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This is to certify that the
thesis entitled

PHYLOGENETIC RELATIONSHIPS IN POLIOMINTHA
AND RELATED GENERA IN THE MENTHEAE (LAMIACEAE)

presented by

GRANT THOMAS GODDEN

has been accepted towards fulfillment
of the requirements for the

M. S. degree in Plant Biology

 

 

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PHYLOGENETIC RELATIONSHIPS IN POLIOMINTHA
AND RELATED GENERA IN THE MENTHEAE (LAMIACEAE)

By

Grant Thomas Godden

A THESIS
Submitted to
Michigan State University

In partial fulfillment of the requirements
for the degree of

MASTERS OF SCIENCE
Plant Biology

2009

ABSTRACT

PHYLOGENETIC RELATIONSHIPS IN POLIOMINTHA
AND RELATED GENERA IN THE MENTHEAE (LAMIACEAE)

By

Grant Thomas Godden

Poliomintha (Lamiaceae; subfamily Nepetoideae; tribe Mentheae) is a genus of
aromatic shrubs from the southwestern United States and northem Mexico. For
over 100 years, the circumscription of Poliomintha has remained uncertain and
taxa have been transferred to and from Poliomintha and related genera on the
basis of various morphological characters. For my thesis research, sequence
data from the nuclear ribosomal ITS and two plastid regions, tmL-tmL-trnF and
mL32-tmL, were used to objectively test morphologically-based hypotheses of
relationship within Poliomintha and among genera in tribe Mentheae.
Poliomintha is not monophyletic as currently circumscribed and represents at
least two distinct lineages: a lineage that includes the type species, P. incana,
and either (or both) Rhododon or Clinopodium; and a second lineage that
includes the remaining species of Poliomintha and species of the allied genus,
Hedeoma. Shimodaira-Hasegawa tests confirm that P. incana is not related to
the remaining Poliomintha species. Therefore, the genus Poliomintha may be
best limited to the type species, P. incana, pending phylogenetic evidence that
determines the relationships of Poliomintha species to the other genera with

confidence.

ACKNOWLEDGEMENTS

This study was made possible through financial support from both the Paul
Taylor Academic Achievement Fund administered by the Department of Plant
Biology as well as the Graduate School, College of Natural Sciences, and the
International Studies Program of Michigan State University (MSU). I wish to
extend my sincerest thanks to my major professor and friend, Alan Prather,
whose encouragement and expert guidance enhanced my research and
graduate school experiences. Additional thanks belong to members of my
guidance committee, Richard Triemer and Andrew Jarosz, who always made
time for me and offered helpful guidance. To my labmate and friend, Orlando
Alvarez-Fuentes, thank you for providing my initial training experiences, for
assisting me in the laboratory, and for your support and advice during

challenging times.

Additional thanks belong to my fellow graduate students, thank you for
your friendship and support throughout my MSU experience. To my family, thank
you for your patience and encouragement. Lastly, I wish to express my eternal
gratitude for the support of my dearest friends. To Jannes Szyren, Natalia Ruiz
Rubio, Eleni Beli, Abe Pollister, Nidal Karim, Isabel Alvarez Sancho, Julian
Taurozzi, and Marisa Rinkus, thank you for believing in me when I couldn't
believe in myself. You are my family and I wouldn’t have made it without your

support, encouragement, and love.

TABLE OF CONTENTS

LIST OF TABLES ................................................................................... v
LIST OF FIGURES ................................................................................ vi
CHAPTER 1: INTRODUCTION ................................................................. 1
Taxonomic History ........................................................................ 2
Morphology ................................................................................. 5
Phylogenetic Relationships ............................................................. 6
CHAPTER 2: MATERIALS AND METHODS
Plant Material and Sampling ............................................................ 9
DNA Extraction ........................................................................... 10
Marker Choice ............................................................................ 11
Amplification and Visualization ....................................................... 12
Purification and Sequencing .......................................................... 15
Sequence Alignment and Data Analyses .......................................... 16
Congruence and Hypothesis Testing ............................................... 18

CHAPTER 3: RESULTS

Marker Choice ............................................................................ 20
Phylogenetic Analyses .............................................................................. 20
ITS Analyses of Poliomintha and Related Genera .............................. 21
Plastid Analyses of Poliomintha and Related Genera .......................... 28
Combined Analyses of Poliomintha and Related Genera ..................... 34
Tests of Alternative Phylogeny ....................................................... 41
CHAPTER 4: DISCUSSION .................................................................... 43
Phylogenetic Relationships among Poliomintha and Related Genera ...... 43
Importance of Morphological Characters in Delineating Genera ............ 46
The Taxonomy of Poliomintha ........................................................ 48
APPENDICES ..................................................................................... 50
APPENDIX A ............................................................................................. 51
Voucher lnforrnation for Material Used in the Phylogenetic Study .......... 52
APPENDIX B ............................................................................................. 61
Aligned Sequence Matrix 1: Nuclear Ribosomal lntemal Transcribed
Spacer (lTSl, 5. BS, ITSZ) ............................................................. 62
Aligned Sequence Matrix 2. trnL lntron and tmL- th Spacer of the
Chloroplast Genome ............................................................................... 101
Aligned Sequence Matrix 3: rpL32 - tmL Spacer of the
Chloroplast Genome ............................................................................... 137
LITERATURE CITED ........................................................................................ 173

LIST OF TABLES

Table 1-1: Recognized taxa and nomenclature in preVious taxonomic treatments
of Poliomintha ....................................................................................... 4

Table 1-2: Characteristics distinguishing Poliomintha from the closely allied
genera Hedeoma and Hesperozygis. Characters adapted from R. S. Irving (1972,
1980) .................................................................................................. 6

Table 2-1. Primer sequences used for PCR amplification and sequencing.
Forward (F :) and reverse (R:) designates are indicated before each primer
name ................................................................................................ 1 3

Table 2-2. Components of polymerase chain reactions. Amplification was carried
out using AmpliTaq Gold® DNA Polymerase with Buffer II and MgCl2 reagents
(Applied Biosystems, Foster City, CA). All plastid regions (trnL—th, tmD-tmT,
th-tme, rpl32-tmL, and nth-rpl32) were amplified using the same quantities

of reaction components ........................................................................ 14
Table 2-3. PCR profiles for amplification of nuclear and plastid markers ......... 14
Table 2-4. Aligned characters excluded from the phylogenetic analyses ......... 16

Table 2-5. Models of sequence evolution and parameters for three datasets
analyzed using the maximum likelihood method. Models were determined by a
hierarchical likelihood ratio test (Goldman 1993; Huelsenbeck and Crandall 1997)
as implemented in ModelTest 3.7 (Posada and Crandall 1998). Parameters
estimated by ModelTest are indicated for each model ................................. 18

Table 3-1. Characteristics and phylogenetic utility of sequences from two plastid
regions acquired from ten taxa in the Mentheae ........................................ 20

Table 3-2. Sequence characteristics of regions used in phylogenetic
analyses ........................................................................................... 22

Table 3-3: Results of SH tests. Numbers indicate P value / difference in negative
log likelihood (-In L) score. Significance at the a = 0.05 level is indicated with an
asterisk ............................................................................................ 43

LIST OF FIGURES

Figures 3-1 A and B: One of 97,217 most parsimonious (MP) trees recovered
from an analysis of ITS sequence data (treelength = 485 steps; CI = 0.518, Rl =
0.722, and RC = 0.374, excluding uninformative characters). The randomly
selected MP tree is shown as a rectangular cladogram with branch lengths
(above) and bootstrap support values (bold, below) indicated. Branches that
collapse in the strict consensus tree are indicated with arrows .................. 24-25

Figures 3-2 A and B: Most likely tree recovered from a maximum likelihood (ML)

analysis of ITS sequence data from 71 accessions in the tribe Mentheae. Shown
here is a phylogram (-In L = 352426454) with branch lengths drawn to scale and
bootstrap support (>50%) indicated above the branches .......................... 26-27

Figures 3-3 A and B: One of 88,000 most parsimonious (MP) trees recovered
from an analysis of combined plastid (tmL-F and rpl32-tmL) sequence data
(treelength = 248 steps; CI = 0.887, RI = 0.841, and RC = 0.746, excluding
uninfonnative characters). The randomly selected MP tree is shown as a
rectangular cladogram with branch lengths (above) and bootstrap support values _
(bold, below) indicated. Branches that collapse in the strict consensus tree are
indicated with'arrows. Accessions used to represent identical groups of
sequences are indicated with an asterisk ............................................. 30-31

Figures 3-4 A and B: Maximum likelihood tree (-In L = 4086.57963) recovered
from an analysis of combined plastid (tmL-F and rpl32-trnL) sequence data for
83 accessions in the tribe Mentheae. Branch lengths are drawn to scale and
bootstrap support (>50%) is indicated above the branches. Accessions used to
represent identical groups of sequences are indicated with an asterisk ...... 32-33

Figures 3-5 A and B: One of 43,966 most parsimonious (MP) trees recovered
from a combined analysis of ITS, trnL-F, and rpl32-tmL sequence data
(treelength = 735 steps; CI = 0.553, RI = 0.749, and RC = 0.414, excluding
uninfonnative characters). The randomly selected MP tree is shown as a
rectangular cladogram with branch lengths (above) and bootstrap support values
(bold, below) indicated. Branches that collapse in the strict consensus tree are
indicated with arrows ........................................... . .......................... 37-38

Figures 3-6 A and B: Maximum likelihood tree (-In L = 7951.01811) recovered
from the combined analysis of nuclear (ITS) and plastid (tmL-F and rpl32-trnL)
sequence data for 71 accessions in the tribe Mentheae. Branch lengths

are drawn to scale and bootstrap support (>50%) is indicated above the
branches ..................................................................................... 39-40

vi

Chapter 1

INTRODUCTION

Poliomintha A. Gray (Lamiaceae) is a small genus comprising seven species with
distributions in the deserts and arid regions of northeastern Mexico and the
southwestern United States. Poliomintha species are characterized by a shrubby
habit, tubular and symmetrical calyces, large tubular corollas with two stamens,
and oblong nutlets (Epling and Stewart 1939; Irving 1972). None of these
characters are unique to Poliomintha; rather the combination of characters can
be used to distinguish Poliomintha from other closely allied genera in family
Lamiaceae.

Poliomintha is a member of subfamily Nepetoideae, which is divided into
four tribes: Escholtzieae, Lavanduleae, Mentheae, and Ocimeae (Cantino et al.
1992). The genus belongs to a clade of New World Mentheae (Wagstaff et al.
1995; Williams, unpublished data), a group historically viewed as taxonomically
difficult because of complex morphological interpretation. Systematists such as
Bentham (1834), Briquet (1897), Epling and Stewart (1939), and Irving (1972,
1980) and others circumscribed groups in the tribe based on habit and other
morphological characters, but discussed difficulty in assigning characters
sufficiently important to distinguish genera. Recognition of some genera in the
Mentheae, including Poliomintha, has been controversial and an ongoing subject
'of taxonomic debate. Consequently, the generic status of Poliomintha has been

questioned multiple times throughout history and its species have been

transferred to and from genera such as Hedeoma Pers. and Hesperozygis

Epling.

Taxonomic History

The genus Poliomintha was first proposed by Asa Gray (1870) to accommodate
two species, P. Iongiflora A. Gray and P. incana (Torr.) A. Gray. However,
suggestion of the group came eleven years earlier with Torrey’s description of
Hedeoma incanum (1859). Regarding the classification of his new species,
Torrey noted, “It may remain in Hedeoma for the present, but, if other species
like it should be found, it may be the type of a new genus.” During the twenty
years after Gray’s formation of Poliomintha, two species (Hedeoma moi/e Torr.
and Keithia [Hesperozygis] man'folia S. Shauer) were transferred by Gray (1872)
to Poliomintha and two additional species, P. glabrescens A. Gray ex. Hemsley
(1882) and P. bicolor Watson (1890), were described. A fifth name, P. greggii,
was referenced in Watson’s publication, but was not formally described (nomen
nudum).

Briquet (1897) recognized the inherent difficulties in the taxonomic
disposition of the taxa and subsumed Poliomintha into Hedeoma in his treatment
of. the Labiateae in Die Naturlichen Pflanzenfamilien. However, the work of
Epling and Stewart (1939) restored the generic status of Poliomintha and
informally established two new sections, section Incanae to accommodate P.
incana and section Saturejoides to accommodate P. glabrescens and P.

Iongr'flora. As for the other species, Epling and Stewart (1939) placed P. bicolor in

synonymy with P. longiflora and transferred P. molle and P. marifolr’a into
Hedeoma and Hesperozygis respectively. Poliomintha conjunctrix was
described by Epling and Wiggins in 1940, bringing the genus to a total of four
species.

In the most recent revision (Irving 1972) four species of Poliomintha were
recognized: P. Iongiflora and P. glabrescens belonging to section Saturejoides R.
S. Irving; and P. conjuntn'x and P. incana belonging to section Poliomintha.
Poliomintha longiflora var. congesta also was described by Irving, but more
recently has been reduced to synonymy with Clinopodium dominguense Urb. and
Ekman. Since Irving’s revision, three additional species were described; P.
madrensis by Henrickson (1982) and P. dendritica and P. bustamanta by Turner
(1993). With the latest additions, a total of seven species were recognized for this

study (Table 1-1).

Table 1-1: Recognized taxa and nomenclature in previous taxonomic treatments

of Poliomintha.

Taxonomist

Recognized Taxa

 

Standley (1923)

Poliomintha A Gray

P. bicolor

P. glabrescens
P. incana

P. longiflora

P. marifolia

P. moi/is

 

Epling and Stewart (1939)

Poliomintha A Gray
Poliomintha sect. Incanae (bold text below)
Poliomintha sect. Saturejoides

P. glabrescens
P. incana
P. Iongiflora

P. bicolor =syn. Poliomintha longiflora
P. man'folia = syn. Hesperozygis marifolia
P. mollis = syn. Hedeoma molle

 

Irving (1972)

Poliomintha A Gray
Poliomintha sect. Poliomintha (bold text below)
Poliomintha sect. Saturejoides R. S. Irving

P. conjunctrix

P. glabrescens

P. incana

P. Iongiflora var. congesta
P. Iongiflora var. Iongr‘flora

P. manfolra= syn Hesperozygis marifolia
P. moi/is = syn. Hedeoma molle

 

Proposed

Poliomintha A Gray
Poliomintha sect. Poliomintha (bold text below)
Poliomintha sect. Saturejoides R. 8. Irving

P. bustamanta
P. conjunctrix
P. dendritica
P. glabrescens
P. incana

P. Iongiflora

P. maderensr's

P. Iongiflora var. congesta = syn. Clinopodium dominguense
P. marifolia = syn. Hesperozygis marifolia
P. moi/is =gyn. Hedeoma molle

Morphology

Poliomintha are small aromatic shrubs with slender, moderately branched, and
ascending-erect four-angled stems with minute and usually appressed
pubescence. The species have small leaves that are oval, elliptiwl, or linear in
shape and with entire margins. The leaves are opposite and attached to the stem
at the base (sessile) or on short petioles. The flowers are solitary or grouped into
two-, three-, or five-seven-flowered cymules in the leaf axils of the upper one-half
to one-third of the plant. The calyces are characteristically tubular and more or
less symmetrical, 13-15-nerved, sometimes hirsute-annulate within, softly
pubescent on the margins and inner surfaces, and with subequal teeth that are
narrowly deltoid, acute, and connivent to close the orifice. The red, orange-red,
purplish, pale lavender, pink, or white corollas of Poliomintha are tubular and
zygomorphic, with a conspicuous emarginated upper lip and a spreading, three-
Iobed lower lip. The corollas are pubescent on the outer surface and have a
dense ring of pubescence (annulus) within, below the middle of the tube. The
flowers have two, slightly exserted fertile stamens that are seated well above the
middle of the tubular corolla with glabrous filaments and widely divergent anther
sacs on well-developed connectives. The gynobasic style is long exserted and
bifid. The fruit is a schizocarp of four oblong nutlets.

Previous taxonomic treatments by Epling and Stewart (1939) and Irving
(1972, 1980) distinguished Poliomintha from the closely allied genera Hedeoma
and Hesperozygis on the basis of a combination of habit, calyx, and nutlet

characters and chromosome counts (Table 1-2).

Table 1-2: Characteristics distinguishing Poliomintha from the closely allied
genera Hedeoma and Hesperozygis. Characters adapted from R. 8. Irving (1972,

 

 

1976,1980)
Hedeoma Poliomintha Hesperozygis
Habit: Herbaceous, Habit: Shrubs Habit: Shrubs

occasionally semishrubs

Cam sham: Gibbous or

saccate, but never funnelforrn

Cain smmetm:
Zygomorphic

Cam teeth: Differentiated into
upper and lower sets; usually
upper deltoid, lower
lanceolate

Calyx annulus: Well-defined
ring at juncture of upper and
lower teeth

Fruit: Nutlets oblong,
mucilaginous when moistened

Chromosome counts’:

2n = 34, 36, 44, 72, 144

 

Calyx shag: Tubular

Calfl symmetm:
Actinomorphic

Calyx teeth: Not differentiated

into upper and lower sets;
deltoid

Calfl annulus: Absent or in an
irregular ring

Fruit: Nutlets oblong,
not mucilaginous when
moistened
Chromosome counts‘:

2n=36

 

Calfl shag: Funnelforrn,
campanulate

W:
Actinomorphic

Calfl teeth: Not differentiated

into upper and lower sets;
deltoid or lanceolate

Calyx annulus: Well-defined
ring near middle of tube

Fruit: No comparison provided

WI:

2n=44

 

1.

Chromosome counts based on cytological studies of 24 species of Hedeoma and one species

each of Poliomintha and Hesperozygis, respectively (Irving 1976).

