Title: Phylogeny and biogeography of Elatostema (Urticaceae) from Mount Kinabalu, Sabah, Malaysia
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Title: Phylogeny and biogeography of Elatostema (Urticaceae) from Mount Kinabalu, Sabah, Malaysia
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Language: English
Creator: Beaman, Reed S
Publisher: State University System of Florida
Place of Publication: Florida
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Publication Date: 2000
Copyright Date: 2000
 Subjects
Subject: Urticaceae -- Kinabalu, Mount (Sabah)   ( lcsh )
Botany thesis, Ph. D   ( lcsh )
Dissertations, Academic -- Botany -- UF   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
bibliography   ( marcgt )
theses   ( marcgt )
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Summary: ABSTRACT: Methods from phylogenetic systematics and cladistic biogeography are combined with geographical information system (GIS) and remote sensing technology to develop a novel approach to biogeography at a landscape scale, bridging the gap between traditional perceptions of ecological and historical biogeography. This study focused on determining phylogenetic and biogeographic relationships between 23 species of Elatostema (Urticaceae), eight of which are new, from Mount Kinabalu, the highest mountain in Borneo at 4094 m. Kinabalu is a biodiversity and endemicity hot spot of global significance with ca. 6000 vascular plants species. Factors contributing to biodiversity and endemicity include the extensive elevation and concomitant climate variation, diverse habitats including those on ultramafic (serpentine) outcrops of ophiolitic origin, the Miocene and late Pliocene uplift of the Kinabalu granitic pluton, Pleistocene glaciation, and Borneo's history of tectonic accretion. Phylogenetic relationships between the taxa were applied to a cladistic biogeography analysis. The approach involved defining the areas in the analysis as satellite image classes using a maximum likelihood supervised image classification and class probability analysis using Landsat TM imagery of the area. Image signatures for each class represented 12 habitats at varying elevations and geology. The assumption was made that the image classes represent ecological entities with a basis in the historical geology of Mount Kinabalu, Borneo, and its neighbors.
Summary: ABSTRACT (cont.): Speciation, extinction, dispersal, and failure to vicariate were accommodated by the cladistic biogeography analysis. Vicariant speciation is hypothesized to have resulted from the geologically recent emplacement and uplift of the Kinabalu pluton, habitat displacement during Pleistocene climatic changes, and population fragmentation due to the cleaving action of glacial melting to form deep river valleys. Dispersal may have taken the form of Bornean tectonic accretion of ophiolitic fragments originating in New Guinea, rafting not just into present day Sulawesi and the Philippines, but also into Borneo. Results indicate that recent speciation has occurred primarily in taxa occurring at mid and high elevation habitats. Species of Elatostema on low elevation habitats, both ultramafic and otherwise, are phylogenetically basal, suggesting that adaptation to ultramafics may have occurred early in the phylogenetic history of Elatostema, long before Kinabalu arose.
Summary: KEYWORDS: Urticaceae, Elatostema, biogeography, phylogeny, GIS
Thesis: Thesis (Ph. D.)--University of Florida, 2000.
Bibliography: Includes bibliographical references (p. 255-262).
System Details: System requirements: World Wide Web browser and PDF reader.
System Details: Mode of access: World Wide Web.
Statement of Responsibility: by Reed S. Beaman.
General Note: Title from first page of PDF file.
General Note: Document formatted into pages; contains ix, 264 p.; also contains graphics.
General Note: Vita.
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Source Institution: University of Florida
Holding Location: University of Florida
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Resource Identifier: oclc - 45825589
alephbibnum - 002566158
notis - AMT2439

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PHYLOGENY AND BIOGEOGRAPHY
OF ELATOSTEMA (URTICACEAE)
FROM MOUNT KINABALU, SABAH, MALAYSIA














By

REED S. BEAMAN


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


2000































Copyright 2000

by

Reed S. Beaman































For Ethan and Sydney
















ACKNOWLEDGMENTS


I thank everyone who provided research material, assistance, helpful advice, or

encouragement during the tenure of this work. Most notable is Walter S. Judd, my

committee chair, who has been my primary mentor for many years. I am also very

grateful to the others who have served on my committee, John Atwood, Dana Griffin, Jon

Reiskind, Scot Smith, and Norris Williams. Stephen Mulkey, although not a member of

my committee, provided food for thought. Alexis Thomas furnished the rectified digital

satellite imagery that is featured prominently throughout this document. The original raw

imagery was kindly donated to him by the EOSAT corporation.

The staff of the University of Florida Herbarium (FLAS), particularly Kent

Perkins and Trudy Lindler, provided essential assistance, without which this project

would have been impossible. The staff at A, BM, GH, K, MSC, SAN, UC, and US have

been extremely helpful, providing access to specimens and some work space. I also thank

those in the Botany Department of the University of Texas, particularly Robert Jansen,

Beryl Simpson, and Todd Barkman, for a semester of training in molecular systematics.

Mark Whitten and Norris Williams have also been extremely helpful and supportive in

my attempts to obtain nucleotide sequence data.

Many individuals have generously provided assistance, transportation, and

lodging during trips to Sabah. Datuk Lamri Ali, Francis Liew, Jamili Nais, Remi Repin,

Ludi Apin, Alim Biun, and many others from Sabah Parks made my work more

productive and enjoyable through their help, hospitality, and companionship.
iv









Special thanks to my wife, Nicoletta Cellinese, for her encouragement and

tolerance, especially in these last few months. I also wish to thank Peter Alcorn, who has

always been ready to help in a pinch. Natasha Akopian provided some much-needed last-

minute help in preparing the manuscript. My employer, Andre Clewell, has been

extremely supportive throughout this entire endeavor, as has my co-worker Marion

Lasley. I also thank my parents, John and Teofila Beaman, for their life-long support and

encouragement.

Finally, I thank the National Science Foundation (Grant DEB-9400888), the

MacArthur foundation, and the American Society of Plant Taxonomists for their generous

financial support.
















TABLE OF CONTENTS

A C K N O W L E D G M E N T S ........................................................................................... iv

ABSTRACT ............ ............................. ............... viii

CHAPTERS

1 M O U N T K IN A B A LU ................................................... .............................. 1

Introduction ..................................................................... . . 1
T he F lora.................................................................. . 5
Geography ........................................ 7
Geomorphology .............. ................... ........ .................... 9
G e o lo g y ................................................................................................................ 1 0
C lim ate ........................................................ ......... ..... 16
Elevational Zonation .. ..... ................................................... .............. 17

2 GEOGRAPHICAL INFORMATION SYSTEM ............................................... 29

S satellite Im ag ery ...................................................................... 3 0
Predicting Species Ranges using Digital Elevation Models ............................. 32
D diversity A nalyses through G IS ........................................ ........................ 33

3 PHYLOGENY OF THE URTICACEAE...................... ..... ......... ..... 42

In tro d u ctio n ................................. ........................................ 4 2
The U rticales .................... ................. ............................... 42
The Urticaceae ................................................... ...................... 45
Methods ............................................. 46
Results ............ ............................... ............... 47
Discussion ............................................ 50

4 PHYLOGENY OF KINABALU ELATOSTEMA ............................................... 89

In tro d u ctio n .......................................................................................................... 8 9
Elatostem a in M alesia .......................................................... 89
Generic Delimitation and Monophyly of Elatostema ................................. 89
M methods ......................................... ... ... ..... ................ .......... 90
Character and Character State Categories and Clarification ...................... 91










R results ................................... ........ ............ .................. ........... 95
Interpretation of Selected Clades. ......................................... .............. 96
Interpretation of Selected Characters ............................................................ 98
D iscu ssion .. ................................................... ............... . 10 1
Conclusions ............ ............................ ............... 104

5 BIOGEOGRAPHY OF KINABALU ELATOSTEMA .................................. 147

In tro d u ctio n ................................ ........................................ 14 7
D elim itation of A reas ................................................................................. 150
Phylogenetic and Cladistic Biogeography ................................. .............. 151
M materials and M methods ................................ .. ......................................... 153
Delimitation of Classes ..................... .............. ...... ......... 153
Im age Classification ...................................... ............. 154
Results and D discussion ............................... ......... .......... ............. 156
Im age Classification ...................................... ............. 156
Cladistic Biogeography ... .................................................................. 160
Speciation ... ....... ............................... ........... 162

6 TAXONOMIC TREATMENT ................................. 176

Introduction ............................................ .......... 176
Key to the Species of Elatostema ..................................... ............... 179

REFEREN CES ......... ............................. ...... .......... 255

BIOGRAPH ICAL SKETCH ....................... ......................................................... 263

















Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

PHYLOGENY AND BIOGEOGRAPHY
OF ELATOSTEMA (URTICACEAE)
FROM MOUNT KINABALU, SABAH, MALAYSIA

By

Reed S. Beaman

May 2000

Chairman: Walter S. Judd
Major Department: Botany

Methods from phylogenetic systematics and cladistic biogeography are combined

with geographical information system (GIS) and remote sensing technology to develop a

novel approach to biogeography at a landscape scale, bridging the gap between traditional

perceptions of ecological and historical biogeography. This study focused on determining

phylogenetic and biogeographic relationships between 23 species of Elatostema

(Urticaceae), eight of which are new, from Mount Kinabalu, the highest mountain in

Borneo at 4094 m. Kinabalu is a biodiversity and endemicity hot spot of global

significance with ca. 6000 vascular plants species. Factors contributing to biodiversity

and endemicity include the extensive elevation and concomitant climate variation, diverse

habitats including those on ultramafic (serpentine) outcrops of ophiolitic origin, the

Miocene and late Pliocene uplift of the Kinabalu granitic pluton, Pleistocene glaciation,

and Borneo's history of tectonic accretion.













Phylogenetic relationships between the taxa were applied to a cladistic

biogeography analysis. The approach involved defining the areas in the analysis as

satellite image classes using a maximum likelihood supervised image classification and

class probability analysis using Landsat TM imagery of the area. Image signatures for

each class represented 12 habitats at varying elevations and geology. The assumption was

made that the image classes represent ecological entities with a basis in the historical

geology of Mount Kinabalu, Borneo, and its neighbors. Speciation, extinction, dispersal,

and failure to vicariate were accommodated by the cladistic biogeography analysis.

Vicariant speciation is hypothesized to have resulted from the geologically recent

emplacement and uplift of the Kinabalu pluton, habitat displacement during Pleistocene

climatic changes, and population fragmentation due to the cleaving action of glacial

melting to form deep river valleys. Dispersal may have taken the form of Bornean

tectonic accretion of ophiolitic fragments originating in New Guinea, rafting not just into

present day Sulawesi and the Philippines, but also into Borneo. Results indicate that

recent speciation has occurred primarily in taxa occurring at mid and high elevation

habitats. Species ofElatostema on low elevation habitats, both ultramafic and otherwise,

are phylogenetically basal, suggesting that adaptation to ultramafics may have occurred

early in the phylogenetic history of Elatostema, long before Kinabalu arose.

















