigs often becoming
reddish late in the
season

 

Lyonia ligustrina var. ligustrina occurs in dry

 

 

acid woods and thickets, hillsides, grassy balds, and acid
swamps and bogs. Mohr (1901) lists 5. ligustrina as part
of the mesophile plant association along spring-fed moist

brooks in the mountains, along with Kalmia latifolia,

 

Ilex opaca, Amelanchier canadensis, Aronia arbutifolia

and Hydrangea arborescens. Harshberger (1911) indicates

 

that it occurs in New England bogs with Clethra alnifolia,

 

Rhododendron viscosum, Vaccinium corymbosum, Ilex sp.,

 

 

yyrica pensylvanica, M. gale, Cassandra calyculata and

 

257

Chamaecyparis thyoides. It is also common in the pine

 

barrens of New Jersey where it occurs in open pine
forests with an undergrowth of oaks and other ericaceous
species (Harshberger, 1911; and personal observations).
It also occurs in the deciduous and cedar swamp communi-

ties, growing as a shrub with Alnus serrulata, Rhododen-

 

dron viscosum, Cephalanthus occidentalis, Cassandra caly-

 

culata, Clethra alnifolia, Ilex glabra, Leucothoé racemosa,

 

Toxicodendron vernix, Lyonia mariana, Myrica pennsylvanica,

 

Aronia arbutifolia, Vaccinium atrococcus and V. corymbosum

 

 

 

(Harshberger, 1961).

I found 5. ligustrina var. ligustrina to be

 

 

exceedingly common in the Pocomoke River swamp in WOr-
chester Co., Md. This swamp resembles the Dismal Swamp

and is usually regarded as the most northern of the exten-
sive southern Cypress swamps characteristic of the Atlantic

Coastal Plain. The major trees of the swamp are Taxodium

 

distichum and Nyssa biflora. Acer rubrum, Nyssa sylvatica

 

Liquidambar spyraciflua and Fraxinus pennsylvanica are

 

also important. Ericaceous shrubs are plentiful; other
characteristic shrubs include Viburnum spp., Cephalanthus
occidentalis, Aronia arbutifolia, Ilex verticillata,

Euonymous americanum and Itea virginica (Shreve, 1910:

 

 

and personal observations).

Lyonia ligustrina var. ligustrina is not usually

 

found along rapidly flowing, rocky mountain streams where

 

258

both Rhododendron maximum and Leucothoé axillaris var.

 

 

editorum abound. Rather it occurs in quieter marshy

 

areas along with Euonymous, Viburnum, Aronia and a few

 

 

other ericaceous shrubs. Shreve (1910) and Braun (1937)
also report it from similar habitats.

A specimen collected by Dr. H. T. Darlington

 

(7 Aug. 1916, MSC) near Hess Lake in Newaygo Co., Mich.

was identified by him as 5yonia ligustrina and was cited

 

bY Billington (1949). This specimen is actually Nyssa ;

 

 

sylvatica. No authenticated occurrences of 5. ligustrina

for Michigan are known.

4b. Lyonia ligustrina (L.) DC. var.foliosiflora (Michx.)

 

 

Fern.

Lyonia ligustrina (L.) DC. var. foliosiflora

 

 

(Michx.) Fern. Rhodora 10: 53. 1908. Andromeda paniculata

 

L. var. foliosiflora Michx., Fl. Bor. Am. 1: 255. 1803.

 

Andromeda frondosa Pursh, Fl. Am. Sept. 295. 1814. Lyonia

 

frondosa (Pursh) Nutt., Gen. N. Am. P1. 1: 267. 1818.

 

Andromeda ligustrina var. frondosa (Pursh) Wood, Class

 

 

Book 488. 1861. Andromeda ligustrina var. pubescens A.

 

 

Gray, Syn. Fl. N. Am. 2: 33. 1878. Andromeda paniculata

 

var. pubescens (A. Gray) Dippel, Handb. Laubholzk. 1:

 

370. 1889. Xolisma ligustrina (L.) Britt. var. foliosiflora

 

 

(Michx.) C. Mohr, Bull. Torrey Bot. Club 24: 24. 1897.

Xolisma foliosiflora (Michx.) Small, F1. Se. U.S. 889.

 

1903. Andromeda ligpstrina (L.) Muhl. var. foliosiflora

 

 

259

(Michx.) Fern. 1p Robins. & Fern., Gray's Man. Ed. 7 635.

 

1908. Andromeda ligustrina var. foliosiflora (Michx.)

 

 

Schneider, Ill. Handb. Laubholzk. 2: 533. 1911. Xolisma

ligustrina var. pubescens (A. Gray) Millsp., Liv. F1.

 

 

W. Virginia 324. 1913. Arsenococcus frondosus (Pursh)

 

Small, Shrubs Fla. 97. 1913. Lyonia ligustrina var.

 

pubescens (A. Gray) Bean, Trees Shrubs Brit. Isles 2:

 

64. 1914. (Lyonia ligustrina pubescens Rehder in Bailey,

 

Cycl. Am. Hort. l: 62. 1900, without citation of basionym
or (as a trinomial) designation of rank.) Type: ”in
sylvis Carolinae inferioris" and ”in stagnosis” Michaux
(P?, not seen).

Andromeda tomentosa Dum. Cours., Bot. Cult. ed. 2,

 

3: 495. 1811. Andromeda paniculata var. tomentosa (Dum.
Cours.) Dippel, Handb. Laubholzk. l: 369. 1889, Type:
(unknown).

5yonia salicifolia P. W. Watson, Dendr. Brit. 1:

 

t. 38. 1825. 5yonia ligustrina var. salicifolia (Wats.)

 

 

DC., Prodr. 7: 600. 1839. Type: ”Place . . . Arboretum,
Kew. Country . . . North America." (not seen).

Lyonia capreaefolia P. W. Watson, Dendr. Brit. 2:

 

t. 127. 1825. Lyonia ligustrina var. capreaefolia (Wats.)

 

DC., Prodr. 7: 600. 1839. Type: “Place . . . Mr. James
Lee's, Hammersmith. Country . . . North America." (not

seen).

260

 

Lyonia multiflora P. W. watson, Dendr. Brit. 2:
t. 128. 1825. Type: "Place . . . Mr. James Lee's,
Hammersmith. Country . . . North America." (not seen).

5yonia ligustrina (L.) DC. f. nanella Fern.

 

Rhodora 163: 158. 1947. Type: VIRGINIA: Mansemond Co.,
sphagnous and peaty bog by Norfolk and Western Railway,
about 1/2 mile west of Kilby. Sept. 8 and 12, 1946.
Fernald, 555g_5_C1ement 15,331 (GH, holotypelz US,

isotype!)

Leaves elliptic to narrowly so, ovate, obovate
or oblanceolate, 2.0-9.5 cm long, 1.0-4.5 cm wide,
deciduous to partly evergreen, often moderately cori—
aceous, serrulate to nearly entire, often lustrous above,
with revolute margins (at least along the lower portion
of the leaf), adaxial surface sparsely to densely pilose-
glandular, abaxial surface sparsely to extremely pilose-
glandular, glabrous to densely nonglandular pubescent on
the veins. Flowers in axillary or pseudoterminal
inflorescences which are small panicles aggregated near
the ends of the growth of the preceding year, and are
conspicuously leafy bracted or with at least the lower
panicles with large bracts. Inflorescence often very
large and lax toward the southern portion of the range.
Calyx with abaxial surface pilose-glandular, rarely also

nonglandular pubescent. Capsule globose to subglobose,

 

261

sparsely to densely pilose-glandular, sometimes also
nonglandular pubescent (Figure 45).

Distribution: mainly Coastal Plain, southeast
Virginia to Florida, west to Arkansas and east Texas
(Figure 46).

Flowering: late April - early July.

Representative specimens: Alabama. Barbour Co.:
near Pig Creek, west of Clio, 555145 Demaree 55555 (FSU,
NCU, VDB*). Bibb Co.: without definite locality, Laessle
5.5., 13 May 1937 (GA). Chilton Co.: 6.6 miles north-
east of Clanton, 51555‘11551 (NCU*). Choctaw Co.: 3
miles northwest of Butler, 5551’19755B (VDB). Clark Co.:
near US. 43, 6 miles south of Thomasville School, 5551
41151 (VDB*). Cleburne Co.: top of Cheaha Mt., 51355
4515 (NCU). Covington Co.: 17 miles south-southeast of
Andalusia, Duncan 15515 (NCU). Cullman Co.: St. Bernard,

Wolf 5.5., 29 May 1922 (NCU). Geneva Co.: 3.5 miles

 

west of Samson, Kral 35072 (VDB). Henry Co.: 8 miles

 

north of Headland, Wiegand 5_Manning 2396 (GH). Jefferson

 

Co.: 10 miles north of Alabaster, Kral 24358 (VDB*).
Lee Co.: Auburn, Earle 5 Underwood s.n., 10 May 1896

(NY*). Mobile Co.: near Mobile, Mohr 5.5., 13 May

 

1893 (A). Perry Co.: along Ala. 183 at Dabine Creek,
Clark 17958 (NCU). Talladege Co.: along Salt Creek,
T165, R8E, near boundary of sections 34 8 35, Clark 2844

(NCU). Tallapoosa Co.: about 6.1 miles northeast of

 

“' ”-7 262

 

 

Figure 45. Representative specimen of Lyonia ligus-

 

trina var. foliosiflora (Nash 528, MSC).

 

 

   

263

   
  

w W (lam.
—' var. toll-una-
M. 1.!- 1961

II.
on.a-oa—.A5.o~pepsoaoucaaleO

PLANTS OF CENTRAL PENINSULAR FLOROOA
COLL‘CVIO m vucmuvv 0! (new unyo muv
I

fit OIU V IA.- ADD-l IC‘” l.‘ . .. / 14.1

528. xm “OHM“!!! ll “lulu”.

Inn-n We." ...“.

264

 

Figure 46. Distribution of 5yonia ligustrina var.

foliosiflora.

 

 

265

   

266

Alexander City, Clark 15220 (NCU*). Winston Co.: Camp

McDowell, along Clear Creek, Clark 12969 (NCU*). Arkansas.