Phylogenetic Relationships

The relationships of Poliomintha to other New World genera in the Mentheae are

uncertain. The only phylogenetic study that included Poliomintha is the molecular

study of Wagstaff et al. (1995), who used a phylogenetic analysis of chNA

restriction site variation to infer relationships among several relevant genera in

subfamily Nepetoideae and tribe Mentheae. They included only one species

 

each of Poliomintha and Hedeoma; Poliomintha longiflora was placed as sister to
Hedeoma drummondii Benth. in an unresolved clade of species from genera that
included Acinos Mill., Blephilia Raf, Calamintha L., Conradina A. Gray,
Minthostachys (Benth.) Spach, Monardella Benth., Piloblephis Raf., and Satureja
L. (of which the sampled species have since been transferred to Clinopodium L.).
Previous morphologically-based studies have closely examined the
relationships among Poliomintha and related genera (Epling and Stewart 1939;
Irving 1972; and Irving 1980), but not in a modern phylogenetic context. The
most recent revision of Poliomintha by Irving (1972) suggested that Poliomintha
is monophyletic and closely allied to the genus Hedeoma and some species of
Hesperozygis. However, it is important to recognize that the definitions of
monophyly and phylogeny have evolved with more recent advances in molecular
biology (Judd et al. 2002) and a rigorous phylogenetic analysis for Poliomintha
and related genera has never been performed. Many contemporary studies of
the Lamiaceae have employed DNA sequence data and phylogenetic methods to
infer relationships among mints at lower taxonomic levels (Wagstaff and
Olmstead 1997; Steane et al. 1999; Prather et al. 2002; Jamzad et al. 2003;
Trusty et al. 2004; Walker et al. 2004; Braiichler et al. 2005; Trusty et al. 2005;
Edwards et al. 2006; Rosello et al. 2006; Walker and Systma 2007; Edwards et
al. 2008; and Schmidt-Lebuhn 2008), facilitating the testing of monophyly for
several genera and allowing for much-needed revision of existing
circumscriptions. The studies have demonstrated that some genera in the

Mentheae are not monophyletic—e.g., Micromen'a Benth. (Brailchler et al. 2005),

Salvr'a L. (Walker et al. 2004), and Satureja (Wagstaff et al. 1995)—and, in some
cases, the discovery of paraphyletic or polyphyletic groups has initiated
taxonomic revisions at the generic level (Cantino and Wagstaff 1998; Steane et
al. 1999; Harley and Granda Paucer 2000; Steane et al. 2004; BraiJchler et al.
2006; Ryding 2006; and Pool 2008).

For my thesis research, a molecular phylogenetic study was conducted to
examine closely the generic status of Poliomintha. The goals of the study were to
test the monophyly of the genus, to elucidate interspecific relationships within
Poliomintha and among taxa from closely related genera in the Mentheae, and to
clarify the taxonomy (of Poliomintha and closely related or ambiguously placed

species using phylogenetic data.

Chapter 2

MATERIALS AND METHODS

Plant Material and Taxonomic Sampling

Herbarium specimens from subfamily Nepetoideae and tribe Mentheae
(Lamiaceae) were selected for morphological review from the following herbaria:
CAS, Ml, MO, MSC, NY, US, and TEX/LL. Specimens were evaluated and
selected for the phylogenetic analyses to represent morphological and
geographical variation within species and major taxonomic groupings within the
genera Poliomintha, Hedeoma, and Hesperozygis and tribe Mentheae.
Permission for destructive sampling was requested and granted from each
herbarium prior to DNA extraction (see Appendix A for voucher information).

Individuals sampled for phylogenetic analyses included six taxa from
Poliomintha; 23 taxa from Hedeoma representing the subgenera Cilata,
Poliominthoides, Saturejoides, and Hedeoma; and four taxa from Hesperozygis.
The seventh Poliomintha species, P. maderensis J. Henrickson, was unavailable
for sampling due to its rarity in collections.

Previous revisions of Hedeoma by Epling and Stewart (1939) and of
Poliomintha and Hedeoma by Irving (1972 and 1980 respectively) suggested that
Poliomintha is not only closely allied to genera such as Hedeoma and
Hesperozygis, but also related to several genera in tribe Mentheae and subtribe
Melissinae, including two- and four-stamened genera such as Satureja section
Gardoquia (R. and P.) Briq. (now Clinopodium), Cunila L., Glechon Spreng.,

Rhabdocaulon (Benth.) Epling, and Rhododon Epling, etc. Therefore, in order to

infer phylogenetic relationships among Poliomintha and closely allied genera
within the Mentheae, 21 additional taxa were sampled (see Appendix A).
Mentha and Thymus were chosen as outgroup genera based on the
phylogenetic results of Wagstaff et al. (1995) and Walker and Systma (2007).
Mentha rotundifolia Huds. and Thymus mastichina L., were selected as
representatives of the two genera to facilitate use of ITS sequence data
previously published by Prather et al. (2002) and plant tissue available at MSC

for re-extraction and amplification of selected plastid genomic regions.

DNA Exu’action

Genomic DNA was extracted from herbarium tissue using the method of Doyle
and Doyle (1987) and the small-scale extraction techniques of Loockerrnan and
Jansen (1996). Approximately 10-20 mg plant tissue from each herbarium
specimen was sampled. Tissue was homogenized in 1.7 mL tubes with 400 uL
extraction buffer comprised of 100 mM Tris-HCI at pH 8.0, 1.3 M NaCl, 20 mM
ethylenediaminetetraacetic acid (EDTA), 2% hexadeclytrimethyl-ammonium
bromide (CTAB) at pH 8.0, 0.5% 2-mercaptoethanol and 4% polyvinyl-
pyrrolidone (PVP). An additional 400 pL extraction buffer was added to each
homogenate, incubated at 60°C for 30 minutes and inverted six times at five-
minute intervals. DNA was extracted from each homogenate with the addition of
550 uL chloroformzoctanol (24:1) followed by centrifugation at 14,000 rpm for
three minutes. The supernatant from each sample was transferred to a new 1.7

mL tube and DNA was precipitated with tvvo-thirds volume ice-cold isopropanol

10

and overnight storage at -20°C. Precipitated DNA was centrifuged into a pellet at

14,000 rpm for 6 minutes, washed with a layer of 800 pl 76% EtOH/0.01M
Nl-l40Ac for ten minutes, and resuspended in 30-40 pL water. DNA samples that

failed to amplify using PCR were suspected of having impurities that inhibited
amplification and subsequently purified using the Qiagen DNeasy Plant Mini Kit

(Qiagen, Ltd) following the manufacturer protocol.

Marker Choice

Two markers, the nuclear ribosomal lntemal transcribed spacer (including ITS1,
5.88, and ITSZ regions) and the combined tmL intron and tmL-th intergenic
spacer (referred to hereafter as trnL-trnF) from the plastid genome, were
selected because they have been used successfully in previous systematic

studies of the Mentheae (Trusty et al. 2004; Trusty et al. 2005; Edwards et al.

2006; and Walker and Systma 2007). Four additional plastid regions, trnDGUC-

tmTGGU, tmsUGA-tmeCAU, rpl32-tmL, and nth-rpl32, recommended by

Shaw et al. 2005 and Shaw et al. 2007 were surveyed to identify one additional
phylogenetically informative plastid region for the analysis. The regions were
compared for their efficient amplification across in-group taxa and the
phylogenetic utility of their sequence content as measured by average sequence
variability and number of potentially phylogenetically informative characters. The
following subset of in-group taxa were used to evaluate and select a region:

Clinopodium mexicanum, Hedeoma costata, Hedeoma drummondii, Hedeoma

11

hispida, Hedeoma molle, Hedeoma palmeri, Hedeoma patrina, Hedeoma pusilla,
Poliomintha incana, and Poliomintha Iongiflora. For reasons identified in the
results, rpl32-tmL was selected as the second plastid marker for the phylogenetic

analyses.

Amplification and Visualization

Polymerase chain reaction (PCR) was carried out using an MJ Research PTC-
100 then'nalcycler and 25 pL reactions containing standard PCR components
(Table 2-2), 1 pl DNA template from one of three prepared 1:1, 1:9, or 1:99
dilutions, and universal primers (Table 2-1). Three PCR profiles were used to
amplify nuclear and plastid regions (Table 2-3). Bands were resolved by
electrophoresis on a 2% agarose gel in 1X Tris-acetate-EDTA (T AE) buffer,
stained with ethidium bromide, and visualized with UV illumination. For samples
that successfully amplified, PCR was repeated two additional times using a 50 pL

reaction to amplify sufficient quantities of DNA for purification.

12

Table 2-1. Primer sequences used for PCR amplification and sequencing.
Forward (F:) and reverse (R:) designates are indicated before each primer name.

 

Marker 1: ITS
(ITS1, 5.88, ITSZ)
Primer Primer sequence (5’-3')
F: ITS4 (White et al. 1990) TCCTCCGC‘I‘I‘ATTGATATGC
R: ITSS (White et al. 1990) GGAAGTAAAAGTCGTAACAAGG
R: ITSSM (Sang et al. 1996) GGAAGGAGAAGTCGTAACAAGG

Marker 2: tmL-trnF
(trnL intron, trnL-trnF intergenic spacer)

 

Primer Primer sequence (5’-3')
F: TabC (T aberlet et al. 1991) CGAAATCGGTAGACGCTACG
R: TabF(Taber1etetal. 1991) ATI’TGAACTGGTGACACGAG

Marker 3: trnDGUc-tmTGGU
(intergenic spacer, embedded trnY and trnE genes)

 

Primer Primer sequence (5’-3’)
F: tmDGUCF (Demesure et al. 1995) ACCAATTGAACTACAATCCC
R: trnTGGU (Demesure et al. 1995) CACCACTGAGTTAAAAGGG

Marker 4: trnSUGA-tmeCAU
(intergenic spacer, embedded trnG, ycf9, and pst genes)

 

Primer Primer sequence (5’-3’)
F: trnSUGA (Demesure et al. 1995) CGAAATCGGTAGACGCTACG
R: trnfMC’“U (Demesure et al. 1995) ATTTGAACTGGTGACACGAG

Marker 5: rpl32-tmL
(intergenic spacer)

 

. Primer Primer sequence (5’-3’)
F: rpL32-F (Shaw et al. 2007) CAGTTCCAAAAAAACGTACTTC
R: trnL‘UAG’ (Shaw et al. 2007) CTGCTI’CCTAAGAGCAGCGT

Marker 6: nth-rpl32
(intergenic spacer)

 

Primer Primer sequence (5’-3’)
F: nth (Shaw et al. 2007) GAAAGGTATKATCCAYGGMATATT
R: rpL32-R (Shaw et al. 2007) CCAATATCCCI WW I I ICCAA

13

Table 2-2. Components of polymerase chain reactions. Amplification was carried
out using AmpliTaq Gold® DNA Polymerase with Buffer II and MgCl2 reagents
(Applied Biosystems, Foster City, CA). All plastid regions (tmL-trnF, tmD-tmT,
th-tme, rpl32—tmL, and nth-rpl32) were amplified using the same quantities
of reaction components.

 

 

Components ITS Plastid
Purified distilled H20 14.75-16.00 pl 16.50 pl
10X PCR buffer 2.50 pl 2.50 pl
MgCl2 2.50 pl 2.00 pl
0.2 mM dNTPs

(lnvitrogenTM Life Technologies) 2'00 “I 2'00 “I
20 pM forward primer 0.25 pl 0.25 pl
20 pM reverse primer 0.25 pl 0.25 pl
Dimethyl sulfoxide (DMSO) 1.25-2.50 pl 1.25 pl
Taq DNA polymerase 0.25 pl 0.25 pl
DNA Template 1.00 pl 1.00 pl
TOTAL 25.00 pl 25.00 pl

 

Table 2-3. PCR profiles for amplification of nuclear and plastid markers.

 

ITS — following the parameters of Prather and Jansen (1998)

 

Step Temperature Time Notes
(00) (minutes)

1 94 5

2 72 2

3 94 1

4 43 1 Cycle to Step 3 for 30 cycles
5 72 7

6 15 - Indefinite hold

14

Table 2-3. Continued

 

tmL-th, trnD-tm T, and th-trn f M

 

Step Temperature Time Notes
(°C) (minutes)

1 80 5

2 94 1

3 50 1

4 72 2 Cycle to Step 3 for 36 cycles
5 72 5

6 4 - Indefinite hold

 

rpl32-trnL and nth-rpl32 — following the parameters of Shaw et al. (2007)

 

Step Temperature Time Notes
(°C) (minutes)

1 80 5

2 95 1

3 50 1

4 65 4 Cycle to Step 3 for 31 cycles
5 65 5

6 4 - Indefinite hold

 

Purification and Sequencing

PCR products were purified using the Qiagen PCR Purification Kit (Qiagen, Ltd)
according to the manufacturer’s recommendations and sequenced in both
directions at the Michigan State University Genomics Technology Support
Facility using either the ABI 3730 Genetic Analyzer or ABI Prism 3700 DNA

Analyzer (Applied Biosystems, Foster City, CA).

15

Sequence Alignment and Data Analyses

DNA sequence data were assembled and edited using MEGA4 (T amura et al.
2007). Sequences were initially aligned using Clustal W and were refined by eye
in MEGA4. End regions from each of the three aligned data matrices were
excluded from consideration in the phylogenetic analyses, as were regions where

sequence homology could not be easily determined (Table 2-4).

Table 2-4. Aligned characters excluded from the phylogenetic analyses.

 

Aligned Characters

 

 

Regions of Total
Marker Excluded end regions ammuous alignment excluded
ITS 1-73; 712-779 474-504 172
trnL-th 1 - 63; 944 - 1010 n/a 130
rpl32-trnL 1 - 14; 946 - 953 691-702 34

 

To maximize computational efficiency by eliminating sequence
redundancy in the data matrices, analyses of pairwise genetic distance were
conducted using the combined plastid and ITS datasets. Multiple accessions with
identical plastid sequences were identified within each of the following nine taxa
(two accessions each, except where noted): Acinos arvensis, Clinopodium
brownei, Clinopodium chandlen', Hedeoma mandoniana, Hedeoma media,
Hedeoma pulegioides (four accessions), Hesperozygis marifolia, Poliomintha

incana, and Rhododon angulatus. Twenty-two other accessions with identical

16

plastid sequences were classified into one of two groups (see Results: groups
defined in Figures 3-3 A and B). In total, 33 accessions were removed from
consideration and a single accession was used in the phylogenetic analysis to
represent each of ten groups of identical sequences. No sequences were
removed from the ITS matrix.

Analyses of the ITS, combined plastid, and combined ITS plus plastid
datasets were conducted in PAUP*4.0.b10 (Swofford 2002) using maximum
parsimony (MP) and maximum likelihood (ML) optimality criteria. Branch support
for the resulting trees from each analysis was evaluated using bootstrap
analyses (Felsenstein, 1985). Gaps and missing data were ignored.

For the MP analysis, a heuristic search was conducted using 1000
random addition replicates with tree bisection and reconnection (TBR) branch
swapping, holding one tree during each stepwise addition and saving only the
best trees. Bootstrap (BS) support values were obtained for the resulting trees
using a heuristic search with TBR branch swapping, 1000 replicates and ten
random addition sequences, saving no more than 100 trees per replicate.

For the ML analyses, a hierarchical likelihood ratio test (Goldman 1993;
Huelsenbeck and Crandall 1997) as implemented in ModelTest 3.7 (Posada and
Crandall 1998) was used to determine the best model of sequence evolution for
the ITS, combined plastid, and combined ITS plus plastid datasets (Table 2-5).
Model parameter values generated by ModelTest were used in the ML analyses
conducted in PAUP and a heuristic search with 100 random replicates and TBR

branch swapping was used, saving no more than 100 trees per replicate.

17

Bootstrap analyses were conducted in PAUP using TBR branch swapping, 100

replicates with ten random additions per replicate, saving no more than 100 trees

per replicate.

Table 2-5. Models of sequence evolution and parameters for three datasets
analyzed using the maximum likelihood method. Models were determined by a
hierarchical likelihood ratio test (Goldman 1993; Huelsenbeck and Crandall 1997)
as implemented in ModelTest 3.7 (Posada and Crandall 1998). Parameters
estimated by ModelTest are indicated for each model.

 

Dataset

Model of
sequence evolution

Estimated parameters

 

ITS

Combined tmL-th
+ rpl32-tmL

Combined tmL-trnF
+ rpl32-trnL + ITS

 

 

TrN + l + I"
(T amura and Nei
1993; Yang 1994)

K81 uf + I‘
(Kimura 1981; Yang
1994)

TrN + l + I'
(Tamura and Nei
1993; Yang 1994)

 

Base= (0.2187 0.3121 0.2833)
Nst=6 Rmat= (1.0000 2.6190
1.0000 1.0000 5.4251)
Rates=gamma Shape=0.6195
Pinvar=0.3664

Base=(0.3771 0.1605 0.1354)
Nst=6 Rmat=(1.0000 1.6402
0.6516 0.6516 1.6402)
Rates=gamma Shape=0.5858
Pinvar=0

Base=(0.3267 0.2085 0.1800)
Nst=6 Rmat=(1.0000 2.1232
1.0000 1.0000 3.2821)
Rates=gamma Shape=0.7141
Pinvar=0.6149

 

Congruence and Hypothesis Testing

To test for congruence between ITS and combined plastid data partitions, an

' hoongruence Length Difference (ILD) Test of Farris et al. (1994) was conducted

a s implemented in PAUP as the Partition Homogeneity Test (Swofford 2002). For

18

 

the lLD analysis, a null hypothesis of congruence was used and the nuclear and
plastid datasets were compared using only parsimony informative characters and
1000 replicates (q = 0.05). A heuristic search was used with TBR branch
swapping, gaps coded as missing data, ten random additions per replicate,
saving no more than 100 trees per replicate.

To test alternative hypotheses of phylogeny, the combined ITS and plastid
dataset was analyzed in PAUP with an enforced constraint using the MP search
strategy outlined above. Trees constrained to one of two topologies (i.e., a
monophyletic Poliomintha [Constraint 1] and a monophyletic Poliomintha that
excluded P. conjunctrix [Constraint 2]), were prepared in MacClade 4.1
(Maddison and Maddison 2001 ). The most likely tree topologies recovered by the
unconstrained MP and ML analyses and the constrained MP analyses of the
combined dataset were compared using a Shimodaira-Hasegawa (SH) test
(Shimodaira and Hasegawa 1999; Shimodaira 2002), as implemented in PAUP
with RELL optimization and 1000 bootstrap replicates. A null hypothesis of no

difference was used and assessed at c = 0.05.