CHAPTER 1
MOUNT KINABALU



Kinabalu is a scenic wonder, a test for mountaineers from the amateur to
the skilled rock-climber, a holiday from the hot lowlands, and botanically
a paradise. The people of Sabah possess on this famous mountain what I
believe is the richest and most remarkable assemblage ofplants in the
world.
E. J. H. Corner (1978, page 113)


Introduction

The purpose at hand is to examine the phylogeny and biogeography of the 23

species ofElatostema Forst. (Urticaceae) known from Mount Kinabalu (Figure 1-1) in

Sabah, Malaysia. Kinabalu is located near the northern tip of Borneo, and is the tallest

mountain in Southeast Asia, between the Himalaya and New Guinea (Figure 1-2). With

an elevation of 4,093.7 m, this isolated massif stands 1000 m taller than any other in

Borneo. Kinabalu is, in a sense, an isolated alpine island. Barkman et al. (1997)

developed this analogy and the concept that the theories of island biogeography may be

applied to isolated montane floras such as that on Kinabalu. Earlier workers, including

Stapf(1894), Corner (1964), and van Steenis (1964, 1967) recognized the significance of

the Kinabalu flora. Van Steenis pioneered biogeographical work in the region by

assessing relationships between Kinabalu and neighboring mountains based on their

distance and number of shared species and genera.









2

Elatostema species are abundant on Kinabalu in herbaceous undergrowth,

especially along rivers and streams and near waterfalls. Some species occur on exposed

ridges and slopes. Friis (1993) estimates there are 300 species of Elatostema, but notes

that the genus is in need of study. Few species of Elatostema have been studied beyond

their original descriptions or listings in floristic treatments. The total number is probably

higher because many areas in the montane paleotropics, particularly in Borneo and New

Guinea, remain poorly explored. Shroter and Winkler (1936) monographed 106 species

of Elatostema, including six Kinabalu taxa. Their monograph included none of the 14

species of Elatostema known only from Kinabalu. The other Kinabalu species appear to

be restricted to Borneo, with the exception of E. acuminatum Brong., which occurs in

Java, Sumatra, and southern India. Descriptions of eight new species of Elatostema from

Kinabalu are included in this dissertation.

This study was conceived as a means to elucidate and quantify evolutionary

mechanisms pertaining to the high floristic diversity on Kinabalu. With as many as 6000

species of vascular plants on its slopes (Beaman and Beaman 1998), Kinabalu ranks

among the global hot-spots for biological diversity. Beaman and Beaman (1990)

hypothesized that the high diversity on Kinabalu is the collective result of several factors.

Geological, geomorphological, and ecological factors addressed herein include (1)

diversity of habitats due to elevational variation from lowland rainforest to alpine scrub,

(2) precipitous topography, often opening up new habitat owing to landslides, (3)

geological diversity and patchiness (particularly of ultramafic outcrops) resulting in

highly localized edaphic conditions, (4) historical geology and tectonic history of

Kinabalu, the rest of Borneo, and the Malesian floristic region, and (5) climatic











oscillations such as those during Pleistocene glacial and inter-glacial cycles, as well as

more recent oscillations influenced by EINiho events (Beaman et al. 1986).

An underlying assumption is that these factors have charted present day species

distributions on a geologically very recent mountain. How and to what extent do they

contribute to speciation and high diversity? While that question may never be answered

explicitly, it is possible to discern patterns in species relationships that bear on the issue

of speciation. The technical goal of this study is to determine whether patterns in the

phylogeny and distribution of species are congruent with discernible spatial attributes and

patterns in the landscape. Of special interest are the phylogenetic and biogeographic

relationships between very localized species and their habitats. The Kinabalu Elatostema

make an appropriate choice as a research taxon primarily because of the occurrence of

endemic species at mid and high elevations and on ultramafic substrates. Table 1-1 lists

the species studied and their general distributions.

Using geographical information system (GIS) and remote sensing tools, analysis

and visualization of phylogenetic and biogeographical data are greatly enhanced.

Figures 1-3 and 1-4, showing a Landsat TM image of the Kinabalu area and a three-

dimensional terrain model combined with an image drape, while visually striking, are

only a glimpse at the digital data contained therein. Remote sensing and three-

dimensional terrain modeling provide an opportunity to make localized analytical spatial

queries on ecological and physical characteristics of the places where species and clades

occur. In Chapter 2, a more detailed description of the Kinabalu GIS that forms the basis

for the spatial data set is provided.











On Kinabalu, endemic and sometimes closely related sympatric species occur in

very local areas. Spatial patterns or correlation between remotely sensed, terrain defined

data and phylogeny are explored for the Kinabalu Elatostema. Barkman et al. (1997)

addressed similar questions for the genus Dendrochilum (Orchidaceae) using a molecular

based phylogeny. An approach based on developing hypotheses of phylogenetic

relationships and cladistic biogeographic relationships between areas on Kinabalu is

developed to address issues of the origin, distribution, and diversity of Kinabalu

Elatostema, and of the mountain's flora in general. Specific questions formulated to

address the issues stated above include the following:

* Did higher elevation endemic species on Kinabalu evolve from lower elevation

species as uplift and Pleistocene glacial/interglacial cycles drastically and dynamically

altered habitat and increased natural selection pressure? Elatostema bulbothrix Stapf

is a Kinabalu endemic found only on some of the highest elevation ultramafic

outcrops. What is the relationship of this rare species to other Kinabalu Elatostema?

* Does precipitous topography hinder gene exchange between isolated populations,

driving rapid adaptive radiation and speciation? If so, one would expect to see closely

related species with micro-disjunct area relationships.

* What role does the occurrence of ultramafic geology play in the large number of

endemics?

To answer these questions, a phylogeny of Kinabalu Elatostema based on

morphological characters was constructed. Elatostema is especially rich in vegetative

characters, particularly in the form and distribution of cystoliths and trichomes on both

vegetative and reproductive structures. Floral and inflorescence characters also proved to









5

be quite informative. The phylogenetic analysis is presented in Chapter 4, with the

higher-level analysis in Chapter 3 providing the appropriate outgroups.

In Chapter 5, a biogeographical analysis to determine relationships between taxa

and areas is presented. Novel techniques for visualizing phylogenetic patterns, levels of

endemicity, and species distributions are employed. A cladistic biogeography approach is

also employed. Congruence between evolution of Elatostema and the geological history

of Kinabalu is evaluated. Modes of speciation via vicariance and long distance dispersal

are considered, as well as the roles that differing geological substrates, changing climatic

conditions and mountain orogeny play in speciation. In addition, the possibility exists that

high biological diversity in the Kinabalu region may be due to the intermingling of floras

resulting from continental accretion. This may be even more significant than long

distance dispersal. Finally, revision and taxonomic treatment of Kinabalu Elatostema is

presented in Chapter 6, including nomenclature, descriptions, diagnoses of new species,

and specimens cited.

General observations on the flora, geography, geology, geomorphology, climate,

and elevational zonation are described in the balance of this first chapter. For further

information on the natural history of Kinabalu the reader is directed to Wong and Phillips

(1996) who provide a recently revised compendium by authors in numerous disciplines.


The Flora

In a summary of global plant diversity, Barthlott et al. (1996) ranked northwest

Borneo, inclusive of Kinabalu, among the top ten biological diversity centers in the

world. They indicated that the area has more than 5,000 vascular plant species per









6

10,000 km2. Kinabalu actually may harbor 5,000-6,000 species in an area of only 1150

km2 (Beaman and Beaman 1998). More than 1,000 genera representing over 200 families

are known in the Kinabalu vascular plant flora. Roos (1997) asserted that the Malesian

vascular plant flora consists of about 42,000 species in an area of 3.1 million km2. Thus

the relatively small area (0.04% of Malesia) encompassed by Kinabalu may include as

much as 14% of the Malesian flora and 2% of the global flora of an estimated 300,000

vascular plant species. To place this in perspective, the flora of Florida probably

comprises about 1.3% of the global flora, and Europe about 3.7%.

Exceptionally high diversity provides a compelling argument for recognizing

Kinabalu as a biodiversity hot-spot. However, numbers alone do not make a botanical

paradise. Kinabalu serves as home to a number of non-urticaceous taxa of special

botanical significance. Brief mention of certain taxa is warranted here.

The largest family of flowering plants on Kinabalu is the Orchidaceae with over

643 species. Many are well known for their horticultural value and rarity (Cribb 1998,

Wood et al. 1993, Wood and Cribb 1994). Notable taxa include several species of the

well known genera Paphiopedilum and Dendrobium. Bulbophyllum, the largest orchid

genus on Kinabalu, includes over 87 species.

No less engaging is Rafflesia keithii Meijer (Rafflesiaceae), an achlorophyllous

root parasite, having flowers measuring over one meter in diameter. The flowers look and

smell like rotting meat which attracts carrion flies as pollinators (Beaman et al. 1988).

The genus Rafflesia is known for having the largest flowers in the world.

Npe,,i/we,\ rajah (Nepenthaceae) is a carnivorous plant whose pitchers are formed

from extensions of the leaf midrib, and the pitchers of this species may hold over a liter of









7

water laden with digestive enzymes, variously digested insects and even small mammals.

Numerous other pitcher plants species occur on Kinabalu, some of them endemic, and

many on ultramafic substrates.

The genus Rhododendron (Ericaceae) deserves special mention, primarily because

80% of the Kinabalu species are endemic (Argent et al. 1988). Other Ericaceae, such as

Diplocosia, are also abundant, with numerous endemic species on Kinabalu.

Other large families on Kinabalu include the Rubiaceae, Euphorbiaceae,

Melastomataceae, Lauraceae, Myrtaceae, and Araceae. The single largest genus on

Kinabalu is Ficus (Moraceae), with ca. 100 species. Floristic enumerations have been

completed for the ferns and fern allies (Parris et al. 1992) and gymnosperms (Beaman and

Beaman 1993, 1998) as well as for the monocotyledons (Beaman and Beaman 1998).

Sipman (1993) provided a checklist of lichens, listing 286 known species, and estimated

the actual number to be about double this value.


Geography

Low's Peak, the highest on Kinabalu, is a mere 40 km from the South China Sea.