 

 

Clark Co.: Arkadelphia, Demaree 22085 (NY). Cleburne Co.:
Tumbling Shoals, Demaree 65652 (VDB). Columbia Co.:
near Waldo, Demaree 39230 (GH). Conway Co.: Petit Jean

Mt., Morrilton, Demaree 36834 (FSU, GH*). Drew Co.: near

 

Monticello, Demaree 13514 (NY*). Faulkner Co.: along

 

Cove Creek, near Martinville, 5. 5. Palmer 26521 (A).
Garland Co.: Cedar Mt., Hot Springs National Park,

Demaree 36827 (GH*, NCU). Giant Co.: near Poyen,

 

Demaree 11551 (FSU, GH). Hempstead Co.: near Columbus,
5. 5. Palmer 15551 (A*). Hot Springs Co.: near Bismark,
Demaree 11415 (GH*, NY); near Malvern, 5. 5. Palmer 55555’
(A*). Howard Co.: Baker Springs, Kellogg 5.5., 6 October
1909 (A). Independence Co.: Sections 27 & 28, Tl4N,

R8W, Thomas 5555 (NCU*). Jefferson Co.: near Pine Bluff,

Demaree 14956 (NY). Johnson Co.: banks of Piney Creek,

 

near Knoxville, Demaree 15554_(NY). Montgomery Co.:

near Caddo Gap, Demaree 5.5., 10 October 1960 (GH*);
Collier Springs, near Norman, Demaree 55455 (FSU*).
Ouachita Co.: near Stephens, Demaree 15551 (NY). Perry
Co.: along Bear Creek, near Hollis, Demaree 55551 (FSU,
GH, NCU). Pike Co.: near Langley, Demaree 9517 (GH, NY).
Polk Co.: near Mena, 5. 5. Palmer 15541_(A). Pope Co.:
Crow Mt., Galla Creek woods, Tucker 1515 (NCU). Pulaski

Co.: along Rock Creek, Boyle Park, near Little Rock,

 

267

Merrill 1204 (A*). Saline Co.: 2 miles east of Benton,

Demaree 35602 (NCU, VDB). Stone Co.: Ozark National

 

Forest, near Fiftysix, Demaree 58226 (NCU, VDB). Van
Buren Co.: bluffs of Little Red River, near Shirley,
5. 5. Palmer 24309 (A). White Co.: along East Cadron

Creek, about 0.5 mile west of Rosebud, Redfearn 19239

 

(NCU). Yell Co.: near Rover, Demaree 53079 (NCU, VDB).
Florida. Alachua Co.: near Devil's Millhopper along
Fla. 23, Wiggins 19754 (NCU). Baker Co.: east of Taylor,

Ashe 5.5., 11 May 1929 (NCU). Brevard Co.: Okeechobee

 

region, Fredholm 5802 (GH*, US). Clay Co.: 2 miles north

 

of Penney Farms, Totten 5.5., 21 October 1945 (NCU*).
Columbia Co.: 20 miles north of Lake City, 5555_5_Arnold
5.5., 14 May 1948 (GA). Duval Co.: near Jacksonville,
Curtiss 1154 (GH, MISSA, NY). Jefferson Co.: near Lloyd,
Godfrey 51455 (FSU*). Hamilton Co.: 6 miles east of
Jasper, Godfrey 55511 (FSU, GH*, VDB). Highlands Co.:

Sebring, Harbison 11 (and 4610) (A*, NCU). Lake Co.:

 

vicinity of Eustis, 5555'555_(A, MSC, US). Leon Co.:

15 miles west of Tallahassee, Godfrey 55415 (FSU*).
Osceola Co.: Old Campbell Station Rd., Singeltagy 5.5.,
29 May 1935 (DUKE). Polk Co.: 7 miles southwest of

Bartow, Williams 5.5., 21 April 1957 (FSU, GH). Putnam

 

Co.: Welaka, University of Florida Conservation Reserve,

Lems 5.5., 17, 18 July 1959 (MSC). Saint Lucie Co.:

 

 

268

8 December 1919 (US). Seminole Co.: north of Goldenrod,
Cooley 55551. 7404 (NCU*). Taylor Co.: 1.3 miles north-

east of Shady Grove, McDaniel 5 Godfrey 4343 (FSU, NCU*).

 

Volusia Co.: about 3 miles east of Deland, Prichard 964
(FSU*). Walton Co.: near Darlington, Godfrey 58608 (FSU,
GH). Georgia. Bartow Co.: about 4.5 miles southeast

of Adairsville, near Mildred Pond, Greear 63104 (GA).

 

Ben Hill Co.: Faircloth 5545 (M0). Berrien Co.: With-
1acoochee River Swamp on Ga. 37 west of Ray City, Bozeman
4145 (NCU). Brantley Co.: about 3 miles north of Nahunta,
5555'5551 (A*, NCU). Brooks Co.: 6.3 miles north-north-
west of Quitman, Faircloth 1515 (NCU). Bullock Co.:
without definite locality, Harper 555 (GH, US). Calhoun
Co.: along Ichawaynochaway Creek, 2 miles east of Morgan,
Thorne 5545 (GH). Camden Co.: without definite locality,

Schallert 5,5., 4 May 1941 (NY). Candler Co.: on Ga. 129,

 

0.4 mile north of Canoochee River, Bozeman 4551 (NCU).
Charlton Co.: west of Chessar's Island, Okefenokee Swamp,
Duncan 5551 (NY). Chatham Co.: Savannah, Nuttall 5.5.
(GH). Coffee Co.: on Rd. to Flat Tub Landing, 0.9 mile
north of Ga. 107, Bozeman 4555 (NCU). Colquitt Co.:

swamp of Ochlocknee Creek, near Moultrie, Harper 1515

(A, GH, NY, US). Cook Co.: 7.6 miles north-northeast

of Adel, Faircloth 4889 (NCU). Decatur Co.: Butlers

 

Creek,‘3 miles west of Faceville, Thorne 4792 (NY).

Early Co.: west of Hilton, Thorne 5350 (US). Echols

 

269

Co.: 6 miles south of Stockton, Faircloth 650 (NCU).

 

Effingham Co.: 3 miles south of Stillwell, Bozeman 4075

 

(NCU). Emanuel Co.: on Ga. 46, 0.6 mile west of Emanuel-

Candler Co. line, Bozeman 4277 (NCU). Fulton Co.: Fulton

 

Co. Botanical Garden near Adamsville, Venard 5 McDowell

 

V-377 (GA). Grady Co.: 9 miles southeast of Cairon on

east side of Ochlockonee River, Kral 15200 (VDB*). Harris

 

Co.: above and south of Callaway Gardens entrance on
Pine Mt., Jones 20781 (GA, VDB*). Irwin Co.: about 5
miles south of Fitzgerald, Wilbur 3075 (GA). Lanier Co.:

2.6 miles southeast of Lakeland, Faircloth 2581 (NCU).

 

Lowndes Co.: 10 miles east-southeast of Valdosta, 5515:
51555 4151 (NCU). McDuffie Co.: vicinity of Thomson,
Bartlett 1692 (NCU, VDB). McIntosh Co.: 1.3 miles north-
east of Fort Barrington, Bozeman 5554 (NCU). Mitchell Co.:

3.9 miles east of Pelham, Faircloth 1345 (M0, NCU*).

 

Montgomery Co.: on Co. Rd. S-1125, 0.4 mile east of
Oconee River, Bozeman 5141_(NCU). Rabun Co.: at Tal-
lulah Falls, 55511‘5,5., 15 August 1893 (GH). Randolph
Co.: southeast of Cuthbert, Harper 1145 (GH, US). Rich-
mond Co.: near Augusta, Cuthbert 1151 (GH*). Screven
Co.: near Cameron, 55555 51555 (GA). Tatnall Co.: 3.9
miles northwest of Reidsville, 55155 54155 (NCU*).
Telfair Co.: about 3 miles northwest of Lumber City,
Wilbur 5155 (GA). Thomas Co.: 12 miles north of

Thomasville, Godfrey 67435 (FSU*, NCU). Turner Co.:

270

4.1 miles east-northeast of Ashburn, Faircloth 4389 (NCU).

 

Upson Co.: Pine Mountain, Higgins 5.5., 20 May 1928 (GA).
Ware Co.: Suwannee Lake, Harper 76a (GH). Wayne Co.:
4.1 miles south of Jessup, Kral 24168 (VDB). Wilcox Co.:

12 miles northwest of Abbeville, Wilbur 5 Webster 2710

 

(NY*, US). Louisiana. Jackson Co.: 5 miles southwest

 

of Chatham, 5515,55'51. 1515 (NCU). Lincoln Co.: 2 miles
Ouachita Co.: Sec. 9, T17N, R2E at Tiger Branch off La.
838, Thomas 11551 (VDB). Union Co.: about 3 miles north-
northwest of Spencer, Thomas 14155 (VDB). Vernon Co.:
along Yellow Branch, about 2.5 miles north of Burr Ferry,
Thieret 51555 (DUKE*, FSU). washington Co.: 3 miles
northeast of Franklinton, 5.5. & 5.5. Correll 5155 (DUKE,

GH*). Webster Co.: 4 miles west of Minden, 5.5. 5’5.5.
Correll 15515 (DUKE*, GH, NY). Winn Co.: 4 miles west
‘of Winnfield, 5.5. 5_5.5. Correll 15555 (DUKE, GH*, NY).
Mississippi. Covington Co.: junction of Hwy. 37 & 84,
south of Hot Coffee, 55555 1554 (GA). Forrest Co.:
junction of Hwy. 11 & 49, Hattiesburg, 55555 5555 (GA).
George Co.: along Hwy. 98, about 0.5 mile west of
.Alabama—Mississippi state line, Lelong 5555 (NCU*).
Lamar Co.: about 12 miles west of Purvis, Rogers 1079-B

(NCU). Lee Co.: Natchez Trace Parkway, McDougall 1834

 

(US). Perry Co.: 9 miles north of Beaumont, Webster 5

Wilbur 3423 (GH, NY, US). Scott Co.: Roosevelt State

271

Park, 3 miles southwest of Morton, Ray 7286 (FSU, GH,
MISSA*, NCU, VDB). Stone Co.: 3 miles south of Wiggins,

McDaniel 3110 (NY). Tishomingo Co.: 6 miles northeast

 

of Iuka, Ray 7522 (GH, MISSA*, NCU). Wayne Co.: 6 miles

 

southeast of Shubuta, ng 7004 (FSU, GH, MISSA*, NCU).

North Carolina. Beaufort Co.: 0.5 mile north of Wilmar,

 

555_4554 (NY*). Bladen Co.: south of Garland, Blomquist
5.5., 9 May 1951 (DUKE*). Brunswick Co.: about 7 miles
north of Southport, Wilbur 5115 (DUKE*, TEX). Camden Co.:
1.3 miles southwest of South Mills, 5555.5.5., 23 May

1959 (MSC). Carteret Co.: Harker's Island, Smith s.n.,

 

10 May 1942 (NCU). Columbus Co.: near Old Rock, Godfrey
5.55555 4188 (GH*, US). Craven Co.: New Bern, Harbison
53,(A). Cumberland Co.: 2.3 miles south-southeast of ‘
Roslin and 0.5 mile east, 55155 55455 (NCU). Currituck

Co.: 1.15 miles west of Aydlett, Lems 5.5., 24 May 1959

 

(MSC). Dare Co.: on US. 264 near Manns Harbor, Radford
.5155 (NCU). Duplin Co.: 1.7 miles south-southeast of
Rose Hill, 55155 55555 (NCU). Gates Co.: 4.6 miles west
of Mintonsville, 55155 45555 (NCU). Greene Co.: 0.6
mules south of Walstonburg, Radford 55515 (NCU). Barnett
Co.: 1 mile southeast of Pineview, Wilbur 5115 (DUKE).
IHoke Co.: about 4 miles southwest of Montrose on Mountain
(:reek, 55155 55551 (NCU). Hyde Co.: 1.7 miles northeast
(of Rose Bay, Radford 55515 (NCU*). Johnston Co.: near

lBemtonsville Rd., 1.5 miles east of US. 301, Fox 4929

 

272

(GH, NY*). Jones Co.: 4.3 miles west-southwest of

Hargetts Store, Radford 36963 (NCU*). Lenoir Co.: 0.5

 

mile south of Deep Run, Radford 55555 (NCU*). Martin Co.:
4 miles south-southeast of Smithwick, Radford 55454 (NCU).
Montgomery Co.: 5 miles east of Pekin, Radford 14554

(NCU). Moore Co.: 1.5 miles from Pinehurst, Wicker 5.5.,
17 July 1942 (NCU). Onslow Co.: 3.7 miles west-southwest

of Haw, Ahles 28174 (NCU). Orange Co.: 12 miles north-

 

west of Chapel Hill, Wiegand 5 Manning 2393 (GH). Pamlico

 

Co.: 0.4 mile west of Olympia, Radford 55555 (NCU). Pas-
quotank Co.: bank beside Pasquotank River, just south

of South Mills, Oosting 5455 (DUKE). Pender Co.: north-
east of Costin, 55155 55115 (NCU). Perquimans Co.:

swamp along Mill Creek 0.25 mile north of Perquimans

River, 5. 5.,5 5. 5. Randolph 637 (GH). Pitt Co.: 5

 

miles south of Greenville, 55155 1145 (NCU). Richmond
Co.: 3.75 miles northwest of Hoffman, Radford 11554
(NCU). Sampson Co.: about 10 miles south of Newton
Grove, Wilbur 5445_(FSU*, DUKE). Scotland Co.: 9 miles
south of Aberdeen, Godfrey 5145_(GH*). Tyrrell Co.:

2 miles south of Columbia, Godfrey 5 5555 5551 (GH).
Washington Co.: near Roper, Godfrey 4555 (GH, US).
Wayne Co.: about 5 miles north of Seven Springs, 555
55‘51. 4545 (GH). Wilson Co.: 3.2 miles southeast of
Lucama, Radford 55155 (NCU). Oklahoma. LeFlore Co.:

near Page, Blakley 1427 (GH*, US). Logan Co.: Jacks

273

Creek camp, Strictland, Demaree 15753 (NY). McCurtain
Co.: along Eagle Fork Creek, about 1.5 miles southwest
of Smithville, Redfearn 19806 (NCU*); 3 miles north of

Tom, Waterfall 16052 (NCU*). Pushmataha Co.: along

 

Little River, Honobia, Waterfall 13023 (GH, US). South

Carolina. Bamberg Co.: northwest of Govan, Ahles 26035

 

(NCU). Beaufort Co.: Hilton Head Island, Butler 5.5.,
16 May 1968 (NCU*). Berkeley Co.: 8 miles southwest

of Monocks Corners, Godfrey 5_5y555_1455 (GH). Charles-
ton Co.: Northampton Rd., 0.25 mile north of Halfway
Creek Rd., EEEE.Z§§Z (GH*). Clarenden Co.: 3 miles
southwest of Manning, Godfrey 5’5y555 555 (DUKE*, US);
north of Santee River, east of US. 15, Leonard 1551.
(FSU, GH, NCU} NY, TEX*). Colleton Co.: 2.4 miles

northeast of US. 17 bridge over Combahee River, Bell 2321

 

(NCU). Darlington Co.: Hartsville, 55155 5.5., 10 June
1942 (NCU). Dillon Co.: 0.8 mile north-northeast of Oak
Grove, 55155 55555 (NCU). Dorchester Co.: 5.9 miles
southeast of St. George, 55155 55551_(NCU*). Florence
Co.: 3 miles west of Cowards, 5511 15554 (NCU*). George-
town Co.: 3 miles southeast of Rhems, Radford 51555_
(NCU). Hampton Co.: 1.8 miles north-northeast of Early

Branch, Ahles 57131 (NCU); 1.8 miles north-northeast of

 

Early Branch, Bell 2546 (NCU). Horry Co.: Myrtle Beach,

Coker 5.5., 12 July 1945 (NCU). Jasper Co.: Ridgeland,

 

Ahles 12306 (NCU*). Kershaw Co.: along Lynces River

 

274

south of McBee, 5. 5. Palmer 45555 (A). Laurens Co.:
{south side of Enoree River, north-northeast of Tylers-
ville, Duncan 15155 (GA). Lee Co.: 0.2 mile north of
Lucknow, Radford 51551 (NCU). Marlboro Co.: 3 miles
south of Blenheim, Radford 15555 (NCU). Sumter Co.:
7.5 miles north-northeast of Pinewood, Radford 24021
(NCU). Williamsburg Co.: 2 miles west of Salters,
Godfrey 5 1y£55 555 (GH). Tennessee. Franklin Co.:

near Tullahoma, Alexander 55'51. 5.5., 24 September 1933

 

(NY*). Madison Co.: marshes, Jackson, 5515 455 (NY).
55555. Bowie Co.: near Texarkana, 5. 5. 5_5. 5. Heller
4115 (GH, NY). Jasper Co.: 23 miles northwest of Jasper,
555y 55551 (A*). Newton Co.: east of Newton, Lundell
11551 (TEX). San Augustine Co.: San Augustine, 5. 5.
Palmer 15545 (A). Shelby Co.: 9.5 miles southwest of

Center, McVaugh 8425 (GH, TEX*). Tyler Co.: 6 miles

 

east of WOodville, 5551_55555 (ENCB, VDB*). Vigginia.
Chesapeake: near entrance to Portsmouth Ditch, Lake
Drummond, Great Dismal Swamp, west of Wallaceton, Fernald
5.555g_15414_(GH). Nansemond Co.: east of Cox Landing,
south of South Quay, Fernald 5 555g 15155_(DUKE*, GH).
Southampton Co.: about 7 miles south of Franklin,
Fernald 5_5g5g_15555_(GH*). Suffolk Co.: Dismal Swamp,
Trudell 5.5., 27 May 1920 (GH). Virginia Beach: north

of Blackwater River, Fernald 5_Long 4118 (A*).

 

275

Because of the great variability in the southern

populations of 5. ligustrina, many varieties have been

 

recognized. These are actually nothing more than extremes
in the total pattern of variation. Forms with elongate,

membranous, lustrous, and usually glabrous leaves with

 

acuminate tips and large persistently leafy-bracted
inflorescences with loose panicles have been called var.

salicifolia (Fernald, 1941; Watson, 1825). Plants showing

 

these characteristics are common in the Dismal Swamp and
in adjacent regions of North Carolina. Individuals show-
ing most of the above characters can be found on the
Coastal Plain as far south as Florida. There, most of

the plants have large lax, extremely foliaceous inflor-
escences; moderately coriaceous elliptic, or obovate to
ovate, lustrous, almost glabrous leaves and reddish twigs.
Such plants can be found on the Coastal Plain from the
Carolinas south to Florida and west to Louisiana. Fernald

(1941) limits his variety foliosiflora to just this form.

 

Probably Watson's 5. multiflora (1825) was also from this

 

group. In southern Florida plants with extremely ovate
and coriaceous leaves occur. Plants often referred to

as var. ppbescens occur in the Carolinas and Georgia,

 

and are especially distinct in South Carolina. These
individuals are small, rigidly ascending shrubs with
coriaceous, moderately to densely pilose, obovate to

elliptic leaves with acuminate, mucronate, or acute tips.

276

The panicles usually have fewer floral bracts than the
forms described above. But some pubescent plants have

a completely different appearance! Scattered individuals
occur in South Carolina and Georgia that have very cori-
aceous, revolute elliptic, densely pilose and tomentose
leaves, with prominent reticulate veins on the lower

surface. Such plants were named var. pubescens by A.

 

Gray (1878). A plant of this type was probably used by

Nuttall for his description of Lyonia frondosa in his

 

Genera of North American Plants (1818). His description
states:
Every part of the plant densely and pulverulently
villous: leaves oblong, or oblong-ovate . . .
under side furfuraceously villous, often fer-
rugineous [sic], coriaceous and opaque, deciduous,
prominently veined beneath, margin revolute, entire,
and scabrous; panical terminal, frondose?
A specimen collected by Nuttall in Savannah, Ga. fits
these characters perfectly, and this may have been one of
the plants he had in mind when writing the above
description! The words "var. pubescens" in A. Gray's
handwriting appear on this sheet. Individuals with

larger leaves have traditionally been called var.

capreaefolia (Watson, 1825; K. Koch, 1872; Fernald, 1941).

 

Watson's excellent illustration shows a plant with large
elliptic to obovate leaves and large leafy bracts scattered
throughout the rather long and lax inflorescence. The

flowers also appear to be larger, possibly 5 mm long?

 

277

Fernald (1941) states that the panicle is naked or more
or less leafy bracted. Actually, the inflorescence always
has at least several large bracts, although they may fall
in the autumn before the leaves. Plants falling into

this category can be found on the Coastal Plain in the
Carolinas, and from southwest Georgia to Alabama, Miss-
issippi, Louisiana, Texas and Arkansas. The Arkansas
populations have especially large obovate leaves, often
shining above, and with twigs that become reddish in the
fall.

The range of 5. ligustrina var. foliosiflora

 

 

overlaps with that of var. ligustrina on the Coastal Plain

 

of southeast Virginia and the Carolinas (Figure 47). The
ranges also overlap in northeast Alabama and adjacent
parts of Tennessee. In these areas plants with inter-
mediate characters occur, and it is in these areas where
most problems in identification occur. The populations
in South Carolina are especially interesting in this

respect, where it appears as if var. ligustrina has

 

invaded the Coastal Plain and hybridized with var. folio-
siflora. The situation is further confused by the pre-

sence of extremely pubescent forms of var. foliosiflora

 

which superficially resemble var. ligustrina. Crossing

 

experiments are probably essential for a complete under-

standing of the situation.

 

Figure 47.

278

Regions of overlap in distributions of

Lyonia ligustrina vars. ligustrina and

 

 

foliosiflora. (Note: var. ligustrina is

 

 

 

to the north; var. foliosiflora is to the

south.)

 

 

 

279

 

 

 

 

 

 

 

 

 

-
‘l 00..
......C.I..
I .-
causes...
I.

 

 

 

 

 

 

 

 

 

280

Disjunct populations of var. foliosiflora occur

 

in Rabun Co., Georgia, and possibly also in Scott Co.,
Tennessee and McCreary Co., Kentucky. The Plants from
Rabun Co., collected in the canyon at Tallulah Falls by
J. K. Small (Aug. 15, 1893), are definitely var. £51157
siflora. This colony is probably extinct due to the .
flooding that followed the construction of Tallulah Dam
in the 1930's.

Kearney (1901) states that Lyonia l1gustrina var.

 

foliosiflora is one of the characteristic shrubs in the

 

Ericaceae shrub association which is important in the
cedar swamp communities of the Dismal Swamp. The more
important species of this association are Clethra alni-

folia, Itea virginica, Lyonia ligustrina var. folio-

 

 

siflora, Lyonia lucida, Ilex glabra, Rhododendron vis-

 

 

 

55555 and Vaccinium corymbosum. This woody undergrowth
is especially well developed along ditches and in clear-
ings. It is an association which is rapidly increasing
in the interior of the swamp, where it is said to have
been once almost unknown (Kearny, 1901).

Lyonia ligustrina var. foliosiflora is also

 

common in moist sandy pine forests where it grows mainly

in association with Leucothoé racemosa, Ilex spp., Myrica

 

cerifera and other species of Lyonia. It is equally at

 

home in shrub bogs (pocosins), cypress swamps or in

sandy pine flatwoods.

281

Lyonia Nutt. sect. Maria (DC.) C. E. Wood
J. Arnold Arb. 42:48. 1961.

Leucothoé D. Don sect. Maria DC., Prodr. 7(2):

 

602. 1839, Andromeda L. sect. Maria (DC.) A. Gray, Man.

 

Bot. North U.S. 266. 1848. Pieris D. Don sect. 55515
(DC.) Benth. & Hook., Gen. Pl. 2:588. 1876, Lyonia sub-
gen. 55515 (DC.) Drude, Nat. Pflanzenfam. IV. 1:44. 1889,
Xolisma Raf. sect. 55515 (DC.) Rehd., J. Arnold Arb.
5:55. 1924.

Deciduous or evergreen shrubs with round or
terete branches. Leaves entire, nearly glabrous with
small glandular hairs, sometimes nonglandular pubescent
on the major veins, 555 lepidote. Flowers in axillary
corymbose fascicles; corolla cylindric to ovoid-cylindric,
glabrous; calyx lobes lanceolate, glabrous to somewhat
pubescent. Filaments pubescent or merely roughened, with
prominent spurs. Capsule glabrous or nearly so, with
pale sutures usually not separating as a unit from the 5
valves in dehiscence. Coastal Plain from Long Island,
New York, south to Florida and west to Oklahoma, Arkansas
and eastern Texas.