19

Chapter 3
RESULTS

Marker Choice

Two plastid regions, thUGA-trnfMCAU and nth-rpl32, failed to amplify

consistently for the surveyed taxa and were not used in the study. Of the
remaining regions surveyed, analyses of painrvise genetic distances among
sampled taxa suggested that rpl32-tmL was neariy twice as variable as tmD-tmT
and more phylogenetically informative (Table 3-1). Therefore, rpl32-tmL was

identified as the best additional plastid marker for the study.

Table 3-1. Characteristics and phylogenetic utility of sequences from two plastid
regions acquired from ten taxa in the Mentheae.

 

All accessions

 

 

Variable,
Parsimony parsimony Average pairwise
Total informative uninformative uncorrected genetic
Region Characters characters characters (p) distance
tmD-tmT 934 0 7 0.00157
[pl32-tmL 857 2 1 1 0.00364

 

Phylogenetic Analyses
Sequences from 83 accessions representing 57 taxa were included in the
phylogenetic analyses (Appendix A). Of the total, 71 accessions amplified and

sequenced successfully for all three regions. The remaining twelve accessions

20

successfully sequenced for both tmL-th and rp132-tmL regions of the plastid

genome, but failed to yield quality ITS sequences.

ITS Analyses of Poliomintha and Related Genera
The data matrix of aligned ITS sequences from 71 accessions included a total of
607 characters, of which 140 were parsimony-informative, 62 were variable but
parsimony-uninforrnative, and 405 were constant (Appendix B; Table 3-2).
Parsimony analysis of the ITS data resulted in 97,217 most parsimonious (MP)
trees of a length of 485 steps, consistency index (CI) of 0.518, retention index
(RI) of 0.722, and rescaled consistency index (RC) of 0.374. In the MP
topologies, the ingroup taxa were weakly supported by the bootstrap analysis as
a monophyletic clade (BS = 75%), with Acinos arvensis weakly supported as
sister to the remaining taxa (BS = 74 %; Figures 3-1 A and B). The monophyly of
Poliomintha was not supported by the ITS data. The type species, P. incana, was
positioned in the MP topologies as sister to the genus Rhododon, as part of a
weakly supported Poliomintha incana / Rhododon clade (BS = 76%). The
remaining Poliomintha species did not form a monophyletic clade in any of the
MP topologies recovered by the heuristic search. The species were distributed
among two clades in the strict consensus of MP trees: an unresolved clade with
P. conjunctn’x and three Hedeoma species (H. apiculata, H. drummondii, and H.
reverchonir); and an unresolved clade with P. Iongiflora as sister to a group that
il'1cluded P. bustamanta and P. dendritica. In the randomly selected MP tree, the

Mo clades were part of a polytomy with H. acinoides that was positioned in a

21

 

large clade referred to here as the “Hedeomintha” clade (for communication
purposes) that included most species from Hedeoma, but excluded the

Poliomintha incana / Rhododon clade (Figures 3-1 A and B).

Table 3—2. Sequence characteristics of regions used in phylogenetic analyses.

 

All accessions

 

 

Variable,
Parsimony parsimony
Included informative uninformative

Region characters characters characters
ITS 607 140 62
tmL-th 880 25 45
rpl32-tmL 91 9 45 94
Combined trnL-trnF + rpl32-tmL 1799 70 139
Combined trnL-trnF + rpl32-tmL + ITS 2406 2271 169

 

For nrne specres, where multiple accessrons had identical sequences, each specres was

represented by a single sequence in the phylogenetic analyses of plastid data matrix (see
Chapter 2). However, they were not combined in phylogenetic analyses of the combined plastid +
ITS data matrix. The increased number of phylogenetically informative characters reported here
for the combined plastid + ITS data is the result of pseudosynapomorphic characters that were
autapomorphic within each species in the analyses of plastid data.

Maximum likelihood (ML) analysis of ITS sequence data recovered one
tree with a negative log likelihood (-In L) of 3524.26454 (Figures 3-2 A and B).
The ingroup taxa were monophyletic, but poorly supported by the bootstrap
analysis (BS = 63%). The ingroup formed a large polytomy that included
Hedeoma pulegioides and two unsupported clades comprised of the remaining
taxa (Figures 3-1 and 3-2). The monophyly of Poliomintha was not supported in

the ML topology. Poliomintha species were distributed among the

22

“Hedeomintha” and Poliomintha incana / Rhododon clades with the same
membership described for the MP topologies. However, bootstrap support (BS =
73%) decreased slightly for the Poliomintha incana / Rhododon clade. While its
position was unresolved in the strict consensus of MP topologies, the
Poliomintha incana / Rhododon clade recovered by the ML analysis was
positioned within a polytomy that included taxa from genera such as Blephilia,
Conradina, Monarda, Pycnanthemum, and some taxa from Hedeoma and
Hesperozygis from eastern South America. This polytomy with the Poliomintha
incana / Rhododon clade was positioned as sister to the “Hedeomintha” clade,

but without bootstrap support (>50%).

23

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27

Plastid Analyses of Poliomintha and Related Genera
Plastid sequences (tmL-th and rpl32-tmL) from 50 accessions were aligned for
the combined plastid analyses, representing a total of 83 accessions from 57
taxa. The plastid data matrix included 1799 aligned characters, of which 70 were
parsimony-informative, 139 were variable but parsimony—uninfonnative, and 1590
were constant (Appendix B; Table 3-2). Parsimony analyses of the combined
plastid datasets recovered 88,000 trees with a length of 248, CI of 0.720, RI of
0.841, and RC of 0.605. In the strict consensus of MP trees (Figures 3-3 A and
B), a sister relationship between Acinos arvensis and a polytomy that included all
remaining ingroup taxa was strongly supported by the bootstrap analysis (BS =
98%). Poliomintha species were distributed among two clades. The first clade
(Clade l) was marginally supported (88 = 69%) and included P. incana and two
Clinopodium species from Baja California (i.e., C..chandlen’ and C. ganden). The
second clade (Clade II) was a weakly supported polytomy (38 = 61%) comprised
of P. bustamanta, P. conjunctrix, P. dendritica, P. glabrescens, P. Iongiflora,
Clinopodium hintoniorum, and eighteen Hedeoma species. While the relationship
between Clade l and Clade II and their position relative to other ingroup taxa was
unresolved in the strict consensus, the topology suggests that Poliomintha is not
monophyletic. However, poor bootstrap support in the topology limits confidence
in a formal rejection of monophyly.

Maximum likelihood (ML) analysis of the combined plastid datasets
recovered one tree with a negative log likelihood (-In L) of 4086.57963 (Figures

3-4 A and B). The ingroup taxa were strongly supported (BS = 97%) as a

28

monophyletic group, with Acinos arvensis positioned basally and strongly
supported (BS = 99%) as sister to the remaining taxa. The monophyly of
Poliomintha was not supported in the ML topology. As described above for the
parsimony results, Poliomintha species were distributed among the same two
clades (Clade I and Clade II) in the ML topology. Bootstrap support increased for
Clade l (BS = 74%), but slightly decreased for Clade ll (BS = 59 %). While
unresolved in the MP topologies, Clade II was positioned here as part of a larger,
unsupported (BS < 50%) polytomy that included Hedeoma hispida, a Rhododon
clade, and an unsupported (BS < 50%) polytomy that included species of
Blephilia, Clinopodium, Conradina, Dicerandra, Monarda, and Pycnanthemum.
The relationship of the above polytomy was unresolved relative to the remaining
ingroup taxa, which included Clade I, two Clinopodium species, two Hedeoma

species and four other clades of ingroup taxa.

29

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33

Combined Analyses of Poliomintha and Related Genera
The partition homogeneity test (PHT) of the combined ITS and plastid (trnL-th
and rpl32-tmL) datasets recovered a highly significant level of incongruence (P =
0.005). Visual comparison of trees recovered by parsimony and maximum
likelihood analyses of the individual ITS and plastid datasets revealed supported
topological differences for two taxa. The most important incongruence was
observed for the focal taxon, Poliomintha incana, which is the type of the genus.
Poliomintha incana was supported as sister to the genus Rhododon in the ITS
topologies (Poliomintha incana / Rhododon Clade; MP BS = 76%; ML BS =
73%), but was supported as sister to a clade that includes the two Clinopodium
species, C. chandlen' and C. ganden', in the plastid topologies (Clade l; MP BS =
69%; ML BS = 74%). The second topological difference was observed for
Conradina etonia, which was well supported as sister to Conradina brevifolia in
the ITS topologies (MP BS = 99%; ML BS = 95), but within a weakly supported
polytomy that included Clinopodium macrostemum and Hedeoma mandoniana in
the plastid topologies (MP BS = 71%; ML BS = 74%).

The combined nuclear (ITS) and plastid (tmL—th and rpl32-trnL) data
matrix for 71 accessions included 2406 characters, of which 227 were
. parsimony-informative, 169 were variable but parsimony-uninformative, and 2010
were constant (Appendix B; Table 3-2). Parsimony analyses of the combined
datasets recovered 43,966 trees with a length of 735 steps, CI of 0.553, Rl of
0.749, and RC of 0.414. The topology in the strict consensus of MP trees

strongly resembled that reported from the ITS data (Figures 3—5 A and B; Figures

3-1 A and B). The monophyly of the ingroup was strongly supported in the MP
trees (BS = 96%), with Acinos arvensis positioned basally as sister to the
remaining ingroup taxa (BS = 98%). Poliomintha was not supported as a
monophyletic group. Species of Poliomintha were distributed among the
“Hedeomintha” and Poliomintha incana / Rhododon clades described for the ITS
topologies. However, bootstrap support for the Poliomintha incana / Rhododon
clade was higher for the combined data (BS = 83%) than it was for the ITS data
alone. The remaining Poliomintha species did not form a monophyletic group
within the “Hedeomintha” clade, but were present as part of two subclades within
all MP topologies. However, increased resolution and bootstrap support (>50%)
among clades with Poliomintha in the “Hedeomintha” clade was not improved
with the addition of plastid data.

The ML analysis of the combined nuclear and plastid data recovered one
tree with a negative log likelihood (-In L) of 7951.01811 (Figures 3-6 A and B). As
described above, the ingroup taxa were supported as a monophyletic group, but
bootstrap values decreased slightly in comparison .to the MP results (ML BS =
91%; MP BS = 96%). Acinos arvensis was positioned basally and strongly
supported as sister to the remaining ingroup taxa (BS = 91%). Consistent with
previous results, the monophyly of Poliomintha was not supported in the ML
topology. Poliomintha species were distributed among the Poliomintha incana /
Rhododon and “Hedeomintha” clades as previously described. However, support
for the Poliomintha incana / Rhododon clade decreased in the ML topology (ML

BS = 75%; MP BS = 83%). The position of the Poliomintha incana / Rhododon

35

clade was resolved as sister to another clade that included taxa from genera
such as Blephilia, Conradina, Monarda, Pycnanthemum, and some taxa from
Hedeoma and Hesperozygis from eastern South America. As for the
“Hedeomintha” clade, it was positioned within a polytomy that included
Clinopodium hintoniorum and Hedeoma jucunda. The addition of the two taxa as
part of a larger group that included the “Hedeomintha” clade was consistent with

relationships observed in ML plastid topology.

36

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Tests of Alternative Phylogeny

Results from the SH tests supported the conclusion that Poliomintha is not
monophyletic, regardless of whether or not P. conjunctn'x was included.
Parsimony searches of the combined nuclear and plastid datasets constrained to
one of two topologies recovered a total of 232 trees with a monophyletic
Poliomintha (Constraint 1) and 72 trees with a monophyletic Poliomintha that
excluded P. conjunctrix (Constraint 2). Based on the TrN + l + l' model of
sequence evolution for the combined data, the most likely tree recovered under
Constraint 1 had a negative log likelihood (-In L) of 7997.70634, length of 745
steps, CI of 0.543, RI of 0.739, and RC of 0.401. The most likely tree recovered
under Constraint 2 had a -In L of 7991 .02576, length of 743 steps, CI of 0.545, Rl
of 0.741, and RC of 0.404. The SH tests comparing the mostly likely constraint
trees with the most likely unconstrained MP tree (-ln L = 795602084) and the ML
tree (-In L = 795101811) detected highly significant differences in likelihoods for
all pairwise comparisons (Table 3-2). Not reported in Table 3-3 are the results of
an SH test comparing the most likely MP tree and the ML tree, of which
significant likelihood differences were not detected among the two topologies at

the a = 0.05 level (P = 0.382; difference in —In L = 5.00273).

41

Table 3-3: Results of SH tests. Numbers indicate P value / difference in negative
log likelihood (-In L) score. Significance at the a = 0.05 level is indicated with an

asterisk.

 

MP no constraint:

Most likely tree
(-In L = 7956.02084)

ML no constraint:

Most likely tree
(-In L = 7951.01811)

 

MP: Constraint 1
Poliomintha monophyly

 

Most likely tree
(-In L = 7991 .02576)

 

 

 

. 0009* / 41.68551 0004* / 46.68824
Most lrkely tree

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MP: Constraint 2

Poliomintha monophyly

exc'“d'"9 P‘ °°”’“"°t”x 0018* / 35.00492 0012* /40.00765

 

42

 

Chapter 4
DISCUSSION

The phylogenetic hypotheses presented herein, inferred from over 2,400 base
pairs of nuclear and chloroplast DNA sequence data, resurrect a number of
systematic issues pertinent to Poliomintha previously discussed by some
researchers and offer much needed insight into long-standing taxonomic
questions and debate over the generic boundaries among Poliomintha and
closely related genera in the tribe Mentheae. Several important findings from the
study and their implications are discussed here, including: (1) phylogenetic
relationships among Poliomintha and related genera; (2) the importance of
morphological characters in delineating genera; and (3) the taxonomy of

Poliomintha.

Phylogenetic Relationships among Poliomintha and Related Genera

The phylogenetic hypotheses consistently suggest that Poliomintha is not
monophyletic. As currently circumscribed, Poliomintha represents at least two
distinct lineages; a lineage that includes the type species, P. incana, and either
(or both) Rhododon or two species of the polyphyletic genus Clinopodium; and a
second lineage that includes the remaining species of Poliomintha and most
species of Hedeoma (e.g., the “Hedeomintha” clade). These results are, in part,
consistent with the findings of Wagstaff et al. (1995), who first reported a sister

relationship between Poliomintha (P. Iongiflora) and Hedeoma (H. drummondii)

43

based on a parsimony analysis of chNA restriction site variation in subfamily
Nepetoideae. The results of my study consistently reveal a close relationship
between most species of Poliomintha and Hedeoma. However, no evidence is
available to suggest that the type species of either genus (P. incana and H.
pulegioides) is included in the same clade as the majority of the species from the
two genera. In fact, the results of the constraint analyses and SH tests suggest
that P. incana is not related to the remaining Poliomintha species, representing a
new and important finding that has taxonomic implications.

Phylogenetic relationships among genera in the Mentheae are difficult to
interpret with confidence from the results presented here for several reasons.
First, insufficient resolution and low bootstrap support within and among most
clades imposes substantial limits on inference of evolutionary relationships
among sampled taxa, especially at the generic level. It is possible that the lack of
resolution and support is a consequence of either incomplete sampling of taxa or
insufficient sequence data, or both (Graybeal 1998). As suggested by my results,
several genera included the study may be polyphyletic (Figures 3-1 to 3—6)——e.g.,
Clinopodium, Hedeoma, Hesperozygis, and Poliomintha—and more extensive
sampling of taxa within these genera as well as from other genera in the
Mentheae such as Acanthomintha, Bystropogon, Cunila, Dicerandra, Eriothymus,
Glechon, Minthostachys, Piloblephis, Pogogyne, Rhabdocaulon, and
Stachydeoma may help to better resolve relationships. As for character sampling,
additional work is needed to identify informative nuclear and plastid markers. The

plastid regions used in this study exhibit low levels of sequence divergence and

contribute less than one-third of the total phylogenetically informative characters
to the analyses (Table 3-1). As a result, insufficient resolution is obtained for
many groups from analyses of plastid data alone (Figures 3-3 A and B; Figures
3—4 A and B) or in combination with lTS data (Figures 3-5 A and B; Figures 3-6 A
and B).

The incongruence among the ITS and plastid datasets represents another
challenge to phylogenetic inference of relationships in the Mentheae. The PHT
has been demonstrated to be overly sensitive in large datasets such as that used
here (Hipp et al. 2004). Therefore, it is possible that the incongruence suggested
by the PHT does not represent phylogenetic differences, but instead is an artifact
of the overly sensitive nature of the test. In this case, it appears that the
incongruence is widespread and involves more than one taxon. The topological
differences observed among individual analyses diminish confidence in some
relationships, particularly for those involving the focal taxon (P. incana). For
example, relationships between P. incana and either Rhododon or species of
Clinopodium are weakly supported in the phylogenetic analyses of ITS and
plastid datasets, as well as in those using total evidence.

Casesof widespread incongruence among ITS and plastid datasets have
been documented for other groups in the Mentheae such as Conradina,
Clinopodium, and related scrub mints from the southeastern United States
(Edwards et al. 2006; Edwards et al. 2008). In fact, incongruence among different
gene trees is a commonly encountered problem in phylogenetic studies,

particularly in those involving recently derived and rapidly radiating species

45

(Buckley et al. 2006). In examples of incongruence between nuclear and plastid
data, the patterns of plastid sequence variation may be explained by processes
such as introgression or shared ancestral polymorphism lineage sorting (Wendel
and Doyle 1998; Schaal and Olsen 2000). However, distinguishing between
processes responsible for the patterns is often difficult because of the large
variance associated with the coalescent process and the fact that the processes
can produce similar phylogenetic patterns (Holder et al. 2001). For some groups
in the Mentheae, it may be difficult to reconstruct phylogenetic relationships using
DNA sequence data, particularly in cases where taxa are recently derived,
hybridization is widespread, or where there is a lack of coalescence (Edwards et

al. 2008).