The massif rises impressively from the surrounding foothills and the coastal plane (Figure

1-3). The oft-quoted 4,101 m (13,455 ft) elevation for this peak was determined by the

East India Shipping Company in 1910 (Jenkins 1996), but several of the peaks on

Kinabalu have been recently surveyed. In 1997, the Sabah Department of Lands and

Surveys, using carrier phase global positioning system (GPS) equipment, measured the

height of Low's peak at 4093.7 m.









8

Kinabalu Park, one of six parks in the Sabah Parks system, encompasses the

Kinabalu massif in an area of about 750 km2. The park also includes Mount Madeleng

(the name Tambuyukon is often misapplied to this peak), the third-highest (ca. 2540 m)

mountain in Sabah. Kinabalu forms the northern terminus of the Crocker Range (also

under protection by Sabah Parks), which parallels the west coast of Sabah.

A Kinabalu location map, reproduced here at reduced scale (Figure 1-5) and

gazetteer (Beaman et al., in prep.) are near completion. Botanical collection sites, as well

as locations of general geographical interest have been mapped. Much of this

geographical knowledge was obtained through interviews with the local Dusun people. A

large number of sites, especially historical collection localities, were known only to some

of the oldest individuals in their communities. Without much of this detailed geographical

knowledge, the micro-geographic specimen mapping employed in this study would not

have been possible.

Kinabalu is the only mountain above 2649 m in Sabah and the only one in Borneo

above 3200 m. Sarawak, Sabah's neighboring East Malaysian state to the south, has a

number of mountains and mountain ranges including Mount Murud, the Dulit Range, and

the Hose Mountains, but none of these approach Kinabalu's stature. Likewise, the interior

of Kalimantan (Indonesian Borneo) is rugged and mountainous (Figure 1-2) but not well

explored botanically. Numerous other mountains between 2500 and 3000 m elevation

occur in the neighboring Philippine islands, including Palawan and Mindanao, and in

Sulawesi (the Celebes), which is close to Borneo but on the other side of Wallace's line.











Geomorphology

Steep slopes abound on Kinabalu; razorback ridges and deep valleys make for

some very rugged terrain. This combination may foster effective geographic and

reproductive isolation of species over short distances.

The summit area is bisected by Low's Gully. Low's Gully formed along a thrust

fault (Jacobson 1970) that was sculpted at its upper end by Pleistocene glaciers. The

gully decends into what becomes the Penataran River. Together, they drop more than

3500 m over a four kilometer distance. Low's Gully exemplifies the precipitousness of

Kinabalu and it remains a serious challenge to explorers. In 1993, ten British Army

soldiers set out on a ten-day descent of Kinabalu through Low's Gully. It was concluded

tardily after a 12-day search and rescue operation used a helicopter to save five soldiers

trapped between waterfalls. The other five made it out on their own.

The highest peaks of the Mountain are west of Low's Gully. King George Peak is

the highest peak east of the gully and forms a pivot point between Eastern Ridge and

Northern Ridge. The northern slopes of Kinabalu are not well explored. Most botanical

exploration has been concentrated on and around the southwest slopes of Kinabalu

(Figure 1-5). This is exemplified by the higher concentration of named places in this area.

The southwest slopes have historically afforded the easiest ascent. Hugh Low was the first

European to reach the summit, in 1852. Granite crags line both of these ridges at their

upper elevations. The Eastern Ridge gradually descends to 600 m at Poring Hot Springs.

The Northern Ridge runs nearly level towards the north.

Five majors river systems originate near the summit of Kinabalu. The Northern

Ridge terminates where the headwaters of the Mekedeu River run to the east, and the











Wariau river runs to the northwest past Sayap and into the western coastal plain. The

Liwagu River originates on the southern slopes and ultimately flows out the eastern coast

of Sabah. The Kadamaian River cascades off the southwest slopes in a 1000 m high

waterfall. Numerous rivers, including the Penataran, join with the Kadamaian as it flows

north through the coastal plain. The Kadamaian becomes the Tampassuk River near Kota

Belud, the city from which the earliest expeditions up Kinabalu commenced.


Geology

Mount Kinabalu is one of earth's most recent major granitic plutons (primarily

composed of ademellite, with porphyritic margins). It was emplaced diapirically about 9

ma, cooling until 4.9 ma and uplifted in the last 1.5 million years (Jacobson 1970). It may

still be rising at a rate of between 3 and 5 mm per year. This granite core is exposed in

most areas above 3000 m. The summit area was much affected by the sculpting action of

glaciers during the Pleistocene, during which a 5 km2 ice cap formed. Deglaciation,

ending about 9,200 years ago, further eroded deep river valleys and deposited alluvial

sediments such as the Pinosuk gravels. Habitats above 3000 m were colonized after the

glacier retreated. Below the summit, a number of other substrates, including highly folded

and faulted Tertiary Crocker and Trusmadi sedimentary formations and Quaternary

Pinosuk gravels support a range of tropical forest communities.

Of particular interest are plant species and communities growing on the ultramafic

(also referred to as ultrabasic or serpentine) outcrops that occur at various elevations on

Mount Kinabalu. A higher proportion of endemic species are found on ultramafic

substrates than on non-ultramafics (Beaman and Beaman 1993, Parris et al. 1992, Wood









11

et al. 1993). Elatostema exemplifies this trend (see Table 1-1). More than half of the 15

Kinabalu plant communities recognized by Kitayama (1991) occur on ultramafics.

Plants growing on ultramafic substrates must adapt to low availability of nitrogen,

phosphorous, potassium, and calcium and high levels of iron, magnesium, nickel,

manganese, chromium, and cobalt. This is a toxic mixture to many plants. While there is

no evidence that any ultramafic adapted plants require high levels of heavy metals, some

species occur only on ultramafic outcrops. Competitive exclusion may be invoked to

explain why some of these ultramafic endemics do not occur on less toxic soils.

The stature of trees and shrubs on ultramafic habitats is often stunted, and is one

of the factors that makes these ultramafic patches quite recognizable with remote sensing

software. Serpentine (ultramafic) vegetation was noted by Kitayama (1991) to be

strikingly different from that of surrounding forests on non-ultramafic substrates. Not all

trees on ultramafics are stunted, however. The commercial value of some enormous trees

in the family Dipterocarpaceae and the genus Agathis (Podocarpaceae) endangers this

community at lower elevations.

Kitayama (1991) also indicated that there are at least three altitudinal subdivisions

of woody serpentine vegetation. The first subdivision is that dominated by Tristania

(Tristaniopsis) elliptica, the second by Leptospermum javanicum and Tristania elliptica,

and the third by Leptospermum recurvum and Dacrydium gibbsiae. Another ultramafic

vegetation type on which Kitayama did not comment are the forests dominated by

Casuarina sumatranum or C. terminal. These are particularly distinctive markers of

lower and mid elevation ultramafic soils. In addition, there are very interesting graminoid











ultramafic communities at Marai Parai and on the summit ofMt. Madeleng. These

graminoid communities often support the highest levels of local endemism.

The occurrence of ultramafics around Kinabalu is not anomalous for the region.

Ultramafic outcrops occur in Palawan, Sulawesi, New Guinea, as well as around the

volcanic belt known as the "Pacific Ring of Fire." These are active or previously active

zones of subduction of the spreading oceanic Pacific Plate beneath the Asian, Australian

and American continental plates.

Van Steenis (1964) concluded that there must have been a former mountain

connection between Asia and Australia, and that Kinabalu was a part of a "transtropical

bridge of temperate genera." Stapf (1894) held that Kinabalu must have shared an

"immediate connection with the highlands of New Guinea, or what was then its

equivalent," for the presence of Australian elements in the Kinabalu flora to be

intelligible.

In light of recent tectonic evidence (Hall 1996, 1999, Metcalfe 1999, Moss and

Wilson 1999), Stapf and van Steenis may have been more prophetic than we realized even

recently (Beaman 1996). Malesia is a tectonically complex region and the biogeography

is as convoluted and complex as that of the Caribbean.

Borneo, the Philippine islands, Sulawesi, New Guinea and other Indonesian

islands are situated in an area of striking tectonic activity at the juncture of the Eurasian

and Indian/Australian cratons, and the Pacific, Caroline and Philippine Sea plates.

Metcalfe (1999, page 25) described the area as "giant 'jigsaw puzzle' of continental

fragments bounded by major geological discontinuities... now huge strike-slip faults,

whereas others are actual suture zones that include remnants of oceanic crust (ophiolites),











oceanic and continental-margin sedimentary rocks, accretionary complexes, melange, and

sometimes volcanic arcs."

Ophiolites are typically ocean floor material thrust up onto and sometimes folded

into volcanic and sedimentary material. Brooks (1987) suggests that the ultramafics on

Kinabalu may be part of the Palawan Ophiolitic Belt. Palawan, a southern Philippine

island just to the north of Borneo is rich in ultramafics. It is a marginal fragment of

Eurasia that rafted eastward as the South China sea opened in the Mid Oligocene through

the Late Miocene, an event that coincided with the final uplift of Kinabalu. It was

submerged until the early Pliocene. Alternatively, the ultramafics, or at least the

ultramafic elements of the flora, on Kinabalu may have more closely shared a history with

those in Mindanao and Western Sulawesi, and as a result with New Guinea and Australia.

Ophiolites in Sabah crop out in a discontinuous S-shaped curve, with the Darvel Bay

Ophiolitic complex at the southeastern end and those on Kinabalu at the northwestern

end. The Darvel Bay ophiolites were already eroding by the Eocene (Omang and Barber

1996) and Brooks (1987) indicates these as related to the Mindanao ophiolites.

Borneo appears to be the result of dispersion of Gondwana marginal slivers and

later accretion of a number of these fragments from Paleozoic to Cenozoic times

(Metcalfe 1999). Continental blocks and fragments believed to contribute to Borneo

include South West Borneo, Semitau, Luconia, and Kelabit/Long Bowan fragments

derived from Cathaysialand, which was formed by the amalgamation of South China,

Indochina, and East Malaya in the Late Devonian to Early Carboniferous. These

fragments and parts of the Philippines basement separated from Cathaysialand opening up

the Proto-South China Sea in the Late Cretaceous. The Mangkalihat and Paternoster









14

fragments found to the east of the Adio and Meratus sutures in Borneo were derived from

Gondwanaland in the Late Triassic to Late Jurassic, along with West Sulawesi. These

fragments rafted from the northern margin of Australia in the Late Jurassic through the

Late Cretaceous. East Sulawesi, once contiguous with New Guinea/Australia (Metcalfe

1999), is shown by Hall (1999) as accreting to West Sulawesi in the Mid to Late Miocene,

a time when there may have also been a discontinuous volcanic arc connection to Borneo

via the eastern Philippine islands and Mindanao. Moss and Wilson (1999) indicate that

microcontinental blocks from the New Guinea-Australian margin that accreted onto

eastern Sulawesi in the Miocene or Pliocene may have remained emergent as they rafted.