Type Species: Lyonia mariana (L.) D. Don

S. Lyonia lucida (Lam.) K. Koch
Lyonia lucida (Lam.) K. Koch, Dendr. 2: 118. 1872.

Andromeda lucida Lam., Encycl. 157. 1783. Desmothamnus

 

 

282

lucidus (Lam.) Small, N. Am. Fl. 29(1): 64. 1914.
Pieris lucida (Lam.) Rehd., Dendr. Ges. 1915: 266. 1915.
Xolisma lucida (Lam.) Rehd., J. Arnold Arb. 5: 50. 1924.
Type: (P-LA?, not seen).

Andromeda nitida Bartr. 3x Marsh., Arbust. Amer.
8. 1785. Pieris nitida (Bartr. ex Marsh.) Benth. & Hook.,

Gen. P1. 2: 588. 1876. Andromeda obovate Raf. g§_A. Gray,

 

Syn. F1. N. Am. 32. 1878, as synonym. Lyonia nitida
(Bartr. ex Marsh.) Fern., Rhodora 10: 53. 1908. Neopieris
nitida (Bartr. ex Marsh.) Britton in Britt. & Brown, Ill.
F1. ed. 2, 2: 690. 1913. Desmothamnus nitidus Small,
Shrubs Fla. 96. 1913. Type: ”grows naturally in Carolina
and Florida" (BM?, not seen).

Andromeda coriacea Ait., Hort. Kew. 2: 70. 1789.

 

ggucothoé coriacea (Ait.) DC., Prodr. 7: 602. 1839.

Type: "Nat. of North Am. Mr. John Cree" (BM?, not seen).
Andromeda myrtifolia Salisb., Prodr. 290. 1796.

Type: (K7, not seen).

Andromeda marginata Viellard, Nouv. Duham. 1:

 

188. 1802. Lyonia marginata (Viellard) D. Don, Edinburgh

 

New Phil. Jour. 17: 159. 1834. Leucothoé marginata
(Viellard) Spach, Hist. Veg. 9: 482. 1840. Type: (not
seen).

Andromeda nitida var. rhombifolia Wood, Class

 

Book (ed. 1861) 488. 1861. Type: (NY?, not seen).

 

283

Andromeda lacustris C. Wright in Sauvage, Anal.

 

Acad. Ci. Habana 6: 250. 1870. Type: CUBA. E. Wright
3664 (US, holotypel).

Andromeda coriacea rubra Loddiges, Bot. Cab. 7:

 

672. 1822. Lyonia marginata var. rubra Lodd. ex Loudon,

 

Arb. Frut. Brit. 2: 1110, fig. 901. 1838. Pieris nitida

 

var rubra (Lodd.) Rehd. in Bailey, Cycl. Am. Hort. 5:

2622. 1916. Xolisma lucida var. rubra (Lodd.) Rehd.,

 

Man. Cult. Trees Shrubs 712. 1927. Lyonia lucida f.
rubra (Lodd.) Rehd., J. Arnold Arb. 20: 425. 1935.
Type: "Carolina and Georgia, growing in sandy forests"

(not seen).

A rhizomatous shrub to 2 (rarely 5 m) tall. Young
branches sharply three angled, glabrous, often arching.
Buds ovoid, glabrous or very sparsely pubescent, 1.0-3.0
mm long. Leaves evergreen, very coriaceous, reddish when
young, alternate, short-petioled, entire with revolute
margins, with intramarginal vein, elliptic to obovate,
1.0-10.5 cm long, 0.5-4.5 cm wide, with tips acuminate,
and bases obtuse or cuneate, adaxial surface lustrous,
very slightly pubescent, becoming glabrous with greater
age, abaxial surface sparsely dotted with glandular hairs,
midvein prominent, lateral veins diverging at an angle
and passing straight to edge of leaf. Flowers in axillary
corymbose fascicles; pedicels glabrous to sparsely

glandular pubescent, becoming glabrous with age, each

 

284

with a pair of bracteoles at the base. Calyx S—lobed,
persistent, lobes lanceolate, 2.0-5.0 mm long, 1.0-2.0 mm
wide, with abaxial surface glabrous, and adaxial surface
slightly nonglandular pubescent. Corolla ovoid-cylindric,
with 5 short lobes, pink, less commonly white or red,
glabrous, 5.0-9.0 mm long, 2.5-5.0 mm wide. Stamens 10,
inserted at the base of the corolla; filaments flattened,
slightly expanded near the base, S-shaped or with several
curves in older flowers, roughened, with two long spurs
on the back near the apex; anthers obtuse, the lobes
parallel, lacking apical awns, dehiscent by large ter-
minal pores, always with a white line of disintegration
tissue on the back of each lobe extending along the apex
of the filament and into the spurs. Style columnar or
fusiform, not exerted; ovary superior, 5-locular, glabrous.
Capsule ovoid to ovoid-globose, slightly S-angled, 3.0-
5.0 mm long, 3.0-5.0 mm wide, loculicidal, with pale
moderately to slightly thickened sutures, usually not
separating as a unit from the 5 valves in dehiscence,
glabrous. Seeds ca 300 per capsule, very narrow,
averaging 0.9 mm long. Chromosome number, n.= 12
(Callan, 1941), Figure 48.

Distribution: Coastal Plain from southeastern
Virginia to Florida, west to Louisiana. Also in Cuba
(Figure 49).

Flowering: late March to early May.

 

 

 

285

Figure 48. Representative specimen of Lyonia lucida

 

(Leonard 1269, NCU).

 

286

 

W0. 0' ‘50 Umvmfiy 0‘ Not!‘ Codi-O

SOUTH CAROLINA
Jasper County

Lvorln lucida (Lam.) K. KOC?‘

Swamp fort-st about 5,: ~1103 wsf-norfhwnn’. of
1.111%!” on unnunbnrac paved res: '.: canal.

F. N. Leonard 1904 '51.. r‘

Pth U! 'hr \‘1I'IUH‘I‘.'Y" lvnl'v.‘ \-.v,.

 

 

-, ml

 

Figure 49.

287

Distribution of Lyonia lucida.

 

 

‘————__—J-k

 

288

 

 

.....
.

 

  
 
 
  

' ‘ o d ‘ ~v' _r‘ ‘v‘-‘ ,
u -—* - - ~ ._ ' -
I—f f“ I f—L4 , l .XI) r‘/
-b’-: -0 ‘r’r “ ~__1 {ii ....
--«’ ‘ £0“ ~ r- 74.-fry?" :13'
. J
- T. ‘ ‘ _l f K

 

 

 

289

Representative specimens: Alabama. Baldwin Co.:
Fort Morgan Peninsula, Demaree 35890 (FSU*, NCU*); between
Gulf Shores and Fort Morgan, Ezanswiggig (NCU*); near
Fort Morgan, Harvill g Stallard s.n., 28 January 1950

(TEX*); 2 miles southwest of Theodore, Iltis g£_§1. 21590

 

(ENCB, NCU*, WIS); 10 miles south of Foley, Shinners 32265
(FSU*). Chilton Co.: 6.3 miles northeast of Clanton,
Clark 11626 (NCU). Coffee Co.: about 0.3 mile west of
Zion Chapel Church, §l§£k_1§§Q§ (NCU). Conecuh Co.:

Old Federal Rd. between US. 84 and Escambia Co. line,
Clark 12240 (NCU*). Covington Co.: 6 miles northwest

of Red Level, El2£§.l£££3.(NCU*)' Dale Co.: banks of

Choctawhatchee River, west of Newton, Godfrey 55480

 

(FSU*). Escambia Co.: vicinity of Atmore, Ahles 7043
(MISSA). Geneva Co.: just south of junction of Ala.
53 & Ala. 153, Clark 7685a (NCU). Henry Co.: near

Abbieville, Harbison 4101 (NCU*); 2.4 miles north of

 

Tumbleton, Wood 9074 (NCU*). Houston Co.: Cypress
Creek Swamp, Clark 7326 (NCU). Lee Co.: Auburn, Earle

§_Baker s.n., 8 July 1897 (NCU). Mobile Co.: 1 mile

 

north of Satuma, Iltis 32 31. 21410 (NCU, WIS). Monroe
Co.: Old Federal Road, between US. 84 and Escambia Co.
line, Clark 129§2_(NCU). Washington Co.: along Bassetts
Creek, near US. 43 bridge, Clark §1§2_(NCU*). Florida.
Alachua Co.: 2 miles west of Citra, 5521.6359 (FSU).

Baker Co.: McClenny, Lighthipe 688 (NCU). Bay Co.:

 

 

290

St. Andrews State Park, Wooten 209 (FSU*). Brevard Co.:

Merritts Island, Cocoa, Coker n.§., 14 January 1947 (NCU).

 

Broward Co.: near Pompano Beach, Lakela 28296 (FSU).

 

Charlotte Co.: Caloosa Forest Experiment Area, Parrott
64 (DUKE). Clay Co.: Gold Head Branch State Park, Lems
s.n., 24 July 1959 (MSC). Columbia Co.: near Lake City,

Rolfs 3.2,, 18 April 1892 (GA). Dixie Co.: about 1.7

 

miles north of Suwannee, Traverse 731 (TEX*). DeSoto

 

Co.: without definite locality, Small_§ DeWinkeler 9605
(US). Duval Co.: 1 mile south of Mayport, Godfrey 61201
(FSU). Escambia Co.: Pensacola, Harbison 4130 (NCU).
Dade Co.: west of Fulford, Moldenke 454 (DUKE). Flagler

Co.: about 6 miles east of Seville, Lems s.n., 19, 24

 

July 1959 (MSC*). Franklin Co.: 2 miles east of East

Point, Lazor 5209 (FSU, TEX*). Gilchrist Co.: 3 miles

 

east of Trenton, Wiggins 19502 (NCU). Glades Co.: south
of Palm Dale Junction, Lakela 31364 (NCU). Gulf Co.: 4

miles west of Wewahitchka, McDaniel 4077 (FSU*). Hamilton

 

Co.: 2 miles north of Bellville, Faircloth 1643 (NCU*).

 

Hernando Co.: Sect. 36, R17E, T218, Cooley 35 31. 6044

 

(FSU, MISSA, NY). Highlands Co.: Archbold Biological
Station, Colley gt_al. 6124 (FSU*). Hillsborough Co.:
about 12 miles southeast of Tampa, Bay_gt_al. 12129 (NCU).
Holmes Co.: 1 mile east of Ponce de Leon, Godfrey 51635.
(DUKE, FSU*, MISSA). Indian River Co.: Vero Beach,

Lemaire 165 (FSU*). Jackson Co.: Round Lake, Harbison

 

291

12559 (NCU). Jefferson Co.: near Wacissa, Demaree 49388
(NCU). Lake Co.: near Eustis, Nash 52 (MSC, US). Lee

Co.: Alva, Hitchcock 191 (US). Leon Co.: Silver Lake

 

Recreation Area, Lems 5.2., 8, 10, 11 July 1959 (MSC).

 

Levy Co.: about 5 miles northeast of Cedar Key, Godfrey
52824 (FSU, NCU). Liberty Co.: 9 miles northeast of

Bristol, McDaniel 4092 (FSU*). Manatee Co.: south out-

 

skirts of Bradenton, Kral 2117 (FSU). Marion Co.: Ocala

 

National Forest, Cooley 5251 (NY*); Juniper Springs, 5225
5.2., 16 July 1959 (MSC*): near Valparaiso, 23215‘11125
(NCU*). Okeechobee Co.: near Okeechobee, Demaree 52252
(FSU*). Orange Co.: 0.6 mile north of Vineland, 521E-
21222.5511 (NCU). Osceola Co.: 9 miles south of Kenans-

ville, Clark 3485 (DUKE*). Palm Beach Co.: Kelsey City,

 

29§g£,§.g., 28 January 1933 (NCU). Pasco Co.: 1 mile
north of Gower's Corners, Cooley 5522 (FSU, NY*).
Pinellas Co.: 1 mile southeast of Snug Harbor on Weedon
Island, BEI.EE.E£° 12221 (FSU, NCU*). Polk Co.: near
Lake Wales, Demaree 22252_(FSU, NCU). Putman Co.: 7
miles southwest of Palatta, Walker 1551 (TEX). St.
Johns Co.: 2 miles south of Moultrie, 2.‘§..§ 2. 2.
Correll 2252 (DUKE). Sarasota Co.: near Osprey, 52132
5.2., 8 March 1904 (DUKE). Seminole Co.: about 2 miles
east of Oviedo, 521.2E.2£3 12122 (FSU, NCU). Sumter Co.:
in Withlacoochee Development Project, Cooley 22 31. 2222

(FSU*, NCU). Taylor Co.: shrub-bog between Mikeville

 

292

and Perry, Godfrey 52854 (DUKE, FLA, MISSA, NCU).
Volusia Co.: Tomoka Game Preserve along tributary to

Little Tomoka River, Prichard 968 (FSU*). Wakulla Co.:

 

St. Marks Wildlife Refuge, Trott 104 (DUKE, FSU*).