Importance of Morphological Characters in Delineating Genera

For over 100 years systematists have been uncertain about the generic
circumscription of Poliomintha. lts similar morphology and close relationship to
genera such as Hedeoma and Hesperozygis have been discussed by several
researchers, some of whom have questioned its distinction at the generic level.
Briquet (1897) first discussed in-depth the inherent complexity of morphological
interpretation for these genera, emphasizing the fact that some taxa exhibit
character states that appear “intermediate” between those traditionally used to
delineate generic boundaries. It seems that Briquet’s decision to subsume
Poliomintha into Hedeoma on the basis of insufficient clear-cut differences was,

as Epling and Stewart (1939) effectively point out, inappropriate within the

46

context of a larger Mentheae and would necessitate the inclusion of other genera
beyond the scope of those discussed here. Epling & Stewart (1939) argued that
“the fact that no single constant point of difference may be brought forward
wherewith to distinguish Hedeoma and Poliomintha is not presumptive of their
unity.” Characters such as stamen number (two versus four), habit (herbaceous
versus woody), calyx morphology (gibbous, saccate, campanulate, or bilabiate
versus tubular and symmetrical), and the presence or absence of a well-defined
or irregular calyx annulus were defined by Epling and Stewart (1939) and Irving
(1972, 1980) for the purposes of delineating boundaries among Hedeoma,
Hesperozygis, and Poliomintha. However, in light of my phylogenetic results, it
appears that the taxonomic importance of at least some of the characters used
by these researchers has been overestimated.

Characters such as habit and calyx morphology do not represent
synapomorphies that characterize monophyletic groupings of taxa, but instead
are homoplasious within the phylogenetic context presented here. The habit, in
particular, is not an ideal diagnostic character by which to distinguish Poliomintha
from Hedeoma. Within the monophyletic “Hedeomintha” clade, one can find
numerous examples of Hedeoma species that exhibit woody or semi-woody
habits. Moreover, species with annual and perennial herbaceous habits as well
as species with woody or semi-woody habits can be found in other monophyletic
genera in the Mentheae—for example, in Monarda (Prather et al. 2002). Without
sufficient resolution and bootstrap support within and among most clades, it

seems premature to undertake a comprehensive morphological study to uncover

47

synapomorphies. However, character analyses may be helpful in determining
what characters best distinguish monophyletic clades, pending the acquisition of

more conclusive phylogenetic evidence.

The Taxonomy of Poliomintha
The phylogenetic evidence from this study consistently suggests that Poliomintha
is not monophyletic as currently circumscribed. In consideration of these results,
the genus Poliomintha may be best limited to the type species, P. incana,
pending future phylogenetic and morphological studies that determine the
relationships of P. incana with confidence. As for the remaining Poliomintha
species, their taxonomic disposition depends on the relationships among the
species of Hedeoma. Based on my results, Hedeoma is polyphyletic and the type
of Hedeoma, H. pulegioides, is most likely unrelated to the remainder of the
genus. However, more evidence is needed to confirm this result. If further
phylogenetic evidence suggests that H. pulegioides is a member of the
“Hedeomintha” clade, the Poliomintha species that comprise the clade are best
subsumed into the genus Hedeoma to reflect their common ancestry.
Altematively, if H. pulegioides is excluded from the “Hedeomintha” clade, the
Poliomintha and Hedeoma species comprising the clade should be reclassified
into a new genus.

Many genera in family Lamiaceae are not monophyletic as currently
circumscribed. In consideration of molecular synapomorphies, many genera are

polyphyletic or paraphyletic relative to other genera, including: Satureja L.

48

(Wagstaff et al. 1995); Clerodendrum (Steane et al. 1999); Salvia L. (Walker and
Systma 2004); Huxleya (Steane et al. 2004); Basilicum Moench, Endostemon N.
E. Br., Fuerstia T. C. E. Fr., Haumaniastrum P. A. Duvign and Plancke,
Hemizygia Briq., Hos/undia Vahl, Hyptis Jacq., Platostoma P. Beauv.,
Plectranthus L. Hér., Puntia Hedge, Ocimum L., Orthosiphon Benth., and
Syncolostemon E. Mey. (Paton et al. 2004); and Micromeria Benth. (Brauchler et
al. 2005). Many of these genera, including Poliomintha, Hedeoma, and
Clinopodium, will likely necessitate (or have been subject to) taxonomic and
nomenclatural revision. Although no nomenclatural changes are recommended
based on my results, any future taxonomic revisions that involve nomenclatural
changes for Poliomintha or other genera should carefully consider the possibility

of broader societal impacts.

49

APPENDICES

50

APPENDIX A

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60

APPENDIX B

61

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HOOPH .HU..¢...00000.0<..............H..04~III.U.......0.0 m MHCWEQHWZQ QCUCHEOHNOQ
HOOK; .BU..¢....UUUUU.U¢...0.....0....H00mudllloU.......mu._U 1N WHCQEMVUWDQ MCUQflEOHNOnN
HOOFH ..mv..<...UUDUU.U¢.............B...UIII.U........WU

00440044 4440444434

95

Hoth
Homba
Homha
Homba
momha
Hoth
Homhg
flomba
fiomba
Homha
fiomnfi
Homba
Homha
womb.
Homhw
Hombfi
Hom50
Hom50
Homba
HombH
momma
40050
Homhw
Homwa
Homha
HombH
HombH
Homba
Hom90
Hom50
Hom50

990000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
111111111111111111111 040000400111111100....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 11111111 0....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
11111111111111111 4090040000400111111100....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00.....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
190000049940000004090040000400 1111111 00....
I90000049940000004090040000400 1111111 00....

mmUHOHQmNDQ
mmbwowomwsa
0004040040Q
0004040043Q
muflthHQ
mefiequ
m0m044Q

m HemEHmQ

4 wemEHmQ
m440440mmeo
m meme

4 meme
memueoE
04408

m m4me

4 mwbmfi

m memweOUemE
4 mem4e00emE
mbezozm
44e00mee0h
440e4>44
0040040

0 44erEEzen
4 HwbeoEEzeb
m mumumoo

4 m0m0000
mumaoflwwo
m0m4304Qm
mmUHOeHUm

411300

mEombmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomnmm
mEombwm
mEombmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEombmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm
mEomnmm
mEomUmm
mEomUmm
mEomUmm
mEombmm
mEomUmm
mEomUmm

m4e00m E3400Q0e440

m440440044 E2540QQQ~FM

96

momma
HomNH
momha
Momba
flom90
flomPH
Hom0_
momma
Hommg
Homha
Hemp“
Hom5_
momma
Hommg
Hombu
mom9H
Hom9H
Homhg
momma
Hom50
fiomba
Homba
40050
homba
momma
Hom50
Homnw

3E .............................
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII DU.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000040940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00 .....
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.
190000049940000004090040000400 1111111 00.

0440440
mefleu

m 0:0m4

e00oe me0e02
HummE szxe5

440 00000004

0 mzumwsoem eouonoem
4 mzumwzoem eOUOerm
EzoweeomHamo EzEmeuemeUAM

m mHOHMHDeOH
4 mHONMHQeO~
0 memuefl
m memoew
4 memoew
m0fluflenemb
x4400e59000
0 muemEmumse
m muemEmusz
4 muemEmumBQ
mmbwoefl
mwaowmme
mmOHD
m0m40e0QO
eononoee
mnwuwe
m mwwomflemfi
4 mNHOMHHmE
HHeoeoe
0 mHN
m mNH
4 mam
0 0004040

meuewEOHHom
meueHEowwom
me0e4804404
me0e4E04404
me0e4E04404
meue4E04404
mnuC4EO4aom
me0e4504404
meue4E04404
me0e4E04404
4 mfiwmwemeoz
0emE mnemeoz
0044 mnemeoz
mwuxmoeQOmm
mfluxmoemammm
mwuxwoeQOmm
mfloxmoememm
040%N040Q004
m>04 mEomUmm
Hmsm mEombmm
wmnm mEombmm
Hmzm mEomUmm

0440. 2003

97

”@550 11111111111111111111111111111

.0550 1111111111 4544054540044555040
40550 11111111111111111111111111111
40550 11111111111111 054540044555040
40550 11111111111111 054540044555040
40550 1111111111111111 4540044555040
40550 11111111111111 054540044555040
40550 111111111111111 54540044555040
40550 1111111111 4544054540044555040
40550 1111111111 4544054540044555040
.0550 40040000144544054540044555040
40550 11111111111111 054540044555040
40550 111111 00144544054540044555040
40550 11111111111 544054540044555040
40550 11111111111111 054540044555040
40550 1111111111111 4054540044555040
40550 11111111111111 054540044555040
40550 11111111111111111111111111111
40550 1111111111111 4054540044555040
_m550 11111111111111 054540044555040
40550 11111111111111 054540044555040
_m550 1111111111111 4054540044555040
00550 11111111111111 054540044555040
.0550 11111111111111111111111111111
00550 11111111111 544054540044555040
40550 11111111111111111111111111111
40550 11111111111 544054540044555040
40550 11111111111111111111111111111
.0550 11111111111111111111111111111
40550 111111 00444544054540044555040

mem0e0E
0440E

m mHUmE

4 m4me

0 memweOUemE
4 memHeOUemE
mbezuzm
44e00mee0h
44004044
004e044

m 440eoEEzeU
4 wwbeoEEseb
m mumumoo

4 m0m0000
m0m404440
mumazoflQm
mmbwoeHUm

mEomUmm
mEombmm
mEombmm
mEomUmm
mEombmm
mEomUmm
mEomUmm
mEombmm
mEomUmm
mEomUmm
mEombmm
mEomwmm
mEombwm
mEomUmm
mEombmm
mEombmm
mEomUmm

m4e00m E0400Q0e4~0
m44054>0ee E0400Q0e440
440E4mQ E0400Q0e440
Esemuwme E0400Q0e440
E0E000040mE E3400Q0e4~0
Ezeofle00e4e E0400Q0e440
m 4embem0 E0400Q0e440
m Hemabemeu E0400Q0e4~0
4 440~Ueme0 E0400Q0e440
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4 40:304Q EszoQ0e440
0000444 044400044

04020044 002404
4

98

 

”mung
2055a
Horne
Hm55_
morn“
fimhba
hombw
20550
Hm55~
Hanna
20550
Hanna
40550
50550
Hanna
”mph“
2055_
”@550
20550
”mung
Hanna
.0550
2055a
mmbha
Hmbna
20550
ambba
fimbha
Hanna
20550
4055_

Iu140000:44544054540044555040
lllllll 0:44544054540044555040
IIIIIIIIII 4544054540044555040
uuuuuuuuuu 4544054540044555040
uuuuuuuuuuuuuuuuu 5400:4555040

nnnnnnnnnnnnnn 054540044555040
uuuuuuuuuuuu 44054540044555040
IIIIIIIIII 45:4094540044555040
nnnnnnnnnnnnn 4054540044555040
uuuuuuuuuuuuuuuuuuu 0044555040
uuuuuuuuuuuuuu 054540044555040
uuuuuuuuuuuuuu 054540044555040
......... 44544054540044555040
IIIIIIIIIII 5:4054540044555040

III4UOUUI44H44UH4B4UQ4<HHBO4D.

4wD4DDUOI44H44UB4H4UD44HHBU4D
IIIIIIIIII 4BI4UH4940044HBBU40
IIIIIIIIII 4H44UB4H4UQ<4HHHD4w
IIIIIIIIII 4944094540044899040
IIIIIIIIII 4B44UB4H4UO44HBHU4Q
IIIIIIIIIIIIIIIIIIIIIII HBBO4U
IIIIIII 0144944084940044998040
IIIIIIIIIIIIII UB4H4UQ<4BBBU4O
IIIIIIIIII 4H44UH4H4UU44HHHU4G

4 020024 02024504402
0040440200 02024504402
44420224200 02224504404

0 0020500004 02024504402
2 0020500004 02024504402

4 0020500004

0004:200Q0
20000024

2 044054405
4 044054405

<EIDUCJLIJ

02024504402

00040244 0440040202

000400045

044050020205 0040202

0040202

04024040Q002
04024040Qm02

004042 04024040Q002

44202040>04
0 04440:2

m 0444022

4 0444032
0004040043Q
0004040040Q
0004040040Q
0004040040Q
00040400402
00440Q4Q
024400Q
0000445

2 440540Q

4 440540Q
04404400420
2 0202

4 0202

040240402002
04024040Q002

0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002
0500002

99

 

0m440 uuuuuuuuuuuuuu 054540044555040
00550 -0040000-44514054540044555040
00540 ............. 4054540044555040
04450 uuuuuuuuuuuuuuuuuuuuuuuuuuuuu
04450 uuuuuuuuuuuuuu 054540044555040
04540 ............ 44054540044555040
04550 ............... 54540044555040

0m440 uuuuuuuuu 544054540044555040

2 00004440 20000022

0 000040020 20000022

4 000040020 20000022
500424054400 505020202022

2 0404540204
4 0404240204
0 020024
2 020024

02024504402
02024504402
02024504402
02024504402

100

Homg
2000
HomH
2000
2000
HomH
HomH
HomH
HomH
Homg
2000
Hom_
2000
2000
00mg
2000
HomH
HomH
Hom_

Homa
20mg
0000
20mg
20mg
20mg
HomH

.oooooBo llllllllllllllllllllllllllllllllllllllllll
. . . . . .H.4H_HU44400 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII
oooooorfio< IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII
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......H.4890444008480088UUU4UHB4UOIH44BHU4OUU4BUOU
ooooooHo<H§HU¢ llllllllllllllllllllllllllllllllllll

40504400 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII

4 0204200205 0500002
0020004 0500002
4420002204 0500002
44024044 0500002
044044000022 0500002
0042042 0500002
000404440 0500002

4 00040

000402400 0500002
0004000554 0402040042

042000
0440540042
0202500040

4405400
502004405
50500004005
0 4400200

4 4400200

2 440402020

50400002440
50400002440
50400002440
50400002440
50400Q0244U
50400002440
50400002440
50400002440
50400002440

5040000400440 .400

4023042
4 4023042
0 40200
4 40200

50400002440
50400002440
50400002440
50400002440

0004445 044442040
04020>40 002404
044044020004 020202
0242040005 005225

0:50:69 “mm—QOhO—P—U 9.: *0 ..00an “Chum-4:5 USN €0.59: 125% "N 5.50: ¢0C¢300w U¢=D=<

101

 

HOOHH
20040
20040
20040
20040
20040

2000
2000
2000
20mg
2000
2000
2000
20mg
20m“
20mg
20mg
Home
How“
20mg
Homa
20mg
20mg
20mg
20mg
HomH
2002
20mg
HomH

0544000UH44444H44HH44OOUUUU444U4U4UHHI444UBHHUI44H

O

0 40200 50400002440
4 40200 50400002440
.0044442 044442040
04020040 002404
044044020004 020202
0242040005 005225

2 00004440 20000022

4 000040020 20000022
500424044400 505020202022
2 020024 02024504402

4 020024 02024504402
0040440200 02024504402

4 0020500002 02024504402
00040244 0440040202

044040020205
000400044

0040202
0040202

0004020020 040200402002
20000024 040200402002
004042 040240402002

2 044044405 040240402002

2 0444002

4 0444002

2 00040400402
00440242

4 4405402

4 0202
0200205

4 04005

0500002
0500002
0500002
0500002
0500002
0500002
0500002

44 00040

0500002

102

 

20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040

20040

 

2 0444002 0500002

4 0444002 0500002

0 00040400402 0500002
00440242 0500002

4 4405402 0500002

4 0202 0500002
0200205 0500002

44 00040

4 04005 0500002

4 0204200205 0500002
0020000 0500002
4420002204 0500002
44024044 0500002
044044200022 0500002
0042042 0500002
000404440 0500002

4 00040

000402400 0500002
0004000554 0402040042

042000
0440440042
0202500040

4405402
502004405
50500004005
2 4400200

4 4400200

0 440402020

50400202440
50400202440
50400002440
50400202440
50400202440
50400202440
50400202440
50400202440
50400202440

5040000400442 .40>

4020042
4 4022042

50400202440
50400202449

103

HomHH
20040
HomHH
Hom4_
20020
20040
HomHH

Homaa
HomHH
20040
Hom4_
HomHH
HomHH
HomHH

HOOHH
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
20040
HOOHH

4440999..
4440999..
4440999..
4440999..
4440999..
4440999..
4440999..

4440999..
4440999..
4440099..
4440999..
4440999..

4440999.....

IIIIIII 004440:InInIII:II44440909999090094440004090

E—‘HBE—‘HHB

°HBEB

....0...............