Moss and Wilson (1999) infer a mountainous land connection between southern

Borneo and mainland SE Asia in the Eocene into the Pliocene, possibly dating as far back

as the Jurassic. They also suggest that dispersal may have occurred along volcanic arcs

between Borneo, Sulawesi, and the Philippine islands. Hall (1999) postulated that an

Eocene collision of India into Asia about 45 ma resulted in major mountain building and

resulting changes in climate and drainage systems in SE Asia. This event also allowed

dispersal of the Gondwana flora into Asia. The northward movement of the Australian

craton brought about contact between north Australia, New Guinea, Sulawesi, Borneo,

and the Philippines ca. 25 ma. The uplift of the central mountains of Borneo also occurred

around this time, and Borneo and the Malay Peninsula maintained connection along the

exposed Sunda Shelf through the Pliocene.

While much of the geological activity described above occurred before the final

uplift of Kinabalu, it was important in setting the stage for high levels of biological

diversity. The active orogeny of Kinabalu only 1.5-9 ma opened new niches, including









15

what is now the only high elevation habitat between the Himalaya and New Guinea. Also

important is the recognition that while the ultramafic flora on Kinabalu is represented by a

large proportion of endemic, possibly neo-endemic species, organisms have been adapting

to ultramafic toxicity since at least the Eocene. A cross-section diagram by Jacobson

(1970) shows the intrusion of the Kinabalu pluton occurring through both ultramafic and

sedimentary formations. Mount Madeleng, 24 km northeast of the Kinabalu summit, is a

predominantly ultramafic mountain. Madeleng may provide an indication of what the

Kinabalu area might be like had the final emplacement and uplift of the Kinabalu pluton

in the Pliocene never occurred.

Wallace's line (Wallace 1898) passes just to the east of Borneo. The position of

Kinabalu west of Wallace's line would suggest a Laurasian origin of the flora. However,

there are many representatives of the Australasian flora on Kinabalu. These include many

of the gymnosperms (i.e., Dacrydium, Agathis, Podocarpus) and many Myrtaceae (i.e.,

Leptospermum, Tristania and Tristaniopsis). Recent knowledge of the timing of tectonic

events along with application of cladistic biogeography in the Malesian region are

beginning to provide answers to the question of why some organisms may have crossed

Wallace's line and why others have not.

Dispersal from Australasia of Dacrycarpus imbricatus (Podocarpaceae) into

Sundaland is cited by Morley (1999) as a classic example of westward migration across

Wallace's line. Pollen of this species was common in the Lower Pliocene from offshore of

Sabah and Sarawak. The genus is typically restricted to SE Asian moist montane habitat,

and is represented by two species on Kinabalu, D. imbricatus and the endemic D.

kinabaluensis (Beaman and Beaman 1998).













Climate

Kinabalu supports extreme climatic ranges from tropical rain forests near sea level

to freezing alpine conditions at the summit. Stapf (1894), in the first floristic account of

the Kinabalu flora, made special note of temperate taxa occurring in the flora such as

Ranuculus lowii (Ranunculaceae) and Potentilla s. (Rosaceae). The closest relatives to

either of these are in the Himalaya and New Guinea.

Pleistocene climatic oscillations may have caused upwards and downwards

migrations of species during glacial and interglacial cycles. Glaciers extended as low as

3000-3100 m (Choi 1996, Jacobson 1978), about 1000 below the summit. A 1000 m

difference in elevation may be correlated with a mean temperature drop of 5.50 C on

Kinabalu (Kitayama 1992, 1994). Below the physical extent of the glaciers, this degree of

change in mean temperature would strongly select against many tropical cloud forest

plants. In addition, downward migrations may have isolating effects on some plant

populations. Besides having elevational restrictions, most plants on Kinabalu tend to be

restricted to ridges, stream valleys or slopes. In the case of a species occurring only on

ridges, when populations are forced downwards, those on adjacent ridges would become

more isolated, since ridges radiate as elevations decrease. The opposite effect would occur

with stream species, since streams conjoin as elevations decrease.

Recent El Niho events have lead to extreme short-term climatic instability

throughout Borneo. Beaman et al. (1986) documented El Niho related forest fires

occurring in 1983 in Sabah. Almost 10 % of the forested land of Sabah burned during this

event. Two other ElNiho events have occurred since, one in 1990 and another in 1998.











Both were associated with fires burning through primary forest within Kinabalu Park.

Many of these fires burned vegetation on ultramafic substrates. This vegetation type

seems particularly susceptible to drought. Many of the plants on ultramafics exhibit

characteristics of xerophytes (thick leathery leaves with waxy cuticles), but their ability to

tolerate fire is dubious. One of the characteristics of the El Niho droughts is that they are

often followed by unusually heavy rains. Numerous landslides have occurred on Kinabalu

following the El Niho events. While the wildfires may be only recently occurring on

Kinabalu because of human disturbance (and arson), ElNiho is not new, nor are

landslides. Many very interesting ultramafic species occur only on landslides.


Elevational Zonation

A vegetation map of Mount Kinabalu Park was published by Kitayama (1991) in

which 21 vegetation map units were recognized. Kinabalu Park does not include all the

area covered by this study, but enough is included to make the map useful for

understanding the vegetation of the entire Kinabalu massif. Kitayama indicated that

diagnostic canopy tree species could be used to distinguish vegetation zones. He

considered these species to be mutually exclusive in distribution and their occurrence

correlated with elevation. The upper boundary of lowland rain forest, where the majority

of emergent trees (mostly Dipterocarpaceae) disappear from the canopy, is at about 1200

m. The upper limit of lower montane forest is at between 2000 and 2350 m, and that of

upper montane forest at between 2800 and 3000 m, the latter particularly marked by the

upper limit of Lithocarpus havilandii. Above this level occurs a lower-subalpine

coniferous forest dominated by Dacrycarpus kinabaluensis, Dacrydium gibbsiae,









18

Podocarpus brevifolius, and angiosperms Photinia davidiana, Styphelia malayana, and

Leptospermum recurvum. The upper limit of this forest is at about 3400 m and

corresponds to the closed forest line. A fragmented upper-subalpine forest extends above

this level to the tree line at about 3700 m. Above this level is a summit area zone of alpine

rock-desert with scattered communities of alpine scrub. Kitayama suggested that the tree

line may coincide with the lowest elevation where nocturnal ground frost is frequent. He

noted that great variations in dominance type, species composition, and forest structure

occur within each zone. These were attributed to altitudinal temperature effects, soil

nutrient status in relation to topography (particularly ridge and valley differences) and

slope aspect.

Elevational zonation is so strong on Kinabalu that it may be used as the primary

basis for recognition of vegetation types. Below 500-600 m is what Kitayama refers to as

"substituted vegetation" and is meant to refer to lowlands that have long been subject to

human management and disturbance. "Hill forest" is a much used term for the upper part

of the dipterocarp-dominated lowland forest, and we have applied this to habitats between

the levels of about 600 and 1400 m. Kitayama considered the lowland forest to be six-

layered with emergent trees characteristic and undergrowth sparse. Lower montane forest

is in the elevational range of about 1200-2200 m, is five-layered, and does not have

emergents; upper montane forest has a dense herb layer. The lower subalpine forest is

sparser in undergrowth and lower in height. Upper montane forest is used here to refer to

all forest vegetation above the lower montane forest. The area above the continuous upper

montane forest is referred to as the summit area.

































Figure 1-1. Mount Kinabalu. Top: View of the upper south face from the West
Mesilau River, above the Pinosuk Plateau. Bottom: View of the southwestern
slopes showing ephemeral waterfalls after a rain storm.


















k-
























-e








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cE -65 -b
O
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~ ~CA
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CA .2 j
-e j
a o









F4 mrt CA



































































































I


1,*- ,


T





L .=
;L

































Figure 1-3. Satellite image map of Mount Kinabalu, based on Landsat 5
Thematic Mapper data from 14 June 1991. Bands 3, 2, and 1 are shown applied
to a red- green-blue (RGB) color model. Raw image data was rectified using
digital elevation model (DEM) data and global positioning system (GPS) ground-
control points. Original pixel values were maintained by this method, thus the
image may be used in further analysis. The Kinabalu Park boundary is shown in
red.

















\ ..un kin I l lu I i' l l I l l i -
..... . I\II

Lr V


















































Figure 1-4. Drape of the Landsat TM image of Mount Kinabalu over the DEM, as seen
from the northwest. This figure was produced using image bands 7, 2, 1. The Kinabalu
Park boundary shown in red is an additional coverage. Black holes in the draped image are
areas where surface visibility was miscalculated owing to three-dimensional surface-
resolution limitations. The surface resolution is 50 x 50 m while the image resolution is
28.5 x 28.5 m.
































Figure 1-5. Reduced scale (1:250,000) reproduction of Mount Kinabalu location
map showing place names of kampungs, peaks, rivers and botanical collecting
localities. The shaded relief map was produced by hill shading and color coding
by elevation the digital elevation model (DEM).






















NIMI I KIN %\ \I I
I O( 1O N N1 1 ,1'


Jtv-


27




















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S "* "r . "'*
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Table 1-1. Species of Elatostema included in this study, their general
distributions, and occurrence on ultramafic substrates. 'Species occurs
exclusively on ultramafic substrates. Undescribed species indicated by single
quotes around the specific epithet.


Taxon

E. acuminatum
E. 'auriculatifolium'
E. bulbothrix
E. 'bullatum'
E. 'dallasense'
E. flavovirens'
E. gibbsae
E. integrifolium
E. kabayense
E. kinabaluense
E. linear
E. lithoneuron
E. 'maraiparaiense'
E. pedicillatum
E. penibukanense
E. pinnatumm'
E. purpurascenss'
E. rubrostipulatum
E. 'serpentinense'
E. tenompokense
E. 1th,/i i it i k'
E. variolaminosum
E. vittatum
E. winkleri-huberti


Ultramafic

X
X
X'










X
X
X
X
X'
X





X
X



X
X


Distribution

Kinabalu, Borneo, Java, India
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu, Sabah
Himalayas
Kinabalu, Sarawak
Kinabalu
Kinabalu, Sabah
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu
Kinabalu, Sabah, Sarawak
Kinabalu, Sarawak
Kinabalu, Borneo
Kinabalu, Sabah
















CHAPTER 2
GEOGRAPHICAL INFORMATION SYSTEM


Development of a geographical information system (GIS) for Mount Kinabalu

was originally conceived to augment the botanical inventory of Mount Kinabalu, with a

location map and gazetteer, and as a basis for mapping species distributions. As the

technology has improved, GIS has become much more than just a mapping tool; it forms

a spatial framework for analysis of biological data. Though beyond the scope of this

dissertation, we are developing GIS tools to (1) predict where species occur in unexplored

areas based on occurrence in known localities, (2) develop diversity indices based on

surface areas and elevation, and (3) identify Itramafic habitat islands as part of an effort to

understand plant speciation patterns on Kinabalu. Identification of ultramafic outcrops is

of particular interest because of the correlation between these outcrops and distribution of

endemic species, particularly those hypothesized to have evolved on Kinabalu.