 

Walton Co.: about 4 miles west of Paxton, Godfrey 61257a
(FSU). Washington Co.: 3 miles south of Chipley 54367
(FSU). Georgia. Atkinson Co.: 5 miles east of Willa-
coochee, §y£92_§ McVaugh 1522 (GA). Bacon Co.: about
7.5 miles east of Alma, Bozeman 2212 (NCU). Baker Co.:
Mossy Pond, Duncan 2255 (GA). Bartow Co.: about 4.5
miles of Adairsville, near Quicksand Pond, Greear 5512
(NCU*). Berrien Co.: 1.6 miles south of Ray City,

Faircloth 1924 (NCU). Brooks Co.: 3 miles east of

 

Quitman, Bozeman 4899 (NCU). Bullock Co.: 5.2 miles

 

southwest of Statesboro, Standley 21 (NCU). Candler Co.:
lS-Mile Creek Swamp, south of Metter, Bozeman 2122 (NCU).
Charlton Co.: about 1 mile west of Camp Cornelia, Oke-
fenokee Swamp, 2, 5,'2 5, 5, 2252.1522_(FSU). Chatham
Co.: near Wormslae Gardens, Savannah, Totten 5.3.,

18 April 1933 (NCU*). Clay Co.: Fort Gaines, Wigging-
£22 5.2., 27 December 1952 (GA). Coffee Co.: Rocky
Creek, about 3 miles south of the Ocmulgee River, Bozeman

4595 (NCU). Colquitt Co.: 3.6 miles southeast of Doerun,

Faircloth 5446 (NCU). Cook Co.: 3.6 miles east of Cecil,

 

 

Faircloth 2022 (NCU). CriSp Co.: 7.1 miles west, 38°

north of Cordele, Lane 102 (GA). Decatur Co.: Climax,

 

293

Harbison 13910 (NCU). Dodge Co.: edge of bayhead east

 

of Little Ocmulgee River, 0.5 miles west of Ga. 165,
Bozeman 5437 (NCU). Echols Co.: 1.5 miles from Lowndes

Co. line, northwest of Statenville, Clewell 2640 (FSU).

 

Effingham Co.: 3 miles south of Stillwell, Bozeman 4095

 

(NCU). Emanuel Co.: on Ga. 46, 0.6 mile west of Emanuel-

Candler Co. line, Bozeman 4273 (NCU). Evans Co.: 6.8

 

miles south of Claxton, Duncan 14658 (GA). Glynn Co.:

5 miles west of Brunswick, Cronguist 4919 (US). Grady Co.:

 

17 miles west of Thomasville, Wooten 1942 (FSU*). Jeff

 

Davis Co.: 7 miles southwest of Hazlehurst, Wilbur 3198

(GA). Jenkins Co.: Big Duke's Pond, Duncan 5635 (GA).

 

Lanier Co.: near Banks Lake Swamp, 4.7 miles west of

junction with US. 129 on Ga. 122, Bozeman 4724 (NCU).

 

Laurens Co.: about 25 miles south of Dublin towards
McRae, Wilbur 5221 (FSU). Lee Co.: near Smithville,
Oosting Z2_(DUKE). Liberty Co.: 11 miles east of
Taylor Creek, Duncan 2221 (GA). Long Co.: 1 mile west

of US. 301 on Ga. 261, Bozeman 2587 (NCU). Lowndes Co.:

 

9.8 miles southeast of Valdosta, Faircloth 1665 (NCU*).

 

McDuffie Co.: on Little Sweetwater Creek 3 miles south-
east of Boneville, Duncan 2125 (GA). McIntosh Co.:

about 1.5 miles south of northern tip of Sapelo Island,
Duncan 22522 (DUKE*, NCU). Montgomery Co.: southwest of
Charlotteville, Bozeman 5122 (NCU). Pierce Co.: at

Patterson, Bozeman 56034 (NCU). Richmond Co.: Fort

 

294

Gordon, Frisbee 2.2., 20 January 1957 (TEX). Screven Co.:
swamp near Cameron, 23322 21522 (GA). Tatnall Co.: 1.9
miles northwest of Reidsville, 22132 54258A (NCU*). Tel-
fair Co.: Sugar Creek Swamp on Co. Rd. S-114l, Bozeman
5522 (NCU). Thomas Co.: Governor Forbes' Swamp near

Thomasville, Coker 5,3., 31 December 1941 (NCU). Tift

 

Co.: 12 miles northeast of Tifton, Faircloth 2828 (NCU).

 

Toombs Co.: 4.5 miles north of Vidalia, Palmer 21 (NCU*).

Treutlen Co.: between Soperton and Adrian, Duncan 4734

 

(GA). Turner Co.: 4.1 miles east-northeast of Ashburn,

Faircloth 4392 (NCU). Ware Co.: 10 miles southwest of

 

Waycross, Murphy 2410 (NCU*). Wayne Co.: about 10 miles

 

north of Odum, Bozeman 3738 (NCU*). Wilcox Co.: 1.1

 

miles north of Alapaha River, southwest of Rochelle,
Bozeman 5590 (NCU). Worth Co.: along Abram's Creek,

1 mile west-southwest of Turner-Worth Co. line, Faircloth

 

3847 (NCU*). Louisiana. St. Tammany Co.: just north

 

of Slidell, Lemaire 796 (FSU*); about 3 miles southeast

 

of Abita Springs, Thieret 22252 (DUKE, FSU*). Tangipahoa
Co.: 13 miles east of Hammond, 2222712£g§.2_Warnock 723
(TEX*). Vernon Co.: Kisatchie National Forest, Cooley 2
22222‘2225 (NCU*). Washington Co.: on La. 16, 2.8 miles
west of Bogue Chitto River, 52222_1225_(DUKE*). 2122f
issippi. Clarke Co.: 7 miles southeast of Quitman, Bay
‘5222 (MISSA). Forrest Co.: 2 miles northwest of Eaton-

ville, Ray 7761 (NCU). George Co.: 5.5 miles south of

 

295

Agricola, McDaniel 4168 (FSU*). Handcock Co.: near Bay

 

St. Louis, Demaree 32974(DUKE*, MISSA, TEX); Clermont

 

Harbor, Rogers 2146-B (NCU*). Harrison Co.: near Miss-

issippi City, Demaree 25541 (FSU, NCU*). Jackson Co.:

 

near Ocean Springs, Demaree 29712 (DUKE*, TEX); Fontaine-

 

bleau Point, near Ocean Springs, Demaree 35695 (FSU*,

 

NCU). Lauderdale Co.: 1 mile east of Chunky, Kral 23692
(DUKE*, FSU). Marion Co.: about 2 miles south of Sandy

Hook, Ray 7911 (MISSA). Pearl River Co.: about 3 miles

 

west of Picayune, Sargent 2 25255 11221 (FSU*, NCU).
Perry Co.: near Beaumont, Webster 2 Wilbur 2212 (US).
Stone Co.: DeSoto National Forest, near McHenry, Demaree
22552 (NCU). Wayne Co.: Sect. 32, R8W, T6N, Piney Woods

Recreation Area, Jones 5516 (NCU). North Carolina. Beau-

 

 

fort Co.: 3 miles south of Chocowinity, Radford 22222
(NCU). Bladen Co.: 5 miles west of White Lake, 5555
525., 29, 30 July 1959 (MSC*). Brunswick Co.: about 12
miles northwest of Supply towards Bolton, Wilbur 12122
(DUKE). Carteret Co.: at the western end of Harker's
Island, Wilbur 2211 (DUKE). Chowan Co.: 1.3 miles east
of Chowan River on US. 17, 25155 22522 (NCU). Columbus
Co.: 2 miles south of Chadbourn, Wilbur 5212 (DUKE,
FSU*). Craven Co.: about 4 miles southeast of Havelock,
Wilbur 12225 (DUKE). Cumberland Co.: about 4 miles
southwest of Stedman, Wilbur 5112 (DUKE, FSU*). Cur-

rituck Co.: Grandy, on US. 158, Ahles 40151 (NCU).

 

296

Dare Co.: 1 mile north of Engelhart, Oosting 21 (DUKE*).
Duplin Co.: 2 miles west of Lenoir-Duplin Co. line on
NC. 11, Ahles 18359 (NCU*). Gates Co.: 4.6 miles west

of Mintonsville, Ahles 40332 (NCU*). Harnett Co.: 2

 

miles west of Spout Springs, Laing 514 (NCU). Hoke Co.:

4.2 miles south-southwest of Ashley Heights, Ahles 29439

 

(NCU). Hyde Co.: 5 miles north of Englehard, O'Connell
5,5., 8 July 1954 (NCU*). Johnston Co.: 2 miles south

of Parkers Hill, Radford 27832 (NCU*). Jones Co.: 1.2

 

miles east of Confort, Radford 21152 (NCU). Lenoir Co.:
3.5 miles southwest of Elm Grove, Radford 22222 (NCU).
Martin Co.: 4 miles northeast of Smithwick, Radford 22212_
(NCU). Moore Co.: south of Vass, Oosting354 (DUKE*).
New Hanover Co.: 15 miles south of Wards Corner, Wilbur
5151 (DUKE*). Onslow Co.: west of Jacksonville, 25155
{22215 (NCU). Pender Co.: 2 miles south-southeast of
NC. 210 intersection on US. 421, Weaver 2252 (DUKE*).
Person Co.: 4 miles west of Roxboro, Bowmer 115 (NCU).
Pitt Co.: 3.5 miles west of Greenville, Radford 22212
(NCU). Richmond Co.: Solomon Creek, 2, 2, 2, 5.5.,

13 November 1917 (NCU). Robenson Co.: about 1 mile
south of Red Springs, Wilbur 5222 (DUKE). Sampson Co.:
1 mile southeast of Midway, 22155 22522 (NCU). Scotland
Co.: 9.1 miles north of Laurinburg, 25155 22525 (NCU*).
Tyrell Co.: 3 miles southeast of Cross Landing, Radford

33792 (NCU). South Carolina. Aiken Co.: 3 miles

 

297

northwest of Beech Island, Radford 2251 (NCU). Allendale
Co.: 2 miles southwest of Barton, 25155_12522 (NCU).
Bamburg Co.: 0.3 miles southwest of junction of Co. Rd.
26 and Little Salkehatchie River, 25155 22125 (NCU*).
Barnwell Co.: near Rosmary Creek, southwest of Williston,
25155_55221 (NCU). Beaufort Co.: Hilton Head Island,
Butler 5,5., 7 May 1968 (NCU). Berkeley Co.: 5.3 miles
south-southeast of junction of US. 176 and SC. 27, 25155
22222 (NCU). Charleston Co.: without definite locality,
Kramer 5,5,, 23 April 1935 (DUKE). Chesterfield Co.:

7.7 miles north of McBee, Radford 12222 (NCU). Clarendon
Co.: 3 miles northwest of Silver, Radford 22555 (NCU*).
Colleton Co.: 2.2 miles northeast of Hampton Co. line

on US. 17A, 25155 12512 (NCU). Darlington Co.: near
Black Creek near Darlington Country Club, 55125 222
(NCU*). Dillon Co.: 1 mile east-southeast of Latta,
25155_21222 (NCU*). Dorchester Co.: vicinity of Harley-
ville, 25155 21252 (NCU*). Florence Co.: 4 miles east
of Olanta, 5511 5125_(NCU). Georgetown Co.: 12 miles
north of Georgetown, Godfrey 2 1y£55 1252 (DUKE).