4405402
502004205
50500004005
2 4400200

50400202440

50400202440

50400202440

50400202440

4 4400200 50400202440

2 440402020 50400202440
5040000400442 .40>

4022042 50400202440

4 4022042 50400202440

2 40200 50400002440

4 40200 50400002440

0020242 044222040

04020040 002404

044044020004 020202

0242040005 005224

2 00004440 20000022
4 000040020 20000022
500424024400 505020202020
2 020024 02024504400

4 020024 02024504402
0040440200 02024504402

4 0020500002 02024504402
00040244 0440040202
044020020205 0040202
000400044 0040202
0004020020 040200402002
20000024 040200402002
004042 040200402002

2 044024405 040200402002

104

 

105

Hom40 4440999...... IIIIIIIIIII ....9..................... 0040440200 02024504402
HOMHH— «gUHHn—u.....o llllll igH....B........o............ 4 QHCQEMUWSQ QCHCHEOflNOQ
HomHH 4440999...... IIIIIIIIIII ....9..................... 00240244 2440242202
HomHH 4440999...... IIIIIIIIIII ....9..................... 244042220225 2242202
HomHH 4440999...... IIIIIIIIIII ....9..................... 200400244 2242202
Hom40 4440999...... IIIIIIIIIII ....9..................... 0004020020 040200402002
Hom4_ 4440999...... IIIIIIIIIII ....9..................... 20000024 040200402002
momHH— «@EUHHH...... IIIIIIIIIII ...orH...................... QUHUH: WmefiNOngmwm
Hom40 4440999...... IIIIIIIIIII ....9..................... 2 044044405 040200402002
Hom4. 4440499...... lllllllllll .. IIIIIIIII ............... 0 0444002 0500002
HODHH EUHHH...... lllllllllll ...oH...................... flmflHHWDQMEOQUQm
HomHH gfiUHHB...... lllllllllll .....H..................... mmmnflOflmmfifiQ QEO®U®44
HODHH— g4UBHB...... lllllllllll .....H..................... MMHHmfiwfiQmEOQUmm
HomHH <gUHBH...... lllllllllll .....H...................... <HH®ENQQQEO®U®W~
momHH 4440999...... IIIIIIIIIII ....9..................... 4 2222 2502222
HomHH 4440999...... IIIIIIIIIII ....9..................... 2220205 2502222
20040 4440499.. IIIIIIIIIII . ..4. 44 00040
20040 4440944.. IIIIIIIIIII . ..9. 4 04005 0500002
Hom4u 4440999...... IIIIIIIIIII ....9..................... 4 2224202225 2500202
HomHH 4440999...... IIIIIIIIIII ....9..................... 2223022 2502202
HOmHH <§UH=HH...... lllllllllll ...oH..................o... HNCOHWCCOW msomnmm
HomHH 4440999...... IIIIIIIIIII ....9..................... 44D24>44 2500222
HODHH <¢¢UHHH...... lllllllllll ...ofi..........L.......... WHNONHQOmeg QEOQUmm
HomHH <44U999...... IIIIIIIIIII ....9..................... 224Q242 2500222
HomHH 4440999...... IIIIIIIIIII ....9..................... 202404440 2502202
HomHH <440999...... IIIIIIIIIII ....9..................... H 22040
HomHH 4440999...... IIIIIIIIIII ....9..................... 202402402 2500222
HomHH 4440999...... IIIIIIIIIII ....9..................... 2024502554 242224204Q
HOWHH— EUBfifiooo... lllllllllll .....H..................... mflqoumfisflboqocufiflb
Hom40 4440999...... IIIIIIIIIII ....9..................... 0440440042 50400202440
Hom40 4440999...... IIIIIIIIIII ....9..................... 0202500042 50400002440

 

HQONH
”coma
Hooma
floomg
”coma
HOONH
HOONH
Hoomg
200mg
HOONH
moomg
HOONH
HOONH
Hooma
HoomH
200mg

”coma
HOONH
HOONH
200mg
”CONE
Hooma
200mg

HomHH
20042
Hom42
Hom4H
HomHH

o ooooooooooooooooooooooooooooooo wD/NIIIMNggmmUUIfimN

.004III44444400I444
.004III44444400I444
.004III444444001444
.004III44444400I444
.004III44444400I444
.004III44444400I444

.004III44444400I444.

.004III44444400I444
.004III44444400I444
.00411144444400I244
.004III44444400I444
.00444444444400I444
.00444444444400I444
.004III444444001444
.004III444444001444

.004III44444400I444
.004III42444400I444

..004III444444001444

.004III44444400I444

oooooooooooooooooooooooooooooooo DO¢I lggwolg

.004II444444400I444

49099090044004440904040400400444 uuuuuuuuuuuuuuuuuu

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¢§UHHH...... lllllllllll .
gUHBH...... lllllllllll .
§<UB.H:H....... IIIIIIIIIII .
g4UBHH...... lllllllllll .

044044200042 0500002
0042042 0500002
000404440.0500002

4 2:040

000402400 0500002
0404300554 0402040042
042040 53400202440
044044>042 53400202440
0202830042 83400202440
4405402 53400202440
532004205 53400202440
53504004005 53400202440
0 4400200 53400202440

4 4400200 53400202440

0 440402020 53400202440

5343003400442 .40>
4022042 53400202440
4 4023042 53400202440
0 40200 53400202440
4 40200 23400202440
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0242044005 035424

0 03004440 20000022

4 034043020 20000022
530424044400 535024202042
0 020024 02424504402

4 020024 02424504402

106

HomNH
Hommg

Hooma
Hoomg
MOON“
”coma
HOONH
HOONH
HOONH
HQONH
meow“
HOONH
HQONH
HOQNH
HOONH
80$
HOONH
803
303
HQONH
Hooma
Hooma
Hooma
80m.
HOONH
HOONH
”coma
HOONH
Hoomw

.........D...................U....................

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.......................... . . . . . .Owdlllggwnvlflg
.............................. ..UU¢III¢¢¢¢¢¢UUI¢¢¢
................................ OU<III<¢<¢¢QOUI¢¢¢
............................... .wwfllllggwolg
............. ............ .......UU¢III<<¢¢¢<OUI<¢¢
................................ .00an I lflggw U<§¢
................................ UU¢III<<<¢<¢UUI<<¢
. ........... . ...... . ..... . . . . . . .Uwfilllggmkulg
................................ UD<III¢4¢¢¢¢UUI¢4¢
................................ OU<III<<¢<<<OUI<¢¢
........................OU<III¢<<¢¢<DUI<<¢
... ........................... ..GU¢III¢¢¢<¢<DUI¢¢¢
. . . . . ...................... . . o . .Uwafllllgnglg
.... ....... ........... ..... .....UO<III¢<<<¢¢UUI¢¢<
.. .......... ....................UU<III<¢¢¢<¢UUI¢¢¢

mHNOManzu0H maucmz
mewauflummE m3E%c&

m mzumwfiwu cowonocm

< magmasucm COUOU0£m
Ezbwcuowwmmu Essw£ucmcoxm
m mcmucw mQHCHEOHNOQ

m memocw mauCHEOHNOQ
mowuwubcmb maucwEoflmom

¢ mucmfimumzn mauQHEOHHom
mmnwocwa mawmnnmcoz
mwaowmmaucmfi mnumcoz
mmoazumflw MUHMQOE
wumwzaummm mwmxmoumammm
COUOanu mekmoqummm
mnfluwc mHDANOHQOmm

m mHHOMmeE mwoxwoummmmm
m wwwwmzm mEomUmm

a maaflmsQ mEombmm

m mmwaflUmwsm mEomUmm
wuHMmQHQ mEomUmm

¢ HMmEHmQ mEomUmm

é mum: mEomUmm

mcmucoE mEombmm

HH onuo

a mHUmE mEombmm

¢ mqmwzoncmfi mEomUmm
mbczozh mEomUmm
NHQOumCCOh mEombmm
HHQQH>MN mEowbmm

107

 

HomNH
momma
HomNH
Hommg
Hommg
momma
momma
momma
HomNH
momma
momma
HomNH
Hommg
momma
momma
momma
momma
momma
momma
momma
Hommg
fiomNH
momma
momNH
Hommg

Hommg
HomNH
Hommg
Hommg
Homma

DUOUOOOUQOOOUUUOU(30000000

000L190

UUUUUUUUUUUUUUUUUUUUU

UUUUU

.d flMmEHmQ mEombmm
d meme mEomUmm
mcmuCOE mEomUmm

HH msouo

m mflUmE mEombmm
< mcmHQOUcmE mEomUmm

mbczuzh
HHCOumccoh
HHQQH>MH
MHHOMHQOmmx£
mnHQch
mum~oH~wU

mmbwocwum

mEomUmm
mEombmm
mEomUmm
mEombmm
mEombmm
mEomUmm
H msouw
mEomUmm

mumwzumfifiw muncmnmbflq

MHQOum
meOMw>mMQ
mcwnasooum

HNmemQ
EzemumeE
EzEmumOMumE
m Humbcmo

4 Humbcmu

m wumwbcmcu

ESHUOQOCHNU
E2HU0QOCH~U
ESHUOQOCHHU
EzfiUoroflwb
ESHUOQOCHHU
EzflonQCflab
ESHUOQOCHHU
EzflonOCHHU
EszOQOCHNU

EzazumSHmoHHQ .Mm>

Hmczoun
< Hmczoha
m Hmcmm
< ngmm

Ezflbomocfiab
Eswbomocflwb
EsflUOQOCHHU
Esflnoaocaau

musmuflc mflHHchNm
mwmcm>um mocwom

108

 

Hoomg
Hooma
”coma

Hoomu
Hoomg
Hoomg
”coma
Hoomg
Hoomu
HoomH

momma
momma
fiomNH
Homma
HomNH
Hommg
momma
Hommg
Hommu
momma
aomNH
Homma
HomNH
Homma
momma
momma
momma
HomNH

B¢B0.. ... ......0. ... .. IIIIIIIIIII
H<H0.. ... ......0. ... .. IIIIIIIIIII
H¢H<.. ... ........ ... .. IIIIIIIIIII
IIII.. ... ........ ... .0 IIIIIIIIIII
IIII.. ... ..0..... ... .. IIIIII <HHHH
IIIIH0¢B<¢0BHdH09¢000¢<¢H<0¢<¢0B<00¢<¢H IIIIIIIIIII
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. 0....... .0. .............
...... .0. 0....... .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ ... .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............
...... .0. ........ .0. .............

¢ Hambcmu Ezwnoaoswwu
m Nnmabcmnu Ezflonocwao
Esazumzwmowwm .Mm>
Hmcgoun annouocwwo
¢ quzohfl Eswonocwab
m Hmnmm ESHUOQOCHHU
m Hmnmm EDHUOQOCHHU
musmuflc mfiaflcmmfim
mwmcm>nm mocflom
MNNOMwbczuou M£ucw2
mcwaowumms msExch

m msumwwflo COUOUocm

d msumqsocm sononoam
ESUHQHOMHNMU EzEmcucmzoxm
m mcmocw maucwsoflwom

a mzmucw maucflEoflwom
wwwuwubcmb maucwEowNom

4 mucmEmumza MSHCHEOHHOm
mmUHOCHH wwwmbumcoz
mwaowmmcucme mbumeox
mmo~sumflw mbumcofi
mumwscummm mflmxmoqummm
COUOUOQH mwbxwoummmmm
Mbwuwc mwoxmoummmmm

m mflwowHHmE mfluxmonmmmmm
.m mawwmzm mEomUmm

« mwawmzm mEomUmm

m mmbonmmH3Q mEomUmm
MUflMmQHQ mEomUmm

109

Hoomg
Hooma
”coma
HOOMH
”coma
Hooma
moomg
Hooma
Hooma
Hoomg
”coma
Hoomg
Hoomg
HoomH
Hoomg
Hoomg
Hoomg
_oomH
HoomH
Hooma
Hooma
Hoomg
Hoomg
Hoomg
Hoomg
”coma
Hooma
”coma
Hooma
Hooma
Hoomg

H<H<.

mumwzequm
eonomoee
mnwuwe

m mHNOMmeE

mweOHm
mHHOMH>meQ
mchESUOHQ
HemEflmQ
Enembflme
EzEmumoeomE
m Humbemm

< meme
memueoE

¢ menms

m memweOUemE

mmesozh

HHCOHmccom
Hamefl>uw
mflHOMHQOmmxe

mneemflc

mumHOwNHU

mWOHnumHm mnemeoz
mwoxwoeQOmm
mfioxmoemmmmm
mHoANOMQOmm
mfluxNoeQOmm
m mawwsz mEommmm

d mmawmzfl mEomUmm

m mmbfloflmmwzq
muHHmQfiQ

4 HemENmQ

mEombmm
mEomUmm
mEomUmm
mEomUmm
mEomUmm

HH asoeo

mEombmm
mEomUmm
mEomUmm
mEombmm
mEomUmm
mEomUmm
mEommmm
mEombmm
H asouw

mmbwoewom mEomUmm
mumwsomEEH mebememofle
Ezwmomoeflab
Ezwbomoewau
Esflboaoeflau
Ezwbomoewwb
Ezwboeoeflab
Eswmoeoeflwo
Ezflboeoewwb

110

 

 

HommH .................... llllllllllllllllllllllllllllll mmbfloewum mEombmm
momma .................... IIIIIIIIIIIIIIIII HomemdeHmBUH mumwsomEEH mebememowe
momma .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII mHeOum EDHUOQoefiHU
Hommg ....................H0¢E<¢0BBdEUBB<HUHU¢H¢¢0BB¢HOH mHNOMH>mHQ ESHUOQOCHHU
momma .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII mewnfieooea Esflbomoeaau
Hommg .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII HHmENmQ EzwonOeHHU
momma .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII EzemoHXmE Ezwbomoeflwo
momma .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII EzEmumoeumE Sufibomoewwo
momma .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII m Hemmemo Ezwbouoewwb
HomMH .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII m Hemmemm EswonoeHHU
momma .................... IIIIIIIIIIIIIIIIIIIIIIIIIIIIII m HHmHUemeU ESHUOQOeHHD
90mm”— .................... llllllllllllllllllllllllllllll EDHBUWSNWOHHQ ..NMb
Hmeaoee EzwonoeHNU
Hommg .... . .. llllllllllllllllllllllllllllll < wmeaoee ESflUoQOefiHU
Homm_ .... . ..aU¢B<<weH<aoae<eueu¢9¢<weeeeue m Hmnmm anemoaocfieo
Hommw .... . ..eo<e<<weemHUHeeaoeume<¢wee<ewe < Hmnmm anemoaocflau
Hommg .... . .. uuuuuuuuuuuuuuuuu Homedeweemeue muzmewe mHHHeQmHm
HomMH .... . .. IIIIIIIIIIIIIIIIIIIIIIIIIIIIII mwmem>em moeflue
HommH .... . .. llllllllllllllllllllllllllllll mwwowflmesuou meuemz
momma 09909<<He<wem<<05¢9< IIIIIIIIIIIIIIIIIIIIIIIIIIIIII meweuwummE szAeh
Hoomg III: . .. uuuuuuuuuuu m mzumwaflu eQUOerm
”coma Ina: . .. uuuuuuuuuuu 4 mzumwzmem eomomoem
floomH emae . .. nnnnnnnnnnn EnuHeeowHHmo EsEmeuemeuxm
Hoomg nus: . e. IIIIIIIIIII m memoew meuewfiowwom
HoomH III: . B. nnnnnnnnnnn 4 memuew meueHEOHwom
”coma III: . .. IIIIIIIIIII mofluflnmemn meuewEoflwom
.oomH nun: . .. uuuuuuuuu m muemEmumze meueflEoHHom
”coma lulu . .. .4 IIIIIIIIIII mmbwoewa mfiambemeoz
”coma B<H< . .. IIIIIIIIIII mflwowmmeuemE mnemeoz

111

 

Hommg
Hommg
momma
Homma
mommH
HommH
Hommg
momma
Hommg
momma
Hommg
Homma
momma
Hommg
HommH
HommH
HommH
momma
Homma
momma
mommg
momma
Hommg
momma
HommH
Hommg
HommH

HomMH £000.00

Hommg
momma
HommH

IIIIIIIIIIIIIIIII H04H440HB4HUB
IIIIIIIIIIIIIIIII 8048440994909

m mzumflmflo eOUOerm

4 mzumfinoem eomonoem
Ezoweeowwwmu EzEmeuemeuxm
m memoew meueHEoHHom

4 memoew meueHEoHHom
muflufienemm meueHEOHaom

4 muemEmpmze meueHEoflHom
mmmwoewa mwwmbemecz

quommmeuems
mmOqumHM

mmumeoz
mmumeoz

mumwzeummm mwoxmoeQOmm
eomoeoee mwmkmoemmmmm
mmwuwe mwoxmoeQOmm

m mHHQMHHmE mflOXNOHQOmm

m maaflmae

4 maeemse

m mmUHoHomHzQ
muHHmQHQ

4 HHmEHmQ

4 meme
memueoE

4 memE
4 memweOUemE

mbesozm.

HfleOHmeeOH
Hemme>ee
mHHOMHQOmmxe
mneemea
mumwowfiflu

mEommmm
mEomUmm
mEomUmm
mEommmm
mEomUmm
mEQmmmm
mEomUmm

HH asouw

mEommmm
mEomUmm
mEommmm
mEommmm
mEommmm
mEommmm
mEommmm
mEommm4
H msoeo

112

 

 

meow“
Hoova
Hoova
Hoova
Hoovg
Hoova
Hoova
Hooqa
Hoova
Hoova
Hoova
Hoova
Hoovg
Hoovg
HoovH
HoovH
Hoovg
Hooqa
Hooqa
Hoowa
meow.
Hoovg

Hoova
Hoovg
Hoova
Hoova
Hoova
Hoovg
Hoova

lllllllll ......... ...........IIIII..............U
......... .....................-I---..............w
uuuuuuuuu .....................----u..............o
......... .....................--u-|..............w
......... .....................uuuun..............w
uuuuuuuuu .....................uuuuu..............w
......... .....................uuunu..............o
uuuuuuuuu .....................:---:..............u
......... .....................n-uln..............w
uuuuuuuuu .....................:I---..............o
IIIIIIIII ......... ...........|||||.......o......0
uuuuuuuuu .....................-Iluu..............o
lllllllll .....................l|l|l..............w
lllllllll .....................||||l..............U
uuuuuuuuu .....................-uun-..............w
uuuuuuuuu .....................----u..............w
......... .....................n----..............w
lllllllll ........o............lllll....o.........0
uuuuuuuuu .....................-u-uu..............o
......... .....................--u--..............o
uuuuuuuuu .....................u--un...... ..... .m
uuuuuuuuu .....................----u..............w
lllllllll ....................lllll..............w
lllllllll ..........o..........|||l|..............U
uuuuuuuuu .....................-----..............w
lllllllll .....................|I|Il...........o..w
nnnnnnnnn .....................--uuu..............w
......... .....................uuuun..............w
uuuuuuuuu 444444w94444w94944eeeuuIIIBHHH4BUB44wouvH

HH msouw

4 mHUmE mEommmm

4 memHeOUemE mEombmm
mnezozh mEombmm
HHeOumeeOM mEomUmm
HHueH>HH mEommmm
mHHOHHQOmmHe mEommmm
mUHQmHe mEomUmm
mumHOHHHU mEomUmm

H asouw

mmUHoeHUm mEomUmm
mumHzomEEH munememoHe

mHeOMm
mHHomwbmeQ
memQESUOMQ
HemEHmQ
EzemonmE
EzEmumoeumE
m Hemeemu

4 Humbemm

m Hemeemeu

EszomoeHHo
EsHmoQoeHHo
ESHUOQOCHHU
EszoeoeHHb
EszomoeHHb
EzwboeoeHHU
EzwboqoeHHU
ESHUoQoeHHU
EzHUoQoeHHU