Coverages included in the Kinabalu GIS to date are topography, hydrography

(rivers and streams), the Kinabalu Park boundary, ground-control points, a vegetation

map published by Kitayama (1991), place names toponymss) relating to specimen

collections made since 1851 as well as by local collectors for the Projek Etnobotani

Kinabalu (PEK), the Landsat TM image, and a digital elevation model (DEM) was

developed from topographic coverages.

Predictions of where a taxon occurs in unexplored areas are useful in planning

expeditions into areas such as the still unexplored north side of Mount Kinabalu.

29









30

Likewise, park management may use this type of data to target certain areas for

protection. Predicting the occurrence of a particular taxon is based on knowing the

characteristics of its habitat. Sheila Collenette (pers. comm.) probably was the first person

to use remote sensing in this manner in her rediscovery of Paphiopedilum

1 i/l, hil li, i 1unl on Mount Kinabalu. Using black and white aerial photographs of

Kinabalu she was able to recognize the Casuarina s.l. forests that are characteristic of

many ultramafic areas. Site visits proved her technique worthwhile, because the rare

slipper orchid, P. nIti',hiihl inlnu, had not been known from the wild in almost 70 years.

The type collection had been (mischievously) attributed to New Guinea.

Remote sensing and GIS software (ERDAS Imagine and ESRI Arc/Info,

respectively) provide image enhancement and classification techniques that make these

sorts of predictions possible in a systematic way. Digital elevation models (DEMs) allow

further refinement of predictive models by the inclusion of elevation, slope and aspect

data. Diversity indices may also be enhanced by developing models to incorporate DEM

data into the diversity index. How species diversity relates to habitat diversity and the

spatial extent of certain plant communities is of particular interest on Mount Kinabalu,

because of the high endemicity of species, particularly in ultramafic areas.


Satellite Imagery

The satellite image used extensively in this study is a Landsat 5 Thematic Mapper

(TM) product, from an orbit ca. 700 km above the earth's surface. The image shown in

Figure 1-3 is a quarter of a quarter scene, i.e., one-sixteenth of the total scene that covers

185 x 185 km. This image, recorded 14 June 1991, is one of only two cloud-free images

available for the area, which is characterized by frequent and heavy cloud cover. The









31

nominal ground resolution is 28.5 x 28.5 m for each pixel. The data are provided from

seven bands of the electromagnetic spectrum, as designated by the EOSAT corporation:

Band 1, visible blue (0.45-0.52 [tm); band 2, visible green (0.52-0.60 [tm); band 3, visible

red (0.63-0.69 ipm); band 4, near infrared (0.76-0.90 ipm); band 5, mid infrared (1.55-1.75

[tm); band 6, thermal infrared (10.4-12.5 [tm); and band 7, a second mid infrared band

(2.08-2.35 [tm). Applying the visible bands 3, 2, and 1 to a red, green and blue (RGB)

color model creates a true-color image (Figure 1-3). Several other band combinations are

shown in Figures 2-1 through 2-4 for comparative purposes. These are pseudo-color

images in which differentiation between vegetation types is often enhanced. As the term

implies, pseudo-color images are less faithful at approximating colors seen by the human

eye. The human eye is not sensitive to infra-red wavelengths. Most of the Landsat TM

bands provide useful information on vegetation (Lillesand and Kiefer 1987). For example,

band 1 is useful in differentiating between soil and vegetation types. Band 2 measures

green reflectance of healthy vegetation. Band 5 is sensitive to the amount of water in the

leaves of plants and penetrates through smoke and haze better than bands 3, 2 or 1.

Some particularly conspicuous elements of the landscape are apparent in all

combinations, including primary forest, secondary forest, and shifting agriculture.

Furthermore, the bare granitic outcrop of the summit area of Kinabalu is readily apparent,

as is the Mamut Copper Mine, another feature without vegetation cover. Other less

conspicuous features with which we are working, such as the occurrence of ultramafic

outcrops, require special techniques to enhance. Enhancement techniques we have

employed so far include tasseled cap analysis (enhances brightness, wetness and

greenness), basic mineral composition, and ferro-magnesium indices (e. g., occurrence of









32

ferrous minerals can be enhanced by ratioing band 5 to band 4). In terms of the vegetation

on Kinabalu, the latter method has proved more useful than the former. Ultramafic

outcrops stand out particularly well in Figure 2-4.

Finally, satellite imagery made the production of the location map for Mount

Kinabalu (Figure 1-5), on which accurate and up-to-date placement of roads and

settlements (kampungs) are important, easier and more accurate. Roads vs. rivers, for

instance, are more conspicuous in different band combinations (Figures 2-2 and 2-3).


Predicting Species Ranges using Digital Elevation Models

It is possible to convert the elevational data from contours on topographic maps

into a three-dimensional digital elevation model (DEM) of a landscape. In a grid-based

DEM, elevation values are stored within cells, each of which represents a discrete unit of

space on earth. The cells are organized into rows and columns forming a Cartesian matrix

in the same way that satellite data are stored. The Kinabalu DEM uses cell sizes of 50 x

50 m. When visualized, the process is like draping an enormous fishnet over the

mountain. Different elevations can be represented by different colors, thus enhancing the

three-dimensional aspects of the image (Figure 2-5). Views from any direction can be

produced. Elevation, slope, aspect, and insolation (intensity of sunlight falling on a slope)

attributes can be easily extracted from a DEM. In an analytical environment, these

attributes can be used to predict where a particular rare species may occur in unexplored

areas, based on elevation, slope, and aspect data for known occurrences of the taxon.

A particularly exciting aspect of the DEM is that it provides a mechanism for

draping other types of coverages over basic details such as elevation. For example, river

systems can be shown three-dimensionally draped over the DEM. An even more dramatic









33

example is in the ability to drape satellite imagery. Thus, a realistic three-dimensional

model of Mount Kinabalu can be produced as a drape over the DEM, and additional

coverages, for example the Kinabalu Park boundary, can be added as shown in

Figure 1-4.


Diversity Analyses through GIS

Plant diversity and distribution patterns on Mount Kinabalu were discussed by

Beaman and Beaman (1990), and the hypotheses explaining the high diversity were

outlined there and somewhat elaborated in Beaman et al. (1997). Significant

environmental variables involved in this hypothesis that are amenable to GIS analysis

include extreme elevational variation from just above sea level to 4,094 m, precipitous

topography, diverse geological substrates, relatively open habitats (e.g., landslides), slope,

aspect, and, as dependable data about species distributions on the mountain become

available, relative isolation of individual populations.

In Parris et al. (1992) and Wood et al. (1993), histograms showing generic and

species diversity for the pteridophytes and Orchidaceae indicate that the highest number

of taxa occur at about 1500 m. This type of diversity index is independent of the actual

surface area at any particular elevation range. Among the analytical uses of the digital

elevation model is that it allows us to calculate planimetric surface area for each of the

elevation ranges on Kinabalu. To compare diversity for areas of unlike size it is necessary

to calculate a diversity index. There is of course less overall surface area on any mountain

at progressively higher elevations. The simplest type of diversity index is derived by

dividing the number of taxa by the surface area. In this instance, elevation ranges in 500

m bands around Mount Kinabalu are the areas for which diversity is compared. The









34

results for pteridophytes are shown in Figure 2-6. Although the highest number of

pteridophyte taxa occurs between 1001 and 1500 meters, the highest number of

pteridophyte taxa per unit area is between 2001 and 3000 meters. This area-based

diversity index is more comparable to diversity indices based on plot sampling.



















































Figure 2-1. Landsat TM pseudocolor image of Mount Kinabalu, produced by
applying bands 7, 6, and 4 to the RGB color model. Low elevation ultramafic areas are
quite evident with this band combination.


















































Figure 2-2. Landsat TM pseudocolor image of Mount Kinabalu, produced by
applying bands 7, 5, and 2 to the RGB color model.



















































Figure 2-3. Landsat TM pseudocolor image of Mount Kinabalu, produced by
applying bands 5, 2, and 1 to the RGB color model. This band combination
penetrates haze better than bands 3, 2, and 1. Water features are very clear.



















































Figure 2-4. Landsat TM pseudocolor image of Mount Kinabalu produced by
applying bands 4, 3, and 2 to the RGB color model. This band combination
produces an image very similar to that of color infrared film.
































>db


CA~

-cG




E-2o


C -
Cti

































CC~j
8- a ~





5He








40





















500

*number of species *species per10 sq km
400



300



200



100 -




500 1000 1500 2000 2500 3000 3500
Upper elevation range (m)




Figure 2-6. Pteridophyte diversity on Mount Kinabalu. Note that although the greatest
number of species occurs at 1500 m, the greatest diversity of species per unit area is
between 2500 and 3000 m.
















CHAPTER 3
PHYLOGENY OF THE URTICACEAE


Introduction

Choice of outgroup for the phylogenetic analysis of Elatostema from Mount

Kinabalu in Chapter 4 is based on an analysis of intergeneric relationships presented in

this chapter. Outgroup choice is often critical to the topology of the tree, and tree

topology embodies the direction of character evolution. Patterns of character evolution

cannot be reconstructed from an unrooted tree. Outgroup analysis is an almost universally

recognized method for determining character polarity (i.e., rooting a cladogram). It has

become more or less automatic using most phylogenetic software packages. The selection

of an outgroup, however, is not automatic. When an appropriate phylogeny is not

available on which to base outgroup choice, an ad hoc approach is often used. The

purpose herein is to provide a phylogenetic basis for outgroup choice upon which to base

the analysis of Elatostema in Chapter 4, thus avoiding the ad hoc approach. A brief

review of pertinent higher level classifications places the analysis in context.


The Urticales

Circumscription of the Urticales has been relatively consistent for over 150 years.

Weddell (1856) included the Ulmaceae, Cannabineae, Artocarpeae, Moreae, and

Urticaceae. Takhtajan (1997) and Thorne (1992) used the same circumscription. Dahlgren









43

(1989) and Cronquist (1981) included the Barbeyaceae, a monotypic family occurring in

north-east tropical Africa and the Arabian Peninsula.