Hampton Co.: 1.8 miles north-northeast of Early Branch,

Ahles 12460 (NCU). Horry Co.: Myrtle Beach, Christen-

 

berry 5,5,, 14 July 1938 (NCU). Jasper Co.: 5.5 miles
west-northwest of Tillman, Leonard 1269 (FSU*). Kershaw
Co.: near Lynche's River northeast of Bethune, Britt

861 (NCU). Lee Co.: 2.7 miles north of Lucknow, Radford

 

298

27384 (NCU*). Lexington Co.: 4 miles northwest of

Edmund, Radford 23401 (NCU*). Marlboro Co.: Beaverdam

 

Creek, 1.5 miles south of McColl, Radford 12637 (NCU).

Orangeburg Co.: vicinity of Springfield, Ahles 21618

 

(NCU*); south of Bowman, Ahles 31802 (NCU*). Richland

 

Co.: along SC. 12, 1.9 miles southwest of Kershaw Co.
line, Duke 1614 (NCU). Sumter Co.: 6 miles northwest

of Pinewood, near Poinsett State Park, Radford 21001

 

(NCU*). Williamsburg Co.: 2 miles northwest of New

Morrisville, Radford 24788 (NCU). Virginia. Chesapeake:

 

 

Dismal Swamp, Kearney 1592 (US). Isle of Wight Co.:

 

south of Zuni Harvill 12043 (NCU). Nansemond Co.: 1

 

mile southeast of Blackwater River on Va. 189, then 2.5
miles south, 22155_52125 (NCU*); near Cherry Grove,
Harvill 12125 (NCU*); south Quay, Harvill 12222 (NCU*);
near Cox Lansing, Harvill 11225 (NCU*). Southampton
Co.: 7 miles south of Franklin, Harvill 12252 (NCU*).

Lyonia lucida (Lam.) K. Koch is based on Andromeda

 

lucida., Encycl. Meth. 1:157. 1783. By Schneidner (1911)

this was considered a later homonym of Andromeda lucida

 

Jacq. which he assumed had been published in 1781,
because a plate with a very brief description appeared

in Jacquin, Icones Plantarum Rariorum, vol. 1, dated
1781-1786. It appears from reasons summarized by Rehder
(1935) that Jacquin's Collectanea Botanica vol. 1 was not

published until 1786, thus the correct citation of 2,

299

lucida should read Jacquin, Coll. Bot. 1:95. 1786; Icon.
P1. Rar. 1(1781-1786): 8, no. 79, t. 1786, which makes
2, lucida Jacq. a later homonym of 2, lucida Lam. and
Lyonia lucida (Lam.) K. Koch as the correct name under
Lyonia. The correct name for 2, lucida Jacq. is 252f

cothoé pgpulifolia (Lam.) Dippel.

 

As mentioned earlier 2, lucida is a common under-

story shrub in the cedar swamps of the Dismal Swamp in

 

Virginia (Kearney, 1901). Mohr (1901) and Harshberger
(1911) also record it from the white cedar swamps, cypress
swamps, and the damp or wet, more or less sandy, banks
of streams in the pine forests of the coastal plain in
Alabama. It occurs with many other ericaceous shrubs,

and with Cliftonia monop2ylla, Cyrilla racemiflora,

 

 

Myrica cerifera and various species of Ilex. I found

 

2, lucida to be very common around the bases of cypress
trees, often sharing this niche with other ericaceous

shrubs--especially Leucothoé racemosa. In moist habitats

 

such as this it usually is quite a large shrub, reaching
as high as 4 meters in some areas, with beautifully
arching branches and large elliptic leaves. Lyonia
lucida is also common in sandy pine flatwoods. In this
dryer habitat it is a dwarf shrub with more rigidly
ascending branches and much smaller, narrower leaves.
Yet even here it is more abundant in the low damper

localities, giving way to 2, fruticosa on the more

 

300

well-drained sites. When a narrow-leaved plant collected
in dry pine-palmetto scrub was transplanted to the moist
environment of a greenhouse it developed large elliptic
leaves, thus indicating that this difference is mainly
under environmental control.

Small (1927) gives a charming summary of the
beauty of this shrub in its natural surroundings:

In swamps when crowded with other greenery and over-
shadowed by trees this shrub is much subdued and its
pale pink flowers are sometimes eclipsed by larger
flowers. In the scrub or desert-like sandhills this
fetterbush is very decorative. The numerous bright-
green leaves are shining and free from parasitic
fungi and insect mutilation and the numerous drooping
clustered cone-like flowers are rosy. The whole
color scheme is enhanced by the background of white
sand, especially where the shrubs occur in pure
growth acres in extent.

Some interesting disjunct p0pulations of this
species occur around several small ponds in Bartow Co.,
Georgia. In several ponds it is the dominant species in
a clearly demarked zone which lies between the average
growing season water level and the spring high water level.
Thus Greear (1967) concluded that 2, lucida must require

a saturated substrate for at least part of the growing

season. It usually occurs with Acer rubrum and Smilax

 

walteri (Greear, 1967; Watts, 1969).

Lyonia lucida is commonly called Fetterbush
because when growing in thick colonies its branches can
make for difficult walking! Kearney (1901) records that

in southeastern Virginia it was known as ”hemlock.”

301

Marshall (1785) gives its name as Carolinian Red-buds

which is also fitting.

6. Lyonia mariana (L.) D. Don

Lyonia mariana (L.) D. Don, Edinb. New Phil.

 

Jour. 17: 159. 1834. Andromeda mariana L., Sp. P1. 393.

 

1753. Leucothoé mariana (L.) DC., Prodr. 7: 602. 1839.

 

Pieris mariana (L.) Benth. & Hook., Gen. P1. 2: 588. 1876.

 

Neopieris Eariana (L.) Britton 12 Britt. & Brown, Ill.

 

F1. ed. 2, 2: 691. 1913. Xolisma mariana (L.) Rehd.,

 

Jour. Arnold Arb. 5: 51. 1924. Type: "habitat in Vir-
ginia" Kalm (LINN, probably sheet no. 563.5, microfiche
photo seen).

Andromeda pulchella Salsb., Prodr. 289. 1796.

 

Type: (K2, not seen).

Andromeda mariana var. ovalis Sims, in Bot. Mag.

 

37 t. 1579. 1813. Type: (not seen).

Andromeda mariana var. oblonga Sims, in Bot. Mag.

 

sub. t. 1579. 1813. Type: (not seen).

Xolisma mariana f. vestita Rehder, J. Arnold Arb.

 

5: 51. 1924. Lyonia mariana f. vestita (Rehd.) Parks,
First Sci. Field Meet. Palmetto State Park, Texas 9.
1936. Type: TEXAS: Hardin Co. sandy pine lands,

Silsbee. April 25, 1916. 2. 2, Palmer 9561. (A, holotypel).

A rhizomatous shrub to 1.5 m tall. Young branches

erect or nearly so, round or only slightly angular,

 

302

glabrous to moderately pubescent; bark on older twigs
gray, shredding. Buds glabrous, ovoid, 1.0-2.5 mm long.

Leaves deciduous to partly evergreen, membranous to

 

slightly coriaceous, alternate, short-petioled, entire,

with nonrevolute or only slightly revolute margins, lack-

 

ing intramarginal vein, with lateral veins arching toward
edge of leaf, elliptic to elliptic-ovate, 3.0-9.0 cm long,

1.0-4.5 cm wide, with tips acute or mucronate, and bases

 

cuneate, adaxial and abaxial surfaces sparsely dotted
with glandular hairs, becoming glabrous with greater age,
often nonglandular pubescent on the major veins. Flowers
in axillary corymbose fascicles, on leafless branches of
the previous season; pedicels glabrous to densely pub-
escent, each with a pair of bracteoles at the base.

Calyx 5-1obed, deciduous with the leaves, lobes elongate-
lanceolate, 3.0-9.5 mm long, 1.0-4.0 mm wide, with adaxial
surface glabrous, and abaxial surface glabrous to densely
pubescent. Corolla broadly cylindric to ovoid-cylindric,
with 5 short lobes, white or rarely pink, glabrous or
nearly so, 7.0-14.0 mm long, 5.0-8.0 mm wide. Stamens 10,
inserted at the base of the corolla; filaments flattened,
expanded near the base, S-shaped, pubescent, with two
spurs on the back near the apex; anthers obtuse, the lobes
parallel, lacking apical awns, dehiscent by large terminal
pores, always with a white line of disintegration tissue

on the back of each lobe extending along the apex of the

303

filament and into the spurs. Style columnar or fusiform,
not exserted; ovary superior, 5-1ocular, glabrous or
slightly pubescent. Capsule ovoid-pyramidal, truncate

at the contracted apex, slightly 5-angled, 4.0-6.5 mm
long, 3.0-6.0 mm wide, loculicidal, with pale, moderately
to much thickened sutures, sometimes separating as a unit
from the 5 valves in dehisence, glabrous or essentially
so. Seeds ca 450 per capsule, very narrow, averaging

1.0 mm long. Chromosome number, 2 = 12, (Callan, 1941),
Figure 50.

Distribution: Long Island, New York, south to
Florida, with disjunct populations in eastern Texas,
Louisiana, Arkansas and Missouri (Figure 51).

Flowering: late April to early June.

Representative specimens: UNITED STATES.

Arkansas. Calhoun Co.: Demaree 22119 (MO). Clark Co.:

 

2 miles south of Hot Springs-Clark Co. line, Tucker 6989

 

(NCU). Dallas Co.: Demaree 15016 (MO). Drew Co.: near

 

Monticello, Demaree 14978 (MO, WIS). Faulkner Co.:

 

Cove Creek, Buchholz 979 (TEX, WIS). Garland Co.: near

 

Mill Creek Rd., Ashe 522., 23 May 1931 (NCU*). Grant Co.:

near Poyen, Demaree 11050 (GA). Hempsted Co.: near

 

Blerins, Moore 450116 (WIS). Hot Springs Co.: near Mal-
vern, Demaree 14991 (MO, WIS). Jeff Co.: Demaree 19100
(MO). Pope Co.: Crow Mt., Galla Creek woods, Tucker

7214 (NCU*). Pike Co.: sandy knolls near Antonine,

 

 

Figure 50.

304

Representative specimen of Lyonia mariana

 

(Ahles 12436, NCU).

 

 

 

_J—np— _._-—_——f__

305

 

 

     

 

~d: ...,

L'umuun‘ .. (in-u.“ Hun-u u
Lyonia autism (1..) D. Den

W H.Dvnu§ 1”)

Wk- 00 Oh Ulivonlt, ol Noni Carolin.
SOUTH CAROLINA

”kit". or. Co.:":-

I L221. min: (1..) D. Lon
\
’ .1-.. 1;. _,.., n , ‘ -
.1? ‘on\. ".L. .I.’ WHOLE. . .k ’ I... l".:'..'.e'.5', .1- -nr '
\ ‘I '-r-nr';, 3.1 r.‘.c: Yxr‘rtrnrsnt of YI-f‘psfiv... n; -. . ."
_ 2,.

lift: b. ."-‘.. Ari:
Hany E AMOI ‘_ 32:9‘

I ¢.lv.l '..| "u ")n r. u' "yr I-.v..l” .