EszUmszoHHQ .Mm>

Hmexoee
4 Hmezoue
m Hwemm
4 ngmm

EszoQOeHHU
EszoQoeHHU
EDHUOroHHU
EDHUOQoeHHU

muameec meewcemHm
mHmem>um moeflo4
mHHOHHUezuOM meuemE
meHeUHummE szAeH

113

Homqw
Homvg
Homva
Hom4_
Homvg
Homvg
Homva

Hoovg
HOOVH
Hoovg
Hooqg
Hoovg
moova
”0040
”Dove
Hoov_
Hoova
HoovH
Hoovg
Hoovu
H0046
Hoova
H004“
Hoovw
Hoov_
Hoova
Hoova
Hoovu

{—1

LDLDLDLDULDLDLDLDLDLDCDLDLDCDLDLDUULDLD

44044099wo4ueae4eee44wfiweww:0944444009 uuuuuuuuuuuu

Hmeaoee
4 Hmezoee
m Hmemm

4 404mm
muzme

mHm
mHHOHHU
meHeu

m mzumH

EzHUoQOeHHu
ESHUoQoeHHU
EsflbomoeHHu
ESHUOQoeHHU
44 mNNHaem4m
em>em moewm4
ezuoe meuemz
HummE szer

440 cononoam

4 mzumHsmem eonouoem
EoneMOHHHmU EDEmeuemeuxm

m memoeH
4 memuefl
mUHuHMUemU
4 muemEmusz
mmUHoeH
mHHOHmme
mmoH:
mumHzequm
eomomoee
mbHuHe
m mHHOHHMmE
m mHH
4 mHH
m mmmfloflm
muHe

4 HemEHmQ

4
mem

meueHEoHHom
meueHEoHHom
meueHEoHHom

meueHEOHHom
H mHHmUemeoz
uemE mnemeoz
umHH mnemeoz
mHDHNOHQOmm
mwmxmoemmmmm
mHUHNOMQOmm
mHQHNOMQOmm
Hmzq mEomUmm
Hsz mEomUmm
mwsq mEOmUmm
mQHQ mEomUmm
mEomUmm
mEomUmm
mEommmm

meme
ueOE

114

 

Homqg
Homvu
Hom4_
Homva
HomVH
Homva
HomvH
Homva
Homvg
HomQH
Homva
Homvg
Homvg
Homvg
Homvg
Homva
Homva
Homvg
Homva
Homva
HomVH
HomvH
HomvH
Homva
Homvy
Homvg
Homvg
Homvg
Homvg
Homva
HomvH

mUHuHe mHmHNoeQOmm

m mHHowHHmE mHuxmoeQOmm

m mHHHm3Q mEomUmm
4 mHHHm2Q mEomUmm

m mmUHoHumHzQ mEomUmm

muHemQHQ mEombwm
4 HMmEHmQ mEomUmm
4 meme mEomUmm
memueoE mEombmm

HH asouw

4 mHUmE mEommmm

4 memHeOUemE mEommmm

mbezuzh mEombmm
HHeOumeeOM mEombmm
HHDeH>MH mEombmm

mHHOHHQOmmAe mEomUmm

mbHQmHe mEombmm
mumHoHHHo mEommmm

H Quouw

mmUHoeHum mEombmm

mHeOMm

mHHOHH>meQ
memQEsuouQ

HMmEHmQ

EzemonmE
EzEmumoeomE
m Hemmemu
4 Humbemm
m HMmHUemeo

MMMNDUQEEH MHUQmHmUHQ

EanoQoeHHU
EzflonoeHHU
ESHUOQOCHHU
EzHUoQOeHHU
ESHUOQOeHHQ
EswnoQoeHHU
ESHUOQoeHHU
EzwonoeHHb
EzHquoeHHb

EszomzwmoHHQ .em>

115

hf.
i5

Hoomg
”coma
Hoomu
Hoom_
Hoomg
Hooma
Hooma
Hoomg
Hooma

Hooma
”coma
Hooma
”coma
HoomH
Hooma
”coma

HomvH
Homvg
Homvw
Homvg
Homqa
Homva
Homqa
HomvH
Homva
HomvH
momva
Homvg

..40........
..40.. ..... .

..404.......
..4w4.......
..404.......
..40........

..40........
..40........
..40........
..40........

LDLDLDLDLDLDLDLDLD
UUUUUUUUU

LDLDLDLDLD
°UUUU

.0..0.

UUUUUUUUU

UUUUUU

HHHHUH404B40UUB44H4UUH0448940?04408409849440494444

mHHOHH>meQ EsHUOQoeHHb

memessooem EDHUOQoeHHU
HemEHmQ EszoQoeHHo
EzemumeE EsHmoQoeHHb
EzfimumoeomE EszomoeHHb
m Humbemo EszomoeHHU

4 Humbemo EzHUOQOeHHU

m HemHUemeQ EzHUOQOeHHw
EszomszoHHQ .em>
Hmeaoee EszoQoeHHU

4 Hmezoue EsflonoeHHb

m Hmewm ESHUOQoeHHU

4 Hmnmm EDHUOQoeHHU

musmufla mflaflcemam

mHmem>em moeHu4

mHHOHHUezuoe meuemz
meHeUHummE szHeH

m msumHHHo eOBOUQem

4 mzumHzmem eomomoem
EsmHeHOHHHmU Ezsmeuemeuxm
m memoefl meueHEoHHom

4 memoew meueHEOHfiom
moHuHeUemb meueHEOHHom

4 muemEmumze meueHEOHHom
mmbHoeHH mHHwUMmeOE
mHHOHmmeuemE mnemeoz
mmoHsumHH mnemeoz
mumHzequm mflmxmoemammm
eomomoee mHDXNOMQOmm

116

 

HoomH— ..do ..... ...0.
Hooma 0o¢w.o0.......
”Dom“ .0<U....000..0
Hoomfl— 00<w.0........
Hoomw .0¢U...0....0.
HoomH— 0.¢0.00.000.0.
Hoomg 0.¢U.0..0.0..0
HoomH 0.¢U.0..0.0...
Hoomfl ..¢w.........0
HoomH— 00¢w0......0.0
HoomH .0¢D.0...0.0..
Hooma 0.<O..0...0...
HoomH— ..flw.o.o0.0...
Hoomg ..flo..0.....0o
”Coma ..«O00ooooo0oo
HoomH— ..«0000....00.
RoomH— ..do.0...0....
”coma 0.4mo0...0.....
”Comm— ..¢O.0 .........
Hoomu ..flU..........
”com“— 0.46.0...oooo.
”Coma .../N0....0....o
”coma ..Anamv ...... ....
Hoomg .0dw.0.0.....0
HOOMH 0.<MU000.0..0..
Hoomw 00¢wo. ........
HoomH .0¢O0.0.......
Hoomw .0fi00....0...0
”coma .0<w.0.......0
HOODH 0.¢0.0...0....
Hoomw ..¢00.0..00..0

m memueH meueHEOHHom
4 memueH meueHEOHHom
mUHuHHUemU meueHEOHHom

4 muemEmumse meueHEOHHom
mmUHoeHH mHHmmemeoz
mHHOHmmeuemE mnemeoz
mmoHsumHH mnemeoz
mumHzequm mHOXNoeQOmm
eomomoee mwoxwoemmwmm
mmHuHe mHmHNoeQOmm

m mHHOHHMmE mwuxmoeQOmm
.m mHHHsz mEommmm

4 mHHHsz mEommwm

m mmmHonmHzQ mEomUmm
muflquHQ mEombmm

4 HemEHmQ mEombmm

4 meme mEomUmm

memueoE mEommmm
............. .... HH QSOHU
. 4 mHUmE mEommmm

4 memHeomemE mEomUmm
......................... QUCDUDW QEombmm
.......... ............... HHQOMWCCON QEOQUQm
kueH>MH mEomUmm
mHHOHHQOmmHe mEommmm
mUHQmHe mEombmm
mumHoHHHu mEombmm

H asouw

mmUHOeHUm mEomUmm
mumqumEEH mememembwe
mHeOum EszoQoeHHU

LDL'JLDLDEDCDUUUUOUUUUOUUUUUUOOUOUULDLDLD
UUUUUUUUUUUUUUUUUUUUUUUUUUUUUUU
UUUUUUUUUUUUUUUUUUUUUUUUUUUUUUU

O
O
I
O
O
O
I
O
O
O
C
O
C
O
C
O
O
O
O
C
O
C
O
O
O
O
O
O
C
O
O
O
O

117

 

HommH
Hommu
Hommg
Homma
Homfl
Homfl
momma
8mm:
momma
8mm:
3mm:
HommH
Homfl
momma
Homm:
Homfl
momma
8mm;

momma
momma
momma
Homma
Homma
momma
momma

Hoomg
802
302

I I I I
I I I I
I I I I

I I I
I I I I

I I I
I I I I
I I I I
I I I
I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I I
I I I
I I I I

I I I I

I I I I

I I I I

I I
I I

I I I I

I I I I

I I I I

I I I

I I I

I I I

I I

I

I I

I I

I I

4444444444444444

444344444444

........<.........

........<.........

0H404HUHH400090404840444H4404000000H44HHHHH0440444

..40 ........... 0.0 ...... 0 ......... .... ............
..40...........0.0......0.........................
..40...........0.0......0........... ..............

HHeOume40h mEomUmm
HHOeH>HH mEomemm
mHHOHHQommHe mEomUmm
mUHQmHe mEomUmm
mumHoHHHo mEommmm
H Q5040
mmUHoeHum mEomUmm
mumHsomEEH mebemMmUHQ
mHeoMm EszoQoeHHb
mHHOHH>mMQ EzwonoeHHU
memeasuoem EDHUOQoeHHU
HMmEHmQ EzwmoqoeHHU
EzemumeE EzwbomoeHHU
EzEmumoeumE ESHUoQoeHHU
m Hemmemo EzHUOQoeHHU
4 Hemmemo EszoQOeHHU
m HemHUemeu ESHUOQoeHHU
EszomszoHHQ .Mmb
Hmeaoee EzHUoQoeHHU
4 HmeaoHQ EzwbomoeHHU
m Hmewm ESHUOQOCHHU
4 Hmemm ESHUOQOCHHU
muzmeHe mHHHeQmHm
mwmem>em moewo4
mHHOHHUezuoe meuemz
meHeUHummE szer

m mzumHHHU e0b0boe4
4 msumquem eononoem
EoneMOHHHmU Ezfimeuemeoxm

118

 

Hoooa
HOO©H
Hoowa
Hoowg

momma
HommH
momma
momma
Hommg
Hommg
momma
Homma
Hommg
HommH
momma
Hommg
HommH
momma
Hommg
Homma
Hommg
momma
Homma
Momma
momma
momma
momma
momma
momma

00054444004044904H4HBH4440H44044000004H44084H404H0

muzmefle mHHflcemem
mwmem>em moeHu4
mHHOHHUezuoe meuemz
meHeoHummE mzsxes

m maumHHHU eononoem

4 mzumHzoem eomoeoem
EmuHeMOHHHmU E3Emeeemeoxm
m memoeH meueHEOHHom

4 memoeH meueHEOHHom
mUHuHebemm meueHEOHHom

4 muemEmumze meueHEoHHom
mmbfloeHH mHHmUMmeOE
mHHOHmmeuemE1mUHme02
mmOHzemHH mmeeoz
mumHzequm mwuxwoemammm
eomomoee mHoHNoeQOmm
mUHuHe WHOANOHQOmm

m mHHOHHHmE mHuHNOMQOmm
m mHHHsz mEomUmm

4 mHHHmsQ mEomemm

m mmUHonmHzQ mEomUmm
meHemQHQ mEomUmm

4 HHmEHmQ mEommmm

4 meme mEomUmm

memueoE mEommmm

HH asoew

4 mHUmE mEomUmm

4 memHeOUemE mEomUmm
mnezuzh mEommmm

119

 

Hoowa
Hooma

82:.

Hooma
Hoooa
flooma
HoomH
Hoooa
Hoowa
H0006
Hooog
Hoowa
HoowH
moomg
”coma
mooog
moowg
Hoowg
Hoomg
HOOQH
Hooog
meow“
”coma
HQO©H
Hoowg
”coma
Hoooa

moowa
”coma
Hooma

m mmbHoHQmHsQ mEomme
muHHmQHQ mEombmm

4 HemEHmQ mEombmm

4 meme mEomUmm
memueoE mEomUmm

HH asoew

4 mHUmE mEommmm

4 memHeomemE mEommmm
mbezuzh mEomUmm
HHeOHmee0h mEombmm
HHUeH>MH mEomUmm
mHHOHHQOWmHe_mEomUmm
mUHQmHe mEomUmm
mumHOHHHU mEommmm

H meoew

mmmHoeHUm mEomUmm
mumHsumEEH mebememowe

mHeOum
mHHOHH>mMQ
memQESerm
HHmEHmQ
EzemonmE
EzEmumouomE
m Humbemu

4 Hembemu

m HemHUemeu

EswbomoeHHb
EzwmoeoeHHb
EDHUOQOCHHU
EsHUomoeHHb
EszoQoeHHu
EzflmomoeHHb
EanoqoeHHb
EzHeroeHHU
EswnomoeHHD

53Hzom2HmOHHQ .em>

Hmeaoee
4 Hmeaoee
m Hwemm
4 Hmemm

ESHUOQoeHHU
EszouoeHHb
EsHanoeHHo
EDHUOQoeHHU

120

Hommg
Homwg
Home“
momma
Homog

Home“
momma
Homwg
Home“
Homog
Homou
Homog

Hooo_
Hooog
”coma
HoowH
Hoowg
Hooma
Hooma
Hoooa
Hoooa
”coma
Hooou
Hoomg
HOO®H
Hooog
Hooog
Hoomu

9000444440000H4H0800050440HB00040H00844444HHH04009

EzEmumoeomE
m Hemmemm

EzHUOQoeHHD

EswmomoeHHU

4 Hemnema EszoQOeHHU

m Hemeemeo EsHboeoeHHU
EszumsHmoHHQ .Mm>

Hmeaoee EzHUOQoeHHQ

4 Hmeaer EzHUomoeHHo

m Hmemm EDHUOQoeHHU

4 Hmemm EDHUOQOCHHU

augmeee 44444Qm4m

mHmem>em moeHo4

mHHOHHUezuoe meeemE
meHeUHummE mzexeh

m maumHHHo eomomoem

4 mzumquem eOU0boe4
EoneMOHHHmu EzEmeuemeuxm
m memueH meueHEOHHOm

4 memueH meueHEOHHom
mUHuHeUemn meueHEoHHom

4 meemEmusz meueHEOHHom
mmUHoeHH mHHmmemeoE
mHHOHmmeeemE mbnmeoz
mmoHsumHH mnemeoz
mumHzequm mHDHNOMQOmm
echoeoee mHmHNoeQOwm
mUHuHe mHoxmoeQOmm

m mHHoHHMmE mflmHNOMQOmm
m mHHHsz mEomUmm

4 mHHHsz mEommmm

121

 

momma
Hommw
Homwfl
momma
momma
Homwa
momma
Homog
Homma
Homwg
Home“
HommH
momma
momma
momma
momma
Homwg
momma
Hommg
Homog
Homma
Hommg
Hom©H
Home
momma
Homog
HomoH
Homma
momma
Homwa
Homo_

mmUHoeHH mHHmUMmeoz

mHHOHmmeuemE mbumeoz

mmOHSumHH mUHmeoE

mumHsequm
eomomoen

mHuxmoeQOmm
mHUANOMQOmm

mUHeHe mHmHNoememm

m mHHOHHHmE

mHUHNOMQOmm

m mHHHsz mEombmm
4 mHHHmSQ mEomUmm

m mmUHOHmmHzm mEommmm

muHemQHQ mEommmm
4 HMmEHmQ mEommmm
4 meme mEombmm
memueoE mEomUmm

HH meoew

4 mHUmE mEombmm

4 memHeOUemE mEomUmm

mmezozh mEomUmm
HHeOemeeow mEommmm
HHUeH>HH mEomUmm

mHHOHHQOmmHe mEombmm

mUHQmHe mEommmm
mumHoHHHU mEommmm

H msoew

mmmfloeHom mEOmUmm

mHeOum

mHHOHH>meQ
memessooem

HemEHmQ

EzemUHXmE

mumHsumEEH memememuHQ

EzHeoQOeHHU
EsHmoQOeHHU
EsHUomoeHHU
EsflmomoeHHo
EzHeroeHHU

122

 

Hoong
HOOFH
HOOPH
Hoong
floor“
HOOFH
”coma
Hoona
meowa
Hoora
”Dong
moona
Hoong
Hoonw

HOOPH
HOOFH
Hoo>_
moona
Hoona
HoowH
”GONE

momma
momma
HommH
Homww
Homwa
momma
Homo_

{90000000000000

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KC
E-iE-*E-1

-4
4

4

4

BHHHBHE‘BE—iE-ifi

4

°BBBBH

H00044HH0098HBBUHB0000094B0004HHH4B44449000040B894

LDCDLDULDLD
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mumHoHHHu mEommmm

H msoeo

mmbwoeHUm mEombmm

mumHzomEEH mebememUHQ

mHeOem EzHUOQOeHHU

mHHOHH>mHQ EzwnomoeHHb

memefisooea ESHUOQoeHHU

HemEHmQ EsHonoeHHo

EzemonmENESHmouoeHHU

EsEmemouomE ESHUOQoeHHU

m Hemnemm ESHUOQoeHHU

4 Hembemm EszoeoeHHb

m HMmHUemeo E:HU0QOeHHU
EszomszoHHQ .em>

Hmezoee EzHUOQOeHHU

4 Hmezoee EszomoeHHb

m Hmewm EDHUOQoeHHU

4 Hmemm EDHUOQOCHHU

musmeflc mflflflcemNm

mHmem>em moeHU4

mHHOHHUezuOM meuemz
meHeuHummE szer

m mzumHHHo eonomoem

4 mzumHzoem eononoem
EzmHeeowHHmo EnEmeuemeuxm
m memuew meueHEoHHom

4 memoeH meueHEoHHom
mUHeHMUemU meueHEOHHom

4 muemEmumze meueHEoHHom

123

 

mocha
mocha
Hoona
”Gong
HOOFH
Hoop.
Hoop“
HoonH
HOOPH
moon“
HOOFH
Hoona
”COPE
Hoona
mocha
HOOPH
mocha
Hoomg
Hoona
HOONH
Hoona
moose
HOQPH
Hoop.
Hoop“
HOOPH
mocha
H0050
Hoong