Placement of the Urticales has been less certain. Cronquist placed the Urticales

within the Hamemelidae, a group that also includes the Fagales, Casuarinales, Myricales,

Juglandales, etc. Berg (1977, 1989) suggested affinities with the Malvales. Dahlgren,

Takhtaj an, and Thorne followed suit. Recently, relationships within the Urticales and with

other orders have been evaluated by Chase et al. (1993), Sytsma et al. (1996) and Qiu et

al. (1998) based on phylogenetic analysis ofrbcL nucleotide sequences. Based in part

upon these analyses, Judd et al. (1999) and the Angiosperm Phylogeny Group (1998)

considered the Urticales (suborder Urticineae sensu Judd) to be nested within the Rosales.

Monophyly of the Rosales is weakly supported by a reduction in endosperm and presence

of a hypanthium. Qiu et al. (1998) also reported that in all Urticales, Elaeagnaceae, and

Rhamnaceae, the nad] mitochondrial gene carries a trans-spliced intron, whereas in most

other dicots, including the Rosaceae, the intron is cis-spliced. This is perhaps the

strongest molecular evidence that the Urticales is nested within the Rosales as sister to the

Rhamnaceae/Elaeagnaceae clade. The tree of Qiu et al. (1998) also shows Barbeya as

basal to the Urticales.

Monophyly of the Urticales is supported by occurrence of cystoliths, reduction in

flower size, reduction in the number of stamens to five or fewer, a two-carpellate

unilocular ovary with a single apical ovule (a basal ovule in the Urticaceae), presence of

urticoid teeth, and at least one prominent prophyllar bud (Judd et al. 1999). The

hypanthium is lost in the Urticales with exception of the Ulmaceae, possibly because of

reduction in flower size. Laticifers are a synapomorphy for the (Moraceae(Cecropiaceae/









44

Urticaceae)) clade, but are lacking or reduced to occurring in the bark in the Cecropiaceae

and Urticaeae (in Urera), or lost or perhaps modified into mucilage cells and ducts

(Renner 1907) in the rest of the Urticaceae.

In the description of the Cecropiaceae as a new family, Berg (1978) included

Poikilospermum, but noted the genus had a number of characters in common with the

Urticaceae, including presence of elongated cystoliths, stamens that are inflexed in bud,

lack of milky latex (laticifers still present but reduced in size), and production of

mucilaginous sap. The mucilage is distinct from that of of Urticaceae, however, in that it

turns black upon exposure to air. Berg (1989) reiterated his doubt that Poikilospermum

belonged to the Cecropiaceae, noting that anatomical features discussed by Bonsen and

ter Welle (1983) suggested a relationship closer to the Urticaceae than other

Cecropiaceae.

Humphries and Blackmore (1989) carried out a phylogenetic analysis at the tribal

level for the Moraceae (Ficeae, Dorstenieae, Castilleae, Moreae, Artocapeae), and

included the genera Poikilospermum, Conocephalus (= Poikilospermum), and

Sparattosyce (Moraceae) and families Urticaceae, Ulmaceae, Cannabaceae, Cecropiaceae,

and Barbeyaceae, with the latter as outgroup. Their tree showed a polytomy between the

Urticaceae, Poikilospermum, and Conocephalus, with the Cecropiaceae as the sister

lineage, and below that, the Moraceae.

Morphological, anatomical and molecular data now support the hypothesis that

Poikilospermum is more closely related to the Urticaceae than the Cecropiaceae. In the

morphology-based analysis of Judd et al. (1994) Poikilospermum and Pilea were placed

as sister taxa (no other Urticaceae s.s. were analyzed), with Cecropia subtending the pair.









45

Phylogenetic analysis of 25 urticalian taxa (Sytsma et al. 1996) based on rbcL sequence

data hypothesized a slightly different relationship, suggesting that the Cecropiaceae is

polyphyletic and that Poikilospermum is actually embedded deeply within the Urticaceae.


The Urticaceae

Judd et. al. (1994) circumscribed the Urticaceae to include the Cecropiaceae,

Moraceae, and Cannabaceae because their analysis showed the Moraceae to be

metaphyletic. Molecular evidence, based on phylogenetic analysis of rbcL nucleotide

sequences by Sytsma et al. (1996) and Qiu et al. (1998) suggests that the Moraceae are

monophyletic. Judd et al. (1999) adopted a narrower circumscription of the Urticaceae

based on synapomorphies of elongate cystoliths, incurved stamens in bud, wood

anatomical characters, and molecular data. Phylogenetic analysis of molecular data from

rbcL sequences (Chase et al. 1993 and Qiu et al. 1998) also supports the monophyly of

the Urticaceae, but neither of these analyses included Poikilospermum.

The Urticaceae sensu Friis (1993) consists of 45 genera and about 1000 species.

He did not include Poikilospermum. Weddell (1854, 1856) recognized five tribes as did

the more recent work of Friis (1989, 1993). The names adopted here follow the tribal

names used by Friis (1993). They are the Urticeae (10 genera), Lecantheae (17 genera,

including Elatostema), Boehmerieae (19 genera), Parietarieae (5 genera), Forsskaoleae (4

genera). Although Friis provided a rich discussion of characters, no rigorous phylogenetic

analysis was conducted. Friis (1989) presented two ad hoc cladograms. These showed

confidence in the circumscription of the tribes, but did not suggest what the relationship

might be between the Urticeae, Lecantheae, and Boehmerieae.









46

The purpose of this analysis is not to determine phylogenetic relationships for the

entire family, but to find an appropriate outgroup for the analysis of Elatostema in the

next chapter. The disposition of Pellionia is one of the more interesting taxonomic

problems within the Lecantheae. Species of Pellionia are often placed in Elatostema.

Robinson (1910) considered them distinct. Schroter and Winkler (1935) treated Pellionia

as a subgenus ofElatostema. Friis (1989) considered them as two entities, but later he

(Friis 1993) combined Pellionia into Elatostema. In the analysis below they are treated as

separate taxa. However, an analysis at the species level will better address this issue in the

next chapter. In this chapter, generic relationships within the Lecantheae and between the

five tribes are evaluated.


Methods

A subset of 36 genera of Urticaceae was selected, representing each of the five tribes. In

addition, four other genera, Cannabis, Ficus, Morus, and Cecropia, were included as

outgroup taxa. Poikilospermum and Conocephalus were included as part of the ingroup

owing to the evidence supporting their placement in the Urticaceae. A total of 28

characters was scored. Friis (1989, 1993) was consulted for much of the data for

Urticaceae. Humphries and Blackmore (1989), Berg (1989), and Judd et al. (1994, 1999)

were consulted for outgroup characters. Characters (Table 3-1) were selected that were

most consistent within genera, but where they were known not to be, they were scored as

variable. Data (Table 3-2) was analyzed with PAUP* 4.0d65 (Swofford 1998) utilizing

the heuristic search algorithm (500 replicates) and using tree bisection reconnection

(TBR) and random sequence addition options.









47

Results

The heuristic search generated 219 equally parsimonious trees, each of 117 steps,

a consistency index (CI) of 0.752, and a retention index (RI) of 0.800. A strict consensus

tree is shown in Figure 3-1 and a 50% majority rule consensus tree in Figure 3-2. The

tribal classification of Friis (1993) is mapped onto the strict consensus tree. In all trees,

the Urticaceae + Cecropiaceae form a monophyletic group. The Urticaceae s.s is

monophyletic in 92% of the 219 trees. The Conocephalus/Poikilospermum clade is sister

to the Urticaceae s.s. in 92% of trees, in those cases suggesting that the Cecropiaceae is

paraphyletic. Based on the strict consensus tree, relationships between the Cecropiaceae

and basal Urticaceae appear unclear. The situation may be clarified by comparing two

alternative tree topologies from the pool of 219 equally parsimonious trees.

The first of the 219 most parsimonious trees is shown in Figure 3-3, with locations

of unambiguous character state changes mapped. In this tree, the Conocephalus/

Poikilospermum clade is sister to the Urticaceae, and Cecropia basal. With respect to the

base of the tree, a competing, equally parsimonious basal topology (shown by the tree in

Figure 3-4) is found in 8% of trees. In this topology, the Cecropiaceae is monophyletic

and nested within part of the Boehmerieae. In this second topology, moving Cecropia to a

position sister to the Urticaceae s.s., while leaving Poikilospermum and Conocephalus

embedded, increases the length of the tree by only two steps. Poikilospermum embedded

in the Urticaceae s.s. was the hypothesis supported by the analyses of Sytsma et al. (1996)

based on rbcL sequences.

Cecropiaceae embedded within the Urticaceae s.s. also requires the independent

evolution of laticifers in two lineages (see char. 6, Figure 3-5), as opposed to a single loss









48

(see char. 6, Figure 3-6), reduction, or modification of laticifers in the Urticaceae.

Laticifers are present in the bark only in the Cecropiaceae and Urera in the Urticaceae.

(Urera was not included in this analysis.) Neraudia often has milky sap, so was scored as

having laticifers. Renner (1907) argued that the mucilage cells and ducts of the

Cecropiaceae and Urticaceae were derived from laticifers. That line of reasoning supports

a tree topology with a paraphyletic Cecropiaceae basal to the the Urticaceae s.s.

For these reasons, the topology of the tree shown in Figure 3-3 is preferred and

was selected to illustrate other character distributions. Five characters support the

monophyly of the Cecropiaceae/Urticaceae in this tree. Mucilage cells or ducts occur in

the Cecropiaceae and throughout the Urticaceae, although some homoplasy is evident

(char. 5, Figure 3-7). There is also a reduction in the number of stigmas (char. 25, Figure

3-8), a change to basal placentation and an orthotropus ovule (char. 27), and reduction to

a pseudomonomerous gynoecium (char. 28). Arachnoid hairs (char. 11, Figure 3-9) are

present in the Cecropiaceae and basal Urticaceae, but are lacking in the Urticeae,

Lecantheae, Parietarieae, and Boehmerieae (p.p.).

Three anatomical wood characters occur in Poikilospermum and Conocephalus

and in the Urticaceae, but not in Cecropia. These are presence of unlignified vessel

elements (char. 1, Figure 3-10), wood fiber dimorphism (char. 2, Figure 3-11), and

presence of tangential and radial fiber pits (char. 3, Figure 3-12). In the Urticaceae,

unlignified vessel elements occur in members of the Urticeae and Lecantheae, and in

Touchardia and Nothocnide in the Boehmerieae.