-' W'vli

 

Figure 51.

306

Distribution of 2yonia mariana.

307

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

9
\o ...
I\ o
. \ .
m 9. \J
"
u. ‘ u
. 0s\ .\
‘\
Am.
/
3'.
1...}.
l-
g ,
.... O

 

 

308

Demaree 9644 (GA, MO). Pulaski Co.: Engelman 5,2,

 

(MO). Saline Co.: near Bauxite, Demaree 53814 (VDB*).

 

Sevier Co.: near Horatio, Brinkley 169 (TEX). Delaware.

 

 

Sussex Co.: 0.8 miles north of Fenwick, Judd 22 (MSC*).

District 22 Columbia. Swamp near Reform school, McCarthy

 

 

 

5,2., June 1885 (US). Florida. Alachua Co.: sec. 4,
TllS, R17E, BTY 421 (NCU*). Columbia Co.: about 5 miles

northwest of Lake City, Ward 5624 (NCU). Duval Co.: near

 

Jacksonville, Curtiss 4717 (US). Gadsden Co.: along US

 

90, 1 mile west of Ochlockonee River, Jackson 535 (FSU).

 

Jefferson Co.: along Lloyd Creek, east of Lloyd, Godfrey
55552 (FSU*, NCU). Lake Co.: Ocala National Forest,
southeast of Lake Dorr, 25y 2225 (NCU). Leon Co.: south-
west of Tallahassee, Potagas (number eligible) 5 May 1964
(FSU). Madison Co.: ‘14 miles southeast of Madison,
Ramsey 52_21, 222 (FSU). Marion Co.: Ocala National
Forest, Mather 2:122 (GA). Manatee Co.: Palma Sala.,

Tracy 6954 (M0, MSC). Putnam Co.: Welaka, University

 

of Florida Conservation Reserve, Lems 5,2., 17, 18 July
1959 (MSC). Seminole Co.: north of Goldenrod, Cooley
52251. 7402 (FSU, NCU). Taylor Co.: 4 miles north of

Perry, McDaniel 2 Godfrey 4312 (FSU*, NCU); 1.3 miles

 

northeast of Shady Grove, McDaniel 5 Godfrey 4346 (FSU*);

 

along Fla. 98, 29 miles east of Jefferson-Taylor Co. line,
Wiggins 20051 (NCU*). Volusia Co.: 1.5 miles east of

Astor bridge, Kral 6566 (FSU*); Tomoka Game Preserve,

 

.‘-
..__

 

309

Prichard 643 (NCU*). Wakulla Co.: near Wakulla, Mol-

 

denke 1121a (DUKE*). Georgia. Appling Co.: 3 miles

 

southwest of Baxley, Duncan 10925 (DUKE, MO, TEX). Ben

 

Hill Co.: 0.8 mile north of Rebecca, Faircloth 4434
(NCU). Berrien Co.: at intersection of Ga. 645 and 168,

Faircloth 3630 (M0, NCU). Brooks Co.: 0.8 mile northeast

 

of Quitman, Faircloth 4246 (M0, NCU*). Bullock Co.:

 

without definite locality, Harpgr 867 (MO, US). Candler

 

Co.: on Ga. 129, 0.4 mile north of the Canoochee River,

 

Bozeman 4233 Chatham Co.: near Savannah, Biltmore Herb.

 

25d (US). Clinch Co.: 17 miles south-southeast of Homer-

ville, Duncan 21692 (GA). Colquitt Co.: 6.5 miles west

 

of Lenox, Faircloth 3368 (M0, NCU*). Cook Co.: 6.6 miles

 

 

west of Adel, Faircloth 2369 (MO, NCU). Dodge Co.: north
of McRae, Duncan 13755 (GA). Echols Co.: 1 mile north

of Mayday, Faircloth 3325 (NCU*). Effingham Co.: south

 

of Stillwell, Bozeman 2122 (NCU). Emanuel Co.: 9 miles
east of Adrian, Wilbur 2212 (FSU*). Irwin Co.: 3.2 miles
southwest of Irwinville, Faircloth 5463 (NCU). Jeff Davis
Co.: 2 miles east-southeast of Hazelhurst, Duncan 11222
(GA). Jenkens Co.: 7.5 miles south of Millen, Duncan
22212 (GA, MO). Lanier Co.: 1.4 miles northeast of the
junction of Ga. 168 and US. 221, Faircloth 2552 (M0, NCU).
Laurens Co.: about 25 miles south of Dublin towards
McRae, Wilbur 2222 (FSU*). Long Co.: 2.5 miles west

of US 301 on Ga. 261, Bozeman 2583 (NCU). Lowndes Co.:

310

 

10 miles north of Valdosta, Faircloth 3739 (M0, NCU).

 

McIntosh Co.: 2.6 miles northwest of Cox, Bozeman 1011

 

(NCU). Pierce Co.: 1.3 miles east-southeast of Bristol,
Duncan 16296 (GA). Screven Co.: 3.5 miles north of

Newington, Kral 24024 (DUKE). Telfair Co.: about 3

 

miles northwest of Lumber City, Wilbur 3134 (GA). Thomas

 

Co.: north of Thomasville, Bozeman 4958 (NCU*). Turner

 

Co.: 3.7 miles east-northeast of Ashburn, Faircloth 4353

 

 

(NCU). Ware Co.: near Suwannee Lake, Harper 868 (NCU*).
Wheeler Co.: without definite locality, Harvey 5.2., ‘

17 May 1939 (GA). Louisiana. Caddo Co.: 4.5 miles

 

northwest of Vivian, Thieret 32265 (DUKE, FSU*). Maryland.

 

Anne Arundel Co.: Bodkin Creek, Smith 5,2., 1 June 1876
(US). Baltimore Co.: Towson, Campus of Goucher College,

Lems 5,2,, 6 June 1961 (MSC*). Cecil Co.: Elks Neck,

 

2, 2, 2 2, 2, Leonard 5993 (US). Charles Co.: 6.9 miles
northwest of Rock Point, Ahles 57835 (NCU*). Hartford

Co.: Oakington, Fessender 5192 (US). Prince Georges

 

Co.: Beltsville, in the last remnant of Powderville

Bogs, 2, 2, Churchill 5,2,, 30 May 1970 (MSC*). St.

 

Marys Co.: 2 miles west of Clements, Smith 5427 (US).

 

Talbot Co.: Easton, Shreve 181 (US). WOrchester Co.:

 

near Snow Hill, Gleason 5,2,, 6 June 1934 (DUKE).

Missouri. Dent Co.: 2 miles south of Lake Spring,

 

T35W, R7W, $.15, 16, Kucera 5,2., 11 May 1953 (GA).

New Jersey. Atlantic Co.: without definite locality,

311

Reed 51_(FSU*). Burlington Co.: Atsion, along the Mullica

River, 2, 2. Churchill 5.2., 15 June 1946 (MSC*); along

 

Rt. 206 just south of Atsion, Grashoff 349 (MSC*). Camden

 

Co.: Clementon, Hawes 310 (US). Cape May Co.: Bennett
Bog, 3.6 miles north of Cape May, Crow 109 (MSC*); near
Goshen, Gillis 5633 (MSC*). Cumberland Co.: 2 miles

east of Milville, Lems 60.05.26.02 (MSC*). Middlesex W

Co.: 2, 2, 2, 5,2,, (MO). Monomouth Co.: near Asbury .

 

 

Park, VanSickle 5,2., 12 July 1894 (US). Ocean Co.: 5 )
miles northeast of Egg Harbor 2525'512., October 1964

(MSC*). 252_2222. Richmond Co.: north shore, Drushel

5225 (US). Suffolk Co.: Ocean Beach, Lighthipe 522.,

September 1913 (TEX*). North Carolina. Alamance Co.:

 

Swepsonville along the Haw River, Ramseur 2222 (NCU).
Anson Co.: without definite locality, 2, 2, 2, 5,2,,

20 July 1917 (NCU). Beaufort Co.: 3 miles south of
Chocowinity, Radford 22225 (NCU). Bladen Co.: 6 miles
north of White Lake, 2251_12522 (FSU*). Brunswick Co.:
about 13 miles northwest of Supply, Wilbur 5522 (DUKE*).
Carteret Co.: west end of Harker's Island, Wilbur 2222
(DUKE*). Chatham Co.: near Seaforth, Radford 22222
(NCU). Chowan Co.: 0.5 mile north of Valhalla, 22155
22225_(NCU). Columbus Co.: near Reaves Ferry,

Schallert 5,2,, 30 April 1929 (DUKE). Craven Co.:

 

14 miles east of Kingston, Radford 2 Steward 1106 (NCU).

Cumberland Co.: 2 miles west of Fayetteville, Wilbur

312

5814 (DUKE). Dare Co.: Roanoke Island, Schallert 877

 

(NCU). Duplin Co.: 1 mile southeast of Pin Hook,
Ahles 33178 (NCU). Durham Co.: about 6 miles north-

northeast of Durham, Wilbur 14978 (DUKE*). Edgecombe

 

Co.: 0.5 mile south of Speed, Radford 36852 (NCU*).

 

Franklin Co.: 5 miles east of Franklenton, Thompson 12

 

(DUKE). Gates Co.: sandridge near White Oak Pocosin,

Duke 780 (NCU). Granville Co.: 0.3 mile south of

 

Bullock, Bell 2222 (NCU). Green Co.: 0.6 mile south

 

of Walstonburg, Radford 36509 (NCU). Halifax Co.:

 

1 mile northwest of Halifax, 22155 12225 (NCU). Harnett
Co.: 1 mile south of Pineview, Wilbur 2125 (DUKE*).

Hoke Co.: 0.4 mile southwest of Ashley Heights, 22155
22225 (NCU*). Johnston Co.: 4 miles north of Edmundson,
Radford 21251 (NCU). Lee Co.: 4 miles north of Sanford,

Bell 194 (NCU). Lenoir Co.: 1 mile west of Deep Run,

 

Radford 22221 (NCU). Martin Co.: 4 miles south-southeast
of Smithwick, Radford 25255 (NCU*). Montgomery Co.:

Mt. Gilead Lp, 2, 2, 2, 522., 14 September 1917 (NCU).
Moore Co.: 2 miles west of Lakeview, Wilbur 12212 (DUKE*).
Nash Co.: 0.4 mile west of Baxley, 22155_21112 (NCU).

New Hanover Co.: 9 miles south of Wilmington, Wilbur

5521 (DUKE). North Hampton Co.: 1 mile east-northeast

of Gumberry, 22155 21252 (NCU). Onslow Co.: 1 mile

east of Catherine Lake towards Chinquapin, Wilbur 2221

(FSU). Orange Co.: about 0.5 mile east of junction of

 

313

NC. 87 & Rt. 1727, Whigham 1222 (NCU). Pamlico Co.:
near Arapahoe, Oosting 22212 (DUKE). Pender Co.: 4
miles south of Wallace, 2121 12222 (FSU). Pitt Co.:
near King's Cross Roads, 22125‘222 (NCU). Randolph Co.:
4.4 miles southeast of Liberty, 2311 12221 (NCU). Rich-
mond Co.: 3.5 miles northwest of Hoffman, Radford 11222

(NCU). Robeson Co.: 2.5 miles south of Red Springs,

Ahles 28916 (NCU*); 0.9 mile east of Rennert, Weaver 2303

(DUKE*). Sampson Co.: 2.8 miles west-northwest of Mid-
way, 22125‘22522 (NCU). Scotland Co.: 1.3 miles north-
east of Silver Hill, 22132.22152 (NCU). Union Co.: 2
miles west-southwest of Marshville, 221g§_21525_(NCU).
Wake Co.: near Macedonia, Godfrey 22222 (NCU). Warren
Co.: 0.5 mile north of Embro, 2211.2215 (NCU). wash-
ington Co.: near Roper, Godfrey 2221 (DUKE). Wayne Co.:
Neuse State Park, Radford 2225 (NCU). Wilson Co.: 1.4
miles south-southwest of Wilbanks, Radford 22115 (NCU).
Oklahoma. McCurtain Co.: 2.7 miles north of Tom,
Waterfall 12222 (US). Pennsy1vania. Chester Co.:
Nottingham Barrens, Pennell 1222 (US). Lancaster Co.:
without definite locality, Garber g,2., June 1864 (MSC).
Philadelphia Co.: without definite locality, 222H2,2.,
1883 (MO). 22222 Carolina. Aiken Co.: without definite
locality, Gabrielson 222., 16 May 1966 (NCU). Allendale
Co.: 2.6 miles east of Sycamore, 2211‘2522 (NCU*).