0000000000000 '00000000000

~424:4:454:4 4

HE—‘E—‘HHE—iE-‘E-IE—IE-iE-IE-‘I

BHHHHBBHBHHHHHHB

m mzumHHHU eomomoem

4 mzumHzmem eomomoem
EoneMOHHHmU EzEweuemeoxm
m memoeH meueHEOHHom

4 memuefl meueHEOHHOQ
mowuwubemb meueHEoHHom

4 muemEmumze meueHEOHHom
mmewoeHH mHHmUMmeOE

mHHOHmmeuemE
mmonumHH

mnemeoz
mnemeoz

mumHzequm mflmxmoememmm
embomoee mHOHNOMQOmm
mUHUHe mwmxmoeQOmm

m mHHOHHHmE mHOANOHQOmm

m mHmeze

4 maawmze

m mmUHOHmmHzQ
muHMmQHQ

4 HHmEHmQ

4 meme
memueoE

4 mHUmE

4 memHeomemE
mUe305h
HHeOHmeeOh
HHmeH>MH
mHHOHHQmeAe
mneemwa

mEomUmm
mEomUmm
mEommmm
mEommmm
mEombmm
mEomUmm
mEommmm

HH Qsoew

mEomUmm
mEomUmm
mEombmm
mEombmm
mEomUmm
mEomUmm
mEommmm

124

 

Homba
Hombw
HombH
Homna
Homwg
Homha
Homha
Homba
Homwa
momma
Hombg
Hemp;
momma
Hombg
HomnH
Homba
Homba
momma
momma
moms;
Hombg
Home
Homba

HomFH
Homhg
Homng
Homba
momma
Hombg

° E4

00000000000000000000000

'00000

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HBE—‘BHHBBBHE—‘HEBBBHBBHBBH

BEBE-+88

Homng 44ew00003994900444040959099409099uoe4eeuouee444uuo

memueoE mEombmm

4 mHUmE

4 memHeOUemE
mbezozh
HHeOumeeoh
44u24>e4
mHHoHHQommxe
mnflemwa
mumHoHHHU

mmUHoeHom

HH aeoew
mEomUmm
mEomUmm
mEomUmm
mEommmm
mEomUmm
mEombmm
mEomUmm
mEombmm
H asoew
mEomUmm

Mu QNSUMEEH MHUCMHmb “HQ

mHeOUm
mHHOHH>meQ
meweEsoer
HMmEHmQ
EzemoHXmE
EzsmumoeumE
m Humbemm

4 Humbemu

m HHmHUemeo

EzfleoQoeHHU
EszouoeHHb
ESHUOQoeHHU
EzwonoeHHU
EzwonoeHHb
EzHUoQOeHHU
EzwonoeHHU
EzwnoQOeHHU
ESHUOQOeHHU

EszumszoHHQ .em>

HmeEOMQ
4 Hmezoee
m.Hmemm
4 Hmemm

EszoQOeHHU
EszomoeHHU
EDHUOQoeHHU
EDHUOQoeHHU

muzmeflc 444444444
mHmem>em moeHU4
mHHOHHUeSHOH meuemz
meHeowummE msEer

125

 

Hoomg

80m:
302
”coma
Hoomg
”coma
meow;
Hoom:

Sm:
Sm:
gonna
mom:
mom:
HomPH
flown;
momma
Home;
8m:
Homh_
Homng
Hombg
HomNH
Homng
Sm:
Homwa
Rom:
Hom>_
momma

 

'00000

0000000000000

000000

0

HBHBHBBE—‘BBHBHBHBBE—‘HH

EszomszoHHQ .em>
Hmeaoee EzHUoQoeHHU
4 Hmeaoee ESHUOQoeHHU
m Hmemm ESHUOQOQHHU
4 Hmemm ESHUOQoeHHU
muzmefle mflawcemmm
mHmem>em moeHo4
mHHOHHUezuoe meuemz
meHeoHummE szHeh

m mzumHHHu eomomoem

4 mzumquem eonomoem
EzuHeMOHHHmU EzEmeuemeoxm
m memoeH meeeHEOHHom

4 memoeH meueHEoHHom
muHuHeUemU meueHEoHHom

4 muemEmumze meueHEoHHom
mmmwoeHH mHHmbemeoz
mHHOHmmeuemE mbumeoz
mmonumHH mmemeoE
mumHzequm mHoANoeQOmm
eomomoee mHmHNoeQOmm
mUHUHe mHOANOMQOmm

m mHHOHHemE mHUHNOMQOmm
m mHHHm3Q mEomUmm

4 mHHHmsQ mEomUmm

m mmUHoHumHzm mEombmm
muHMmQHQ mEomUmm

4 HMmEHmQ mEommmm

4 meme mEommmm

126

 

Hooma
HoomH
HoomH
Hooma
Hoomg
”coma
”coma
Hoomfi
Hoomg
Hoomfl
Hoomg
”com“
HoomH
”coma
Hoom_
”coma
”coma
Hoomw
”coma
Hoomg
Hoomg
HoomH
Hoomg
Hooma
”coma
Hooma
Hoom.
Hoomg
HoomH
Hooma
Hoomw

0000000000000000000000000000000

 

0000000000000000000000000

 

echoeoee mHuxNoeQOmm
mUHuHe mHORNOMQOmm

m mHHOHHMmE mmeNOHQOmm
m mHHHan mEomUmm

4 mHHHsz mEomUmm

m mmUHoHomHzm mEomUmm
muHemQHQ mEomUmm

4 HHmEHmQ mEombmm

4 meme mEomUmm

memueoE mEombmm

HH asoem

4 mHUmE mEomUmm

4 memHeOUemE mEomUmm
mbesuzh mEomUmm
HHeOumeeON mEombmm
HHmeH>eH mEombmm
mHHOHHQOmmAe mEomUmm
mUHQmHe mEomemm
mumHOHHHU mEomUmm

H meoew

mmbHoeHom mEombmm
mumHzomEEH memememuHQ

mHeOHm
mHHOHH>meQ
memefieooea
HemEHmQ
EzemumeE
Essmumoeums
m Humbemm

4 Humbemo

m HMmHmemeu

EDHUOQoeHHU
EszoQoeHHb
EDHUOQOCHHU
EsHUoQOeHHU
EzwbomoeHHU
EszoeoeHHb
EszoeoeHHU
EszoQOeHHU
EzHUoQOeHHO

127

 

Homma
HommH
momma
Homma
Homma
momma
HommH
Hommg
Homwu
Homwg

momma
Homma
momma
HommH
Home
Homwg
momma

meowH
HoomH
Hoowa
Hooma
HoomH
Hoowa
”coma
Hoomfi
”coma
Hooma
HoomH

.....4..

440980004H<40H440H<H4000094409IIII4400440H44400HHI

00000000000

00000000000

mHeONm EszoQoeHHU
mHHOHHbmeQ ESHUoQoeHHU
merEDUOMQ ESHUOQoeHHo
HMmEHmQ Esz0Q0eHHU
EaemonmE EDHU0QoeHHU
EzEmumoeumE ESHUOQOeHHU
m Hemmemm EszouoeHHb

4 Hemmemo EzHUOQoeHHU

m HewHUemeo SSHmoQoeHHU

EszomzflmoHHQ .emb
Hmezoee EzHUoQoeHHU

4 Hmezoee EzHUOQOeHHU
m Hmemm EDHUOQoeHHU
4 Hmemm EDHUOQoeHHU
musmeec mwfiflaemam
mHmem>em moeHo4
mHHOHHUeseOM meeemz
meHeuHummE mzfier

m mzumHHHo eomomoem
4 mzumHzmem embomoem
EsoHeMOHHHmU Ezfimeuemeuxm

m memueH meueHEoHHom
4 memoeH meueHEoHHom
mUHuHeUemU meueHEOHHom

4 muemEmumze meueHEOHHom
mmbfloeHH mHHmUemeOE
mHHOHmmeuemE mnemeoz
mmoHsumHH mbemeoz
mumHsequm mHm4NOHQOm4

128

EzuHeHOHHHmU EzEmeuemeoxm
m memueH meueHEOHHom

4 memuew meueHEOHHom
mUHuHememU meueHEOHHom

4 muemEmumze meueHEoHHom
mmmHoeHH mHHmbemeoz

mHHOHmmeuemE
mmOqumHH

mnemeoz
mmemeox

mumHzequm mHUHNOMQOmm
eonomoee mHUANoeQOmm
mbHuHe mHDHNOMQOmm

m mHHoHHMmE mwoxmoeQOmm

m mNNHmze

4 «4444.4

m mmUHoHomHzQ
muHMmQHQ

4 HemEHmQ

4 meme
memueoE

4 mHUmE

4 memHeomemE
mUe305h
HHeOHmeeOh
HHueH>MH
mHHOHHQOmmAe
mUHQmHe
mumHOHHHu

mmmwoeHum

mEombmm
mEombmm
mEomUmm
mEommmm
mEomUmm
mEommmm
mEommmm

HH msoew

mEombmm
mEomUmm
mEomUmm
mEommmm
mEommmm
mEomUmm
mEombmm
mEomUmm
H Qsoew
mEombmm

mumHzUmEEH memememowe

129

HoomH— ..0.l.|lll.l.0....< fl
HOOQH ...III'I......A& <
Hoomu .0.II'II-oo....«.....0....0¢....0......0.....0..Illl||
4 4
4 4

”com“ ...llIl-ooooo
HOOQH ...Illul|......
HoomH ...:uul ......
HoomH— ...IIIIII- ......
HOOQH ...IIIII...-.
”coma ...-uu-.....
400mg ...-nlu.....
Hooma .. I---
HOOGH ...l'lnl.....
Hoomg ...Illlulooo.
HoomH ...:uuu..e.
40040 ...:n.......
HoomH ...11'1.....
_oom0 ...-u--.....
”com“ ...---l.....
HOOQH ....lllalooooo

~4
4:4:4:4:4 4 4.4 4 4 4 4 4 4

d . I .
414141424 4 4 4 4 4'4 4 4 4

..........04....................

moom_ ...IIII......4

HoomH ...IIII ...... 4 ..... ......4

HoomH ...IIII ...... 4 ...... .....4....................
4

HoomH— ...Illloonoo.<oooooo.....
HOOQH— ...llllooooo.¢o..........

HOOGH ...llll...llll....................................

Hoom_ 449--Ineauoueoewoouuoe44eweeeu4444w4wwoeu4ww4u4ee4

Hommg llllll oo.oo..o....o...o.......I'Ilonoooooooooooool

momma IIIIII ..........o.oo..........IIII.......ooooooool.

mbesozm mEombmm
HHeOMmeeON mEombmm
fiwoew>uw mEombmm
mwwomflQommxe mEomUmm
mUHQmHe mEomUmm
mumHoHNHo mEomUmm

H @5040

meHoeHUm mEomUmm
mumasomEEH mHUemMmUflQ

mHeOMm
mflwoww>mun
memQESUOeQ
HMmEHmQ
Esemowme
EzfimumoeomE
m Hemmemm

4 whmbemm

m flemwmemeo

ESHUOQoeHHU
EzwonoeHNU
ESHUOQoeHHU
Ezwbomoewwo
EsweoQoeHHU
EzwonoeHHU
Ezwmomoeflwo
EswUOQoeHHU
EzwnoqoeHHo

EszomzwmoHHQ .Mm>

wmeEOMQ
4 Hmeaoufl
m Hwemm
4 Hmcmm

Ezwboaoeflmo
EzflmomoeHHb
EefiboaoeHHo
EsflvomoeHHU

muzmewe mHHHeQmHm
mwmem>em moewo4
mflflowflmezuoe meuemz
meweuwummE mzfixeh

m msumwwflo eOU0boem
4 mzumwzoem eOUOerm

130

 

Hommg
Hommw
HommH
momma
HommH

HoomH
Hoomg
HoomH
Room“
Hoomg
”coma
”coma
Hooma
HoomH
moomH
Hoomw
moomg
Hoomg
”coma
Hooma
”coma
Hooma
Hoomg
HoomH
”coma
”coma
Hooma
Hooma
”coma

...........0......

 

 

994045!U094000B00040H4444H44B094080400H040094040HB

444444444444444444444444

I I I (D I I

4 4 4’4 4'4 4 4'4 4 4.4 414 4 4.4 4 4 4 4 4 4 4

4 Hmemm EDHUOQoeflHU
mumm444 m44444044
mwmem>em moeflo4
mwwowflbesuon meuemz
mefleowummE msExeh

m msumewo eomomoem

4 mzumwzmem eomomoem
EsofleHOMHamo EzEmeuemeoxm
m memoew maueHEONaom

4 memoew meuewEOHHom
mowuwebemm meuewfiowfiom

4 muemEmumBQ meueHEowHom
mmbwoeflw mwammumeoz
mwwommmeuems mUHmeoz
mm0~zumflw mnemeoz
mumwzeummw mfluxmoeQOmm
eomomoeu mflUXNOHQOmm
mbfluwe mwoxmoummmmm

m mHHOMHHmE mfloxwoeQOmm
m mawflmzm mEombmm

4 mHmezQ mEomUmm

m mmUHoHumazm mEomUmm
muwemQHQ mEommmm

4 NemENmQ mEombmm

4 meme mEombmm

memueoE mEommmm

HH Q2040

4 mHUmE mEomUmm

4 memHeOUemE mEomUmm

131

 

HommH
momma
momma
momma
momma
Homma
momma
Hommg
momma
Homma
momma
Homma
momma
Homma
Hommw
40mma
momma
Hommg
mommg
HommH
Homma
40mm_
40mm“
momma
Homo.
momma
momma
flommg

Hommg
HommH

.0.

4 mwwwmzq mEomUmm

m mmnwowmmwsm mEombmm
mumeQfiQ mEomUmm

4 flumEHmQ mEommmm

4 meme mEomUmm
memueoE mEommmm

HH @5040

4 mHUmE mEomnmm
4 memHeOUemE mEomUmm
mmezuzh mEomUmm
wweOHmeeom mEomUmm
Hfibew>ew mEomUmm
mflwomfiqommxe mEombmm
mUHQch mEombmm
mumaowwflo mEomUmm

H Qsoew

mmbwoewom mEombmm
mumwzumEEw mebemumowQ

mHeOHm
mwwoww>mefl
mcmnesooem
HemEHmQ
EzemUmeE
EzEwumogumE
m Hemmemo

4 Hemmemo

m Hhmweemeo

EzflUOQOeHHU
ESHUOQOeHNU
EDHUOQoeHHU
EszoQoeHHU
Ezwboqoeflwo
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Ezwnomoewwb
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EswmomoeHHD

EszumzwmonQ .em>

Hmezoun
4 HmeROMQ
m Hmnmm

EzwmoQoewwb
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EDHUOQOCHHU

132

 

 

HoooHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ............HH4440 Ezemowme Ezflmoqoewwu
HOOOHH 4440BU40040400404H0090040900484. . . . . . . . . . . . . IIIIII EzEmumOnHumE E: HUOQOCHHD
HOQOHH nnnnnnnnnnnnnnnnnnnnnnnnnnnnnnn ............. llllll m Hemmemb Ezwmoaoewwo
HOOOHH uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuu ............ IIIIII 4 Hemmemu Ezflbouoeflwb
MOOOHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ............ IIIIII m Humwmemeo E24U0Q0ewwu
floooHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ............ IIIIII EzfizomzwmoHHQ .em>
Hmezer SufionoewNU
flOOOHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ............ IIIIII 4 wmezoen EsonQOeHHU
HOOOHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ....U nnnnnn m stmm EsfivomoeflHo
HOQOHH uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuu ..... uuuuuu 4 Hwnmm EsflnoaoeHHu
HoooHH llllllllllllllllllllllllllllllll ............ IIIIII muzmuwz mwwwszHm
HOOOHH lllllllllllllllllllllllllllllll ............. IIIIII mwmem>em moeflu4
HoooHH uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuu ............ 111111 mwfiomHmezuon meuemz
400040 uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuu wwwugwwe444wo nnnnnn meHeUHummE 425445
mommH ...... .................. m WBMMflNHU COUOUOCm

mOmmH ...... ......o........... « WDUQNBUCQ COUOUOSK
momma ...... .................. ESUHCNONHHQU EDEmfiuCMCUNW
HOmmH ...... .................. m QCQUCN MQHCHEOHHOQ
mommH ...... .................. 4 MCQUQH QCuQfiEOflHOm
momma ...... .................. QUHHHHUCQU MQHCHEOHNOQ
momma ...... .................. < MMQQEMHWSQ MQHCHEOHNOW
Hommg ...... .................. WQUHOQHN MHNWUHMQOE
HODQH ...... .......... ........ QflfiowmmfiucmE mUHMCQz
momma ...... .................. MWOHDHWHM MUMMCOE
mommg ...... .................. MMQHDCUMQW WHOXNOHQQQQW
mommg ...... .................. COUOUOCH WHORNOHQQMQE
momma ...... .................. MUHUHQ WfluxNOqummm
HOmmH ...... .................. m QHHONWHWE mflmeOHmmmmm
mommg ...... .................. m QHflHWSQ meombmm

133

 

 

HOQOHH
HQOOHH
HOQOHH
HOQOHH
HOOOHH
floooHH
moooHH
fiOOOHH
HOOOHH
HOQOHH
HoooHH
HoooHH
floooHH
HOOOHH
HQOOHH
HoooHH
HOQOHH
floooHH
floooHH
HQOOHH
HoooHH
HoooHH
HOOOHH
HOQOHH
floooHH
HoooHH
HOOOHH
HOQOHH
HOOOHH
HOOOHH
HoooHH

4440H040040400404BO0HU040H004H4.........

uuuuuuuuuuuuuuuuuuuuuu woue444woa-......
uuuuuuuuuuuuuuuuu waeuo4ueou4e4........

IllIHU40040400404H00H0040900494........
lllllllllllllllllllllll UHg<®UBI . . o a . .
444mgo4ow4w4oo4w4eooa0040500494........