The relationships between the Boehmerieae, Forsskaoleae, and Parietarieae remain

unresolved. In all trees, however, the Lecantheae together with the Urticeae form a









49

monophyletic group (Figure 3-13). Two floral characters support this clade. All members

of this clade have a glabrous male pistillode (char 19, Figure 3-14), while the pistillode is

either lanate or absent in other taxa. Free or lobed pistillate perianth parts (char. 24,

Figure 3-15) are also consistent for this clade. The perianth in pistillate flowers is either

lacking (as in the Forsskaolaep.p., Phenax, and Leucoscyse), or strongly fused into a tube

(as in the Cecropiaceae, Moraceaep.p., and Boehmerieae p.p.).

The Lecantheae, as circumscribed by Friis (1989, 1993), is also monophyletic.

The Lecantheae, itself, are supported by presence of staminodia (char. 20, Figure 3-16) in

the pistillate flowers. These staminodia function in explosive achene dispersal. Pellionia

and Elatostema are sister taxa and are sister to the remaining genera of Lecantheae. The

Elatostema and Pellionia clade is supported by anisophyllous phyllotaxy as opposed to

opposite or spiral phyllotaxy (char. 7). Alternate leaves in Elatostema (including

Pellionia) are considered to be derived via anisophyllous reduction (see Chapter 4).

Taxa belonging to the Urticeae sensu Friis form clade with the exception of

Gyrotaenia. Gyrotaenia lacks the stinging hairs (char. 9, Figure 3-17) characteristic of the

Urticeae, and is sister to the Lecantheae in the tree shown, but lacks unambiguous branch

support.

The Forsskaoleae are always monophyletic but may be nested within part of the

Boehmerieae or may alternatively be a basal clade of the Urticaceae. Monophyly of the

this tribe is supported by a reduction in the number of stamens to one.

The Boehmerieae appear to consist of a number of small clades that tend to be

basal within the family. In the majority rule tree, two small genera of the Boehmerieae are

basal within the Urticaceae. Neraudia consists of five species found on the Hawaiian









50

islands (Cowan 1949) and Debregeasia with four species occurring in Northeast Africa

and the Arabian peninsula, Pakistan, India, in the Himalaya, Japan, and Southeast Asia

throughout much of Malesia, but not in New Guinea (Wilmot-Dear 1989). Debregeasia

velutina Gaud. and D. squamata Kina ex Hook. f occur on Mount Kinabalu.


Discussion

Delimitation of tribes by Weddell (1854, 1856) and Friis (1989, 1993) are in many

instances well supported by this analysis, with the exception of taxa in the Boehmerieae.

In presenting character distributions on a phenetic map of genera, Friis (1989) placed the

Boehmerieae in a central location relative to the four other tribes, visually suggesting that

members of the other tribes may have been derived from elements within the

Boehmerieae. The Boehmerieae is the largest tribe in terms of number of genera, and is

defined at least partially on pleisiomorphic characters for the Urticaceae, such as a

tubular pistillate perianth and filiform stigmas (see Weddell 1854), both character states

found also in the Cecropiaceae. In addition, many character states are polymorphic at the

tribal level for the Boehmerieae, including phyllotaxy, cystolith and indumentum

morphology, and pollen grain characteristics. Thus, it is not surprising that inter- and

intra-tribal relationships concerning the Boehmerieae are still not well resolved. The tribe

is surely non-monophyletic.

The monophyly of the Urticaceae s.s. is tentatively supported by the single loss of

laticifers (or conversion into mucilage cells/ducts) in almost all genera of the Urticaceae

(except Urera and possibly Neraudia, where they are present and functional). This is

preferable to the equally parsimonious alternative that laticifers evolved independently in

the Cecropiaceae (in the bark only) and the Moraceae (throughout the plant). The









51

transformation proposed by Renner (1907) that the mucilage cells and ducts found in

many taxa in the Urticaceae are derived from laticifers seems plausible.

The suggestion by Sytsma et al. (1996) that Poikilospermum but not Cecropia

may be embedded in the Urticaceae still requires either the independent evolution of

laticifers, or the less parsimonious solution of laticifer loss in multiple lineages.

Constraining the analysis to force Cecropia into a position as the sister group to the

Urticaceae, and leaving Poikilospermum and Conocephalus embedded in the Urticaceae,

produced a tree only two steps longer. Thus, morphology does not strongly reject this

hypothesis. The present analysis suggests that the Cecropiaceae may indeed be

paraphyletic (as currently circumscribed), but probably not polyphyletic. The doubt

expressed by Berg (1989) that Poikilospermum belongs in the Cecropiaceae is reinforced

by the analysis of Sytsma et al. (1996) and by the results of the present analysis, which

place Poikilospermum as sister to the Urticaceae s.s. Additional sampling of taxa for both

molecular and morphological characters may place Poikilospermum slightly or even

deeply nested within the Urticaceae. More study is needed.

The task of choosing an outgroup for the Elatostema analysis becomes more

objective based on the foregoing hypothesized phylogenetic relationships. In a

preliminary analysis, representatives of Pilea and Boehmeria were used based on ad hoc

reasoning. Relationships within the Lecantheae/Urticeae clade are now better resolved

and the Elatostema/Pellionia clade is shown to be sister to the remaining Lecantheae,

with Pilea a basal member of the sister group. Thus Pilea continues to be an appropriate

outgroup taxon. On the other hand, Boehmeria appears to be relatively distantly related.









52

Therefore, in the ensuing analysis, Pilea and Gyrotaenia are used to root the resulting

trees. Gyrotaenia is either basal to the rest of the Lecantheae, or basal in the Urticeae.

Representatives of Pellionia would not make an appropriate outgroup, since it

may actually be part of the ingroup. A number of species of Elastostema have at some

point or another been described as or transferred to Pellionia. Pellionia was included as

distinct in the generic analysis only because some authors have recognized it (Weddell

1856, Friis 1989) at the generic level. However, others have not (Schroter and Winkler

1935, Friis 1993). Fortunately, these two genera consistently show up as sister taxa. The

next chapter treats taxa at the species level, and the monophyly of Pellionia and

Elatostema is addressed.














Cannabis
Morus
Ficus
Cecropia
Boehmeria
Neraudia
Leucoscyce
Pipturus
Nothocnide
Debregeasia
Phenax
Poikilospermu
Conocephalus
Touchardia
Gyrotaenia
Urtica
Nanocnide
Laportea
Discocnide
Girardinia
Dendrocnide
Sarcopilea
Pilea
Achudemia
Procris
Lecanthus
Meniscogyne
Pellionia
Elatostema
Neodistemon
Parietaria
Soleirolia
Didymodoxa
Australina
Forsskaolea
Drogeuetia


o


.
ft








m


Boehmerieae













r

a

ft





Boehmerieae

Parietarieae


o

'


Figure 3-1. Strict consensus of 219 equally parsimonious trees, showing generic
relationships between 30 genera of the Urticaceae and six genera in the Cannabinaceae,
Cecropiaceae, and Moraceae. Cannabis, Ficus, and Morus were designated as outgroup
taxa. Tribes, sensu Friis (1993), are mapped for the Urticaceae.





























Figure 3-2. Majority rule (50%) consensus tree of 219 equally parsimonious trees
showing generic relationships between 30 genera of the Urticaceae and six genera in
the Cannabaceae, Cecropiaceae, and Moraceae. Cannabis, Ficus, and Morus were
designated as outgroup taxa.











Majority rule
100 E Morus
L_- Ficus
Cecropia
100 Poikilospermum
Conocephalus
100 Urtica
Nanocnide
100
10 Laportea
10 Discocnide
10C Girardinia
100 Dendrocnide
Gyrotaenia
Pilea
100
0 Achudemia
74 Procris
100 67
5g 0 Lecanthus
92 10 Meniscogyne
Sarcopilea
100 Pellionia
100 Elatostema
100 Neodistemon
S- Parietaria
Soleirolia
Touchardia
Boehmeria
84 1Leucoscyce

84
92 84 Phenax
100 Forsskaolea
51 10 Drogeuetia
84 Didymodoxa
Australina
84 Pipturus
Nothocnide
Debregeasia
Neraudia
Cannabis





























Figure 3-3. Preferred cladogram showing generic relationships between 30 genera of
the Urticaceae and six genera in the Cannabinaceae, Cecropiaceae, and Moraceae.
Cannabis, Ficus, and Morus were designated as outgroup taxa. Positions where
unambiguous character state changes occur are mapped onto the tree. The
Cecropiaceae is paraphyletic and subtends the Urticaceae in this tree.






















7c 15 u*1 2 2





212
713 15 8 1 2 24 1





235 .
15 3 16 11


..13
..22 10



..7 14




..22 7 26



15 21 22
9 10 10





























Figure 3-4. Representative cladogram showing generic relationships between 30
genera of the Urticaceae and six genera in the Cannabinaceae, Cecropiaceae, and
Moraceae. Cannabis, Ficus, and Morus were designated as outgroup taxa. Positions
where unambiguous character state changes occur are mapped onto the tree. The
Cecropiaceae is embedded within the Urticaceae in this tree.
























Oz 0
C-) C-0





























Figure 3-5. Representative most parsimonious tree. Branch tracing indicates
occurrence of laticifers. This topology requires independent evolution of laticifers or
multiple loss. Laticifers absent (0), Laticifers present (1).














U )

Cz C)z
Cz 0s
B o0 5 g.
5 Cz 5 0
70) 0 U -j 0 0. 75,
:) ~ ~ ~ m 4 &P. P w P4U 4 ):


Character 6
2 steps
unordered
0
E1





























Figure 3-6. Preferred most parsimonious tree. Branch tracing indicates occurrence of
laticifers. This topology involves a single loss of laticifers. Laticifers absent (0),
Laticifers present (1).












C)z
C) C)
C) z 0 B 0
-a U El 0


Character 6
1 step
unordered
0


PELL1t equivocal


vld
Q24






























Figure 3-7. Preferred most parsimonious tree. Branch tracing indicates occurrence of
mucilage cells or ducts. Mucilage cells or ducts absent (0), mucilage cells or ducts
present (1).










































































Character 5
1 step
unordered
0
W1


Y


Ua~





























Figure 3-8. Preferred most parsimonious tree. Branch tracing shows number of
stigmas. Two (0), one (1).





















































Character 25
1 step
unordered
0

UIE1t equivocal


vli





























Figure 3-9. Preferred most parsimonious tree. Branch tracing indicates occurrence of
arachnoid hairs. Arachnoid hairs absent (0), arachnoid hairs present (1).