Bamberg Co.: west of Denmark, Ahles 26010 (NCU).

 

 

314

Barnwell Co.: northwest of Cline, Ahles 57051 (NCU).

 

Beaufort Co.: 1.1 miles north of Pritchardville, Ahles
12374 (NCU*); 1 mile south of Prichardville, Judd 319
(MSC*). Berkeley Co.: 1.6 miles south of Wadboo Creek,

Bozeman 9145A (NCU*). Calhoun Co.: 1.8 miles south of

 

junction of Co. Rds. 71 and 31, Ahles 30344 (NCU).

 

Charleston Co.: 6 miles northwest of McClellanville, l
Judd 22 (MSC*). Chesterfield Co.: 1 mile south of

Ruby, Radford 12334 (NCU). Clarendon Co.: 5 miles east

 

of Rimini, Badford 24399 (NCU). Colleton Co.: 0.6 mile

 

southwest of US. 21 on Co. Rd. 48, Bell 2273 (TEX, NCU*).

 

Darlington Co.: north of Darlington, Harbison 14147

 

(NCU). Dillon Co.: 0.6 mile northeast of Floydale,
22152'21222 (NCU). Dorchester Co.: 0.4 mile north-
northwest of Colleton-Dorchester Co. line on US. 15,
22122‘25152 (NCU). Edgefield Co.: 3 miles south-south-
west of Trenton, Radford 25222 (NCU). Florence Co.:

northwest of Florence, Bell 2,2., 30 June 1958 (NCU*).

 

Georgetown Co.: 5.5 miles south of Georgetown, Godfrey

2 Tryon 199 (DUKE). Hampton Co.: 3.1 miles northwest

 

of Yemassee, 22122 12225 (NCU*). Horry Co.: Myrtle
Beach, 29225_2 Totten 2,2., 10 July 1932 (NCU). Lan-
caster Co.: 0.4 mile north-northwest of Little Lynches
Creek near US. 601, 22135.21225 (NCU). Lee Co.: 2.5
miles southeast of Manville, Radford 24132 (NCU). Lex-

ington Co.: 3.5 miles northeast of Gaston, Radford 23379

315

(NCU). Marion Co.: south of Britton Neck by Co. Rd. 49,
Bell 7866 (NCU). Marlboro Co.: 10 miles north of Ben-

nettsville, Radford 18945 (NCU). Orangeburg Co.: Orange-

 

 

burg, Freeman 56160 (NCU). Richland Co.: near Columbis,
Chilsen 2,2,, 30 April 1936 (DUKE). Sumter Co.: 7.5

miles north-northeast of Pinewood, Radford 24019 (NCU*).

 

Williamsburg Co.: 3 miles northwest of Greenleyville,

Radford 21305 (NCU). Texas. Gonzales Co.: Palmetto

 

 

State Park, Reis 106 (DUKE*); Saefje Bog, Tharp 2271244

 

(DUKE*). Hardin Co.: Silsbee, Avenue L extension,

Correll 29392 (ENCB, FSU*, NCU, TEX, US). Handerson Co.:

 

11 miles southwest of Athens, Spring Lake, Correll 29612

(FSU*). Jefferson Co.: Beaumont, Hooks 2.2., 30 May

 

1934 (TEX). Marion Co.: near Jefferson, Biltmore

Herbarium 252 (US). Nacogdoches Co.: without definite

 

locality, Wallace 2,2,, 22 April 1938 (TEX). Tyler Co.:
Hyatt Lakes Estates, about 2 miles south of Warren,

Correll 35830 (ENCB, FSU, NCU). Virginia. Arlington

 

 

Co.: Arlington Forest, Balls 7803 (US). Caroline Co.:

 

at Mattaponi River off Co. Rd. 722, 22222.12211 (NCU).
Chesterfield Co.: just east of junction of Co. Rd. 619
& US. 95, 22222’5251 (NCU). Dinwiddie Co.: 1 mile
north of Burgess, Harvill 12252 (NCU). Fairfax Co.:

Buel 102 (MO). Greenville Co. northeast of Emporia,

 

Ahles 58485 (NCU). Gloucester Co.: 3 miles northwest

of Gloucester Point, Harvill 12440 (NCU*). Henrico Co.:

316

west of Elk Station, Harvill 16290 (NCU*). Isle of
Wright Co.: 2 miles southeast of Franklin, Harvill 19807
(NCU). James City Co.: College of William and Mary
Woods, Barans 122 (NCU). King George Co.: near Cash

Corner, Harvill 16157 (NCU*). King and Queen Co.: 0.2

 

mile south of Middlesex-King & Queen Co, line, James 5243
(NCU). Louisa Co.: 1.5 miles west of junction of Va. 22

& US. 522, James 13832 (NCU). Lunenburg Co.: south of

 

Kenbridge, James 3053 (NCU). Middlesex Co.: 5 miles

 

northwest of junction of US. 17 & Co. Rd. 610, 22222 5552
(NCU). Nansemond Co.: south of South Quay, Harvill 12222
(NCU*); Cox Landing, Horton 252:2 (NCU*); near Cox Land-
ing, 5212212 Hodgdon 211 (DUKE, MISSA*, NCU, TEX). New
Kent Co.: 1.4 miles south of junction of Co. Rds. 627 &

1002, Gillespie 624b (NCU*). Northumberland Co.: 0.9

 

mile north of Northumberland-Lancaster Co. line, James
5872 (NCU). Powhatan Co.: southeast of Powhatan, James

6185 (NCU*). Prince Georges Co.: 1.2 miles northeast

 

of junction of Co. Rds. 612 and 611, 22222'2122 (NCU).
Southampton Co.: 4.2 miles southwest of junction of US.
460 & Co. Rd. 635, 22222 2122 (NCU). Sussex Co.: 3.7
miles south of Littleton, 22122‘52522 (NCU*). Spotsyl-
vania Co.: south of Fredricksburg, 22222 5122 (NCU*).
In the Linnean Herbarium (as seen from micro-
fiche) there are four sheets identified as Andromeda

mariana. The first three of these were collected by

 

 

317

P. Kalm and are indeed 2, mariana, but the fourth, col-
lected by James Gordon of Mile End, is 2, lucida. Sheet
no. 563.5 is probably the type and was evidently collected
in August or September as it is flowering with the leaves
present. This specimen has widely elliptic, moderately
coriaceous leaves, and appears to be from the southern
part of the range (i.e., Virginia or southward).

2yonia mariana is often confused with Zenobia

 

pulverulenta from which it may be separated by its
awnless anthers, longer sepals, and unique urn-shaped
capsule with thickened sutures.

2yonia mariana occurs in sunny habitats with

 

sandy acid soil, and is often found at the edge of
forests, along roadsides and on railroad grades. Delzie
Demaree (oral communication) has said that in Arkansas
it occurs on sand hills and old pastures. Kearney (1901)
only reports it from a railway embankment near Suffolk,
Virginia. Duke (1961) records it as occurring at several
localities on the xeric sandhills of the Carolinas.
These sandhills once supported vast stands of long-leaf
pine but are now mainly covered with scrubby oaks. Fire
resistance is of considerable importance to sandhill
species (Duke, 1961). Lyonia mariana is well adapted

to surviving fires because of its extensive network of
underground rhizomes and even seems to increase in

abundance after fires! I have seen many vigorous young

 

 

318

shoots around charred older branches where the plant had
evidently been burned a year or two previously. Lyonia
mariana is also common in the palmetto-scrub flatlands

of southeastern Georgia and is a common component in the
scrub vegetation occurring after the larger trees have
been removed for lumber, etc. Farther north, it abounds
in the pine barrens of New Jersey. It is a common species
in the deciduous swamps where it grows with Kalmia

angustifolia, Lyonia ligustrina, Toxicodendron vernix,

 

 

 

Leucothoé racemosa, Vaccinium atrococcum, Clethra alni-

 

 

folia, Ilex glabra, Alnus serrulata and Rhododendron

 

 

 

 

viscosum. Lyonia mariana is also abundant in the moist

 

 

savannas which are flat grassy plains dotted with clumps
of trees and which are often flooded for part of the year.

Here it occurs with various sedges and grasses, Sarracenia

 

purpurea, Leucothoé racemosa, Iris pgismatica, Eriocaulon

 

 

 

 

 

29cangu1are and Tofieldia racemosa. It is also found in

 

 

the New Jersey pine plains. There some of the common
associated species are Comptonia peregrina, Leiophy11um

buxifolium, Gaylussacia frondosa, G. resinosa, Hudsonia

 

 

 

ericoides, Ilex glabra, Kalmia angustifolia, Myrica

 

 

 

 

carolinensis, Smilax glauca, Vaccinium pennsylvanicum,

Quercus marylandica, Kalmia latifolia and Pinus rigida

 

 

(Harshberger, 1916). The causes of this stunted vege-

tation are well summarized by Harshberger (1916):

 

319

We may state that the dwarfed character of the
pines, oaks, and other plants of the plains is
primarily due to the stiff, impervious subsoil,
and the light easily dried-out, sandy surface soil
which may freeze to the depth of a meter in winter
and dry out in midsummer until the subsoil is hard
and stiff and the surface soil almost devoid of ‘
moisture. The elevated character of the country
facilitates rapid drainage by run-off and seepage,
and thus influences of elevation, coupled with the
strong winds that blow, has a strong dwarfing
effect. . . . Fire has had an influence on the [A
native vegetation, but its effect has been secondary as
to the soil factors, for forest fires have burned
over as severely and as repeatedly the nearby

forest areas without appreciably stunting the growth
of the pine trees.4 L“

 

 

Thus 2. mariana is not only a beautiful little shrub but
hardy as well!

The western populations tend to be much more
pubescent and have been described as forma vestita
(Rehder, 1924). The branchlets, calyx and leaves of
this form are covered with a soft villous pubescence.

Yet plants do occur in the eastern parts of the range
that are densely villous on the major veins of the leaves.
It is probable that this trait is controlled by a certain
complex of closely linked genes which occur at a higher
frequency in the western populations. In all other char-

acters the plants of the two regions are identical.

 

4However, Andresen (1959) has concluded that the
primary cause of these dwarf forests of Pinus ri ida,
Quercus ilicifolia and Q. marylandica is repeateg Burning.
He states that I'from thE results of the chemical and
physical analyses of the soils sampled that no appre-
ciable differences exist in any property between the
Plains and the Barrens areas."

 

 

320

2, mariana is commonly called Stagger-bush
because of its poisonous properties, especially to young

animals such as lambs or calves.

 

 

LITERATURE CITED

LITERATURE CITED

Adams, C. D. 1972. Flowering Plants of Jamaica. Robert
MacLehose and Co., Ltd. The University Press.
Glasgow.

Aiton, W. 1799. Hortus Kewensis. 3 vols. London.

Alston, R. E. and B. L. Turner. 1963. Biochemical Sys-
tematics. Prentice Hall, Inc. Englewood Cliffs,
New Jersey.

Alston, R. E. 1966. Chemotaxonomy or biochemical sys-
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