................. aoweuu4ueuw4e4........
444ueu4mw4w4uo4o4ewoaow4oeoo4e4........
Iuuueu4oo4w4uo4u4emmeou4ue00494........

lllllllllllllllllllllllllll D<H<...0.0..

. IIIIII 4 muemEmumsn meueHEOHHOQ
. nnnnnn mmmwoefla mNHmmMmeoz
. IIIIII mwwommmeuemE mnemeoz
. tttttt mmowsumHM mbemeoz
. IIIIII mumHsequm mwoxwoemummm
. IIIIII eomomoee mflmxwoemmmmm
. IIIIII mnwuwe mflu4NOHQOm4
. IIIIII m mHHOMHMmE mekmoeQOmm
. IIIIII m mHHHmDQ mEombmm
. IIIIII 4 mwwflmsm mEomUmm
. IIIIII m mmUHOHomHzQ mEomUmm
. nnnnnn muwemQHQ mEommmm
. IIIIII 4 HumENmQ mEomUmm
4 meme mEomUmm
. IIIIII memueoE mEomUmm
. IIIIII HH Q5040
. IIIIII 4 mHUmE mEommmm
. IIIIII 4 memHeOUemE mEommmm
. IIIIII mbezuzh mEomUmm
. IIIIII wweOumeeOh mEombmm
. IIIIII meew>ew mEomUmm
. IIIIII mw~OMHQommxe mEomUmm
. IIIIII mUHQmHe mEomUmm
. llllll mumHoHHHu mEomUmm
. IIIIII H asoeo
. uuuu mmmfloeflom mEomnmm
. IIIIII mumwzomEEH mebememowq
. nnnnnn mweOum EzwmoQoeHHU
. IIIIII mfiaomH>meQ EzfionoeHHD

memQEeoer EDHUOQOCHHQ
HHmENmQ E .
D HUOQOCHND

134

 

 

_oHOHH IIIIIIIIII mUmeHe mEomUmm

HOHOHH IIIIIIIIII mumHoHHwU mEomUmm
”OHOHH IUHGUHUUH4 . H asoeo
HQHQHH uuuuuuuuuu mmbwoeflom mEomUmm
HOHQHH 1111111111 mumwsumEEH mebememUwQ
HOHOHH_ IIIIIIIIII mweOMm Sawmomoeflao
HOHOHH uuuuuuuuuu mflwowfl>mea Ezwnouoewao
HOHOH_ IIIIIIIIII memeESUOHQ ESHUOQoeHHU
HOHOHH IIIIIIIIII HHmEHmQ ESwUOQoeHHU
HOHOHH llllllllll Enemowme EzflmoQoewfib
HoHOHH IIIIIII 0H4 EzEmumouomE Ezwboaoeflwb
HoHoHH IIIIIIIIII m Humbemm ESHUOQOeHNU
moHoHH uuuuuuuuuu 4 wembemo EnHmOQoeHHU
HOHOHH llllllllll m Hemwbemeu Eswmoaoeflwb
HQHOHH IIIIIIIIII EzwzomswmonQ .em>
flmezoen EzwmoQoeHHb
HOHQHH IIIIIIIIII 4 HmeEOMQ ESHUOQOCNND
flQHOHH uuuuuuuuuu 4 chmm EDHUOQOCHHU
HOHOHH IIIIIIIIII 4 Hmnmm EzflvoaoeHHo
hoHOHH IIIIIIIIII musmewe mwwflszNm
HQHOHH IIIIIIIIII mflmem>em moewu4
HOHQHH IIIIIIIIII mHHowwbezuou meuemz
fioHoHH IIIIIIIIII meweoHummE m:E%e&
HOOOHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ............ llllll m MDMMflHHU COUOUOCK
HOOOHH llllllllllllllllllllllllllllllll ............ llllll fl WSMQHSOQM COUOUOCK
HOOOHH IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ............ IIIIII Ezoweeowflwmu EzEmfiuemeoxm
HoooS IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII ........... IIIIII m memuew meueHEOHwom
H000.: IIIIIIIIIIIIIIIIIIIIIIIIIIII 4B4............. IIIIII 4 memoefl mfiuQHEOHMOm

HQQOHH uuuuuuuuuuuuuuuuuuuuuuuuuuuuuu oe............ nnnnnn muHuHeUemm meuefiEonom

135

 

 

HQHOHH IIIIIIIIII m msumwwflo embobocm

HQHOH_ IIIIIIIIII 4 mzumazuem eonomoem
HOHOHH IIIIIIIIII EzofleeomHHmo E3Eme0emeoxm
HoHoHH IIIIIIIIII m memuefl meueHEowflom
HQHOHH IIIIIIIIII 4 memuefl meueHEOHHOm
HOHOHH uuuuuuuuuu muwuwememb meueHEowaom
404040 uuuuuuuuuu 4 muemEmumsn m40e4504404
HoHoHH nnnnnnnnnn mmmwoeflw mwwmmemeoz
HoHoHH IIIIIIIIII meowmmeuemE muemeoz
HOHQHH IIIIIIIIII mmowzumwm mbumeoz
HQHOHH 1111111111 mumwzequm mwoxwoemnmmm
HOHOHH_ IIIIIIIIII eonomoeh mwoxmoeQOmm
HOHOHH IIIIIIIIII mbfiuwe mwuxmoqummm
HQHQHH IIIIIIIIII m mflHOMHMmE mflmxmoqummm
HOHOHH 9090090094 m mHHHsz mEomUmm
HOHOHH IIIIIIIIII 4 mHflfisz mEomUmm
HoHoHH IIIIIIIIII m mmbwowmmwsm mEombmm
HOHOHH llllllllll mumeQwQ mEomUmm
HOHOHH IIIIIIIIII 4 HemEHmQ mEomUmm
HOHQHH IIIIIIIIII 4 meme mEomUmm
HOHOHH nnnnnnnnnn memueoE mEommmm
HOHOHH IIIIIIIIII HH msoe0
HOHOHH IIIIIIIIII 4 mHUmE mEombmm
HOHOHH llllllllll 4 memweobemE mEmemm
HQHOHH uuuuuuuuuu mmezusm mEommmm
HoHoHH 9090090094 “weepmeeom meombmm
HoHoH. uuuuuuuuuu kuew>ew mEomUmm

HOHOHH nnnnnnnnnn mHHomNQOmmAe mEombmm

136

 

 

Hm40
4440
Hmvg
4440
4440
Hm4_
4440
4440
4040

4440
mmvg
Hmva
Hmvg
Hmv_
H440
Hmwa
HmvH
HmvH

4440
4440
4440
4440
fimva
Hmvg
4440

I I I I I I I I

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I
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9400444499000000009949900444004440099999444 .....

mbezuzh
44e00mee0h
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mwfiowfiuommxe
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mmbfloewom

mEommmm
mEomUmm
mEombmm
mEomUmm
mEomUmm
mEomUmm
H @5040
mEombmm

mumwzumEEH mubememowe

mwe00m
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HMmEHmQ
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m Humbemm

4 Nemeemm

m flHmeemeo

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Esflbomoeflw0

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0:50:00 $0320.30 05 no Scam 4.54423 "n 5..qu mucmzaom 02.9.4

137

 

meH
40mg
Home
~0m_
_0m0

Hmvg
4440
4440
4440
Hmva
4440
Hmva
4440
4440
Hmvg
Hmva
4440
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000000000000000000000000

 

4 405mm EDHUOQoeHHO
muzme4c 44444em44
mflmem>em moewo4
mHHOMNUezuok mguemE
mefleowummE szAe9

m mzumeHu eOUOUQem
4 mzumwzmem eonouoem
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m memoew meueHEoHflom

4 memoew meuewEOHflom
mowuwebemb maueHEowflom

4 muemEmumze maueHEOHHOm

mmbwoeww mwwmbemeoz

mwaowmmeuemE mUHmeOZ
mmo~zumww mbnmeoz
mumwzeumum mHDXNOMQOmm
eOUOerH mfloxmoummmmm
mmwuwe mfloxmoemmmmm

m mHHOMHHmE mwmxmoeQOmm
m mwwflmzm mEomUmm

4 mwwwmzm mEomUmm

m mmmwoNomwzm mEomUmm
muwemQHQ mEommmm

4 HHmENmQ mEomUmm

4 meme mEomUmm

memueoE mEomUmm

HH Q5040

4 memE mEomUmm

4 memHeOUemE mEombmm

138

40mg
40mg
90mg
40mg
40mg
40mg
40mg
40mg
40mg
40mg
40mg
40mg
Roma
Homa
HomH
40mg
40mg
40mg
40mg
40mg
momH
40m.
40mg
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4000
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40mg

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4 4444mze

m mmbwowomwsq
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4 meme
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mEombmm
mEombmm
mEomUmm
mEomUmm

HH msoe0

4 mfime

4 memHeomemE
mnezozh
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mfiwomflaommxe
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mmbwoewum

mEombmm
mEomUmm
mEombmm
mEommmm
mEomUmm
mEombmm
mEombmm
mEomUmm
H asoe0
mEomUmm

mumHzomEEH mebememoHQ
mHe00m Eswbomoeww0

mflwomfl>mefl
memeESUOHQ
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m Humbemb

4 Humbemu

m Hemwbemeu

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4 Hmeaohn
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139

 

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vaH0
vaH0
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40mg
40mg
40mg
Hmmw
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me0
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40mg
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Ezemowme ESHUOQoeflH0

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m Humbemm Ezflmoqoewq0

4 HMmUemu Ezwbouoeww0

m Hemwbemeu Ezwbomoew~0
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4 Hmezoen EswnoQoewH0

m 405mm ESHUOQoeHH0

4 Hmemm EDHUOQOCHH0

muzmefle mHNHeQmNm

mwmem>em moeH04

mwwomHUezqu meuemE
mefleuwummE szAe9

m mzumwwflo eOUOerm

4 mzumwzoem eomonoem
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m memoefl meuewEOwwom

4 memuefl meueHEOHHOm
mowuflebemn meueHEoflHom

4 muemEmumzn meueHEONHom
mmbwoewa mNHmmeeoz
meommmeuemE mnemeoz
mmowzumwm mbnmeoz
mumwzequm mwbkmoeQOmm
eononoee mHDXNoeQOmm
mbwuwe mwomwoeQOmm

m mHNOMmeE mflOANOMQOmm
m mwwwmzm mEomUmm

140

viH_
vaH_
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4 muemEmumsa me0e4E04404
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m0404e 44044040Q004

m m440444mE m404404mmmmm
m 4444mse

4 4.444434

m m004040040Q
m0440Q4Q

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mE00004
mEomUmm
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m500004
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HH Q5040

mEomUmm
mEombmm
mEombmm
msombmm
mEomUmm
mEombmm
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mE00004
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00040e40m mEombmm
mumwnumEEH m40em4004Q
m4e00m 82400Q0e440
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141

 

HNmHH
HNmHH
HNmHH
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4mm:
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m04Q044 mEombmm
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mmbwoewom mEombmm
m0m450mEE4 m40em4004Q
m4e000 E5400Q0e440
m44044>044 E5400Q0e440
mchE5004Q E5400QOCH40
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m 4400em0 5:400Q0e440
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m 44040eme0 85400Q0e440
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4 40e2044 E5400Q0e440
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m mzum4440 e0000044

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142

 

 

HOVNH

”mmHH
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me4£040mmE 05E4e9

m m50m4440 e0000044
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m mem0e4
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mEomUmm
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HH Q5040

mEomUmm
mE00004
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143

 

Hovm_
Hovmg
Hovmg
Hova
mova
HOVNH
HOVNH
HovNH
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404N0
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404m0
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4 meme mEombmm
mem0e0E mEombmm

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4 m4005 mEombmm
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H Q5040

mmnwoewom mEomUmm
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144

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fiovNH
Hovma
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m flumwbcmzo Eswonocwfib
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Hmcaoun Ezwonocwab

4 wmczoun EzflonOCHHQ
m Hmnmm ESHUOQOCHHU

4 Hmcmm ESHUOQOCflHU
muzmuflc mflaflaqmam
mwmcm>um mocflo4
wwwomfibczuOH mnucmz
.mcflgowummE mzsxzh

m mzumwwwo zebonozm

4 mzumwzocm comebozm
Ezoflcuowwamo EzEmcucmcuxm
m mcmuzfl maucwEOHwom

4 mcmocw mzuCHEOHHOm
mowuflubcmb mcucwEowwom

4 wucmEmumzfl mxucwsoflaom
mmbflozflw mwambhmcoz
mwaowmmcucwE mbumcoz
mmo~zumwm mbumcoz
wumazaummm mwuxmoqummm
COUOUocM mfibxmoanmmm
mbwUH: mfluxmonmmwmm

m wwwowwums mwmxmoummmmm
m mwwfisz mEombmm

4 mwawmzq mEombmm

m meHOHomst mEomUmm
muHMwaQ mEomUmm

4 meEflmQ mEowUmm

145

Hmmmg
wamu
Hmmma
ammmg
Hmwmg
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Hmwmu
mema
Hmmmg
meNH
Hmmmg
HmwNH
HmwNH
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ammma
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m mmbHoHomfizQ mEomUmm

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4
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mam: mEombmm
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HH msouw
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4 memHCOU:mE mEomUmm

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HHEOHm
ku:

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mHHOMwQOmmxn mEomUmm

mUHQmwc mEomUmm

mumwo

mmbflo

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146

mmmma
momma
flmmmg
mwmma
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m mzuMfiHHo COUOUocm

4 msumwzucm :0U0b044
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m memocfl mzucwEoflwom

4 mcmucfl maucwEofiwom
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4 wucmEMHmzn mzuCflEOHHOQ
mmbflocwa mHHmUMmcoz
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147

HommH
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4 mzumwzmcm :0U0boam
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m memocw mzu2HEoHHom

4 mzmocfl mauCHEOHHom
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4 mnemEmusz maucHEOHHom
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mmOqumwm mbnmcoz
wumH34qum mwuxmommmmmm
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mbwuw: mwuxwoammwmm

m mHHOMHMmE mfloxmoummmwm
m MHHHmSQ mEomUmm

4 mHHHsz mEombmm

m mmbwowomHzm mEomUmm
muHHmQHQ mEomUmm

4 HumEHmQ mEomUmm

4 mem: mEombmm

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4 mHUmE mEomUmm

4 mcmHCOUCmE mEomUmm
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148

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4 mHUmE

4 mcmwzoncms
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..........4O@40....UU...UH.H IIIIIII ..........
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149

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vamH
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vamH ..........4004U....UU...UH.H IIIIIII .....
..........4UD<U....UU...UB.H IIIIIII ..........
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..........40040....UU...UB.B IIIIIII ..........

EszumswmoHHQ .Mm>
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4 Hmcaomn EuflonOCHHU
m Hmnmm ESHUOQOCHHU
4 Hmnmm ESHUOQOCHHU
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mwmcmbum mocwum
mHHOHHbczuon maucmz
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m msumHHHu cononocm
4 mSHMHsbcm GOUOUOQm
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m MCMUQH MQHCHEOHHOH

4 mamUCH maucwEOHHom
wwwuwubcmb mcucHEOHHom

4 mucmEmwmzn mnucflEoHHom

mwnwocfla mHHmbumcoz
mHHommmaucmE‘mbumcoz
wumH34qum mwuxwoummmmm
:0U0b04h mwoxwoqummm
mbfluw: mwuxmoummmmm

m meOHHMmE mwuxmoumummm
m mHHHsz mEomUmm

4 mHHHsz mEombwm

m mmbwowbmHzQ mEombmm
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4 HHmEHmQ mEomUmm

4 mam: mfiomnmm

mcmucoE mEombmm

150

 

Hmmva
Hmmv.
Hmmva
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HvaH
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mmmvH
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COUOUOSM mHoANoqummm
mnwuflc mwu9N0HQOmm

m mflHOMmeE mwoxmoqummm
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4 mwaflmnm

m mmUHOHOmHBQ
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4 HHmEHmQ

4 meme
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4 mHUmE

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mUCzUSh
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mmbwozwom

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wumwzomEEw muncmnmowm

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4 Humbcmb

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151

 

 

HOWVH lllll ...........o.....o.......o................. QHQOUQ ESHUOQOCHHU
Homva lllll ........................................... MflNONH>mNQ ESHUOQOCNNU
How?“ lllll ........................................... WC®QESUOHQ EDHUOQOCHHU
mowVH lllll ...........................o............... HHmsfimQ EDWUOQQQHNU
momvH lllll ........................................... ESCQUHX®E ESHUOQQCHNU
H0m¢g lllll ........................................... ESEGuWOHUQE EZflUOQOQHHU
HOQVH lllll ......o.................................... m flHQUQQU ESHUOQOCHHU
momva lllll ........................................... m flHmUCWU ESHUOQOCHHU
momvH lllll .................o....................o.... m flHmHUQWCU EDHUOQOCHNU
HOmvH lllll ........................................... ESNDUWBHWONHQ .HQ>
flmczonn Ezwnomocwfib
HOQVH lllll ................o.......................... fl N®Q§OHQ ESNUOQOQNNU
HowvH lllll ........................................... m Hmnmm EsflnoaocflHu
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152

 

 

Esuflcmowwfimo Ezmeucmcoxm
m mEMUCH maucflEowwom

4 mCMUCw mCMCHEOflHOm
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4 wucmEmumnn maucwEOHwom
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mwwommmxucmE
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mbummoz
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mumaznummm mwoxmoqummm
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153

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mEomUmm
mEomUmm

HH Q5040

mEomUmm
mEomUmm

155

 

445mg
Honmw
$92
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$92
409mg
Hobmg
Hmbm_
Hmbma
Hmnmg
Hwhmg
Hobmg
409mg
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4 HMmEHmQ mEomUmm

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HH asouo

4 mHUmE mEomUmm

4 mcmHQOUEmE mEombmm
mU23039 mEomUmm
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156

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Honmg
Hmbmg
mmbmg
HohmH
Hohma
409mg
Hobma
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449mg
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449mg
409mg
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4

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157

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mEomUmm
mEombmm
mEomUmm
msombmm
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158

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162

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165

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166

 

 

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172

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