0. S 0

0 00
0 32
~ o S T'
0 3
7A0000a2 cri4H7


Character 11
7 steps
unordered
0


polymorphic
1EL1 equivocal


vld
Q24


rJ"
0ci
00Q
07-





























Figure 3-10. Preferred most parsimonious tree. Branch tracing indicates occurrence
ofunlignified vessel elements. Vessel elements lignified (0), vessel elements
unlignified (1).














01
C) ) U C)
z0 C) z
CzmF~~F~


Character 1
3 steps
unordered
0


UIE1t equivocal


8 5,





























Figure 3-11. Preferred most parsimonious tree. Branch tracing indicates occurrence
of dimorphic wood fibers. Wood fibers monomorphic (0), wood fibers dimorphic (1).













C)z



91 U"
33aIj & --0
:) PL
~~~r 3FZ 6 a, v EC


Character 2
2 steps
unordered
0


UIE1t equivocal


8l 5





























Figure 3-12. Preferred most parsimonious tree. Branch tracing indicates occurrence
of tangential and radial fiber pits. Fiber pits radial only (0), fiber pits radial and
tangential (1).





















































Character 3
1 step
unordered
0

UIE1t equivocal


vli





























Figure 3-13. Preferred most parsimonious tree, showing relationships within the
Lecantheae/Urticeae clade. Characters and character state changes are mapped.













Caj C)
Ct CA CA
C.) 15 I 0 0 0 I
7 0000 *I ~~C I ^ 1 I^ s
g~ c ^ dQ^g^




























Figure 3-14. Preferred most parsimonious tree. Branch tracing indicates occurrence
of the male pistillode and vestiture. Pistillode glabrous (0), pistillode lanate or pilose
(1) pistillode absent (2).
















Cz 0
3 3 I . S 11 I
s g-ii 9~i isi


vld
Q24


0

* a 8
*^ S 23 2
SIDp


."
rJi

00Q
07-


Character 19
3 steps
unordered
0


2


equivocal


LMi





























Figure 3-15. Preferred most parsimonious tree. Branch tracing indicates degree of
fusion of female perianth. Perianth absent (0), perianth free or lobed (1), perianth
tubular (2).






















































Character 24
6 steps
unordered
0

2
polymorphic
11111 equivocal


vli





























Figure 3-16. Preferred most parsimonious tree. Branch tracing indicates occurrence
of staminodia in pistillate flowers. Staminodia absent (0), staminodia present (1).












4 Cz~~3~
C) 00 5



n~n~~nnnnh


5m

o o~ 8 :~a~
Sz
0 ~ F 00~) ~0 -


Character 20
1 step
unordered
0

UIE1t equivocal





























Figure 3-17. Preferred most parsimonious tree. Branch tracing indicates occurrence
of stinging hairs. Stinging hairs absent (0), stinging hairs present (1).














5 z 0 t
0z -z -'
0 Z
Cz Cn
Cz 0;
vl 8 'l ECn


Character 9
1 step
unordered
0


UIE1t equivocal


r)O














Table 3.1. Characters scored for phylogenetic analysis of the Urticales. All considered
unordered.


1) Vessel elements:
2) Wood fibers:
3) Fiber pits:
4) Storied fibers:
5) Mucilage cells:
6) Laticifers:
7) Phyllotaxis:
8) Cystoliths:
9) Stinging hairs:
10) Hooked hairs:
11) Arachnoid hairs:
12) Stipules:
13) Stipules:
14) Female receptacles:
15) Inflorescence:
16) Inflorescence:
17) Number of stamens:
18) Stamens filaments:
19) Male pistilode:
20) Staminodia:
21) Pollen shape:

22) Pollen spinules:


23) Achenes:
24) Female perianth:
25) Stigma number:
26) Stigma:


27) Ovules:
28) Gynoecium:


lignified (0), unlignified (1)
monomorphic (0), dimorphic (1)
radial (0), tangential and radial (1)
absent (0), present (1)
absent (0), present (1)
absent (0), present (1)
spiral (0), opposite (1), anisophyllus (2)
absent (0), punctate (1), elongate (2)
absent (0), present (1)
absent (0), present (1)
absent (0), present (1)
absent (0), present (1)
intrapetiolar (0), interpetiolar (1), lateral and free (2)
not fleshy (0), fleshy (1)
no involucrate disk (0), involucrate disk present (1)
not glomerulate (0), glomerulate (1)
one (1), two (2), four (4), five (5)
+ straight (0), incurved (1)
glabrous (0), lanate or pilose (1), absent (2)
absent (0), present (1)
sphaeroidal (0), subprolate (1), intermediate (2),
suboblate (3)
small, evenly dispered, and dense (0),
larger, arranged in groups (1),
larger, cone-shaped, in groups (2)
curved (0), straight (1)
absent (0), free or lobed (1), tubular (2)
two (0), one (1)
capitate-penicillate (0), oblong (1), linear (2),
elongate/filiform (3)
apical and anatropus (0), basal and orthotropus (1)
2 carpellate (0), pseudomonomerous (1)























O 3













































0 0 c0
0o0 0


0











































O w


O s



0 0





0 0


0 0

0 0











0 0


0 0



0 0


0 0







0 0



0 0
(N
I-1






0 0

3- C^-


0 0




0 0

c^. ^-


(N (N 0








0 0 0


(N ^


0 0 0


0 00

0 0 0




O OO











0 0 O









0 0


0 0 0
O O (


0 0







0 0 0
i- C -


~ 0


00 0

C ^ I i -


0 0 0








0 0 0


(N (N (N


m C- (N





0 0 0






't 7t 7

0 0 0

0 0 0


0 0 0


0 0 0





0 0 0


0 0 0

0 0 0

(N (N (N





0 0 0
OO






































0 0 0
CD D C











CD CD CD


0 0

0 o -S a

c l *4 | | I |

0 t3 Q 2: 0e Q O S. <; u c;M 3


clin


OO





OO
0 0








0 0


(N (N


m N



OO
00








0 0

0 ^


0 0


0 0





0 0


0 0

0 0





0 ^


0 0

c^. c^.


0 0



0 0

c - c -




















00 - -










-(N N - - 'N - -






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CHAPTER 4
PHYLOGENY OF KINABALU ELA TOSTEMA


Introduction

This chapter focuses on phylogenetic relationships among the Elatostema s.l.

from Mount Kinabalu. In Chapter 5, the phylogenetic reconstruction is applied to a

biogeographical analysis with special detail given to micro-geographical areas and

ecological associations within the Kinabalu floristic community.


Elatostema in Malesia

Based on a review of Index Kewensis entries, the genus Elatostema in the

Malesian floristic region includes ca. 250 species. About 120 of these were originally

described from the Philippines, about 75 from New Guinea and approximately 40 from

Borneo. There are 19 species of Elatostema on Mount Kinabalu already published, and an

additional seven species that are newly described here (see Chapter 6). Twelve species are

known only from Mount Kinabalu.


Generic Delimitation and Monophyly of Elatostema

Schr6ter and Winkler (1935, 1936) recognized four subgenera of Elatostema s.l.,

i.e., E. subg. Pellionia (Gaudich.) Hall. f., E. subg. Elatostematoides (C. B. Rob.) H.

Schroter, E. subg. Weddellia H. Schroter, and E. subg. Elatostema (as subg.

Euelatostema H. Schr6ter). They monographed 106 species in the first three of these

subgenera. Their published work never treated E. subg. Elatostema. However, they list

taxa from Borneo












that they recognized within this subgenus, including a number of Kinabalu taxa. There

are no taxa listed from Kinabalu in E. subg. Weddellia. Of the four subgenera, E. subg.

Pellionia is most frequently recognized at the generic level, and it still is in the

horticultural trade. Pellionia sensu Robinson (1910) has cymose or fasciculate staminate

and pistillate inflorescences (sometimes very condensed, but never involucrate), while the

pistillate inflorescences ofElatostema s.s. are fused into involucral heads.

In a revision of Philippine Urticaceae, Robinson (1910) described the genus

Elatostematoides as a segregate of the Elatostema/Pellionia alliance. He treated

Elatostema s.s. as distinct based on two characters, the presence of the involucrate

inflorescences mentioned above, and what he referred to as a cup-shaped perianth on the

pistillate flowers. Robinson doubted that this cup-shaped perianth was really a whorl of

staminodia as suggested by Stapf (1894). This issue, as well as the monophyly of

Elatostema s.s., Pellionia, and Elatostematoides, is considered further in the discussion

section of this chapter. Elatostema s.l. is distinct among the Urticaceae on the basis of

anisophyllous leaves and alternate leaves through reduction via anisophylly.


Methods

The phylogenetic analysis included 26 taxa and was based on the observation of

76 morphological characters. Outgroup taxa included two species ofPilea, P. leptocardia

and P. leptograma, and Gyrotaenia myriocarpa. The choice of these outgroups was based

on a preliminary phylogeny of the Urticaceae presented in Chapter 3.

Characters and character states used are listed in Table 4-1. While some of the

characters are self explanatory, others require clarification. These are discussed below.

Some suites of characters are used to describe the overall pattern of distribution of hair or

cystolith types over the plant surface. For example, cystoliths sometimes occur on just












one leaf surface or they may line the veins but not the interstices of a leaf surface. By

coding these as separate characters, the overall pattern is defined in binary format. Care

was taken to avoid unintentional weighting due to character correlation in these suites of

characters. Characters 44, 46, 52, and 66 were deleted for this reason. Characters 6, 23,

41, and 62 were found to be parsimony uninformative, and were also deleted.

Data were analyzed with PAUP* 4.0d65 (Swofford 1998) utilizing the heuristic

search algorithm (500 replicates) and using tree bisection reconnection (TBR) and

random sequence addition options.


Character and Character State Categories and Clarification

Sexual dimorphism. Both monoecious and dioecious plants (char. 1) are

common in Elatostema. Only three species were recorded as polymorphic for this

character.

Leaf venation. Venation types (char. 2) are as defined by Schroter and Winkler

(1935). Leaf venation in the Urticaceae is likely to be derived from palmately veined

ancestors. Related families (i.e., Cecropiaceae and Moraceae) often have palmately

veined leaves. In Elatostema, leaf venation appears to be derived from camptodromus

ancestors (common in Pilea). A transformation series occurs to brochidodromus and

craspedromus states in which the two main arching secondaries appear to be compressed

down toward the base of the leaf to varying degrees. This occurs to different degrees even

on either side of the midvein, making the venation pattern asymmetrical across the

midvein. Venation is then camptodromus on one side of the midvein and brochidodromus

on the other.

Type 1 venation is more or less symmetrical, and camptodromus. Type 2 is more

or less symmetrical, camptodromus, but apically brochidodromus. Type 3 and 4a-b




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